Baylands Ecosystem Species and Community Profiles

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S
Baylands Ecosystem
pecies and
Community Profiles
Life Histories and Environmental Requirements
of Key Plants, Fish and Wildlife
Prepared by the San Francisco Bay Area
Wetlands Ecosystem Goals Project
Baylands Ecosystem
Species
and
Community Profiles
Life Histories and Environmental
Requirements of Key Plants, Fish and Wildlife
Prepared by the San Francisco Bay Area Wetlands Ecosystem Goals Project
To order additional copies of this report ($25.00 each), please contact:
San Francisco Estuary Project
c/o S.F. Bay Regional Water Quality Control Board
1515 Clay Street, Suite 1400
Oakland, CA 94612
(510) 622-2465
Please cite this report as:
Goals Project. 2000. Baylands Ecosystem Species and Community Profiles: Life histories and
environmental requirements of key plants, fish and wildlife. Prepared by the San Francisco Bay Area
Wetlands Ecosystem Goals Project. P.R. Olofson, editor. San Francisco Bay Regional Water Quality
Control Board, Oakland, Calif.
Also available from the Goals Project:
Baylands Ecosystem Habitat Goals: A report of habitat recommendations. March 1999. Reprint
with minor corrections, June 2000.
Printing of the Species Profiles was made possible, in part, by a grant from the CALFED Bay-Delta Program through
the U.S. Bureau of Reclamation to Friends of the Estuary:
CALFED Project #99-B10
U.S. Department of the Interior
Bureau of Reclamation Grant #00FC200183
O
n behalf of the Resource Managers Group and all of the other Goals Project
participants, we offer our sincere thanks and appreciation to the authors of
these profiles. The authors willingly volunteered their time to construct a good
part of the biological foundation of the Habitat Goals. Little did they know when
they started that their work eventually would comprise this fine report.
We also want to recognize the role of the San Francisco Bay Regional Water
Quality Control Board in the preparation of this document. Without the support
of the Regional Board’s top managers—Loretta Barsamian, Larry Kolb, and Bruce
Wolf—the profiles would likely still be a pile of interesting papers on the flora and
fauna of the San Francisco Bay area, rather than a unified compendium. By encouraging their staff to compile and polish the profiles, they helped provide the
public with a scientific document that will lead to the improvement of habitat conditions and water quality throughout the Bay and along its tributaries.
Above all, we would like to extend a special thanks to Peggy Olofson of the
Regional Board. Peggy not only helped to manage the Goals Project through its
four-year life, but she spent the better part of the past year preparing this document for press. Her work far exceeded merely editing the draft profiles. Rather,
she collaborated with many of the report authors in acquiring data and expanding
sections of text, cajoled authors into clarifying certain points, obtained photographs,
hunted down references, created the report design, and laid out each page. And she
did all this while attending to her regular duties as a Water Resources Engineer. We
all owe Peggy our gratitude for a job well done.
Michael Monroe
Carl Wilcox
Resource Managers Group Co-Chairs
San Francisco Bay Area Wetlands
Ecosystem Goals Project
Credits
Editor
Peggy R. Olofson — San Francisco Bay Regional Water Quality Control Board
Authors
David G. Ainley
Joy D. Albertson
Janice M. Alexander
Peter R. Baye
Dennis R Becker
William G. Bousman
Andrée M Breaux
Michael L Casazza
Steven C. Chappell
Howard L. Cogswell
Ron Duke
Jules G. Evens
Phyllis M. Faber
Leora Feeney
Steve Foreman
Stephen L. Granholm
Brenda J. Grewell
Janet T. Hanson
Laura A. Hanson
Elaine K. Harding
Bruce Herbold
Catherine M. Hickey
Kathryn A. Hieb
Glen Holstein
Mark R. Jennings
Michael L. Johnson
Robert E. Jones
Kurt F. Kline
Brita C. Larsson
Robert A. Leidy
William Z. Lidicker, Jr.
Kevin MacKay
Wesley A. Maffei
Lt. Dante B. Maragni
Carolyn M. Marn
Michael F. McGowan
A. Keith Miles
Michael R. Miller
Gary W. Page
Thomas P. Ryan
Michael K. Saiki
Howard S. Shellhammer
Ted R. Sommer
Lynne E. Stenzel
John Y. Takekawa
Robert N. Tasto
Scott Terrill
Lynne A. Trulio
David C. VanBaren
Nils Warnock
Sarah E. Warnock
Frank G. Wernette
Report Production
Report Design &Layout: Peggy Olofson
Cover Design: Nina Lisowski
Computer & GISSupport: Jeff Kapellas
Text & Copy Editing: Cristina Grosso (lead)
Elisa Gill
Terra Hendrich
Harini Madhavan
Michael Monroe
Dewey Schwarzenberg
Jill Sunahara
Credits
i
Additional Assistance
In addition to the authors and production staff, we also thank the many people who provided
support and assistance to make this publication possible.
We thank Marcia Brockbank and her staff at the San Francisco Estuary Project for ongoing help
with funding and coordination, and the staff at CALFED and the Bureau of Reclamation for funding assistance. For generous and timely support for preliminary editing and document design, we
thank the California Coastal Conservancy, and particularly Nadine Hitchcock.
Special thanks to Kathy Hieb, who provided extensive organizational and editing assistance
with the fish profiles, in addition to authoring several of them herself. Several other authors, including Peter Baye, Bill Bousman, Howard Cogswell, Jules Evens, and Glen Holstein worked patiently
with the editor over many months to provide additional tables and figures for the report.
For their guidance in publication planning and preparation, we thank Michelle Yang, Mark
Rodgers, and the rest of the staff at Alonzo Environmental. Also, we thank Debbi Nichols for
helping to keep things together when it was needed most.
Photography and Artwork
We would especially like to thank the photographers and artists who so graciously donated the use
of their images for this publication:
Peter Baye
Berkeley Digital Library
Nancy Black, Monterey Bay Whale Watch
www.gowhales.com
Ted Brown, California Acadamy of Sciences
Mia Bruksman
California Dept. of Fish and Game (CDFG)
Les Chibana [email protected]
Citizens’ Committee to Complete the Refuge
Jack Kelly Clark, Statewide IPM Project. Permission by the Regents of the University
of California
Howard L. Cogswell
Josh Collins
Don DesJardin
www.camacdonald.com/birding/DesJardin
Joe DiDonato, bioQuest Wildlife Photography and Consulting (510)769-9209
Dr. Richard B. Forbes
Dr. Allan Francis [email protected]
Rick A. Fridell [email protected]
S.H. Hinshaw
Dan Holland
Jimmy Hu
www.silcom.com/~njhua/otter/otter.html
Marshall Iliff [email protected]
ii
Baylands Ecosystem Species and Communities
Dr. J.L. King
Peter LaTourrette www.birdphotography.com
Denise Loving [email protected]
Wesley Maffei [email protected]
Peter Moyle, from Inland Fishes of California,
copyright 1976. Permission by the Regents of the University of California
Ruth Pratt
T. Douglas Rodda [email protected]
M. Roper, Aquatic Research Organisms,
Hampton, NH (1991)
Tom Rountree
Brad Shaffer
Ted Sommer
Rick Stallcup
Dr. Daniel Sudia [email protected]
U.S. Fish and Wildlife Service (USFWS)
U.S. Geological Survey, National Biological
Service (USGS/NBS)
Staffan Vilcans
Jens V. Vindum, California Acadamy of Sciences
Bob Walker
Peter S. Weber www.wildbirdphotos.com
Zoological Society of Milwaukee County,
Wisconsin www.zoosociety.org
Contents
Credits .................................................................................................................. i
Contents ............................................................................................................. iii
List of Tables ....................................................................................................... ix
List of Figures ..................................................................................................... xi
1. Plant Communities
a.
b.
c.
d.
e.
Plants of Shallow and Subtidal Habitat and Tidal Flats
(with an emphasis on eelgrass)
by Laura A. Hanson .............................................................................................. 1
Tidal Marsh Plants of the San Francisco Estuary
by Peter R. Baye, Phyllis M. Faber, and Brenda Grewell ....................................... 9
Plants and Environments of Diked Baylands
by Peter R. Baye .................................................................................................. 33
Plants of San Francisco Bay Salt Ponds
by Peter R. Baye .................................................................................................. 43
Plant Communities Ecotonal to the Baylands
by Glen Holstein ................................................................................................ 49
2. Estuarine Fish and Associated Invertebrates
a.
b.
c.
d.
e.
f.
g.
h.
Opossum Shrimp (Neomysis mercedis)
by Bruce Herbold ...............................................................................................
Dungeness Crab (Cancer magister)
by Robert N. Tasto .............................................................................................
Rock Crabs (Cancer antennarius and Cancer productus)
by Robert N. Tasto .............................................................................................
Bat Ray (Myliobatus californica)
by Kurt F. Kline ..................................................................................................
Leopard Shark (Triakis semifasciata)
by Michael F. McGowan .....................................................................................
Pacific Herring (Clupea pallasi)
by Robert N. Tasto .............................................................................................
Northern Anchovy (Engraulis mordax)
by Michael F. McGowan .....................................................................................
Sacramento Splittail (Pogonichthys macrolepidotus)
by Ted R. Sommer ..............................................................................................
69
71
76
79
81
81
85
87
Contents
iii
i.
Chinook Salmon (Oncorhynchus tshawytscha)
by Lt. Dante B. Maragni .................................................................................... 91
j. Steelhead (Oncorhynchus mykiss irideus)
by Robert A. Leidy ........................................................................................... 101
k. Delta Smelt (Hypomesus transpacificus)
by Ted R. Sommer and Bruce Herbold ............................................................. 104
l. Longfin Smelt (Spirinchus thaleichthys)
by Frank G. Wernette ....................................................................................... 109
m. Jacksmelt (Atherinopsis californiensis)
by Michael K. Saiki .......................................................................................... 113
n. Topsmelt (Atherinops affinis)
by Michael K. Saiki .......................................................................................... 115
o. Threespine Stickleback (Gasterosteus aculeatus)
by Robert A. Leidy ........................................................................................... 118
p. Brown Rockfish (Sebastes auriculatus)
by Kurt F. Kline ................................................................................................ 121
q. Pacific Staghorn Sculpin (Leptocottus armatus armatus)
by Robert N. Tasto ........................................................................................... 123
r. Prickly Sculpin (Cottus asper)
by Bruce Herbold ............................................................................................. 126
s. Striped Bass (Morone saxatilis)
by Ted R. Sommer ............................................................................................ 129
t. White Croaker (Genyonemus lineatus)
by Kurt F. Kline ................................................................................................ 130
u. Shiner Perch (Cymatogaster aggregata)
by Michael F. McGowan ................................................................................... 132
v. Tule Perch (Hysterocarpus traskii)
by Robert A. Leidy ........................................................................................... 134
w. Arrow Goby (Clevelandia ios)
by Kathryn A. Hieb .......................................................................................... 136
x. Bay Goby (Lepidogobius lepidus)
by Kathryn A. Hieb .......................................................................................... 139
y. Longjaw Mudsucker (Gillichthys mirabilis)
by Kathryn A. Hieb .......................................................................................... 142
z. California Halibut (Paralichthys californicus)
by Michael K. Saiki .......................................................................................... 144
aa. Starry Flounder (Platichthys stellatus)
by Kurt F. Kline ................................................................................................ 148
3.
I nv
er
tebrates
nver
ertebrates
a.
b.
c.
d.
e.
iv
Franciscan Brine Shrimp (Artemia franciscana Kellogg)
by Brita C. Larsson ...........................................................................................
California Vernal Pool Tadpole Shrimp (Lepidurus packardi Simon)
by Brita C. Larsson ...........................................................................................
Reticulate Water Boatman (Trichocorixa reticulata Guerin)
by Wesley A. Maffei ..........................................................................................
Tiger Beetles (Cincindela senilis senilis, C. oregona, and C. haemorrhagica)
by Wesley A. Maffei ..........................................................................................
Western Tanarthrus Beetle (Tanarthrus occidentalis Chandler)
by Wesley A. Maffei ..........................................................................................
Baylands Ecosystem Species and Community Profiles
151
153
154
156
161
f.
Inchworm Moth (Perizoma custodiata)
by Wesley A. Maffei ..........................................................................................
g. Pygmy Blue Butterfly (Brephidium exilis Boisduval)
by Wesley A. Maffei ..........................................................................................
h. Summer Salt Marsh Mosquito (Aedes dorsalis (Meigen))
by Wesley A. Maffei ..........................................................................................
i. Winter Salt Marsh Mosquito (Aedes squamiger (Coquillett))
by Wesley A. Maffei ..........................................................................................
j. Washino’s Mosquito (Aedes washinoi Lansaro and Eldridge)
by Wesley A. Maffei ..........................................................................................
k. Western Encephalitis Mosquito (Culex tarsalis Coquillett)
by Wesley A. Maffei ..........................................................................................
l. Winter Marsh Mosquito (Culiseta inornata (Williston))
by Wesley A. Maffei ..........................................................................................
m. Brine Flies (Diptera: Ephydridae)
by Wesley A. Maffei ..........................................................................................
n. Jamieson’s Compsocryptus Wasp (Compsocryptus jamiesoni Nolfo)
by Wesley A. Maffei ..........................................................................................
o. A Note on Invertebrate Populations of the San Francisco Estuary
by Wesley A. Maffei ..........................................................................................
163
165
167
169
172
173
177
179
183
184
4. Amphibians and Reptiles
a.
b.
c.
d.
e.
f.
g.
h.
i.
California Tiger Salamander (Ambystoma californiense)
by Mark R. Jennings .........................................................................................
California Toad (Bufo boreas halophilus)
by Mark R. Jennings .........................................................................................
Pacific Treefrog (Hyla regilla)
by Mark R. Jennings .........................................................................................
California Red-Legged Frog (Rana aurora draytonii)
by Mark R. Jennings .........................................................................................
Western Pond Turtle (Clemmys marmorata)
by Mark R. Jennings .........................................................................................
California Alligator Lizard (Elgaria multicarinata multicarinata)
by Mark R. Jennings .........................................................................................
Central Coast Garter Snake (Thamnophis atratus atratus)
by Kevin MacKay and Mark R. Jennings ..........................................................
Coast Garter Snake (Thamnophis elegans terrestris)
by Kevin MacKay and Mark R. Jennings ..........................................................
San Francisco Garter Snake (Thamnophis sirtalis tetrataenia)
by Mark R. Jennings .........................................................................................
193
196
198
201
204
208
210
212
214
5. Mammals
a.
b.
c.
Salt Marsh Harvest Mouse (Reithrodontomys raviventris)
by Howard S. Shellhammer .............................................................................. 219
California Vole (Microtus californicus)
by William Z. Lidicker, Jr. ................................................................................ 229
Salt Marsh Wandering Shrew (Sorex vagrans haliocoetes)
by Howard S. Shellhammer .............................................................................. 231
Contents
v
d.
e.
f.
g.
h.
i.
j.
k.
6.
Suisun Shrew (Sorex ornatus sinuosis)
by Kevin MacKay .............................................................................................
Ornate Shrew (Sorex ornatus californicus)
by Elaine K. Harding ........................................................................................
North American River Otter (Lutra canadensis)
by Michael L. Johnson ......................................................................................
Southern Sea Otter (Enhydra lutris nereis)
by David G. Ainley and Robert E. Jones ..........................................................
Harbor Seal (Phoca vitulina richardsi)
by William Z. Lidicker, Jr. and David G. Ainley ..............................................
California Sea Lion (Zalophus californianus)
by David G. Ainley and Robert E. Jones ..........................................................
Non-Native Predators: Norway Rat and Roof Rat
(Rattus norvegicus and Rattus rattus)
by Andrée M. Breaux ........................................................................................
Non-Native Predator: Red Fox (Vulpes vulpes regalis)
by Elaine K. Harding ........................................................................................
233
236
238
241
243
246
249
251
Water
fo
wl and SShor
hor
ebir
ds
aterfo
fowl
horebir
ebirds
a.
Tule Greater White-Fronted Goose (Anser albifrons gambelli)
by Dennis R. Becker .........................................................................................
b. Mallard (Anas platyrhynchos)
by Steven C. Chappell and David C. Van Baren ...............................................
c. Northern Pintail (Anas acuta)
by Michael L. Casazza and Michael R. Miller ..................................................
d. Canvasback (Aythya valisineria)
by John Y. Takekawa and Carolyn M. Marn .....................................................
e. Surf Scoter (Melanitta perspicillata)
by A. Keith Miles ..............................................................................................
f. Ruddy Duck (Oxyura jamaicensis)
by A. Keith Miles ..............................................................................................
g. Western Snowy Plover (Chardrius alexandrinus)
by Gary W. Page, Catherine M. Hickey, and Lynne E. Stenzel .........................
h. Marbled Godwit (Limosa fedoa)
by Gary W. Page, Catherine M. Hickey, and Lynne E. Stenzel .........................
i. Black Turnstone (Arenaria melanocephala)
by Stephen L. Granholm ..................................................................................
j. Red Knot (Calidris canutus)
by Catherine M. Hickey, Gary W. Page, and Lynne E. Stenzel .........................
k. Western Sandpiper (Calidris mauri)
by Nils Warnock and Sarah E. Warnock ...........................................................
l. Long-Billed Dowitcher (Limnodromus scolopaceus)
by John Y. Takekawa and Sarah E. Warnock .....................................................
m. Wilson’s Phalarope (Phalaropus tricolor)
by Janet T. Hanson ...........................................................................................
n. Waterfowl and Shorebirds of the San Francisco Estuary
by John Y. Takekawa, Gary W. Page, Janice M. Alexander,
and Dennis R. Becker .......................................................................................
vi
Baylands Ecosystem Species and Community Profiles
253
259
263
268
273
277
281
284
289
292
296
301
306
309
7. Bayland Birds Other than Shorebirds and Waterfowl
a.
Eared Grebe (Podiceps nigricollis)
by Howard L. Cogswell ....................................................................................
b. Western and Clark’s Grebes (Aechmophorus occidentalis and A. clarkii)
by David G. Ainley ...........................................................................................
c. American White Pelican (Pelecanus erythrorhynchus)
by David G. Ainley ...........................................................................................
d. Brown Pelican (Pelecanus occidentalis)
by David G. Ainley ...........................................................................................
e. Double-Crested Cormorant (Phalacrocorax auritus)
by David G. Ainley ...........................................................................................
f. Snowy Egret (Egretta thula)
by William G. Bousman ...................................................................................
g. Black-Crowned Night Heron (Nycticorax nycticorax)
by William G. Bousman ...................................................................................
h. California Clapper Rail (Rallus longirostris obsoletus)
by Joy D. Albertson and Jules G. Evens ............................................................
i. California Black Rail (Laterallus jamaicensis coturniculus)
by Lynne A. Trulio and Jules G. Evens .............................................................
j. Common Moorhen (Gallinula chloropus)
by William G. Bousman ...................................................................................
k. California Gull (Larus californicus)
by Thomas P. Ryan ...........................................................................................
l. Forster’s Tern (Sterna forsteri)
by Thomas P. Ryan ...........................................................................................
m. Caspian Tern (Sterna caspia)
by Thomas P. Ryan ...........................................................................................
n. California Least Tern (Sterna antillarum browni)
by Leora Feeney ................................................................................................
o. Western Burrowing Owl (Athene (Speotyto) cunicularia hypugaea)
by Lynne A. Trulio ............................................................................................
p. Salt Marsh Common Yellowthroat (Geothlypis trichas sinuosa)
by Scott Terrill ..................................................................................................
q. Savannah Sparrow (Passerculus sandwichensis)
by Howard L. Cogswell ....................................................................................
r. Song Sparrow (Melospiza melodia samuelis, M. m. pusillula,
and M. m. maxillaris)
by Howard L. Cogswell ....................................................................................
s. Response of Birds to Managed Water Levels at
Charleston Slough – A Case Study
by William G. Bousman ...................................................................................
t. The Use of Salt Ponds by Some Selected Birds
Other than Shorebirds and Waterfowl
by Howard L. Cogswell ....................................................................................
317
320
321
322
323
325
328
332
341
346
349
351
355
359
362
366
369
374
386
390
Appendix A: Author Contact Information ..................................................... 403
Contents
vii
viii
Baylands Ecosystem Species and Community Profiles
List of Tables
1. Plant Communities
Table 1.1
Acreage of Individual Eelgrass Beds in San Francisco/San Pablo Bay in 1989 ............................. 3
Table 1.2
Comparison of Three Northern California Estuaries Relative to Size of Estuary
and Total Acres of Eelgrass (Zostera marina) ............................................................................. 3
Table 1.3
Historic Changes in the Distribution and Abundance of Selected Native
Vascular Plant Species Occurring in Tidal Marshes of the San Francisco Estuary ...................... 23
Table 1.4
Rare Plant Species Found in the Nine Counties Adjacent to the San Francisco
Bay Estuary, by Plant Community or Ecotone ....................................................................... 61
2. Estuarine Fish and Associated Invertebrates
Table 2.1
Annual Abundance of Rock Crabs Caught by Otter Trawl (crabs/tow) in
the San Francisco Estuary ...................................................................................................... 78
Table 2.2
Annual Abundance of Rock Crabs Caught by Ring Net (crabs/tow) in
the San Francisco Estuary ....................................................................................................... 78
Table 2.3
Historical and Recent Collections of Splittail ........................................................................... 88
Table 2.4
Migration Characteristics of Sacramento-San Joaquin Chinook Salmon Runs .......................... 25
Table 2.5
Estimated Number of Sacramento-San Joaquin Chinook Salmon Returning
to Spawn: 1967-1991 .......................................................................................................... 95
Table 2.6
Bay Goby Salinity and Temperature Statistics: 1980-92 ....................................................... 140
3. Invertebrates
Table 3.1
Known Collection Sites For Trichocorixa reticulata ................................................................ 155
Table 3.2
Known Collection Sites For Tiger Beetle Populations ............................................................ 158
Table 3.3
Known Collection Sites For Tanarthrus occidentalis ................................................................. 16
Table 3.4
Known Collection Sites For Perizoma custodiata .................................................................... 164
Table 3.5
Known Collection Sites For Brephidium exilis ....................................................................... 166
Table 3.6
Known Collection Sites For Brine Flies ................................................................................. 181
Table 3.7
Known Collection Sites For Compsocryptus jamiesoni .............................................................. 183
Table 3.8
Partial Summary of Organisms Associated with Alkali Heath ................................................. 186
Table 3.9
Food Web Taxa by Major Common Name Category ............................................................ 190
Table 3.10 Known Terrestrial or Semi-aquatic Invertebrate Surveys or Studies of
Selected Invertebrate Taxa ................................................................................................... 191
List of Tables
ix
5. Mammals
Appendix 5.1
Important Data Sets for Salt Marsh Harvest Mouse (1971-1991) .................................... 222
6. Waterfowl and Shorebirds
Table 6.1
Peak Monthly Population Indices for Tule Greater White-fronted Goose on
Migration, Stopover, and Wintering areas in Oregon and California for 1978-79
through 1981-82, 1988-89, and 1989-90 .......................................................................... 256
Table 6.2
Regional Recommendations to Support Western Snowy Plover ............................................. 283
Table 6.3
Recommendations to Support Western Snowy Plover in the South Bay ................................. 283
Table 6.4
Regional Recommendations to Support Marbled Godwit ..................................................... 288
Table 6.5
Recommendations to Support Marbled Godwit in the North Bay ......................................... 288
7. Other Birds of the Baylands Ecosystem
Table 7.1
Estimated Breeding Pairs of Snowy Egrets on West Marin Island .......................................... 327
Table 7.2
Estimated Breeding Pairs of Black- Crowned Night Herons on West Marin Island ................. 331
Table 7.3
California Gull Breeding Sites in the South Bay .................................................................... 350
Table 7.4
Forster’s Tern Breeding Sites in the North Bay ..................................................................... 352
Table 7.5
Forster’s Tern Breeding Sites in the South Bay ...................................................................... 353
Table 7.6
Caspian Tern Breeding Sites in the San Francisco Bay ........................................................... 357
Table 7.7
Censuses in California Bayside Marsh or Adjacent Grasslands Where Savannah
Sparrows Were Reported ...................................................................................................... 371
Table 7.8
Key Habitats Usage by Three Salt Marsh Song Sparrow Races ............................................... 379
Table 7.9
Census Data from 1980 and 1981 for Charleston Slough ..................................................... 387
Table 7.10 Fifteen Most Common Species Censused at Charleston Slough in 1980 –
sorted by 1980 rank order .................................................................................................. 388
Table 7.11 Eighteen Most Common Species Censused at Charleston Slough in 1981 –
sorted by 1981 rank order ................................................................................................... 388
x
Baylands Ecosystem Species and Community Profiles
List of Figures
1. Plant Communities
Figure 1.1
Comparison of Percent Eelgrass Coverage in Three West Coast Estuaries ................................... 4
2. Estuarine Fish and Associated Invertebrates
Figure 2.1
Seasonal Distribution of Juvenile Dungeness Crab Within San Francisco Bay ......................... 73
Figure 2.2
Annual Distribution of Juvenile Dungeness Crab Within the San Francisco Bay –
Caught by Otter Trawl, May-December ............................................................................ 74
Figure 2.3
Traditional Pacific Herring Spawning Areas in Central San Francisco Bay ........................ 84
Figure 2.4
Trends in Age-0 Splittail Abundance for 1975-1995 as Indexed by Eight
Independent Surveys ............................................................................................................ 89
Figure 2.5
Trends in Adult Splittail Abundance for 1976-1995 as Indexed by Six
Independent Surveys ........................................................................................................... 90
Figure 2.6
Life History of Chinook Salmon ......................................................................................... 92
Figure 2.5
Spatial and Temporal Distribution of Young-of-the-Year Pacific Staghorn Sculpin ............. 124
Figure 2.6
Spatial and Temporal Distribution of Adult Pacific Staghorn Sculpin .............................. 124
Figure 2.7
Annual Abundance Indices of White Croaker ................................................................... 131
Figure 2.8
Annual Abundance Indices of Arrow Goby from San Francisco Bay, Beach Seine ................ 137
Figure 2.9
Annual Abundance Indices of All Sizes of Bay Goby, Otter Trawl ................................... 141
Figure 2.10 Annual Abundance Indices of Starry Flounder .................................................................. 149
3. Invertebrates
Figure 3.1
Reticulate Water Boatman – Trichocorixa reticulata ........................................................... 154
Figure 3.2
Known Trichocorixa reticulata Localities Within San Francisco Bay Tidal
and Diked Marshes ............................................................................................................ 155
Figure 3.3
Tiger Beetle – Cicindela senilis senilis ................................................................................. 156
Figure 3.4
Known Tiger Beetle Localities Within San Francisco Bay Tidal
and Diked Marshes ............................................................................................................ 157
Figure 3.5
Cicindela senilis senilis Larva in Burrow .............................................................................. 159
Figure 3.6
Western Tanarthrus Beetle – Tanarthrus occidentalis ......................................................... 160
Figure 3.7
Known Tanarthrus occidentalis Localities Within San Francisco Bay Tidal
and Diked Marshes ............................................................................................................ 161
Figure 3.8
Inchworm Moth – Perizoma custodiata .............................................................................. 163
Figure 3.9
Known Perizoma custodiata Localities Within San Francisco Bay Tidal
and Diked Marshes ............................................................................................................ 163
List of Figures
xi
Figure 3.10 Adult Pygmy Blue Butterfly – Brephidium exilis ................................................................... 165
Figure 3.11 Brephidium exilis Egg and larva ............................................................................................ 165
Figure 3.12 Known Brephidium exilis Localities Within San Francisco Bay Tidal
and Diked Marshes ............................................................................................................ 165
Figure 3.13 Adult Winter Salt Marsh Mosquito – Aedes squamiger ...................................................... 169
Figure 3.14 Terminal Abdominal Segment of a Fourth Instar Larva .................................................... 178
Figure 3.15 Aedes Squamiger Distribution in the San Francisco Bay Area, 1950 ................................ 179
Figure 3.16 Terminal Abdominal Segment of C. tarsalis larva .............................................................. 173
Figure 3.17 Wing of an Adult Cs. inornata ........................................................................................... 177
Figure 3.18 Terminal Abdominal Segment of a Fourth Instar Larva .................................................... 177
Figure 3.19 Adult Ephydra millbrae ...................................................................................................... 179
Figure 3.20 Adult Lipochaeta slossonae ................................................................................................ 180
Figure 3.21 Ephydra millbrae Larva and Pupa ...................................................................................... 180
Figure 3.22 Known Brine Fly Localities Within San Francisco Bay Tidal and Diked Marshes ................. 180
Figure 3.23 Jamieson’s Compsocryptus Wasp – Compso+cryptus jamiesoni .......................................... 183
Figure 3.24 Known Compsocryptus jamiesoni Localities Within San Francisco Bay Tidal
and Diked Marshes ............................................................................................................. 183
Figure 3.25 A Partial Web of the Organisms Associated With Alkali Heath (Frankenia salina)
in San Francisco Tidal Marshes ......................................................................................... 185
Figure 3.26 A Partial Web of the Organisms Associated With Common Pickleweed
(Salicornia virginica) in San Francisco Bay Lower High Tidal Marshes ............................ 187
Figure 3.27 A Partial Web of the Organisms Associated With Willow (Salix lasiolepis) ....................... 188
Figure 3.28 Partial Web of Organisms Associated With Mid Tidal Marsh Pans .................................. 188
Figure 3.29 A Partial Web of the Organisms in the Baumberg and Oliver Brothers Salt
Crystallizer Ponds, Hayward, California ............................................................................ 189
4. Anphibians and Reptiles
Figure 4.1
California Tiger Salamander – Some Current Locations ................................................... 194
Figure 4.2
California Toad – Presumed Bay Area Distribution .......................................................... 197
Figure 4.3
Pacific Treefrog – Presumed Bay Area Distribution .......................................................... 199
Figure 4.4
California Red-Legged Frog – Some Current Locations .................................................... 202
Figure 4.5
Western Pond Turtle – Presumed Bay Area Distribution ................................................. 206
Figure 4.6
California Alligator Lizard – Presumed Bay Area Distribution .......................................... 209
Figure 4.7
Central Coast Garter Snake – Presumed Bay Area Distribution ....................................... 211
Figure 4.8
Coast Garter Snake – Presumed Bay Area Distribution .................................................... 213
Figure 4.9
San Francisco Garter Snake – Current Known Location ................................................... 215
5. Mammals
xii
Figure 5.1
Salt Marsh Harvest Mouse – Some Current Locations and Suitable Habitat .................... 210
Figure 5.2
California Vole – Some Current Locations and Suitable Habitat ...................................... 230
Figure 5.3
Salt Marsh Wandering Shrew – Some Current Locations and Suitable Habitat ............... 232
Figure 5.4
Suisun Shrew – Some Current Locations and Suitable Habitat ........................................ 235
Figure 5.5
Ornate Shrew – Some Current Locations and Suitable Habitat ........................................ 237
Baylands Ecosystem Species and Community Profiles
Figure 5.6
North American River Otter – Some Current Locations and Suitable Habitat ....................... 239
Figure 5.7
Southern Sea Otter – Some Current Locations ...................................................................... 242
Figure 5.8
Harbor Seal – Some Current Haul-out Locations and Suitable Habitat .................................. 244
Figure 5.9
California Sea Lion – Current Locations and Suitable Habitat ............................................... 247
6. Waterfowl and Shorebirds
Figure 6.1
Distribution of Tule Greater White-Fronted Goose in San Francisco Bay .............................. 255
Figure 6.2
Maximum Counts of Mallard .............................................................................................. 260
Figure 6.3
Mean Mid-Winter Survey Counts of Pintails by Decade in Suisun Marsh
and San Francisco Bay ......................................................................................................... 264
Figure 6.4
Maximum Counts of Northern Pintail .............................................................................. 265
Figure 6.5
Radio-Marked Pintail Locations in Suisun Marsh During the Fall and Winter
of 1991-92 and 1992-93 .................................................................................................... 265
Figure 6.6
Common Day and Night Movement Patterns for Pintails Wintering in Suisun Marsh ........ 266
Figure 6.7
Maximum Counts of Canvasback ...................................................................................... 270
Figure 6.8
Maximum Counts of Surf Scoter ....................................................................................... 275
Figure 6.9
Maximum Counts of Ruddy Duck .................................................................................... 278
Figure 6.10 Relative Use of Different Baylands Areas by Marbled Godwit, Willet, Whimbrel,
and Long-billed Curlew Combined .................................................................................... 286
Figure 6.12 Relative Use of Various Tidal Flat and Adjacent Shoreline Areas by Rocky
Substrate Species (Black Turnstone, Ruddy Turnstone, Surfbird, Spotted
Sandpiper, Black Oystercatcher, and Wandering Tattler) ................................................. 290
Figure 6.13 Relative Use of Different Mudflat Areas by Red Knots ..................................................... 293
Figure 6.14 Maximum Counts of Red Knot ......................................................................................... 294
Figure 6.15 Maximum counts of Western Sandpiper, Least Sandpiper, and Dunlin Combined ......... 297
Figure 6.16 Maximum Counts of Western Sandpiper .......................................................................... 298
Figure 6.17 Maximum Counts of Dowitcher Species ........................................................................... 302
Figure 6.18 Maximum Counts of Long-billed Dowitcher .................................................................... 303
Figure 6.19 Maximum Counts of Phalarope Species ............................................................................ 307
Figure 6.20 Relative Use of Salt Ponds by American Avocet, Snowy Plover, Black-necked
Stilt, and Phalaropes ........................................................................................................... 308
Figure 6.21 Relative Use (High, Medium, Low) of Different Mudflat Areas by Tidal Flat
Specialists, as Indicated from the Proportion of Shorebirds Counted in
Different Survey Areas ....................................................................................................... 313
Figure 6.22 Relative Use (High, Medium, Low) of Different South Bay Salt Pond Areas
by Salt Pond Specialists as Indicated from the Proportion of Shorebirds Counted
in Different Survey Areas ..................................................................................................... 314
Figure 6.23 Waterfowl Use of Salt Ponds in the North Bay ................................................................. 315
Figure 6.24 Waterfowl Use of Salt Ponds in the South Bay .................................................................. 315
7. Other Birds of the Baylands Ecosystem
Figure 7.1
Christmas Bird Count data for Snowy Egret ..................................................................... 326
Figure 7.2
Christmas Bird Count data for Black-Crowned Night Heron ........................................... 330
List of Figures
xiii
xiv
Figure 7.3
Known Distribution of the California Clapper Rail ............................................................... 335
Figure 7.4
Distribution and Relative Abundance of Black Rails (Laterallus jamaicensis coturniculus)
in the San Francisco Bay Region ........................................................................................ 343
Figure 7.5
Christmas Bird Count data for Common Moorhen .......................................................... 347
Figure 7.6
Summary of Data from Six Bird Count Studies Conducted Between
Late Fall 1984 and Winter 1985 ........................................................................................ 372
Figure 7.7
Numbering System for the Salt Ponds of South San Francisco Bay .................................. 392
Baylands Ecosystem Species and Community Profiles
Introduction
T
he San Francisco Bay Area Wetlands Ecosystem Goals Project began in
1995 as a cooperative effort among nine state and federal agencies and
nearly 100 Bay Area scientists. The Project’s purpose was to develop a vision
of the kinds, amounts, and distribution of habitats needed to sustain healthy
populations of fish and wildlife in and around San Francisco Bay (Figure 1).
This vision was presented to the public in the Goals Project’s final report, the
Baylands Ecosystem Habitat Goals1.
Developing the Habitat Goals involved several steps, many of which were
carried out by teams of scientists. First, each team selected “key” animal species (or plant communities, in the case of the Plants Focus Team). Then they
compiled available information regarding each species’ historic and modern distribution, use of habitats, migration, relationship and interaction with other
species, conservation and management issues, and research needs. When time
and data were available, some team members compiled additional information
on life history and other relevant topics. The teams then discussed the habitat
needs of their species and developed initial habitat recommendations. Ultimately, the habitat recommendations of all the teams were integrated to form
the Project’s final recommendations.
Compiling the information on key species and plant communities into
“profiles” was a crucial step in developing the Habitat Goals. Sharing these
profiles enabled team members to better understand the habitat needs of a large
proportion of the bayland’s flora and fauna. It also facilitated the development
of more balanced and diverse habitat recommendations.
When Project participants began sharing the profiles, they realized that
much of the information had never before been compiled They also recognized
that, although some of the profiles were not comprehensive, other researchers
interested in the Bay and its watersheds might find them useful.
The intent of this report is to provide useful information to those working to restore the baylands ecosystem. However, because the profiles were compiled to inform a specific process, this report should not be considered a complete treatise. Rather, it should be seen as a reference and a starting point.
Contact information for the profile authors is included in Appendix A for
the reader who would like additional information or clarification, or who would
just like to continue the process of scientific discussion and discovery.
1
Copies of the Baylands Ecosystem Habitat Goals report may be obtained from the
San Francisco Estuary Project at the address indicated in the front of this report.
Introduction
xv
Figure 1. The Project Area of the San Francisco Bay Area Wetlands Ecosystem Goals Project – The
baylands and adjacent and associated habitats of the San Francisco Bay-Delta Estuary, downstream of the Sacramento-San Joaquin Delta
xvi
Baylands Ecosystem Species and Communities
Plants
1
Plant Communities
Plants of Shallow Subtidal Habitat
and Tidal Flats
(with an emphasis on eelgrass)
Laura A. Hanson
Introduction
There are about 200,000 acres of shallow subtidal habitat and tidal flats in San Francisco Bay, San Pablo Bay,
and Suisun Bay. Of this area, approximately 171,000
acres are subtidal habitat and about 29,000 acres are tidal
flats. While relatively simple in terms of species diversity, the plant communities that occur in these areas are
important components of the estuarine ecosystem.
Although this paper describes the plant communities of shallow subtidal habitat and tidal flats, it focuses
on the eelgrass (Zostera marina) community. For more
detailed information on the other plant communities
(primarily microalgae and macroalgae) that occur in the
shallow subtidal areas and on tidal flats of the San Francisco Bay Estuary, please refer to Silva (1979), Nichols
and Pamatmat (1988), Meiorin et al. (1991), and Herbold et al. (1992).
Environmental Setting
Shallow subtidal areas and tidal flats are defined by their
elevation in relation to tidal height. The shallow subtidal
range includes the areas between mean lower low water
(MLLW) and the approximate bathymetric contour 18
feet below MLLW. Tidal flats generally occur between
the mean tide level (MTL), or the lower elevation limit
of Spartina (cordgrass) flats, to about 2.5 feet below
MLLW. Tidal flat composition can include various combinations of clay, silt, sand, shell fragments, and organic
debris. Daily tidal cycles submerge and expose tidal flat
surfaces twice every 24.8 hours. During each tidal cycle,
tidal flats are also exposed to fluctuating wave action,
current velocities, and nutrient supply. Where tidal
marshes still exist, incoming tides flood into the upper
marsh areas. As these tidal waters recede, organic materials are transported downslope to tidal flats where they
become food sources for millions of detritus-feeding invertebrates.
The environmental conditions of shallow subtidal
areas and tidal flats are stongly influenced by suspended
sediments. In general, the San Francisco Bay Estuary has
high concentrations of suspended sediments (Hanson
and Walton 1988). This suspended particulate matter
is comprised of 70 - 97% non-organic sediment made
up of silty clay; the remaining content is comprised of
living and other organic matter (Conomos and Peterson
1977). Suspended sediment concentrations are influenced by wind speed, substrate, particle size, wave action, current velocity, tidal action, water depth and seasonal runoff (Cyrus and Blaber 1987). Human activities
such as type of land use (Kemp et al. 1983), channel
dredging (LaSalle 1988, Hanson and Walton 1988),
construction and use of marinas and ferry terminals, and
propeller wash (Walker et al. 1989, Thom and Shreffler
1995) can also affect water clarity.
Total suspended solids (TSS) in Suisun and San
Pablo bays average between 50 mg/l in the summer to
200 mg/l in the winter (Nichols and Pamatmat 1988).
In North Bay and Central Bay, tides can have a significant influence on sediment resuspension, particularly
during spring tides and during the ebbs preceding
lower low water when the current speeds are highest.
Central Bay – characterized by cold, saline, and relatively clear ocean water – has the lowest TSS concentrations, at 10 to 60 mg/l. South San Francisco Bay
has slightly higher TSS concentrations than Central
Bay (O’Connor 1991).
Salinity levels vary depending on season, weather,
amount of diverted fresh water, and location in the Bay.
In general, salinity levels within the water column and
within tidal flats increase along a gradient from the Delta
to the Golden Gate. For example, the salinity in Suisun
Bay averages about seven parts per thousand (ppt), and
in Central Bay it averages about 30 ppt (Fox et al. 1991).
During dry years, South Bay averages salinity levels up
to 35 ppt.
Chapter 1 — Plant Communities
1
The shallow subtidal areas and tidal flats of the San Francisco Bay Estuary support relatively few plant communities. These communities include diatoms and other
microalgae, macroalgae, and eelgrass.
Microalgae form the basis for the estuarine food
web. These algae, consisting of diatoms and blue-green
algae, often form dense patches on tidal flats, creating a
brown hue to the substrate surface during low tide. Microalgae and settled phytoplankton represent a readily
available food source for creatures, such as worms and
clams, within the mudflats (Nichols and Pamatmat
1988). Shorebirds and waterfowl then consume these
creatures.
Macroalgae (seaweeds) are also found throughout
the Estuary, particularly in the more saline areas. Few
macroalgae can make the necessary adjustments in internal water and mineral content to survive at low salinity levels. The exceptions include Gracilaria sjoestedtii,
Enteromorpha spp. and the closely related Ulva spp. G.
sjoestedtii is usually found from the mid-intertidal to the
shallow subtidal zone attached to rocks partially buried
in coarse sand. It also grows attached to small bits of clam
and oyster shell in muddy portions of the Bay. In such
situations, the plants and associated substrata are easily
moved by currents and wave action. Enteromorpha and
Ulva form bright green patches and can occur in great
abundance throughout the intertidal zone, often growing on any available hard substrate. Enteromorpha can
be found occupying higher tidal zones where shade is
available. It is especially prevalent on boat hulls, buoys,
docks, and woodwork. Ulva occupies the lower tidal
zones, completing its life cycle in a few weeks and varying its distribution over a short time period. These kinds
of macroalgae often undergo seasonal cycles of abundance, becoming common in the warmer months and
virtually disappearing in colder months. Maximum
abundance occurs in late summer and early fall (Jones
and Stokes Associates, Inc. 1981). Many species of Ulva
are often common in heavily polluted areas because they
can use ammonia as a nitrogen source and are generally
tolerant of organic and metal pollution (Dawson and
Foster 1982). In the absence of eelgrass, Ulva can provide a preferred habitat for several invertebrate species
(Sogard and Able 1991).
Eelgrass (Zostera marina) is currently the only
seagrass found in San Francisco Bay. Belying its common name, it is not a grass but is a flowering plant that
has adapted to living submerged in the shallow waters
of protected bays and estuaries in temperate regions of
the world (Den Hartog 1970, Phillips and Menez 1988).
Z. marina reproduces both sexually through pollination
of seeds, and asexually through a rhizome meristem that
extends through the sediments (Setchell 1929). Where
abundant, Z. marina’s dense, matted root and rhizome
2
Baylands Ecosystem Species and Community Profiles
Laura Hanson
Plants
Intertidal and Subtidal Plant Communities
Uprooted Zostera marina from intertidal zone off of
Alameda shroreline. Leaves may be 1.5–12 mm wide
and up to 15 meters in length.
system functions to stabilize the soft bottom. Its leaves
slow currents and dampen wave action, causing sediment and organic material to accumulate. Z. marina
is found in intertidal areas, becoming exposed during
the lower spring tides; it also occurs in subtidal areas
at depths less than one to two meters below MLW
(Kitting 1994).
Historic and Modern Distribution (of
Eelgrass)
Information on historic distribution of Zostera marina
in the San Francisco Bay Estuary is very limited. San
Francisco Bay may have supported extensive Z. marina
meadows in the past. (Setchell 1929, Wyllie-Echeverria and Rutten 1989). Low light availability within
the water column has been found to limit the development of extensive eelgrass meadows and may be the
principal cause of eelgrass decline in San Francisco
Bay (Alpine and Cloern 1988, Zimmerman et al.
1991).
In 1989, Wyllie-Echeverria and Rutten published the first survey on the distribution of Zostera marina in San Francisco Bay (including San Pablo Bay) and
mapped a total of 316 acres (Table 1.1). As Table 1.2
and Figure 1.1 show, the per area abundance of eelgrass
within San Francisco Bay is much less than that of
Humboldt Bay or Tomales Bay, two other northern California estuaries.
The 1989 Wyllie-Echeverria and Rutten survey described the Zostera marina populations as “ patchy” and
some as “ stressed.” Since that time a few of these beds
have increased in size, and new patches have been sited
(Kitting 1993 and pers. comm.).
Table 1.1 Acreage of Individual Eelgrass Beds in
San Francisco/San Pablo Bay in 1989
Acres
San Pablo Bay
124
Point Orient
3
Naval Supply Depot
12
Point Molate Beach
26
Toll Plaza, East
0.5
Toll Plaza, West
0.5
Point Richmond, North
7
Point Richmond, South
Richmond Breakwater, North
4
18
Richmond Breakwater, South
7
Emeryville
13
Alameda
55
Bay Farm, North
2
Bay Farm, South
4
Coyote Point
1
Richardson Bay
13
Angel Island
3
Belvedere Cove
5
Point Tiburon
1
Keil Cove
10
Paradise Cove, North
4
Paradise Cove, South
3
TOTAL ACRES
316
Table from NMFS SW Region. Wyllie-Echeverria and Rutten 1989 Administrative Report SWE-89-05
Associated Fauna Including Rare and
Sensitive Species
Tidal flats include a living system of diatoms, microalgae, and protozoa that are fed upon by suspension or
surface deposit feeding invertebrates. The bottom invertebrates are in turn fed upon by larger consumers such
as fish, shrimp, and crabs. During low tide, these primary and secondary consumers are exploited by millions
of migratory shorebirds. The extensive intertidal mudflats of San Francisco Bay are considered a key migratory staging and refueling area for over-wintering shorebirds of the Pacific Flyway (Harvey et al. 1992).
Location
Humbolt Bay
Tomales Bay
San Francisco Bay
Table 1.2 Comparison
of Three Northern
California Estuaries
Relative to Size of
Estuary and Total
Acres of Eelgrass
(Zostera marina)
(km2)
Extent of
Eelgrass
(Bottom coverage, acres)
62.4
3,053
30.0
965
Spratt 1985
1,140.0
316
Wyllie-Echeverria 1990
Reference
Phillips 1984
Table from NMFS SW Region. Wyllie-Echeverria (1990)
Chapter 1 —
Plant Communities
3
Plants
Location
Macroalgae and eelgrass provide food, shelter, and
spawning grounds for many Bay fish and invertebrates.
The major subtidal spawning areas for Clupea harengus
(Pacific herring), recently the most valuable fishery in
California, are Richardson Bay and the large shallow area
between Richmond and Oakland. In these areas, spawning occurs predominantly on Gracilaria ssp. and small
patches of Zostera marina (Spratt 1981). When available,
C. harengus preferentially uses Z. marina habitats for
spawning (Taylor 1964, Spratt 1981).
Zostera marina beds support a variety of organisms,
more than that of non-vegetated areas (van Montfrans
et al. 1984, Kitting 1993, Hanson 1997). Z. marina
roots and leaves provide habitat for many plants and
animals. For example, the long blade-like shoots provide
shelter and serve as a nursery ground for many fish species. Small plants (epiphytes) and animals (epizoites) attach to the leaves, motile animals find cover between the
leaves, and burrowing animals live among the roots.
Epiphytes are an important part of the eelgrass community, contributing up to 22% of the total primary productivity (Jones 1968, Marshal 1970, Penhale 1977).
They, in turn, provide food for resident invertebrate
grazers (Kitting et al. 1984). Within the rich organic
sediment, anaerobic processes of microorganisms regenerate and recycle nutrients and carbon (Kenworthy et al.
1982).
Because Zostera marina contains noxious sulfated
phenolic compounds that can inhibit bacterial degradation and animal grazing, few animals consume it (Tenore
1977, Harrison and Chan 1980, McMillan et al. 1980).
Notable exceptions include several species of waterfowl
such as Anas americana (wigeon), Anas strepera (gadwall),
Anas acuta (pintail), Branta canadensis (Canada goose),
and Branta nigricans (black brant) (Phillips 1984). Z.
marina has been an obligate food for black brant along
its flyway (Einarsen 1965). Black brant populations are
in great decline along the Pacific Flyway, possibly due
to this species’ dependence on dwindling eelgrass resources (Einarsen 1965).
Some bird species also forage on the fauna associated with Zostera marina. An example is the Sterna
albifrons browni (California least tern) that was listed as
an endangered species in 1970. Least terns are known
to forage on juvenile and small fishes (Magenheim and
Rubissow 1993) that inhabit Z. marina beds, particu-
0.20
Plants
0.18
0.16
0.14
0.12
0.10
0.08
0.06
0.04
0.02
0
Humboldt
Bay
Tomales San Francisco
Bay
Bay
Figure 1.1 Comparison of Percent Eelgrass
Coverage in Three West Coast Estuaries (Based
on Wyllie-Echeverria (1990))
larly at a major nesting site near the Oakland International Airport and the Alameda Naval Air Station
(Collins and Feeney 1983-6, Feeney 1988 and 1989,
Harvey et al. 1992).
Invertebrates such as juvenile Cancer magister
(Dungeness crab) appear to grow up most successfully
in the nursery-like habitat that Zostera marina provides,
particularly in the northern reaches of the Bay. The isopod, Synidotea laticauda is periodically found in high
numbers (up to 200/m2) among Z. marina beds in Central San Francisco Bay (Hanson 1998). They are an
important food item for economically valuable sport
fishes such as young striped bass, starry flounder, steelhead trout, king salmon, white sturgeon, plus other
fishes in San Francisco Bay (Morris et al. 1980)
The transport of Zostera marina fragments acts as
a vector for animal dispersal (Highsmith 1985, Worcester 1994). Kitting (1993) found several fish species and
a variety of invertebrates usually associated with Z. marina on dead blades found at depths greater than four
meters below MLW.
Conservation Issues
Exotic Plants – There is some potential for two
exotic Zostera species to invade San Francisco Bay. The
Asian seagrass, Zostera japonica, introduced to British
Columbia, Washington, and Oregon has not yet been
reported in San Francisco Bay. Z. japonica has a differ-
4
Baylands Ecosystem Species and Community Profiles
ent life history, morphology, and preferred habitat than
Z. marina (Harrison and Bigley 1982). Culture experiments determined that Z. japonica is not likely to displace existing Z. marina beds (Harrison 1982). This may
not be the case in San Francisco Bay. Z. japonica favors
the intertidal zones, the areas where Z. marina has been
limited to in San Francisco Bay. Thus far, Z.japonica occupies only a small fraction of its potential habitat in
North America, threatening significant changes in the
ecology of the intertidal sediments as this seagrass spreads
(Harrison and Bigley 1982).
Zostera asiatica is found from Tomales Bay in the
north, to Santa Monica Bay in the south. Phillips and
Wyllie-Echeverria (1990) published the first record of
this species in the Eastern Pacific. It is a wide bladed
Zostera that occurs sub-tidally from five meters below
MLLW to 17 meters below MLLW. Z. asiatica has not
yet been identified in San Francisco Bay. This is probably due to its deeper water distribution where photosynthetic processes could be limited in San Francisco
Bay.
Factors Limiting Eelgrass Distribution – Under
suitable conditions, Zostera marina can form dense, continuous, and extensive carpets as seen in Tomales and
Humboldt bays. Light, temperature, salinity, tidal range
and water motion all affect growth and productivity of
Z. marina (Thayer et al. 1984, Fonseca et al. 1985,
Fonseca and Kenworthy 1987). The amount of time it
is exposed to air during low tides determines the upper
limits of Z. marina, and the amount of available light
determines the lower limits (Backman and Barilotti
1976; Dennison and Alberte 1982, 1985, 1986; Bulthuis
1983; Bulthuis and Woelkerling 1983; Wetzel and
Penhale 1983; Lewis et al. 1985; Josselyn et al. 1986;
Duarte 1991). The primary factor responsible for a
worldwide decline in Z. marina and other submerged
aquatic vegetation is reduced light availability (Giesen
et al. 1990, Dennison et al. 1993).
In San Francisco Bay, Zostera marina requires
somewhere between three and five hours of Hsat (length
of irradiance-saturated photosynthesis) each day (Zimmerman et al. 1991). In areas with favorable light conditions, Z. marina plants have adequate carbon reserves
to withstand at least 30 days of light limitation (Zimmerman et al. 1991); however, due to frequent and persistent periods of high turbidity, it is unlikely that plants
at the deeper edge of eelgrass meadows in San Francisco
Bay can accumulate large carbon reserves (Zimmerman
et al. 1991). Average turbidity of the Bay and, more critically, brief periods of high turbidity limit Z. marina distribution in deeper water and limit establishment of seedlings and vegetative propagules (Zimmerman et al.
1991). If daily, monthly, and seasonal Hsat requirements
are not met, long-term survival of the plants may be limited (Zimmerman et al. 1991). Any activities that increase turbidity within Bay waters, whether natural or
Laura Hanson
anthropogenic, have detrimental effects on existing eelgrass populations and associated food webs.
Current Restoration Success – The technology
for successfully establishing seagrass beds has been unreliable (Phillips 1974, 1980; Lewis 1987), although, in
1989 Zimmerman et al. (1995) successfully transplanted
Zostera marina at two locations in San Francisco Bay.
According to Fonseca et al. (1988), waning interest in
Z. marina restoration was due to a net loss of habitat
through seagrass mitigation projects. Planting projects
have often failed as a result of poor selection of planting
sites or plant material and incorrect use of planting methods. Factors that limited success include a general lack
of knowledge of physiological requirements and unknown local environmental factors controlling Z. marina
growth (Lewis 1987, Merkel 1990). For example, in
1984, an eelgrass transplant was initiated in San Francisco Bay. Limited transplant success was attributed to
a lack of data on local eelgrass autecology coupled with
nearby dredging operations and diminished water quality (Fredette et al. 1988).
Conclusions and Recommendations
There has been considerable interest in protecting and
expanding existing Zostera marina beds in San Francisco
and San Pablo bays (Fredette et al. 1988). Since the 1989
survey, sitings have indicated a marked change in the
distribution and abundance of this species. Better conserving this species in the Estuary will require more frequent monitoring of individual populations.
It also is imperative to protect the current eelgrass
beds from further decline. Because of the inherent difficulties in establishing eelgrass, plantings conducted
in exchange for permitted losses (mitigation projects)
could result in a greater loss of habitat and should not
be allowed. The current Zostera marina populations
may be the last remnants in San Francisco Bay and
are extremely vulnerable to local extinction (Kitting
and Wyllie-Echeverria 1991); therefore, plantings
should be used to enhance current beds or to create
new beds.
References
Alpine, A.E. and J.E. Cloern. 1988. Phytoplankton
growth rates in a light limited environment, San
Francisco Bay. Mar. Ecol. Prog. Ser. 44: 167-173.
Backman, T.W. and D.C. Barilotti. 1976. Irradiance
reduction: effects on standing crops of the eelgrass
Zostera marina in a coastal lagoon. Mar. Biol. 34:
33-40.
Bulthuis, D.A. 1983. Effects of in situ light reduction
on density and growth of the seagrass Heterozostera
tasmanica (Martens ex Aschers.) den Hartog in
Western Port, Victoria, Australia. J. Exp. Mar. Biol.
Ecol. 67: 91-103.
Chapter 1 —
Plant Communities
5
Plants
Zostera marina plants, discernable as dark blotches in
the foreground, near Belvedere Cove, Marin County.
There are several actions that should be undertaken
when designing potential restoration or enhancement
projects:
1. Conduct a thorough survey to assess physical
conditions of the site. Collect and evaluate
environmental data and/or pilot test the planting at
a particular site before commitment of a full
restoration project. The success of any seagrass revegetation effort, including long-term plant
growth, is strictly dependent upon a physical
environment suitable for initial establishment
(Zimmerman et al. 1991).
2. Carefully evaluate light availability before proceeding with any major transplant effort. Water column
turbidity is sufficiently high throughout much of
the Central Bay, limiting the euphotic zone (depth
where irradiance falls to 1% of surface irradiance)
to less than 1 m (Alpine and Cloern 1988).
3. Use stocks for planting from a site with conditions
as similar as possible to the planting site. There
should be similar or equal water depths, salinity,
temperature, tidal currents, wave exposure, and
sediment composition (Fonseca 1994). Until we
learn more about the genetic structure of this
species, matching of phenotypes among restoration
and donor sites remains the best guide for stock
selection.
4. Limit planting to areas with small tidal ranges
rather than high tidal ranges to provide greater
light availability (Koch and Beer 1996), thus
increasing survival success.
5. Plant in areas where parameters for deeper vertical
distribution are available make the bed less
vulnerable to adverse conditions (such as storm
events or desiccation) due to availability of energy
from the neighboring deeper shoots (Tomasko and
Dawes 1989).
6. Plant in late spring and summer. Periods of high Z.
marina growth and production coincide with
warmer temperatures and greater light availability
(Ewanchuk and Williams 1996).
Plants
Bulthuis, D.A. and W.J. Woelkerling. 1983. Seasonal
variations in standing crop, density and leaf growth
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______. 1985. Role of daily light period in the depth
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______. 1986. Photoadaptation and growth of Zostera
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at the Metropolitan Oakland International Airport.
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______. 1989. California least tern breeding season at
the Metropolitan Oakland International Airport.
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6
Baylands Ecosystem Species and Community Profiles
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McMillan, C. P.L. Parker and B. Fry. 1980. 13C/12C ratios in seagrasses. Aquat. Bot. 9: 237-249.
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Merkel, F.W. 1990. Eelgrass transplanting in south San
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Chapter 1 —
Plant Communities
7
Plants
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Setchell, W.A. 1929. Morphological and phenological
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Baylands Ecosystem Species and Community Profiles
Van Montfrans, J., R.L. Wetzel and R.J Orth. 1984.
Epiphyte-grazer relationships in seagrass meadows:
Consequences for seagrass growth and production.
Estuaries. 7: 289-309.
Walker, D.I., R.J. Lukatelich, G. Bastyan and A.J.
McComb. 1989. Effect of boat moorings on
seagrass beds near Perth, Western Australia.
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Wetzel, R.G. and P. Penhale. 1983. Production ecology
of seagrass communities in the lower Chesapeake
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Wyllie-Echeverria, S. and P.J. Rutten. 1989. Inventory
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Wyllie-Echeverria, S. 1990. Distribution and geographic
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37-86.
Introduction
The general ecology San Francisco Bay has been reviewed by Josselyn (1983), who included a brief treatment of its tidal marsh plant community composition
and structure. Macdonald (1977, 1988) reviewed the
vegetation of California salt marshes, including San Francisco Bay, with emphasis on sedimentation, drainage, topography, salinity, flooding, community structure, and
summaries of autecology of selected dominant species.
Newcombe and Mason (1972) made descriptive accounts of the Suisun Marsh area vegetation. Atwater et
al. (1979) summarized and interpreted the relationships
between tidal marsh vegetation of the San Francisco
Estuary and its landforms and geomorphic processes.
Wells and Goman (1994) reviewed and expanded the
quaternary history of the San Francisco Estuary. The
purpose of this plant community profile is to supplement
previous reviews, and provide additional information on
historic changes in the composition, distribution, and
abundance of tidal marsh plants of the Estuary.
Environmental Setting
Prehistoric Tidal Marsh Development – Tidal
marshes of the modern San Francisco Estuary formed
around 10,000 years ago during the Holocene submergence when the rate of sea-level rise slowed sufficiently
for tidal marsh sediments to accrete near sea-level
(Atwater et al. 1979). Prior to that time, during the Pleis-
Tidal Marsh along
Petaluma River shows
complex channels and
natural salt pans.
Josh Collins
Peter R. Baye
Phyllis M. Faber
Brenda Grewell
Chapter 1 —
Plant Communities
9
Plants
Tidal Marsh Plants of the
San Francisco Estuary
tocene epoch, the site of the modern Estuary consisted
of broad stream valleys far above glacial low sea level.
Pleistocene tidal marsh plant communities were probably associated with either stream mouths or backbarrier lagoons at the edge of an emergent broad coastal
plain, now submerged and eroded or buried offshore
from the modern Golden Gate. Tidal marsh plant species probably migrated upstream in valleys and embayments as sea level rose. Ancestral Pleistocene populations
of tidal marsh plant species in today’s estuaries may not
have been as discontinuously distributed as they are today: coastal plain shorelines (e.g., East Coast of North
America) often provide widespread tidal inlets and tidal
marsh (Davies 1980). Holocene fragmentation of salt
marshes from more extensive or continuous Pleistocene
coastal plain salt marsh distributions may account for historic disjunct, relict populations of species in San Francisco
Bay which are now found only in south-central or southern California tidal marshes (e.g., Solidago confinis (southern goldenrod), Suaeda californica (California sea-blite)).
Tidal marshes of the Estuary fluctuated in vegetation composition during the Holocene epoch, apparently
in relation to changes in long-term climate. This is indicated by stratified deposits of fossil pollen and plant
fragments which indicate periods of accumulation of
plants associated with near-freshwater marsh conditions
with species associated the more saline (brackish) conditions (Wells and Goman 1994). These findings are
consistent with independent evidence on climate changes
during the last 6,000 years which show prolonged periods of drought and high rainfall compared with historic
conditions (Ingram et al. 1996, Peterson et al. 1989).
The tidal marshes in San Francisco Bay were also not
static prior to European influence. Some marsh shoreline configurations indicate long-term scarp retreat across
marshes with large sinuous tidal creeks and growth of
berms and sand spits (Atwater et al. 1979). Areas of rapid
marsh growth in some parts of south San Francisco Bay,
Peter Baye
Plants
outside the influence of Sierran gold mining and prior
to extensive diking, were evident in maps of the Bay prepared in the 1870s (U.S. Coast Survey maps).
Marsh Sediments and Plants – Depositional environments of tidal marshes in the San Francisco Estuary are variable and are significant for the distribution
of uncommon plant species. In most of the San Francisco Estuary, the sediments of the middle-marsh marsh
plain consists of bay mud (fine silt and clay) with significant percentage of organic matter in mature marshes.
Local coarse sediment deposits, often beach ridges
(marsh berms, or marsh-beach ecotones) composed of
sand, shell fragments, organic debris, or mixtures, create physically mobile (periodically eroded and redeposited), well-drained high marsh habitats with affinity for
some common high marsh species (e.g., Grindelia stricta
var. angustifolia, gumplant) and probably also species
now locally extinct or rare, such as Suaeda californica,
Atriplex californica (California saltbush), and Castilleja
ambigua (salt marsh owl’s clover or Johny-nip). Marsh
berms are associated with relatively high wave energy environments in the Estuary, located near coarse sediment
sources such as eroding bluffs, submerged fossil sand and
shell deposits, stream mouths, and eroding marsh edges.
Such features were commonly represented on U.S. Coast
Survey maps of the mid-1800s, but persist today in very
few localities of the Estuary (e.g., Point Pinole, Redwood
City area, San Leandro area, and northern San Francisco
peninsula). Similar coarse-sediment features probably occurred as natural levees of upstream reaches of large tidal
sloughs with significant coarse sediment loads, as is observed today in Morro Bay. Alluvial fans also create
gradually sloping ecotones with uplands, with variably
textured sediments and freshwater runoff and seeps. Few
small alluvial fans exist at tidal marsh edges of the Estuary today (e.g., Point Pinole, Whittell Marsh), but were
historically abundant in parts of the Estuary, supporting diverse ecotonal plant communities (Cooper 1926).
Analogous alluvial fan-tidal marsh ecotones occur in
maritime salt marshes of Point Reyes and Tomales Bay
areas, where they support distinctive local plant assemblages, including uncommon to rare species.
Tidal Marsh Plant Communities
Regionall rare salt marsh owl’s clover, or Johnny-nip
(Castilleja ambigua ssp. ambigua). (Tidal marsh,
Whittell Marsh, Point Pinole)
10
Comparison With Other Estuaries – The tidal
marshes of the San Francisco Estuary are the most extensive on the central coast of California, and their plant
communities are distinct from other central coast tidal
marshes in many respects. Most other large central coast
tidal marshes are associated with shallow embayments
with large open tidal inlets (e.g., Tomales Bay, Drakes
Estero and Bolinas Lagoon in Marin County; Bodega
Bay in Sonoma County; Elkhorn Slough in Monterey
County; Morro Bay in San Luis Obispo County) which
impose strong marine influence on the character of their
sediments, salinities, and vegetation. Central coast tidal
marshes tend to be isolated and few because of the steep
modern shoreline with few valleys or wave-sheltered
bays. These tidal marshes have extensive sandy substrates, relatively small, local inputs of fine sediment and
freshwater discharges and brackish (mesohaline) conditions, and are inundated by water approaching marine
salinity (34 ppt) during most of the growing season.
Some tidal marshes associated with stream mouths have
relatively more freshwater influence and brackish marsh
vegetation (e.g., pre-historic Elkhorn Slough and Salinas River, Monterey County; Russian River estuary,
Sonoma County), but in association with seasonal reduction in tidal influence because of partial or complete closure of coastal inlets at river mouths (dammed by sand
beach ridges during periods of relatively low river discharge). In contrast, the tidal marsh plant communities
of the San Francisco Bay Estuary developed under conditions of abundant and predominantly fine sediment
(bay mud, clayey silts and silty clays with high nutrientholding capacity), relatively large tidal range, and extensive brackish marshes associated with relatively large
freshwater discharges, distributed over broad, fluctuating salinity gradients (Atwater et al. 1979)
Historically, salt pans (unvegetated, seasonally inundated depressions or flats within the tidal marsh) and
local salt ponds (perennial deposits of crystalline salt in
hypersaline ponds) were well-developed in San Francisco
Bay tidal marshes (U.S. Coast Survey T-charts, 1850s),
supporting distinctive vegetation (widgeongrass, Ruppia
maritima, in some pans) or microalgal floras (in salt
ponds). Pans are relatively infrequent in other central
coast tidal marshes compared with the historic conditions of the San Francisco Estuary, and natural salt ponds
were not known to occur in other central coast tidal
marshes. Today, edges of high marsh pans are associated
with at least two regionally rare species (Cordylanthus maritimus ssp. palustris and Castilleja ambigua ssp. ambigua),
and may have been associated with many others in the past
(e.g., Lepidium latipes, L. oxycarpum; Table 1.3)
Baylands Ecosystem Species and Community Profiles
The distribution of tidal marsh plants is strongly (but
not exclusively) influenced by tidal elevation and salin-
Peter Baye
Peter Baye
ity (Hinde 1954, Atwater and Hedel 1976). Following
Peinado et al. (1994), three elevation “ zones” of the tidal
marsh can be objectively distinguished (and are visually
conspicuous): (1) the low marsh zone, occurs from approximately mean sea level to mean high water; (2) the
middle marsh zone, occurs from approximately mean
high water to mean higher high water; and (3) the high
marsh zone (colloquially also called the “ upland transition” or “ peripheral halophyte” zone; “ upper salt marsh
zone” of Peinado et al. 1994), occurs near and above
mean higher high water up to several meters above extreme high water line (Peinado et al. 1994). The typical
species composition of these zones is described below for
tidal salt marsh and tidal brackish marsh. Unlike beach,
dune, and bluff communities (Barbour and Johnson
1977, Barbour et al. 1973), there is no empirical evidence of salt marsh zonation attributable to salt spray;
estuaries and embayments are relatively low-energy wave
environments (Davies 1980, Carter 1988). Other potentially significant influencing factors have not yet been
well studied.
There are significant floristic differences between
the tidal marshes of San Francisco Estuary and other
central coast tidal marsh systems. These include:
1. The dominance of Spartina foliosa (Pacific cordgrass), either absent today or historically absent
from most or all other central coast tidal marshes
(Macdonald 1977);
2. The presence of rare species of disjunct distribution, such as Suaeda californica (native only to
Morro Bay and San Francisco Bay); and
3. The presence of local endemic species such as soft
bird’s-beak (Cordylanthus mollis ssp. mollis) and
Suisun thistle (Cirsium hydrophilum var. hydrophilum).
Conversely, some uncommon tidal marsh species
which have either declined severely or become extirpated
in the San Francisco Estuary still occur in local abun-
Soft bird’s-beak (Cordylanthus mollis ssp. mollis). The
hairy bracts of the flowering stems are jeweled with
salt-encrusted glands. (Brackish tidal marsh, Southhampton Marsh, Benecia)
Chapter 1 —
Plant Communities
11
Plants
Pickleweed (Salicornia virginica), a dominant within
the salt marsh middle marsh zone, shown enshrouded
by parasitic dodder (Cuscuta salina).
dance in some maritime salt marshes of the region (e.g.,
Atriplex californica, Castilleja ambigua, Puccinellia nutkanensis). Few species associated with high marsh zones
of maritime salt marshes in the region were historically
absent from the San Francisco Estuary (e.g., Astragalus
pycnostachyus var. pycnostachyus (coastal marsh milkvetch), Castilleja ambigua ssp. humboldtiensis (Humboldt
Bay owl’s clover), Leymus x vancouveriensis (Vancouver’s
ryegrass), and Grindelia stricta var. stricta (gumplant)).
Differences exist also between the structure of vegetation found in predominantly marine-influenced salt
marshes of the central coast and tidal marshes of the San
Francisco Estuary. Although the middle marsh zone of
San Francisco Bay salt marshes has been described as
supporting “ prostrate” growth forms of pickleweed
(Macdonald 1977), the middle marsh plains of sandy or
sandy peat salt marshes of Bolinas Lagoon (Allison 1992),
Point Reyes, Tomales Bay, and Morro Bay often support
very thin, low (< 10 cm) turf-like vegetation mosaics with
extremely short, sparse, or prostrate pickleweed as a relatively minor component, or at most co-dominant with
species such as Triglochin concinna (slender sea arrowgrass; uncommon to rare in San Francisco Estuary).
These salt marsh turfs often support high plant species
diversity compared with San Francisco Bay salt marsh
plains, which tend to be dominated by pickleweed,
which often grows in dense stands (usually over 20 cm
thick; up to 50-60 cm in some fringing marshes of San
Pablo Bay). Low, turf-like middle marsh vegetation is
very uncommon in San Francisco Bay, both in brackish
and salt marshes.
Salt Marsh Plants and Their Associations – Salt
marsh here refers to tidal marsh plant associations that
approximate the species composition typical of nearmarine salinity during the growing season (34 ppt). Few
if any salt marshes in the San Francisco Estuary are ac-
Peter Baye
Locally rare Point Reyes bird’s-beak (Cordylanthus
maritimus ssp. palustris), is abundant in salt marshes of
Tomales Bay, Bolinas Lagoon, and Limantour Estero.
(Salt marsh, Marin City shoreline)
12
Baylands Ecosystem Species and Community Profiles
Peter Baye
Plants
tually regularly exposed to near-marine salinity, but in
the upper estuarine salinity range (roughly 20 ppt and
above), they are effectively salt marsh in vegetation character. The salt marsh plant community is typical of San
Francisco Bay and the outer marshes of most of San
Pablo Bay.
The low salt marsh zone in San Francisco Bay is
usually dominated by a single species, Spartina foliosa
(Pacific cordgrass), but is increasingly becoming dominated by the invasive introduced Atlantic species, Spartina alterniflora and its highly variable hybrids and novel
“ ecotypes” (Callaway and Josselyn 1992; Daehler and
Strong 1994, 1997; Daehler et al. 1999). S. foliosa stands
occur as uniform fringes along tidal creek banks or as
broad uniform plains on prograding marshes at the edges
of broad tidal mudflats. They extend from approximately
mean high water to mean sea level (Hinde 1954, Atwater
et al. 1979). On gentle elevation gradients, they intergrade with middle marsh plains in mixed stands of Salicornia virginica (pickleweed), as at Dumbarton-Mowry
marsh and eastern San Pablo Bay fringe marshes. They
may also occur as abrupt zones at the edge between tidal
mudflats and wave-cut peat scarps. Pioneer colonies of
Pacific and smooth cordgrasses on mudflats are abundant in some years, particularly in years of high or late
rainfall. They apparently establish by seedlings and regenerated rhizome fragments, but the relative proportion of these of propagule types is unknown. Pioneer
colonies of S. alterniflora were observed on open mudflats of the San Lorenzo Creek delta in 1991, and are visible in aerial photographs of the Alameda Creek area
around 1980. Seedlings and pioneer colonies of S. foliosa
were common on high mudflats of San Pablo Bay and
its tributaries in the late 1990s. The taller S. alterniflora
appears to be able to spread clonally below mean sea level,
but long-term comparisons of colonial spread between
Annual pickleweed (Salicornia europaea)— Occasionally found in conspicuous colonies on higher mudflats
between upper cordgrass and lower pickleweed
zones, it more commonly grows as a short, dense single
plant. It turns brilliant crimson in fall, in contrast with the
dominant dull green-brown Salicornia virginica.
native and introduced cordgrasses have not yet been conducted. The only other species of the low marsh is Salicornia europaea (annual pickleweed), which occasionally
occurs in the upper edge of the zone, often in accreting
high mudflats in transition between low and middle
marsh zones.
The middle salt marsh zone composes the extensive salt marsh plains of San Francisco Bay (Hinde 1954,
Atwater et al. 1979). Younger marshes tend to be characterized by low-diversity vegetation dominated by Salicornia virginica (Cuneo 1987), but some older marsh
remnants (e.g., Greenbrae and Heerdt Marsh; upper
Newark slough marsh) may comprise complex and annually variable mosaics of S. virginica, Distichlis spicata
(saltgrass), Cuscuta salina (salt marsh dodder), Jaumea
carnosa (fleshy jaumea), Frankenia salina (alkali-heath)
and Atriplex triangularis (spearscale or fat-hen). Species
diversity in the middle salt marsh is not necessarily correlated with marsh age: old marshes at China Camp and
Whittell Marsh (Point Pinole) also support relatively lowdiversity vegetation dominated by S. virginica. The parasitic Cuscuta salina (dodder) can become conspicuously
co-dominant or even dominant in the middle marsh zone
by mid-summer in some marshes in some years, turning the middle marsh into an orange and green mosaic
visible at great distances (Dumbarton-Mowry marsh, San
Pablo Bay fringe marshes). Colonization and species recovery dynamics associated with dodder-induced dieback
of marsh vegetation have not been investigated.
Relatively uncommon species of the middle marsh
zone of San Francisco Bay include Triglochin maritima
(sea arrow-grass), Limonium californicum (sea-lavender),
and Polygonum prolificum and P. patulum (non-native
knotweeds). Reports of the rare Point Reyes endemic
Polygonum marinense (Marin knotweed) in San Francisco
Bay require taxonomic verification. Species which sel-
Josh Collins
Tiidal salt marsh low marsh zone dominated by Pacific
cordgrass (Spartina foliosa).
coides or L. x multiflorus (creeping wildrye), or Juncus
lesueurii (salt rush, wire rush), as still occurs commonly
in maritime salt marshes of the region.
Cooper (1926) described a broad high salt marsh
zone along the Palo Alto shoreline dominated by Distichlis spicata and Grindelia stricta var. angustifolia (an association still evident in reduced extent today), and a high
salt marsh - alluvial transition zone which no longer
exists. Cooper’s reconstructed high salt marsh ecotone
community was dominated by native composites—
Hemizonia pungens ssp. maritima, H. congesta (tarweeds),
Helianthus bolanderi (Bolander’s sunflower), Aster subulatus (as “ A. exilis” ; slim or salt marsh aster), Aster chilensis
(Chilean aster; possibly also including the rare A. lentus),
Baccharis douglasii (salt marsh baccharis), Euthamia
occidentalis (western goldenrod), and Iva axillaris (poverty weed). Of these, I. axillaris, A. chilensis, H. pungens,
B. douglasii, and E. occidentalis still occur in high tidal
brackish marsh of San Pablo Bay and Suisun Marsh. It
therefore appears likely that historic upper edges of some
salt marshes were at least locally brackish or subsaline
rather than hypersaline in character, influenced by surface and subsurface freshwater discharges. This is also
indicated by Cooper’s description of water table-dependent, salt-intolerant tall (to 9 m) thickets of willow, cottonwood, box-elder, ash, blackberry, ninebark, and California rose at the high marsh edge (Salix lasiolepis,
Populus trichocarpa, Acer negundo, Fraxinus oregona,
Rubus ursinus, Physocarpus capitatus, Rosa californica).
Cooper (1926) interpreted this community from isolated
remnants of what he assumed was undisturbed vegetation, but the disturbance history of the South Bay marsh
edge at the time of his observations, and older reports
he collected, is uncertain.
The high salt marsh zone also historically included
many other native species, which are now uncommon,
rare, or extirpated in San Francisco Bay (Table 1.3).
Most of these still persist at other California salt marsh
localities. Most high salt marsh zones in San Francisco
Bay today occur on artificial slopes and substrates at the
upper marsh edge, and include many non-native species
that sometimes dominate the zone. Common non-native plants of the high salt marsh zone include Lepidium
latifolium (broadleaf peppercress, perennial peppergrass),
Bassia hyssopifolia (bassia), Salsola soda (saltwort), Beta
vulgaris (wild beet), Mesembryanthemum nodiflorum (annual iceplant), Carpobrotus edulis and its hybrids (iceplant), Atriplex semibaccata (Australian saltbush), Bromus
diandrus (ripgut brome), Hainardia cylindrica and
Parapholis incurva (sicklegrasses), and Polypogon monspeliensis (rabbit’s-foot grass).
Brackish Marsh Plants and Their Associations
– Brackish tidal marshes prevail over northern San Pablo
Bay (slough systems of the Petaluma River, Tolay Creek,
Sonoma Creek, and Napa River), the Suisun Marsh area,
and the Contra Costa marshes (North Bay marshes).
Chapter 1 —
Plant Communities
13
Plants
dom occur in the middle salt marsh zone of San Francisco Bay include Cordylanthus maritimus ssp. palustris
(Pt. Reyes bird’s-beak; Richardson Bay, Heerdt Marsh),
Puccinelia nutkaensis (Pacific alkali grass; Ravenswood
fringe marshes and Newark), Plantago maritima (seaplantain) and Triglochin concinna (slender arrow-grass).
These latter species are locally abundant in maritime salt
marshes of Marin County. Invasive exotic species of the
middle salt marsh include Spartina densiflora (Chilean
cordgrass; Richardson Bay and Point Pinole), Spartina
patens (saltmeadow cordgrass; near Burlingame and in
brackish middle marsh at Southhampton Bay) and
Cotula coronopifolia (brass buttons; early introduction,
widespread but never persistent as a dominant in tidal
marsh). The invasive exotic Salsola soda (Mediterranean
saltwort) also is spreading from high salt marsh to the
middle marsh zone (Dumbarton-Mowry marsh).
High or upper salt marsh may occur as topographic
highs within the marsh plain (e.g., channel bank levees,
wave-deposited ridges or mounds) or along the upland
or alluvial edges of the marsh. This zone today commonly includes natives such as Grindelia stricta var. angustifolia (frequently a dominant in this zone), Distichlis spicata, erect-ascending phenotypes of Salicornia
virginica, Cuscuta salina, Frankenia salina, Limonium
californicum (sea-lavender), and Atriplex triangularis
(spearscale, fat-hen). Where the upper marsh intergrades
with low-lying alluvial soils and high groundwater (a condition today very rare in San Francisco Bay), the high
marsh zone is dominated by dense stands of Leymus triti-
14
Baylands Ecosystem Species and Community Profiles
Peter Baye
Plants
They also occur in transition with San Francisco Bay salt
marshes where significant freshwater discharges occur
(e.g., fringing marshes of Mud Slough, Coyote Creek,
Artesian Slough, Alviso Slough, and Guadalupe Slough).
The distinction between “ salt marsh” and “ brackish
marsh” is a recent convention in descriptions of San
Francisco Bay Area tidal marshes: brackish marshes were
indiscriminately described as “ salt marshes” by early California botanists, making it difficult to separate distinct
elements of “ salt” and “ brackish” marsh associations.
The description and demarcation of brackish marsh
plant communities is essentially a matter of convenience
and convention: there is no precise, stable salinity threshold at which tidal marshes are known to switch from one
“ type” to another (Adam 1990). Instead, brackish marsh
vegetation in the San Francisco Estuary is typically a dynamic continuum between salt marshes of San Francisco
Bay and freshwater tidal marshes of its major tributary
rivers, fluctuating with variable influence of rainfall and
freshwater discharges which alter marsh salinity and vegetation gradients geographically and over time. Associated changes in local tidal elevations (related to freshwater discharges) may also possibly interact with salinity
variations in altering the character of brackish marsh
vegetation. Changes in brackish marsh vegetation between dry and wet years at the same location may be
dramatic: cover can change from that typical of San Francisco Bay salt marsh (dominant pickleweed) to that typical of Suisun Marsh (mosaic of rushes, bulrushes, alkalibulrush, cattails, saltgrass, and many broad-leaved
herbaceous species) in very few years. The causes of these
dramatic changes in brackish tidal marsh vegetation are
presumably related to plant interactions (competition,
facilitation, and parasitism) which are influenced by seasonal and annual variation in salinity and drainage
(Pearcy and Ustin 1984), but are poorly understood
beyond descriptive observation.
The most extensive tidal brackish marshes occur
in the Petaluma Marsh, but relatively large relict tidal
brackish marshes also occur along the Napa River (Fagan
Slough marsh) and in the Hill Slough/Rush Ranch area
in Suisun Marsh. Relatively young but large and welldeveloped brackish marshes also occur bayward of dikes
constructed after the 1870s, particularly in the NapaSonoma marsh complex and Suisun Marsh, including
marsh islands of Suisun Bay. The Contra Costa marshes
are predominantly intermediate between fully tidal
marsh and diked (reduced tidal range) brackish marshes.
The extensive wave-influenced, prograded pickleweeddominated marsh plain and low natural marsh levee
along northern San Pablo Bay are transitional between
salt marsh and brackish marsh, exhibiting increases in
brackish-associated species (particularly Scirpus maritimus at the east end of the Bay) in series of wet years.
Plant species richness and diversity markedly increase in brackish marshes of the San Francisco Estuary
Sea-milkwort (Glaux maritima) is found in tidal marshes
on the northern Pacific Coast, and on the Arctic,
American, and European Atlantic coasts. (Tidal marsh,
Rush Ranch, Suisun Marsh)
compared with salt marsh. Grewell (1993 et seq.) compiled extensive vascular plant species lists of the Suisun
Marsh (including uplands of dikes and artificial uplands),
and presented the only comprehensive and contemporary synthesis of Suisun Marsh plant ecology and its history (Grewell et al. 1999). Mason (Newcombe and
Mason 1972) described plant community composition
of brackish tidal marshes extending into the SacramentoSan Joaquin Delta.
The low brackish marsh zone differs from the corresponding zone in the San Francisco Estuary salt
marshes in several respects: it supports multiple dominant species in variable mixtures or monospecific stands;
it extends to the low end of intertidal zone, and it regularly develops tall, dense vegetation. In San Pablo Bay
and western Suisun Marsh, alkali-bulrush (predominantly Scirpus maritimus around San Francisco and San
Pablo bays and western Suisun Marsh, but also including S. robustus, a taxon formerly misapplied to S. maritimus in floras of the region) occurs in the upper portion of the low marsh, often dominant in the saline end
of the brackish marsh gradient. The tallest graminoid
species, tules and cattails, dominate where freshwater
influence is relatively strong; these include Typha angustifolia, T. latifolia, T. dominguensis and hybrids; Scirpus
californicus (California tule), S. acutus (hardstem tule)
and hybrids. These graminoid species can also establish
within poorly drained portions of the middle marsh
plain.
The middle brackish marsh zone was historically
dominated by Distichlis spicata (saltgrass), as it commonly
is today (Newcombe and Mason 1972). Other native
species of the high marsh which occur in variable abun-
Peter Baye
Ambrosia psilostachya (western ragweed), Euthamia
occidentalis (western goldenrod), Epilobium brachycarpum, E. ciliatum (willow-herbs), Polygonum spp. (smartweeds, knotweeds), Triglochin maritima (sea arrow-grass)
and Eryngium articulatum (coyote-thistle). Uncommon
to rare species such as Lathyrus jepsonii var. jepsonii
(Delta tule pea), Aster lentus (Suisun aster), A. subulatus
var. ligulatus (slim aster), Plantago elongata (dwarf
plaintain), Rumex occidentalis (western dock), Eleocharis parvula (spikerush), and endangered Cordylanthus
mollis ssp. mollis (soft bird’s beak) and Cirsium hydrophilum var. hydrophilum (Suisun thistle) typically occur locally in the lower end of well-drained high marsh gradient, often on slight topographic relief above the marsh
plain. Salicornia virginica (common pickleweed) and
occasionally S. subterminalis (Parish’s glasswort) can also
be abundant elements of high brackish marsh near Suisun. The composition of high brackish marsh vegetation
appears to vary with slope, drainage, and local surface
or subsurface freshwater influence, but no studies have
yet analyzed vegetation patterns or related environmental factors in brackish marshes of the region.
Invasive non-native species (weeds of mesic and
wetland habitats with slight salt tolerance) of the high
brackish marsh zone are numerous, particularly in years
of high rainfall, but the most aggressive and successful
is again Lepidium latifolium. Lotus corniculatus (bird’sfoot trefoil) and Lolium multiflorum (ryegrass) are other
exotics which are locally abundant along portions of the
upper brackish marsh edge some years. Elytrigia pontica
ssp. pontica (tall wheatgrass, currently local around
Alameda Creek and Mare Island), Rumex crispus and R.
pulcher (curly and fiddle docks), Asparagus officinalis (locally abundant near Napa-Sonoma marshes) have also
naturalized along brackish marsh edges, but are seldom
invasive.
Ditch-carrot (Oenanthe sarmentosa), a common
freshwater marsh plant, also occurs in fresher phases of
brackish tidal marshes. (Southhampton Marsh, Benicia)
Chapter 1 —
Plant Communities
15
Plants
dance (common to co-dominant) include Salicornia virginica, Atriplex triangularis, the Juncus balticus-lesueurii
complex, Jaumea carnosa, Frankenia salina and Cuscuta
salina. Locally common natives include Limonium californicum (sea-lavender), Glaux maritima (sea-milkwort),
and Scirpus koilolepis cernuus and S. cernuus (clubrush;
also in high brackish marsh), Eleocharis macrostachya
(creeping spikerush), Helenium bigelovii (Bigelow’s
sneezeweed), and Deschampsia cespitosa ssp. holciformis
(tufted hairgrass; especially eastern Suisun Marsh). Infrequent to rare species of this zone include Lilaeopsis
masonii and L. occidentalis (Mason’s and western lilaeopsis; on exposed eroding channel bank edges as far west
as Tolay Creek), Triglochin maritima (locally common),
T. concinna, T. striata, Sium suave (water parsnip),
Oenanthe sarmentosa (ditch-carrot), Cicuta maculata ssp.
bolanderi (water hemlock), Eleocharis parvula (slender
spikerush), Pluchea odorata (salt marsh fleabane), and
Lythrum californicum (California loosestrife; eastern Suisun Marsh and Delta). In wet years, depressions in the
middle marsh plain support increased abundance of Scirpus americanus (Olney’s bulrush) or S. maritimus (alkalibulrush; western Suisun and San Pablo Bay) and Phragmites australis (common reed; eastern Suisun Marsh, also
in the low-middle marsh zone). The dominant non-native species of the middle brackish marsh is again Lepidium latifolium, which rapidly forms dense monotypic
clonal populations, spreading into the marsh plain.
Other exotic species which have established in the brackish middle marsh zone include Apium graveolens (wild
celery, widespread and abundant in Suisun Marsh),
Lythrum hyssopifolium (annual loosestrife), Cotula coronopifolia (brass-buttons) and Chenopodium chenopodioides (fleshy goosefoot; Napa-Sonoma marshes).
The high brackish marsh zone is today typically
altered by artificial dikes and invasive plants (particularly
Lepidium latifolium (perennial or broadleaf peppercress),
Conium maculatum (poison hemlock), Foeniculum
vulgare (fennel), and Mediterranean grasses. However,
many native remnants of the brackish high marsh community have regenerated on old, stable, relatively undisturbed levees, or have persisted locally along undiked
tidal marsh edges. They include Achillea millefolium (yarrow), Baccharis douglasii (salt marsh baccharis), B.
pilularis (coyote-brush), Leymus triticoides and L. x
multiflorus (creeping wildrye), Scrophularia californica
(California bee-plant), Rubus ursinus (blackberry, in the
upland ecotone) Rosa californica (California rose, also in
the upland ecotone), Iva axillaris (poverty-weed), Atriplex triangularis (fat-hen or spearscale), Grindelia stricta
var. angustifolia (and intermediates with G. camporum),
Calystegia sepium ssp. limnophila (morning-glory), Cressa
truxillensis (alkali-weed), Frankenia salina (alkali-heath),
Lathyrus jepsonii var. californicus (California tule pea),
Juncus balticus - lesueurii complex (salt or wire rush),
Juncus mexicanus (Mexican rush), J. bufonius (toad rush),
High marsh pan in
Whittell Marsh (Point
Pinole, Contra Costa
County), fringed with
salt-marsh owl’s clover
(Castilleja ambigua).
Whittell Marsh is the last
known tidal marsh
locality of this species in
the San Francisco
Estuary.
Peter Baye
Plants
Tidal Marsh Pans and Vegetation – Poorly
drained flats, depressions, and barrier-impounded areas
of tidal marsh lacking emergent vascular vegetation,
called pans (alternatively spelled “ pannes” ), range from
nearly planar unvegetated marsh areas subject to shallow periodic ponding, to steep-sided or cliff-edged shallow ponds which are persistently inundated (Pestrong
1965, Pethick 1974, Atwater et al. 1979). Pans have
various modes of origin and development, which have
not been completely clarified (Adam 1990, Carter 1988,
Pethick 1974, Chapman 1960). In San Francisco Bay
Area marshes, pan variation includes nearly circular
ponds between drainage channels (Pestrong 1965), historic long ponds parallel with impounding bayfront
marsh berms (Atwater et al. 1979), shore-parallel pans
historically present along portions of the back edge (upland or lowland margin) of tidal marsh (depicted in
1880s U.S. Coast Survey Maps), and natural historic salt
ponds impounded by low estuarine ridges (Atwater et
al. 1979). Some sloped to planar pans in the high marsh
(bare flats, rarely submerged) may be related to wrack
deposition and smothering, or local substrate conditions.
Little is known of the ecology of pan types that are no
longer represented in the altered modern Estuary.
Many pans are reported to become seasonally hypersaline (Pestrong 1965) or even salt-crystallizing
(Atwater et al. 1979; see also salt pond profile, this volume) and lack vascular plants, but some pans along the
landward edge of the tidal marsh develop marginal vegetation typical of brackish or fresh marshes (e.g., China
Camp). Ponded pans within the marsh plain have been
described as “ unvegetated” (Pestrong 1965), but they
often support a dense submerged mixed vascular and
non-vascular vegetation variously composed of widgeongrass (Ruppia maritima) and membranous green algae
16
Baylands Ecosystem Species and Community Profiles
(particularly Enteromorpha and Ulva spp.). According to
Mason (Newcomb and Mason 1972), brackish ponds in
Suisun Marsh also support Zannichellia palustris and
Potamogeton pectinatus, submerged species typical of
freshwater ponds. The halophilic microflora of salt ponds
is discussed in the salt pond profile(this volume). Shallow, relatively planar and ephemeral pans in San Pablo
Bay are either periodically or marginally colonized by
pickleweed, which dies back during years of frequent
flooding or high rainfall. The steep-sided edges of welldefined, nearly circular old pans sometimes develop small
natural levees of locally improved drainage, and sometimes support certain species at frequencies more typical of high marsh vegetation. In the high marsh, on gently sloping alluvial fans, “ dry pans” (small playa-like flats
with very short flooding periods and superficial salt films)
also develop, often on relatively coarse (sandy, silty, or
even gravel-silt mixtures) sediments. These features are
very rare today because of diking, but fine examples
persist at Point Pinole (Whittell Marsh). Here, as at similar pans on alluvial fans at tidal marsh edges in maritime
Marin County, the pan-marsh edges are associated with
local abundance of the regionally rare salt marsh owl’s
clover (Castilleja ambigua ssp. ambigua; salt-tolerant
ecotypes). Salt marsh bird’s-beak (Cordylanthus maritimus ssp. palustris) also exhibits a pan-margin local distribution pattern in western San Pablo Bay (e.g.,
JEPS83457). Analogous artificial features (gently sloping, formerly disturbed silty to sandy high marsh fills
with residual vegetation gaps) elsewhere in the Estuary
have also become colonized with rare plants such as
Cordylanthus mollis (Hill Slough near Lawler Ranch; B.
Grewell, pers. obs.) and Cordylanthus maritimus (near
Marin City). Natural and artificial high marsh pans of
this type, associated with alluvial or deltaic deposition
Peter Baye
Regionally rare smooth goldfields (Lasthenia glabrata).
(Whittell Marsh, Point Pinole, Contra Costa County)
or erosion, have not been identified in the regional literature on salt marsh ecology, and require study.
The number of species from former alkali-subsaline vernal pools around San Francisco Bay which were
historically reported from local salt marshes as well (see
diked wetlands profile, this volume) suggests that ecologically equivalent habitat occurred in both ecosystems.
Although there are very few intact remnants of the elongate pans which occurred along tidal marsh edges (represented clearly in historic U.S. Coast Survey maps of
the 1850s), it is possible that some of these seasonally
ponded depressions in the upper marsh ecotone were
partial ecological equivalents of subsaline vernal pools.
Strong historic evidence for this conclusion is found in
Jepson’s (1911) range and habitat descriptions for the
typical vernal pool species, Downingia pulchella, which
he described as “ abundant and of rank growth in the salt
marshes near Alvarado” [now Union City]. Other species indicative of vernal pools and similar seasonally
ponded/desiccated alkaline/subsaline environments,
such as Lasthenia conjugens (JEPS25099), L. platycarpha
(DS695549, Greene 1894) and L. glabrata (CAS897444,
DS73122, DS286573) have been collected from the
edges of San Francisco Bay.
Although pans are often presumed to be generally
hypersaline, some appear to have occurred historically
in alluvial lowlands with probable groundwater or surface discharges that could maintain brackish conditions
in pans along tidal marsh edges. A number of characteristic freshwater marsh species were reported by Jepson
(1911) and others from historic salt marsh habitat (e.g.,
Agrostis exarata, Carex aquatilis, C. densa, Lycopus asper),
suggesting that freshwater sub-habitats occurred marginally along tidal salt marshes. Unpublished historic writings of southeastern San Francisco Bay marsh borders
by 19th century botanist Joseph Burtt-Davy, archived
at the Jepson Herbarium, University of California, describe extensive colorful wildflower meadows with spe-
Uncommon, Rare, Declining, and Extirpated Plant Species
There is a widespread impression, even among ecologists
familiar with the San Francisco Estuary, that native plant
species richness of tidal marshes (particularly salt marsh)
is relatively low, and that rare species in the Estuary are
principally wildlife taxa, not plants. This impression is
due in part to reviews of species richness in tidal marshes
based solely on modern surveys: for example, Atwater et
al. (1979) reported only 15 vascular plant species for San
Francisco Bay, based on modern reports. Josselyn (1983)
discussed only a small representation of the San Francisco Estuary flora, and did not address either its historic
or modern species richness. In addition, very few plants
native to the San Francisco Estuary are federally listed
as endangered or threatened, and only two of these (soft
bird’s-beak, Cordylanthus mollis ssp. mollis, and Suisun
thistle, Cirsium hydrophilum var. hydrophilum) currently
inhabit this Estuary. The modern lack of attention to rare
plants in the Estuary is probably due to unfamiliarity
with plant species which were known only to early botanists, but are either now entirely extinct (or even extirpated) in the Estuary. Plant species that were historically
recorded in the tidal marshes of the Estuary, or along
its edges (high marsh), but have become uncommon,
rare, regionally extirpated, or extinct, are summarized in
Table 1.3. Most of these species were known from tidal
marsh edges, transitional habitats of high ecological diversity. This is significant, because original remnants of
this ecotone are almost completely eliminated from the
Estuary, and their modern counterparts are mostly
weedy, disturbed habitats like dikes.
Extinct species of the Estuary include California
sea-blite, Suaeda californica, a federally endangered
shrubby true halophyte (salt-tolerant plant) which today
inhabits relatively well-drained marshy beach ridges
along relatively high-energy shorelines with coarse sediment in Morro Bay. According to Jepson (1911) and
Greene (1894), it was never abundant in San Francisco
Bay even in the late 19th century. The distribution of
its sandy marsh habitats was unfortunately in areas of
the greatest urbanization: San Francisco, Oakland,
Alameda, and San Leandro were its core populations,
Chapter 1 —
Plant Communities
17
Plants
cies typical of vernal pools and wet grassland (R. Grossinger, pers. comm. 1999). Examples of brackish and
even freshwater vegetation at edges of salt marsh with
pans near zones of groundwater discharge can be observed today at China Camp (Marin County) and Point
Pinole (Contra Costa County), and in maritime Marin
County tidal marshes. Diked seasonal wetlands in historic tidal marsh (this volume) may also approximate this
type of lost habitat, since numerous seasonal wetland
species of vernal pools and alkali basins have colonized
diked Baylands.
Peter Baye
Plants
although it was also collected in Palo Alto (where shell
hash beaches today occur) and at the former San Pablo
Landing (Richmond, where local sand beaches still persist). The species today is restricted to sandy salt marsh
edges of Morro Bay, San Luis Obispo County, and also
exists in cultivation. It was last collected in San Francisco Bay in 1958 in San Leandro (JEPS25020) More
recent local reports are based on misidentification of the
similar species, S. moquinii, in diked Baylands.
Many other salt marsh species that have affinity
for high sandy salt marsh were also reported from San
Francisco Bay, but are now extinct or rare in the Bay
(Jepson 1911, Greene 1894). They include California
saltbush (Atriplex californica), still found in Tomales Bay
and Point Reyes sandy salt marshes, but extinct in the
Bay, and Plantago maritima, common in sandy maritime
salt marshes, uncommon to rare in the Bay. The sea-pink
(Armeria maritima) a showy pink spring wildflower
which still occurs locally along sandy edges of Point
Reyes salt marshes, was cited by Jepson (1911) to range
within San Francisco Bay. (This may possibly have been
along former sandy beaches, sandy salt marsh, or stabilized former bayside dunes. There are no historic herbarium specimens from San Francisco Bay salt marshes
to corroborate Jepson’s report, however.) Other rare species, such as Cordylanthus maritimus ssp. palustris and
Castilleja ambigua ssp. ambigua, are less uncommon in
sandy maritime salt marshes, but are rare in San Francisco Bay. The decline or demise of these species in the
Bay is very likely a result of the near-complete elimination of its sandy estuarine barrier beaches.
Two other species which are probably extinct in
San Francisco Bay, but occur elsewhere, include two
members of the Aster family: southern goldenrod (Solidago confinis) and Pyrrocoma racemosa (=Haplopappus
racemosa). Southern goldenrod was formerly reported as
rare only by Henry Bolander in 1863 (Jepson 1911),
Federally listed as endangered, California sea-blite
(Suaeda Californica) is extinct in San Francisco Bay.
(Morro Bay)
18
Baylands Ecosystem Species and Community Profiles
when it was misidentified as seaside goldenrod (S.
sempervirens). In California tidal marshes today, S.
confinis is known only locally from the high brackish
marsh zone of southern Morro Bay. P. racemosa was formerly reported from the edges of salt marshes and saline soils at Cooley’s Landing and near Alviso (Thomas
1961), but has not been reported from salt marsh edges
in recent decades. Another species, Adobe sanicle
(Sanicula maritima), was found locally in lowlands adjacent to salt marshes at Alameda (Behr 1888, Greene
1894, Jepson 1911) and in San Francisco (Brandegee
1892). It is now extinct in the Bay Area, and is very rare
elsewhere (known from fewer than 10 sites in Monterey
and San Luis Obispo Counties today; Skinner and
Pavlick 1994).
Two popcornflower species (genus Plagiobothrys,
well represented in vernal pools) that were found in saline soils near the edge of the Estuary are now presumed
to be extinct (although it is possible that buried dormant
seed may persist somewhere in diked Baylands, awaiting resurrection). They include Petaluma popcornflower
(Plagiobothrys mollis var. vestitus), which was probably
distributed in alkaline or subsaline seasonally wet depressions (vernal pools) in grasslands and lowlands adjacent
to tidal marsh in the Petaluma Valley, and Hairless
popcornflower (P. glaber), a species of seasonally wet alkaline/subsaline soils of tidal marshes of the south San
Francisco Bay (reported by Jepson (1911) from Alvarado
[now Union City]), as well as some interior valleys. Almost nothing is known of the ecology of these species
because of their early historic extinction.
Other species that are known to occur in subsaline
to alkaline vernal pools, and which historically occurred
in salt marshes (presumably along lowland edges), include several species of goldfields (Lasthenia spp.). Fleshy
goldfields, Lasthenia platycarpha (presumed extinct in the
Estuary) was known from salt marshes near Vallejo
(Greene 1894), and smooth goldfields (L. glabrata ssp.
glabrata) was reported from edges of salt marshes (Thomas 1961, Jepson 1911, Greene 1894). L. glabrata was
recently confirmed to occur naturally at Whittell Marsh,
Point Pinole, and a population of undetermined origin
occured briefly in 1998 on a hydroseeded levee at the
Sonoma Baylands tidal marsh restoration project’s pilot
unit. Behr (1888) listed L. glabberima as a species occurring “ near salt marshes,” but is not otherwise reported
from tidal marshes in the region. The federally endangered vernal pool goldfields species, Contra Costa goldfields (L. conjugens) was reported by Jepson (1911) from
“ subsaline soils” near Antioch and Newark, and was recently discovered in subsaline vernal pools in Fremont
near the diked edge of the at the Warm Springs Unit of
the National Wildlife Refuge in Fremont and adjacent
derelict fields. L. conjugens was also observed along high
tidal marsh edges of Hill Slough in the early 1990s. Another well-known vernal pool species, the showy Down-
Peter Baye
ingia pulchella (producing spring masses of blue, white,
and yellow flowers resembling lobelias) was described by
Jepson (1911) to occur abundantly in South Bay salt
marshes. It still occurs in the subsaline vernal pools adjacent to tidal marsh at the Warm Springs Unit of the
Refuge in Fremont, and in diked agricultural Baylands
(former tidal marsh) near Fairfield. The rare annual
milkvetch (locoweed), Astragalus tener var. tener, was formerly collected from “ saline areas along San Francisco
Bay” as far south as Mayfield (Mountain View area;
Thomas 1961). Once found in alkali vernal pools, it was
collected in the Bay Area in 1959 (Skinner and Pavlick
1994) and was recently rediscovered near the historic Bay
edge in Fremont (G. Holstein, pers. comm. 1999).
Two hemiparasitic annual snapdragon family herbs
are extinct in the salt marshes of south San Francisco
Bay, but occur elsewhere in the Estuary or region. The
Point Reyes bird’s-beak (Cordylanthus maritimus ssp.
palustris), a close relative of the endangered salt marsh
bird’s beak of Southern California (C. m. ssp. maritimus),
was formerly found almost throughout San Francisco
Bay. It is now restricted to very few populations in the
Central Bay, with small remnant populations probably
persisting in Petaluma Marsh and near Gallinas Creek,
Marin County. The remaining San Francisco Bay populations of Marin County are typically showier (usually
more conspicuous, rosy purple flowers and purplish
herbage) than most of the core populations of Point
Reyes, which typically have gray-green foliage and whiteand-maroon flowers. Another annual Snapdragon family herb, Johnny-nip or salt marsh owl’s clover (Castilleja
ambigua ssp. ambigua) was formerly found in the salt
marshes of San Francisco Bay (Berkeley, Oakland,
Alameda, Bay Farm Island, Burlingame), but is nearly
extinct there now. The only salt marsh population of this
colorful annual herb in the San Francisco Estuary is from
Point Pinole, which supports a form with purple-tinged
foliage, bracts, and flowers (atypical of the subspecies
Chapter 1 —
Plant Communities
19
Plants
The southern-most population of Point Reyes bird’sbeak (Cordylanthus maritimus ssp. palustris), in a small
marsh on the Marin City shoreline.
ambigua, but typical of ssp. insalutata of Monterey
County). Salt-tolerant locally adapted populations of this
subspecies also occur at Rodeo Lagoon and Bolinas Lagoon, but are otherwise rare in central coast tidal marshes
(very local in Limantour estero and Tomales Bay). A related salt marsh endemic subspecies, C. a. ssp. humboldtiensis, occurs only in Humboldt Bay and Tomales Bay.
Non-halophyte populations of C. a. ssp. ambigua occur
somewhat more widely in coastal grasslands, headlands,
and bluffs.
Still surviving but rare within its historic range in
brackish tidal marshes from Petaluma Marsh to Antioch
is another annual Snapdragon family herb, soft bird’sbeak (Cordylanthus mollis ssp. mollis). This white-yellow
flowered herb is covered with salt-encrusted secretory
glands. It is listed as federally endangered, and is restricted mostly to the Suisun Marsh area, especially in
old relict tidal brackish marsh. It formerly ranged as far
west as Petaluma Marsh (Howell 1949). Like the other
annual hemiparasitic salt marsh Snapdragon relatives, its
numbers fluctuate tremendously from year to year
(Rugyt 1994), sometimes disappearing for a year or more
before regenerating from dormant seed banks.
Numerous other species, particularly grasses and
sedge species, were cited by early California botanists as
commonly occurring in salt marshes, but are scarce or
absent today in the San Francisco Bay Area. By analogy
with relatively intact tidal marshes of Point Reyes to the
north and Elkhorn Slough to the south, it appears very
likely that these “ missing” salt marsh species occurred
along upland or lowland (alluvial) margins of tidal
marshes. Some, like Agrostis exarata (= A. asperifolia),
Juncus xiphioides, J. lesueurii, and J. effusus var. brunneus
were described as common in Bay Area salt marshes
(Jepson 1911, Brewer et al. 1880, Howell 1949), although they occur only very locally in Bay Area tidal
marshes today. Other grass species, like Leymus triticoides
(including L. x multiflorus), are presumed to be former
marsh edge dominants based on relict occurrences at
intact lowland tidal marsh edges (e.g., Rush Ranch,
Point Pinole) and colonizing behavior on levees which
have not been maintained (Dutchman Slough and Mare
Island, San Pablo Bay). The salt marsh grass Puccinelia
nutkaensis, in contrast, occurred in periodically inundated middle salt marsh zones in the South Bay as well
as on levees (Thomas 1961). It is rarely found in San
Francisco Bay today, such as near Ravenswood, Palo Alto
and Newark. Other grasslike plants, such as Plantago
elongata, were reported as common in Bay Area tidal salt
marshes (Brewer et al 1880, Greene 1894) but have become uncommon or rare here. Other grasslike plants of
uncertain former abundance in tidal marshes, which are
scarce or absent in Bay Area tidal marshes today, include
Carex aquatilis var. dives, C. densa, and C. praegracilis
(Thomas 1961, Jepson 1911); C. praegracilis occurs infrequently in tidal brackish marshes of the Suisun Marsh
stature of S. alterniflora enables it to endure high tides
with relatively little submersion of its foliage, even when
rooted below mean sea level. Turf-forming S. patens (salt
meadow cordgrass) and dwarf strains of S. alterniflora
(Daehler et al. 1999) present in the Bay may be latent
invaders of salt marsh plains. The tendency for S.
alterniflora pollen to swamp the pollen of the native S.
foliosa and produce hybrids and introgressants threatens
to genetically assimilate the native Pacific cordgrass over
a significant portion of its geographic range (D. Ayers
and D. Strong, pers. comm. 1999). The higher densities, larger plant size, and greater colonizing ability of
S. alterniflora at lower tidal elevations also suggest that
its spread may have significant geomorphic impacts on
the Estuary, particularly on channel stability, sedimentation, and mudflat colonization, and their indirect effect on wildlife habitat (Grossinger et al. 1998).
Lepidium latifolium invasion is particularly a concern for the conservation and recovery of rare or endangered plant species of the San Francisco Estuary, most
of which occur in the high marsh zone where L. latifolium is dominant. L. latifolium actively encroaches on
populations of endangered Cordylanthus mollis ssp. mollis
and Cirsium hydrophilum var. hydrophilum in Suisun
Marsh (B. Grewell, pers. obs. 1998) and Southhampton
Bay (P. Baye, pers. obs. 1998). The impact of exotic plant
invasions in the high marsh zone is magnified by the truncation and degradation of this habitat by widespread diking, which compresses the high marsh zone into a relatively
invariant, steep slope of disturbed Bay mud.
Tidal Marsh Restoration Design – Tidal marsh
restoration in the San Francisco Estuary has convention-
Conservation Issues
Exotic Plants – There are many exotic plants that
have become established within, or along the edges of,
the San Francisco Estuary, but only a few are aggressive
invaders that have become widespread and dominant, or
threaten to do so (Grossinger et al. 1998). Of these, Lepidium latifolium, Spartina alterniflora (and hybrids), and
Salsola soda have demonstrated ability for rapid, extensive invasion and development of monodominant stands
in the San Francisco Estuary. Spartina densiflora, an exotic cordgrass from Chile with a bunchgrass growth
habit, has become a dominant species in Humboldt Bay,
and is expected to be able to achieve the same dominance
if its spread is unchecked in San Francisco Bay. The taller
20
Baylands Ecosystem Species and Community Profiles
Peter Baye
Plants
area. Sedges such as Carex subbracteata, and C. obnupta
would also be expected to have occurred in former salt
marsh edges, as they do in other estuaries of the Central Coast, especially northward.
Many broadleaved herbs were also more plentiful
along tidal marsh edges, but have become localized or
rare today. They include Aster lentus (Greene 1984,
Jepson 1911), a species now generally rare in any estuarine habitat; Chilean aster (Aster chilensis) (Howell 1949,
Thomas 1961), a common species of non-saline habitats which has nearly disappeared from salt marsh edges
but persists occasionally in Suisun, Petaluma, and NapaSonoma marshes. Salt marsh baccharis (Baccharis douglasii) was formerly abundant in salt marshes (Jepson
1911) but is now uncommon to rare in brackish
marshes, mostly in the North Bay (Best et al. 1996, Thomas 1961). Two species which were inferred by Cooper (1926) to be major elements of his reconstructed
“ willow-composite” community at South Bay salt marsh
edges, slim aster (Aster subulatus var. ligulatus), and
spikeweed (Hemizonia pungens var. maritima) are now
scarce in tidal marshes, and occur mainly in the North
Bay (Best et al. 1961; B. Grewell, pers. obs. 1997). Other
spikeweeds, H. parryi sspp. parryi and congdonii, were
locally common in the South Bay salt marshes (Munz
1959), but are generally rare today. Species that were
formerly frequent in North Bay brackish and salt marshes
(Greene 1894) include morning-glory (Calystegia sepium
var. limnophila) and sea-milkwort (Glaux maritima),
which are now uncommon to rare. Other herbs which
have historically declined to a significant extent in frequency, distribution, and abundance in Bay Area tidal
marshes and their edges include Hutchinsia procumbens
(Greene 1894, Thomas 1961), tidy-tips, Layia
chrysanthemoides (Howell 1949, Thomas 1961), native
annual peppercress species Lepidium dictyotum, L. latipes,
and L. oxycarpum (Thomas 1961, Munz 1959, Howell
1949, Greene 1894), salt marsh fleabane, Pluchea odorata
(Jepson 1911), and butterweed, Senecio hydrophilus
(Greene 1894, Jepson 1911).
Invasive exotic Lepidium latifolium (background)
looms over the endangered Cordylanthus mollis ssp.
mollis at the high marsh edge. (brackish tidal marsh,
Southampton Marsh, Benecia.)
Many natural resource agencies are cautious about
restoration and reintroduction of rare plants, probably
because this has conventionally been considered in a
mitigation context (Berg 1996). Restrictive generalized
policies on geographic specificity of reintroduction to
documented historic localities, regardless of natural temporal and spatial scales of plant population dynamics and
ecosystem processes, in some cases has narrowed opportunities for re-establishment of rare plants (White 1996).
In situations where the range of rare plants is extremly
reduced, historic collection data are sparse and vague
(which is generally the case), and relatively few potential source populations for founders exist, an experimental approach may be most appropriate for reintroduction
planning. Successful reintroduction will likely require
much replication over years (variable climate conditions)
and at many localities. Caution is appropriate, however,
when the taxonomic interpretation or population variability is at issue when determining suitable populations
for reintroduction.
Artificial Salinity Manipulation – In Suisun
Marsh, salinity control gates on Montezuma Slough were
installed to enforce standards for salinity based on the
perceived needs of waterfowl marsh management in
diked wetlands, aimed at maintaining low channel water salinity. The impacts of sustained low marsh salinity
on the progression of exotic plant invasions and the natural dynamics of brackish tidal marsh vegetation (particularly rare and endangered species) were not considered
in the design and operation of the salinity control gates,
and no long-term monitoring of rare plant populations
during gate operation was authorized. The reduction of
periodic high salinity events during drought cycles, and
Peter Baye
Pioneer plants (1st year
seedling) of native
Spartina foliosa and
Salicornia virginica
colonize the well-consolidated upper mudflats
bayward of the marsh
edge at Mare Island,
eastern San Pablo Bay. The
erosional scour pools and
drainages adjacent to the
plants indicate the
relatively hight wave
energy estuarine environment in which they are
able to establish, given
stable microhabitats.
Chapter 1 —
Plant Communities
21
Plants
ally been designed for wildlife species, treating plants
only as habitat for wildlife species rather than as the
subject of restoration aims. Restoration designs have
generally afforded little or no consideration for soils or
slopes of the high marsh zone, variations in sediment
texture, surface or subsurface freshwater flows, and variation in incident wave energy that influence the microenvironmental variables which are significant for plant diversity. Highly managed estuarine wetlands (e.g.,
artificial salt ponds, extremely microtidal or non-tidal salt
marshes) generally support an artificially low diversity of
native tidal marsh plant species. Plans for rare tidal marsh
plant reintroduction have only recently been proposed
(e.g., Pier 98, Port of San Francisco; Crissy Field,
Presidio/Golden Gate National Recreation Area), and
none has yet been implemented. Of the rare plant refugia in relict tidal marshes of the Estuary (e.g., Hill
Slough, Fagan Marsh, Rush Ranch, Peytonia Slough,
and Whittell Marsh), none has site-specific rare plant
management plans or programs, despite imminent
threats by invasive species. There is no Estuary-wide
program to survey and map rare plant species populations; plant inventories are biased towards species with
special legal status, and are typically driven by environmental impact assessment for projects rather than regional conservation. Other surveys consist of voluntary
and opportunistic reports. Conservation of plant diversity in the Estuary will require both active protection
of remnant rare plant refugia, active management of
conserved areas, systematic inventory of the Estuary’s
botanical resources, and large-scale, scientifically
sound tidal marsh restoration and reintroduction
projects.
sion threatens to encroach into restorable former tidal
marsh sites. Single-purpose management of other diked
wetland types at large scales (salt production, waterfowl
production) also restricts opportunities for tidal marsh
plant community restoration. Large-scale tidal marsh
restoration near centers of relict tidal marsh plant populations (e.g., Cullinan Ranch, Hamilton Airfield, Redwood Landfill, and Skaggs Island) offer some hope for
long-term recovery of tidal marsh plant species in decline.
Conclusions and Recommendations
The San Francisco Estuary tidal marshes are poorly understood in terms of modern and historic plant species
composition, the dynamics of the vegetation, and the
interaction between vegetation and geomorphic and hydrologic processes. Many plant species have become extirpated or nearly so with little or no attention from
botanists or ecologists, and many more species have declined significantly. The Estuary’s historic and modern
flora is considerably richer than has been generally recognized. Further attrition of native plant diversity in the
Estuary is likely because of the uncontrolled spread of
invasive exotic plants, and insufficient planning, management, and restoration of the Estuary’s plant community. Carefully designed tidal marsh restoration
projects that promote native plant species diversity
and recovery are needed to conserve the Estuary’s
flora. Recommendations for the conservation of the
Estuary’s plant communities are presented in the
Baylands Ecosystem Habitat Goals Report (Goals
Project 1999, Appendix A).
Peter Baye
Plants
the subtle changes in tide elevations caused by gate operation, could potentially have significant adverse longterm impacts on rare plant persistence. Scientific investigations of the effects of gate operation on plant
communities and rare plant populations of Suisun tidal
marshes are urgently needed, as recommended by the
Brackish Marsh Subcommittee of the Suisun Ecological Workshop (CWRCB 1999).
In the South Bay, perennial urban wastewater discharges in confined, diked tidal sloughs have caused conversion of salt marsh to brackish marsh (Harvey and Associates 1997). The Alviso and Milpitas area marshes
were the sites of historic rare plant populations (Table
1.3) which could not be re-establish naturally or be reintroduced in marsh vegetation dominated by perennial
pepperweed, bulrushes and tules which are stimulated
by augmented and confined freshwater flows and elevated nutrient concentrations throughout the growing
season.
Loss of Restorable Habitat – Economic pressure
to convert diked Baylands to land uses that are incompatible with potential tidal marsh restoration over large
contiguous tracts (particularly in connection with uplands and alluvial areas) remains high today. Developments in diked Baylands for extensive housing (Redwood
Shores, San Mateo County), golf courses (Black Point,
Marin County), business parks (old Fremont Airport,
Alameda County) have proceeded into the 1990s, and
other large scale land use conversions for dredged material disposal and rehandling (Napa salt crystallizers) have
been considered. The largest tracts of undeveloped diked
Baylands are in San Pablo Bay, where vineyard expan-
An example of marsh progradation — Seedling plants of Salicornia virginica and Spartina foliosa are frequently
comingled without clear zonation, as in these exceptionally firm upper mudflats in eastern San Pablo Bay. (Mare
Island, north of the jetty)
22
Baylands Ecosystem Species and Community Profiles
Table 1.3 Historic Changes in the Distribution and Abundance of Selected Native Vascular Plant
Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Historic References
Contemporary Distribution
[A. asperifolia Trin.]
Jepson 1911: Common in the San
Francisco Bay region in salt marshes
and other wet places: Berkeley… San
Francisco; Martinez.”
Not currently reported from tidal
marsh ecotone in San Francisco Bay
Estuary, but common and widespread in non-tidal moist habitats
[Hickman et al. 1993].
Armeria maritima (Miller) Willd.
ssp. californica (Boiss. ) Pors.
Greene 1894: “ Along sandy beaches
in wet ground…”
[Armeria vulgaris Willd.]
[Statice armeria L.]
Jepson 1911: “ common on the sandy
beaches or fields near the sea… or
about San Francisco Bay.”
Apparently extirpated in San Francisco Bay Estuary; otherwise restricted
to maritime coastal salt marshes,
dunes, bluffs.
Agrostis exarata
Trin.
Aster chilensis Nees.
Cooper 1926: [presumed species of
reconstructed “ willow-composite”
community at salt marsh edges, Palo
Alto vicinity]
Howell 1949: [Marin Co.] Common
and widespread from salt marshes
and coastal swales to low valleys…”
Thomas 1961: “… edges of salt
marshes…”
Aster lentus E. Greene
[A. chilensis Nees. var. lentus Jepson]
[A. chilensis var. sonomensis (E. Greene)
Jepson]
Greene 1894: [A.c. var. lentus] “ Plentiful along tidal streams in the western part of the Suisun Marsh…”
[A. c. var. sonomensis ]“ In open
plains of the Sonoma Valley, in low
subsaline ground.”
Few current reports known from
edges of San Francisco Bay or San
Pablo Bay tidal marshes; local in
Suisun Marsh edges. Presumed rare
from tidal marshes.
Rare; restricted primarily to Suisun
Marsh. Some herbarium collections
known from San Francisco Estuary
prior to 1960 (Berkeley, Alviso, Napa).
Recent status uncertain in San Pablo
Bay area tidal marshes.
Jepson 1911: [A.c. var. lentus]“ very
common and conspicuous in the
Suisun Marshes.” [A. c. var. sonomensis] “ subsaline lands: Petaluma,
Napa”
Munz 1959: [A.c. var. sonomensis]:
Coastal Salt Marsh; saline ground
around San Francisco Bay. Sonoma,
Napa…”
Aster subulatus Michaux
var. ligulatus Shinn.
[Aster exilis Ell.]
[Aster divaricatus Nutt.]
Behr 1888: [A. divaricatus] “ Salt
marshes.”
Greene 1894: “ Borders of Suisun Marshes
and elsewhere on subsaline land”
No current reports known from edges
of San Francisco Bay . Uncommon
to rare in San Pablo Bay and Suisun
tidal marshes.
Jepson 1911: “ Saline soil, not common … .Alvarado.”
Cooper 1926: [presumed species of
reconstructed “ willow-composite”
community at salt marsh edges, Palo
Alto vicinity]
Thomas 1961: “ Salt marshes along
San Francisco Bay and occasionally
elsewhere. San Francisco, Palo Alto,
Alviso…”
Astragalus tener Gray
var. tener
Jepson 1911: “ Alkaline fields, mostly
in moist places.”
Thomas 1961: Known locally only
from saline areas along San Francisco
Bay. San Francisco and Mayfield.”
Recently rediscovered near historic
Bay edge in Fremont, Alameda
County. Known in region from alkali
vernal pools, Solano County.
Chapter 1 —
Plant Communities
23
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Atriplex californica
Historic References
Moq.
Greene 1894: “… along the edges of
salt marshes, from near San Francisco and Alameda, southward.”
Jepson 1911: “ Sandy beaches along
the ocean and about San Francisco
Bay.”
Baccharis douglasii DC.
Jepson 1911: “… abundant in the salt
marshes about San Francisco Bay.”
Thomas 1961: [SW San Francisco Bay]
“… occasionally along the edges of
salt marshes… Alviso…”
Best et al. 1996: “ Uncommon. Damp
thickets, salt marshes.”
Carex aquatilis
var. dives (Holm)
Wahlenb.
[C. sitchensis Prescott]
Brewer et al. 1880: “ In salt marshes,
about San Francisco Bay
(Bolander)…”
Jepson 1911: “ Salt-marshes about
San Francisco Bay and northward
along the coast” .
Contemporary Distribution
Extirpated in San Francisco Bay Estuary margins. Small relict populations
occur on bluffs of Golden Gate in
San Francisco. Maritime salt marsh
populations occur at Limantour
Estero and Tomales Bay (Marin Co.).
Now uncommon to rare in alluvial
high marsh and upland ecotone,
San Pablo Bay area and and Suisun
Marsh; one colony occurs along salt
pond edge at a seep in Coyote Hills,
Alameda Co., possibly rare elsewhere in San Francisco Bay.
No current reports known from edges
of San Francisco Bay or San Pablo
Bay tidal marshes. Presumed rare or
extirpated from tidal marshes.
Munz 1959: “ Rare, swampy places,
usually near the coast…”
Jepson 1911: [C. b. var. densa] “ Salt
marshes near San Francisco…”
No current reports known from edges
of San Francisco Bay or San Pablo
Bay tidal marshes. Presumed rare or
extirpated from tidal marshes.
Thomas 1961: “ Boggy areas along
the edges of salt marshes; San Francisco, Woodside, Mayfield…”
Rare in Suisun area tidal marshes,
west to Southampton Bay. Common
in alkaline, moist places in California
floristic province.
Hook and
Behr 1888: “ Marsh near Tamalpais.”
[Orthocarpus castillejoides Benth.]
Greene 1894: “ Common along the
borders of salt marshes.”
Currently reported only from Point
Pinole salt marsh and pan edges;
other historic records at Greenbrae,
Tamalpais (Mill Valley), Hamilton
Field, Burlingame, Oakland. Halophytic populations rarely occur in
brackish marsh and salt marsh at
Rodeo Lagoon, Tomales Bay,
Drakes Estero, Limantour Estero
(maritime Marin Co. marshes)
Carex densa
Bailey
[C. brogniartii Kunth. var. densa
Bailey]
Carex praegracilis W. Boott
[Carex Douglasii var. brunnea Olney]
[C. usta Bailey]
Castilleja ambigua
Arn.
Jepson 1911: “ Marshy ground near
the coast. Alameda; W. Berkeley;
Napa Valley; Sonoma Co.”
Howell 1949: “ low ground along the
upper reaches of the salt marshes,
occasional:..Mount Tamalpais; Greenbrae Marshes; Hamilton Field…”
Centaurium trichanthum
(Griseb.) Robinson
[Erythrea trichantha (Griseb.)]
Howell 1949: “ in typical form… known
in Marin only from low ground bordering the salt marsh near Burdell
Station” .
Munz 1959: “ Moist often saline
places… edge of Coastal Salt Marsh…
San Mateo Co. to Siskyo Co.”
24
Baylands Ecosystem Species and Community Profiles
No current reports known from edges
of estuarine tidal marshes. Similar
species C. muehlenbergii occurs in
subsaline diked wetlands, NapaSonoma marsh, and tidal marsh
edge at China Camp.
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Historic References
Calystegia sepium (L.) R.Br.
ssp. limnophila (E. Greene)
Brummit
[Convolvulus sepium L.]
Contemporary Distribution
Greene 1894: “ Plentiful in brackish
marshes toward the mouth of the
Napa River and about Suisun Bay; its
roots within reach of tide water; its
stems twining upon rushes and sedges.
Occasional in Suisun Marsh area
west to Southhampton Bay; rare in
San Pablo Bay edges.
Munz 1959: “ Occasional in swampy
saline places; Coastal Salt Marsh;
Marin, Solano and Contra Costa Cos.”
Cicuta maculata L.
var. bolanderi (S. Watson)
Jepson 1911: “ Suisun marshes, abundant and conspicuous.”
Mulligan
Munz 1959: “ Salt marshes, Marin to
Solano and Contra Costa cos.”
[Cicuta bolanderi Watson]
Cirsium hydrophilum (E. Greene)
Jepson var. hydrophilum
[Carduus hydrophilus Greene]
Cordylanthus maritimus Benth.
ssp. palustris (Behr) Chuang and
Heckard
[Cordylanthus maritimus Nutt.]
[Adenostegia maritima (Nutt.)
Greene]
Extremely rare (federally endangered) in Suisun Marsh.
Jepson 1911: “ Suisun marshes”
Munz 1959: “ Brackish marshes about
Suisun Bay” .
Brewer et al. 1880: “ Sandy saltmarshes along the coast, from San
Francisco Bay to San Diego.”
Behr 1888: “ Salt marshes, San Francisco.”
Greene 1894: “ Sandy salt marshes
from near San Francisco southward.”
Jepson 1911: ‘Salt marshes near the
coast from San Francisco Bay south…”
Howell 1949: “ Salicornia flats in salt
marshes along the bay..:.Almonte,
Greenbrae…”
Uncommon to rare in Suisun Marsh;
not currently reported elsewhere in
the Estuary.
Currently reported only from Richardson Bay, Greenbrae, and Petaluma marsh (Marin Co.). Recently
reported from Gallinas Creek area
marsh. Extirpated in central and
southern San Francisco Bay. Major
populations occur in maritime tidal
salt marshes of Tomales Bay, Bolinas
Lagoon, and Limantour Estero (Marin
Co.). San Francisco Estuary populations have purplish foliage, and rosy,
well-exerted inflated flowers.
Thomas 1961: “ Salt marshes along
the borders of San Francisco Bay;
San Francisco, Redwood City, Palo
Alto, and near Alviso.”
Cordylanthus mollis Gray
ssp. mollis
Brewer et al. 1880. “ Salt-marshes of
San Francisco Bay, at Mare Island
and Vallejo, C. Wright, E.L. Greene.”
Behr 1888: “ Salt marshes. Vallejo.”
Greene 1894: “ Brackish marshes
about Vallejo and Suisun.”
Howell 1949: [Marin Co.] “… San
Rafael, acc. Ferris; Burdell Station,
San Antonio Creek… .”
Rare (federally endangered): local
in tidal brackish marsh around Napa
River, Carquinez Straits tidal marsh,
Suisun Marsh area. Presumed extirpated in Petaluma River marshes.
Putative San Francisco (city) record
is erroneous interpretation of early
San Francisco Bay Area collection
acc. L. Heckard.
Best et al. 1996: [Sonoma Co.]: Rare,
estuarine… Petaluma Marsh between San Antonio and Mudhen
Slough… (1978)…”
Downingia pulchella
(Lindley)
Torrey
[Bolelia pulchella E. Greene]
Jepson 1901: “… Abundant and of
rank growth in salt marshes near
Alvarado [Union City]” .
Munz 1959: “… Coastal Salt Marsh.”
Extirpated in Union City. Occurs in
alkaline/saline vernal pools at Warm
Springs, Fremont, Alameda Co., and
in some diked baylands near Fairfield,
Solano Co.
Chapter 1 —
Plant Communities
25
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Historic References
Eleocharis parvula (Roemer
and Shultes) Link
Festuca rubra L.
Glaux maritima L.
Contemporary Distribution
[not reported in early floras; Munz
1959 reported only from coastal salt
marshes of San Luis Obispo and
Humboldt Cos.]
Rare in brackish tidal marshes of San
Pablo and Suisun Bay area. Local in
diked baylands, lower Napa River.
[reported only from generalized
habitats in early floras; halophytic
populations not distinguished. Speculative likely component of historic
sandy salt marsh edges of Central
Bay.]
Not currently reported from San Francisco Bay estuarine tidal marsh edges;
halophytic populations presumed
extirpated. Halophytic populations
occur along edges of maritime salt
marsh and brackish marsh at Rodeo
Lagoon, Limantour Estero, Tomales
Bay (Marin Co).]
Behr 1888: “ Salt marshes.”
Few recent reports known from San
Francisco Bay or San Pablo Bay salt
marshes; reported as infrequent in
Petaluma Marsh; local in tidal marsh
near mouth of Tolay Creek, Sonoma
Co.; occasional to locally frequent in
Suisun Marsh area and Fagan Slough
(Napa River).
Greene 1894: “ Frequent both along
the seabord and in subsaline soils in
the interior”
Jepson 1911: “ Marshy shores of …
San Francisco and Suisun bays.”
Howell 1949: [Marin Co.] “ salt
marshes… Burdell…”
Thomas 1961: “… Palo Alto, but expected elsewhere in salt marshes”
Atwater et al. 1979: [recorded as
present in San Pablo Bay]
Best et al. 1996: [Sonoma Co.] “ Rare,
salt marshes: Petaluma, Davy (1893
UC).”
Howell et al. 1958: “ salt marsh near
Visitacion Valley [southeastern San
Francisco].”
No current reports known from San
Francisco Bay. Recently reported
from Suisun Marsh area.
Hemizonia pungens
(Hook
and Arn.) Torrey and A. Gray
Greene 1894: “ Borders of salt
marshes about San Francisco Bay.”
ssp. maritima (E. Greene)
Cooper 1926: [dominant species of
reconstructed “ willow-composite”
community at salt marsh edges, Palo
Alto vicinity.]
Local, infrequent species along tidal
marsh edge around the San Francisco Estuary.
Heliotropium curassavicum
[Centromadia maritima Greene]
L.
(Robinson and Greenman) Keck
Munz 1959: [ ssp. congdonii] “ Locally
common… s. end of San Francisco
Bay, mostly Alameda Co.” [ssp.
parryi] “ Coastal Salt Marsh… to N. San
Mateo Co…” [not reported from salt
marsh in Jepson 1901, Greene 1894)
No current reports known from San
Francisco Bay Estuary tidal marsh
edges. Rare.
Hutchinsia procumbens (L.)
Greene 1894: “ Borders of salt
marshes.”
No current reports known from San
Francisco Bay Estuary tidal marsh
edges. Occurs in high marsh ecotone of central CA coast salt marsh,
and in other alkaline or subsaline
habitats in California floristic province.
Hemizonia parryi E. Greene
ssp. parryi, ssp. congdonii
Desv.
[Bursa divaricata (O. Ktze) Nutt.]
[Capsella divaricata Walp.]
[Capsella procumbens Fries.]
[Capsella elliptica C.A. Mey.]
[Lepidium procumbens L.]
[Hutchinsia californica,
H. desertorum A. Davids]
26
Jepson 1911: Alkaline soil from
Vallejo (acc. Bot. Cal.), Alameda…”
Thomas 1961: Known locally from
saline areas along San Francisco
Bay; Palo Alto and Mayfield.”
Baylands Ecosystem Species and Community Profiles
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Historic References
Contemporary Distribution
Brewer et al. 1880: “… common in the
salt-marshes about San Francisco
Bay…”
Juncus effusus L. var.
brunneus Engelm.
Brandegee 1892: “ Salt marshes about
the bay shore.”
Rare, local at edges of salt marsh
and brackish marsh ecotones in San
Pablo Bay (China Camp). No known
reports from San Francisco Bay tidal
marshes.
Jepson 1911: “ Common in marshy
ground: Monterey to San Francisco
and Bolinas Bays and northward.”
Howell 1949: “ Swamps and swales
generally near the ocean… Tiburon;
Sausalito…”
Thomas 1961: “ Usually along or near
the coast… Palo Alto, near Alviso…”
Brandegee 1892: “ Salt marshes at
Visitacion Bay. South San Francisco.”
Juncus lesueurii Boland.
Howell 1949. “ Common along the
upper reaches of salt marshes…
Tiburon; Tamalpais Valley… . In Marin
County,… [J. balticus] is not readily
distinguished from J. Leseurii… ]”
Juncus xiphioides E. Meyer
Jepson 1901: “ A common species of
salt marshes… Berkeley; Belmont…
Suisun Marshes…”
Thomas 1961: “ Occasional in sloughs
… Palo Alto, nr. Alviso…”
Lasthenia glaberrima
D.C.
Lasthenia conjugens E. Greene
[Baeria fremontii (Torr.) A. Gray in
part]
No other reports, historic or current,
are known from San Francisco Bay
estuarine marshes.
Greene 1894: “ Subsaline soil near
Antioch…”
Occurs in alkaline/saline vernal pools
bordering salt pond 22 in Fremont,
Alameda Co., and in diked baylands at upper end of Hill Slough
(Potrero Hills), Solano Co. Historic
localities near Mt. Eden along bay
shore and near Newark. Rare; federally endangered.
Jepson 1911: “ Subsaline fields in the
Bay region; Antioch; Newark, etc.”
Greene 1894: “ Border of salt marsh
north of Vallejo: rare or local.”
[Baeria carnosa E. Greene]
[B. platycarpha A. Gray]
Jepson 1911: “ Salt marshes at Vallejo
(Greene).”
Lasthenia glabrata Lindley ssp.
glabrata
Behr 1888: “ Common.”
(A.
Not recently reported; presumed
rare or possibly extirpated in most
tidal salt marshes of San Francisco
Estuary.
Behr 1888: “ Near salt marshes.”
Gray) E. Greene
Lasthenia platycarpha
Apparently associated with sandy
salt marsh edges of maritime coast.
Intermediates with J. balticus not
uncommon in San Francisco Bay
Area tidal marshes; difficult to separate. Rare in south San Francisco Bay
tidal marshes; one colony in seep at
salt pond edge, Coyote Hills.
Greene 1894: “ Borders of salt marshes
only; not common.”
Jepson 1911: “ Borders of salt marshes.”
Thomas 1961: Edges of salt marshes
along San Francisco Bay… Millbrae…
Belmont, Redwood City, Mayfield.”
Historic locality at Redwood City
shoreline. Apparently extirpated
from San Francisco Bay estuarine
marshes. Occurs infrequently in
alkaline vernal pools, Solano Co.
Currently reported within San Francisco Bay Estuary only from Point
Pinole (Whittell marsh) salt marsh
and new seeded levee slope at
Sonoma Baylands. Many historic salt
marsh collections known from Burdell, Alvarado, Mt. Eden, Alameda,
Mowry’s Landing, Denverton. Maritime salt marsh population occurs in
Limantour Estero, Marin Co.
Chapter 1 —
Plant Communities
27
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Historic References
Contemporary Distribution
Brandegee 1891: “ About the borders
of marshes, Islais Creek, Visitacion
Valley, Presidio, South San Francisco.”
Apparently extirpated from San
Francisco Bay estuarine marshes.
Lathyrus jepsonii E. Greene var.
jepsonii Jepson
Greene 1894: “ Suisun marshes.”
Occasional to rare in Suisun Marsh.
Also occurs locally in tidal brackish
marshes along Napa River (Dutchman Slough). May be under-reported
in drought years.
Layia chrysanthemoides (DC.)
Howell 1949: “ Locally common on
flats bordering the salt marshes:
Ignacio;… Chileno Valley.”
Lasthenia minor (DC.) Ornd.
[Baeria minor (DC.) Ferris
[Baeria uliginosa Nutt.]
[Lasthenia uliginosa (Nutt.) E. Greene]
A. Gray
[Blepharipappus chrysanthemoides
Greene]
Jepson 1911: “ Suisun marshes.”
No current reports known from San
Francisco Estuary tidal marsh edges.
Thomas 1961: “ occasionally in low
alkaline soils of San FranciscoBay…
Millbrae, Redwood City.”
[general grassland habitats reported
historically. Presumed abundant or
dominant species of historic tidal
marsh edges.]
Occurs locally (abundant) at salt
marsh edges at Newark, Alameda
Co.; Rush Ranch, Solano Co; Petaluma Marsh, Marin Co.; China Camp,
Marin Co.; Dutchman Slough, Solano
Co.
Lepidium dictyotum A. Gray
Greene 1894: “ Along the borders of
marshes at Alameda.”
No current reports known from San
Francisco Bay tidal marsh edges.
Presumed extirpated or rare in Estuary.
Lepidium latipes
Greene 1894: “ in saline soil at
Martinez, Alameda, etc.” Jepson
1901: “… alkali flats… Martinez…”
No current reports known from San
Francisco Bay tidal marsh edges.
Reported rarely in diked baylands
and tidal marsh edges, Solano Co.
(Suisun Marsh area).
Greene 1894: “ Borders of salt marshes
at Vallejo… also in subsaline soils…
near Alameda.”
No current reports known from San
Francisco Bay tidal marsh edges.
Rare, Suisun Marsh edges.
Leymus triticoides (Buckley)
Pilger
[incl. Leymus X multiflorus (Gould)
Barkworth and D.R. Dewey
[Elymus triticoides Buckley]
Hook.
Lepidium oxycarpum Torrey
and A. Gray
Howell 1949: “ A rare peppercress of
alkaline valley floors and of saline flats
adjacent to coastal salt marshes, in
Marin Co. known only from low pastures bordering San Francisco Bay
near Novato.”
Munz 1959: “ V. Grassland and edge
of Coastal Salt Marsh; largely about
San Francisco Bay…”
Thomas 1961: “ Saline and alkaline
flats along San Francisco Bay and
Santa Clara Valley: Redwood City,
Cooley’s Landing, Palo Alto, Mayfield…”
Lilaeopsis masonii Mathias and
Constance
[Lilaeopsis lineata (Michx.) Greene,
in part
28
Jepson 1911: [ as L. lineata, in part]
“ Salt marshes or brackish mud flats:…
Port Costa to Antioch; Robert’s Island” .
Baylands Ecosystem Species and Community Profiles
Rare in tidal brackish tidal marshes,
Napa Marsh, Suisun Marsh area, to
Tolay Creek, San Pablo Bay. Uncommon in western Sacramento river
delta fresh-brackish marshes.
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Lycopus asper
E. Greene
[Lycopus lucidus Turcz. ]
[L. lucidus Benth. misapplied]
Historic References
Contemporary Distribution
Jepson 1911: “ Salt marshes: Suisun;
Benicia; San Francisco.”
No current reports known from San
Francisco tidal marsh edges; presumed rare or extirpated in Estuary
there.
Hickman et al. 1993: Uncommon.
Moist areas, marsehs, streambanks…
Deltaic GV, SnFrB, GB; to w Can,
Great Plains.
I.M. Johnston
Jepson 1911: “ Alvarado [Union City],
margin of salt marshes.”
[Allocarya salina Jepson]
[Allocarya glabra Macbr.]
Munz 1959: “ Coastal Salt Marsh; s.
shore of San Francisco Bay…”
Plagiobothrys mollis (A. Gray)
I.M. Johnston var. vestitus
Jepson 1911: Petaluma, Congdon,
1880; not since collected.
Hickman 1993: “ PRESUMED EXTINCT.
Wet sites in grassland, possibly
coastal marsh margins…”
Best et al. 1996: “ salt marsh near
Sears Point, Keck 1935”
Reported at Sonoma Baylands, high
tide line, 1996. Otherwise no current
reports known from San Francisco
Estuary tidal marsh edges
Brewer et al. 1880: “ Salt-marshes, San
Pablo Bay, at Benicia and Vallejo,
Bigelow, E.L. Greene.”
Rarely reported from San Francisco
Bay Area high tidal marshes (Suisun
Marsh). No recent localities in tidal
marsh edges confirmed.
Plagiobothrys glaber (A. Gray)
(E. Greene) I.M. Johnston
Hickman 1993: “ PRESUMED EXTINCT.
Wet, alkaline soils in valleys, coastal
marshes… CCo, s SnFrB… Perhaps a
var. of P. stipitatus.”
[Allocarya mollis A. Gray var. vestita
E. Greene
[A. vestita E. Greene]
Plagiobothrys stipitatus
Greene) var. stipitatus
(E.
Plantago elongata
Pursh
[Plantago bigelovii Gray]
Greene 1894: “ Borders of saline or
brackish marshes; quite common
about the Bay…”
Howell 1949: [Marin Co.; not reported
from estuarine stations]
Thomas 1961: [SW San Francisco Bay]
“… edges of salt marshes… Mayfield,
Alviso… ]
Best et al. 1996: [Sonoma Co.] “ uncommon, salt marshes… Petaluma,
Congdon (1880); 5 mi. n. of Sear’s
Point, Rubzoff (1970).
Plantago maritima L.
Greene 1894: “… sandy salt marshes”
[P. maritima L. ssp. juncoides (Lamk.)
Hulten, P. juncoides Lamk.
var. juncoides]
Jepson 1911: “… West Berkeley…”
Howell 1949: [Marin Co.] “ occasional
in salt marshes bordering the bay or
ocean: Almonte…”
Infrequent to rare in San Francisco
Bay tidal marshes, mostly Richardson
Bay. Relatively common in maritime
salt marshes, and occaisional in
Suisun Marsh (Hill Slough).
Thomas 1961: “ occasional in salt
marshes and on coastal bluffs as far
south as San Mateo County: San
Francisco, Redwood City, and
Mayfield.”
Best et al. 1996: [Not reported from
estuarine Sonoma Co. stations.]
Chapter 1 —
Plant Communities
29
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Pluchea odorata
Historic References
(L.) Cass.
[P. camphorata (L.) DC. and
P. purpurascens (Sw.) DC. misapplied ]
Behr 1888: Salt marshes.
Greene 1894: “ Borders of brackish
marshes about Suisun Bay, etc.”
Contemporary Distribution
No current reports known from San
Francisco or San Pablo Bays; uncommon in Suisun marshes.
Jepson 1911: “ Common in the salt
marshes about Suisun and San Francisco Bays…”
Howell 1949: [Marin Co.; not cited]
Thomas 1961: [SW San Francisco
Bay; not cited]
Puccinelia nutkaensis
(J.S.
Presl.) Fern. and Weath.
Thomas 1961: “ levees and salt marsh
along San Francisco Bay” .
Rare, local, south San Francisco Bay.
No records of San Pablo Bay collections in Howell 1949, Best et al. 1996.
Greene 1894: “ A somewhat rare
plant of subsaline soils at Calistoga
and near San Jose.”
No current reports known from San
Francisco Bay high tidal marsh. Presumed extirpated in San Francisco
Bay.
[P. grandis Swallen]
Pyrrocoma racemosa (Nutt.)
Torrey and A. Gray var. racemosa
[Haplopappus racemosa (Nutt.)
Torr.; P. elata E. Greene]
Rumex occidentalis
S. Watson
[R. fenestratus E. Greene]
Thomas 1961: “ edges of salt marshes,
saline soils, and occasionaly disturbed areas. Cooley’s Landing,
Near Alviso, Agnews, and San Jose.”
Greene 1894: “ Frequent in marshy
places.”
Infrequent to rare in North Bay, Suisun
Marsh area brackish tidal marshes.
Jepson 1911: “ Marshes bordering
San Francisco Bay.”
Munz 1959: “ Coastal, often brackish
marshes, San Francisco Bay…”
Hickman et al. 1993: “ Uncommon.
wet +/- salty places.”
Salicornia subterminalis
Parish
[Arthrocnemum subterminale (Parish) Standley]
Sanicula maritima Wats.
(S.
maritima Kellogg)
[Not reported from estuarine stations
in early floras.]
Local, rare in South Bay, south Fremont, Milpitas (in diked wetlands,
former Fremont Airport), and at Hill
Slough, Suisun Marsh.
Behr 1888: “ Alameda marshes.”
Extinct in San Francisco Bay; known
from fewer than 10 stations in 1988,
Monterey and San Luis Obispo
Counties.
Greene 1894: “ In lowlands adjacent
to salt marshes near Alameda, San
Francisco, etc.”
Jepson 1911: “ Local species of low
and wet adobe lands in the vicinity
of salt marshes bordering San Francisco Bay; near Alameda… and
Potrero Hills, San Francisco, the only
recorded localities.”
Senecio hydrophilus Nutt.
Brewer et al. 1880: “… salt marsh at
Vallejo (Greene)…”
Greene 1894: “ Brackish marshes;
formerly plentiful at West Berkeley,
and on the lower Napa River; still
abundant in the Suisun marshes.”
Jepson 1911: Abundant in the Suisun
Marshes and found in other marshes
about San Francisco Bay”
30
Baylands Ecosystem Species and Community Profiles
Apparently extirpated in San Francisco, San Pablo Bay (incl. Petaluma R.); infrequent but locally
common in Suisun Marsh area and
Carquinez Strait tidal marshes; possibly Napa R..”
Hickman 1993: “ Reduced from wetland development.”
Table 1.3 (continued) Historic Changes in the Distribution and Abundance of Selected Native Vascular
Plant Species Occurring in Tidal Marshes of the San Francisco Estuary
Taxon
Sium suave Walter
[Sium cicutaefolium Gmel.
var. heterophyllum Jepson]
Suaeda californica Wats.
Historic References
Contemporary Distribution
Jepson 1911: “ Suisun marshes; Stockton”
Rare in Suisun Marsh, primarily in wet
years. Recently observed near Rush
Ranch, Hill Slough, and Brown’s Island.
Brewer et al. 1880: “ In salt-marshes
on the coast, about San Francisco.”
Extinct in San Francisco Bay Estuary,
probably since ca. 1950. Restricted
to one large population at Morro
Bay, where it occurs primarily along
sandy high marsh edges. Planned for
reintroduction. Misreported occurrences often due to confusion with
Suaeda calceoliformis, S. moquinii,
Salsola soda, and Bassia hyssopifolia.
Behr 1888: “ Salt marshes on an island
near Alameda.”
Greene 1894: “ Vicinity of sand
beaches about San Francisco Bay,
but seldom seen.”
Jepson 1911: “ Sandy beaches bordering San Francisco Bay, the known
stations few: San Pablo Landing; Bay
Farm Island.”
Thomas 1961: “ Occasional in salt
marshes along San Francisco Bay;
San Francisco and Palo Alto.”
Solidago confinis Nutt.
[S. sempervirens L. misapplied]
[S. confinis var. luxurians Jepson]
Jepson 1911: “ Salt marshes, San
Francisco Bay, Bolander [1863].
Rarely collected.
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Greene, E.L. 1894. Manual of the botany of the region
of San Francisco Bay. Cubery and Company, San
Francisco.
Grewell, B. 1993 (et seq.). Unpublished vascular plant
species list, Suisun Marsh.
[References continued on page 29]
Grewell, B., D. Hickson and P. Baye. 1999. SEW (Suisun Ecological Workgroup) Brackish marsh vegetation subcommittee report. Report to the California State Water Resources Control Board. Sacramento, Calif.
Harvey and Associates. 1997. Marsh plant associations
of south San Francisco Bay: 1997 comparative
study. Unpublished report to the City of San Jose.
H.T. Harvey and Associates, Alviso, Calif.
Hinde, H. 1954 . The vertical distribution of salt marsh
phanerogams inrelation to tide levels. Ecol.
Monogr. 24: 209-225.
Howell, J.T. 1949. Marin Flora. Univ. of Calif. Press.
319 pp.
______. 1969. Marin Flora Supplement. Univ. of Calif. Press.
32
Baylands Ecosystem Species and Community Profiles
Howell, J.T., P.H. Raven and P.R. Rubtzoff. 1958. A
flora of San Francisco, California. Wasmann J. of
Biol. 16: 1-157.
Ingram, B.L., J.C. Ingle and M.E. Conrad. 1996. A 2000
year record of Sacramento-San Joaquin river inflow to San Francisco Bay Estuary, California.
Geology 24: 331-334.
Jepson, W.L. 1911. A Flora of middle western California. Cunningham, Curtiss and Welch, San Francisco. (Revised edition of 1901). 515 pp.
______. 1925. Manual of the flowering plants of California. Univ. of Calif. Press. 1238 pp.
Josselyn, M. 1983. The ecology of San Francisco Bay
tidal marshes: A Community Profile. US Fish
Wildl. Serv. OBS-83/23, Oct. 1983. 102 pp.
Macdonald K.B. 1977. Coastal salt marsh. In: M.G.
Barbour and J. Major (eds). 1988. Terrestrial vegetation of California. Calif. Native Plant Society
Publ. No. 9, 1030 pp.
______. 1988. Supplement: coastal salt marsh. In: M.G.
Barbour and J. Major (eds). 1988. Terrestrial vegetation of California. Calif. Native Plant Society
Publ. No. 9, 1030 pp.
Munz, P.A. 1959. A California flora. Univ. of Calif. Press.
1681 pp.
______. 1968. Supplement to a California flora. Univ.
of Calif. Press. 224 pp.
Newcombe, C.L. and H. Mason. 1972. An environmental
inventory of the North Bay-Stockton ship channel
area. San Francisco Bay Marine Research Center. 2 vols.
Pearcy R. W. and S. L. Ustin. 1984. Effects of salinity
on growth and photosynthesis of three California
tidal marsh species. Oecologia 62: 68-73.
Peinado, M., F. Alcaraz, J. Delgadillo, M. De La Cruz,
J. Alvarez and J.L. Aguirre. 1994. The coastal salt
marshes of California and Baja California: phytosociological typology and zonation. Vegetatio 110: 5566.
Pestrong, R. 1965. The development of drainage patterns of tidal marshes. Stanford Univ. Publications
in Geolog. Sciences 10: 1-87.
Pethick, J.S. 1974. The distribution of salt pans on tidal
salt marshes. J. Biogeog. 1: 57-62.
Peterson, D.H., D.R. Cayan, J.F. Festa, F.H. Nichols,
R. A. Walters, J.V. Slack, S.E. Hager and L.E.
Shemel. 1989. Climate variability in an estuary:
effects of riverflow on San Francisco Bay. In: D.H.
Peterson (ed). Aspects of climate variability in the
Pacific and the western Americas. Geophys. Union.
Geophys. Monogr. 55: 41-442.
Rugyt, J. 1994. Ecological studies and demographic
monitoring of soft bird’s-beak (Cordylanthus mollis
ssp. mollis), a California listed rare plant species
(and habitat recommendations). Report to Calif.
Dept. Fish and Game, Natural Heritage Div., Sacramento, Calif. 173 pp.
Plants and Environments of
Diked Baylands
Plants
Skinner, M.W. and B.M. Pavlick. 1994. California Native Plant Society’s inventory of rare and endangered vascular plants of California, 5th edition.
CNPS Publications, Sacramento, Calif. 338 pp.
Thomas, J.H. 1961. Flora of the Santa Cruz Mountains of California. Stanford Univ. Press. 434 pp.
Wells, L.E. and M. Goman. 1994. Late Holocene environmental variability in the upper San Francisco
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11th Ann. Pacific Climate Workshop, Technical
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White, P. S. 1996. Spatial and biological scales in reintroduction. pp. 49-86 In: D. Falk, C. I. Millar,
and M. Olwell (eds). 1996. Restoring diversity:
strategies for reintroduction of endangered plants.
Island Press. 505 pp.
Peter R. Baye
Introduction
This report focuses on wetland areas within historic tidal
marshes that have been isolated from tidal action by dikes
(levees) and converted to non-tidal salt marsh, non-tidal
brackish marsh, or subsaline to freshwater seasonal wetlands. These areas are referred to herein interchangebly
as “ diked wetlands” or “ diked Baylands.” Because instantaneous salinity (or even average annual salinity) of diked
wetland soils does not consistently correspond with plant
community composition, and varies over time, these salinity categories are intended to be broadly descriptive
of plant associations rather than quantitative threshold
values of soil salinity. Accordingly, the marsh types described are not discrete, but intergrade continuously and
may vary over time at any site. Diked wetlands as treated
below do not include artificial salt ponds (treated separately) or “ muted tidal” managed marshes (marshes with
reduced tidal range controlled by tidegates), and cover
only wetlands with non-tidal hydrologic inputs (rainfall,
groundwater, surface runoff, streamflow, engineered
water control structures, or very infrequent overtopping
of dikes by extreme tides).
Published and unpublished sources of useful, precise data and other information on the vegetation and
flora of diked Baylands are very scarce. Most usually are
limited to short-term observations and coarse descriptive accounts (such as lists of dominant species) at a particular time of year, or generalized accounts of resource
management plans (e.g., Eicher 1988, Hudson 1980).
Vegetation was usually described for wildlife habitat
evaluation, rather than for floristic analysis or quantative
plant community description. Relatively more detailed
information about some individual diked Bayland sites
is sometimes available for sites which are proposed for
major development projects, and become the subject of
detailed wetland delineations and field studies for environmental evaluations (e.g., Rugyt 1991, Kaufman and
Harvey 1987). The level of detail in vegetation analysis
of diked Baylands even for site-specific studies was still
low until the mid 1980s when technical vegetation criteria for wetland jurisdictional delineations were promulgated (WES 1997). There are no long-term studies of
changes in vegetation in diked Baylands. Some coarse
information about vegetation change in diked wetlands
is available through inspection of historic aerial photographs, particularly color infrared photos from the 1980s
to the present. Based on recent information from some
of the more intensively surveyed diked wetlands (e.g.,
Montezuma Wetlands, Solano County; Cullinan Ranch,
Chapter 1 —
Plant Communities
33
Environmental Setting
The physical origins of diked wetlands are similar
throughout the San Francisco Estuary. Most of the tidal
marshes were reclaimed for agricultural use in the late
19th century when the use of mechanical dredges became commercially available to landowners (after ca.
1870), although many dikes were constructed manually
(Madrone Associates 1977). Tidal marshes were diked
for reclamation either as pasture, hayfields, salt ponds,
or (rarely) cropland. Reclamation involved construction
of dikes (earthen levees made of locally excavated Bay
mud) along the margins of marsh plains (middle marsh
between approximately MWH and MHHW) where they
bordered mudflats or major tidal creeks. The borrow
ditches for dike construction were typically located inside of the dike, creating narrow canals about 20 ft from
the foot of the dikes. Enclosure of tidal marshes by
dikes, and resulting fluctuation between winter flooding and summer desiccation of saline basins, would
have rapidly killed most standing tidal marsh vegetation. When levees were stabilized after several lifts (sequential layers of dewatered dredged Bay muds) tidegates were installed to enable the enclosed basins to
drain on low tides. After stabilization, dikes typically
stood about 3 (to 4) ft above the marsh plain (VerPlanck 1958).
Agricultural areas
within the diked historic
Baylands can pond water
and exhibit seasonal
wetland plant associations.
(North San Pablo Bay diked
Baylands after a storm
event)
Ruth Pratt
Plants
Solano County; Renaissance Faire site, Marin County),
it appears that the diversity and dynamics of diked wetland vegetation have been substantially underestimated
in past assessments.
Historic information on the diking of San Francisco Bay tidal marshes is based on numerous sources,
particularly U.S. Coast Survey maps (multiple series);
historic accounts of salt pond levee development (Ver
Planck 1958); and field observations of modern levee
maintenance and repair methods and agricultural drain
systems.
34
Baylands Ecosystem Species and Community Profiles
Environmental Changes From Diking – Following the initial phase of dike construction, several changes
occurred. Mature tidal marsh soils accumulated peaty organic matter under anaerobic conditions, which minimizes decomposition. Drained marsh soils high in peaty
organic material underwent aerobic decomposition and
dewatering, causing land elevations to subside. Dikes also
caused compression of underlying plastic clayey silts and
peats, and subsided (Madrone Associates et al. 1983).
Differential subsidence of the marsh surface tended to
exaggerate relict marsh topographic relief, causing natural levees (containing coarser silts) to stand out against
isolated depressions where peat content was relatively
great, and the effects of aerobic peat decomposition were
greatest. Tidal creek topography typically persisted as depressional sinuous swales. Early-succession diked marsh
plant communities, typically dominated by perennial
pickleweed (Salicornia virginica; drier, more saline conditions) or alkali bulrush (Scirpus maritimus), bulrush
(Scirpus californicus; less often S. acutus) or cattails
(mostly Typha angustifolia; wetter brackish to subsaline
ditches) tend to be best developed in relict swales and
depressions. As salts were drained from the diked basins
and lands were managed for agriculture, these pioneer
diked salt marsh communities were reduced or eliminated (Madrone Associates et al. 1983, Harvey 1987).
Marsh Progradation and “Second Generation”
Diked Wetlands – The strong reduction in tidal flows
caused by diking all but the largest tidal creeks in the
marsh system caused significant increases in sedimentation outside of diked marshes, causing rapid marsh
progradation on sloping mudflats. In addition, slow migration of the pulse of hydraulic mining outwash from
the Sacramento River contributed to marsh progradation
in San Pablo and Suisun Bays (Doane 1999, Jaffe et al.
1998). In some areas (e.g., south of Novato Creek),
marsh progradation was so extensive that a second phase
of diking occurred in the newly accreted marshes. These
progradational marshes are typically broad pickleweed-
brush (Baccharis pilularis; South Bay) or mixed coyote
brush and bee-plant (Scrophularia californica; North
Bay), often with sub-dominant mustard, poison-hemlock, and radish. High marsh halophytes (pickleweed,
alkali-heath, gumplant, spearscale) tend to dominate the
lower portion of the outboard dike slopes adjacent to salt
marshes, although weedy species can persist for many
years after a levee has been disturbed by maintenance and
repair activities.
The ecological significance of the dike disturbance
cycle for wetland plants is that it has provided corridors
through tidal marshes and diked marshes for a weedy
flora (both exotic halophytes and glycophytes) to disperse, and places weed seed sources along a topographically superior location for dispersal into adjacent diked
and tidal wetlands. The rapid local spread of weedy halophytes on dredge spoils along recently maintained/repaired dikes (especially Salsola soda, Lepidium latifolium,
Mesembryanthemum nodiflorum) can be observed
throughout the Bay. Similar halophyte weed dispersal
occurs along side-cast spoils in diked marshes where
drainage ditches are created or maintained. Dike disturbance corridors may accelerate the spread of exotic halophyte population outposts into uninvaded wetland habitats. In particular, Lepidium latifolium’s invasion of
brackish marshes appears to have tracked patterns of dike
disturbance, invading first along dredge spoil at levee
edges, subsequently spreading into diked and tidal wetlands.
Hydrologic Changes in Diked Wetlands – Patterns of soil waterlogging and inundation in diked conditions differ fundamentally from tidal marsh. They depend principally on the efficiency of artificial drainage,
the permeability of substrate (related to soil clay content),
and the amount and seasonal distribution of rainfall. The
efficiency of the early drainage systems in diked marshes
was based on the amount of ditching and the pattern of
ditching in relation to subsided marsh topography. Because of the great extent of the areas diked, density of
drainage ditches was relatively low. Ditches were mostly
confined to the borders of farmed parcels, but sometimes
Josh Collins
Plants
dominated plains with fringes of cordgrass, cordgrass/
alkali bulrush mixtures, or erosional scarps in pickleweed
peats. Like early-succession diked salt marsh, they support relatively low salt marsh species diversity, and low
densities of narrow, sparsely branched shore-perpendicular tidal creeks. Because of the influence of wave deposition of sediment and coarse organic debris, the tidal
elevations of these dike-fringing salt marshes is often
above MHHW in some areas, particularly where incipient natural levees form at the edge of mudflats. These
secondary prograded high marshes with little antecedent topography were readily converted to diked agricultural land, as in the Baylands of Novato (Hamilton, Bel
Marin Keys).
Dike Disturbance Cycle and Vegetation – Subsidence of dikes themselves caused a need to maintain
dike crest elevations by dredging borrow ditches to resupply material. This established a periodic disturbance
regime to dike vegetation and adjacent ditches. In areas
of high wave energy (long fetch distance, narrow mudflats), maintenance by topping dikes with dredged muds
and repairing erosional slopes may occur in cycles as
short as five years or less. Many bayfront dikes unsheltered by fringing marsh require armoring by placement
of rock or concrete fragments. Well-protected dikes behind extensive salt marsh on firmer peats may have maintenance cycles longer than a decade or two. Repaired dike
slopes provide bare mineral substrate which is gradually
leached of salts and open to colonization by upland
weeds.
The dike disturbance cycle has favored a ruderal
flora along the upper slopes and crests of dikes (including many native and exotic halophytic weeds as well as
glycophytes; e.g., mustard (Hirschfeldia incana, Brassica
spp.), radish (Raphanus sativus), fennel (Foeniculum
vulgare), plantain (Plantago coronopus, P. major, P.
lanceolata), annual ice-plant (Mesembryanthemum nodiflorum; mostly South Bay), sea-fig (Carpobrotus chilense),
hottentot fig (Carpobrotus edulis and hybrids with C.
chilense), poison hemlock (Conium maculatum), Mediterranean brome species (Bromus spp.), wild barley (Hordeum murinum ssp. gussonianum,), ryegrass (Lolium
multiflorum, L. perenne). Lower portions of disturbed
outboard (bayward) dike slopes are typically more saline
and wetter, and support brackish marsh or salt marsh
species, often with an exaggerated proportion of weedy
halophytes (e.g., spearscale, Atriplex triangularis; perennial peppercress, Lepidium latifolium, sicklegrasses
Parapholis incurva, Hainardia cylindrica; bassia, Bassia
hyssopifolia; saltwort, Salsola soda; wild beet, Beta vulgaris). Interior slopes of dikes which face salt ponds, and
contiguous fringing nontidal saltmarsh, are either bare
or vegetated with saltgrass (Distichlis spicata), Salicornia
virginica, dodder (Cuscuta salina), and alkali-heath (Frankenia salina). Dikes with very infrequent maintenance
tend to become dominated by dense stands of coyote
Diked wetlands in Suisun are managed primarily
for waterfowl production.
Chapter 1 —
Plant Communities
35
Plants
reached across extensive marsh depressions. The drains
were originally driven by gravity, drawing drainage water downslope to one-way flapgates which discharged to
adjacent tidal marshes at low tide. This original gravitydriven drainage system had limited efficiency. Topographic lows in the diked basins (swales of relict tidal
creeks, relict marsh pans) remained poorly drained into
the crop growing season, while relict creek levees and
higher relict tidal marsh became better drained. Relictual
tidal marsh patterns of wetland and upland are evident
in black and white photographs of diked hayfields in the
mid-20th century. Even with modern pump-driven
drainage systems, persistent soil waterlogging and inundation in depressions occurs following rainstorms (Granholm 1986, Madrone Associates et al. 1983).
The proportions of poorly drained (waterlogged or
inundated in spring) and well-drained (aerobic soils in
spring) soils in diked Baylands vary with precipitation
amounts and patterns. Years of normal or above normal
rainfall, particularly those with large storms late in the
precipitation season, cause expansion of wetland areas
in diked conditions. These contract during years of below-normal precipitation, especially with a lack of spring
storms. The proportions of effective wetland and upland
also vary with drainage efficiency and the degree of subsidence.
Long-term Drainage of Diked Wetlands – As
subsidence increased, wetland areas increased behind
dikes, particularly in peaty soils. In the early 20th century, many diked farmlands failed because the costs of
compensating for increased subsidence and dike degeneration at times exceeded the return on agricultural benefits. Many derelict agricultural parcels with degenerated
dikes are evident in aerial photographs of San Pablo Bay
in the 1940s. After abandonment of diked farmlands,
partial levee and drain failures increased, causing reversion of agricultural lands to brackish or salt marsh conditions. For example, prior to conversion to salt ponds,
many of the Napa Marsh area’s derelict hayfields in the
1940s had partially reverted to wetland (Madrone Associates 1977).
Contemporary Drainage of Diked Wetlands –
Today, subsidence of diked active agricultural lands has
increased to the point at which it cannot be compensated
by passive gravity drainage through flapgates alone;
drainage sufficient for oat hay farming depends primarily on active pumping of water in ditches for discharge
to the Bay. It is common for elevations in diked Baylands
of San Pablo Bay to average as low as 0 - 1.0 ft N.G.V.D,
and some average below -3.0 ft or more over extensive
areas, as at Bel Marin Keys and Hamilton Field (USACE
1988). In south San Francisco Bay, which was affected
by past subsidence due to long-term groundwater extraction, diked wetland elevations may be even lower
(Moffett and Nichols and Phil Williams Associates
1988). These subsided diked marsh surfaces are often
36
Baylands Ecosystem Species and Community Profiles
very close to the groundwater surface. Accordingly, the
proportion of wetland and upland in contemporary conditions depends on the intensity of pumping and ditch
maintenance. These conditions vary significantly among
diked parcels under different ownership and management. Therefore, the mosaic of wetland and upland in
diked agricultural lands is relatively variable and unpredictable among years and between parcels.
Variability of Artificial Hydrologic Conditions –
The patchiness and instability of diked wetlands is evident, for example, in recent land-use changes in San
Pablo Bay. Cullinan Ranch, actively drained and farmed
oat hayfield until the early 1990s, supported a matrix of
upland cropland and many seasonally wet depressions
with wetland weeds. After cessation of pumping by the
mid-1990s (a period of above average rainfall), the Ranch
rapidly (within 2 years) and spontaneously converted to
a seasonal freshwater marsh dominated by cattails and
flats of Eleocharis parvula (Takekawa et al. 1999).
Nearby, between Tolay Creek and the Petaluma River,
adjacent hayfields with differing schedules of ditch maintenance changed from very similar extensive winterponded swale patterns to striking contrasts of ponded
and drained fields. At another location near Sears Point,
San Pablo Bay, cessation of pumping in relict hayfields
caused conversion to seasonal wetlands dominated by
annual plant species typical of vernal pool communities
(many of which are present in the ephemeral weed floras of depressions within hayfields; Downingia pulchella,
Plagiobothrys bracteatus, Eryngium aristulatum, Callitriche
spp., Eleocharis macrostachya). Thus, the extent of diked
wetlands and their character today are very much artifacts of drainage pump activity.
Similar artificial drainage controls wetland plant
communities in the diked basins of the Suisun Marsh,
which is managed mostly for waterfowl production.
There, relatively low-salinity tidewaters are admitted to
the basins selectively to sustain fresh-brackish perennial
and seasonal marshes (Jones and Stokes 1976, Mall 1969,
Meiorin et al. 1991). The proportion of ponded to vegetated marsh may be controlled by modifying managed
hydroperiods, so that prolonged flooding causes dieback
of vegetation in areas of relatively lower substrate elevation. The seasonal variations in tidewater salinity enable
the timing of flooding to control substrate salinity, also.
Managed marsh hydroperiods are usually designed to
favor mixtures of shallow submerged mud, bulrushes
(Scirpus maritimus, S. americanus, S. pungens), tules (S.
californica, S. acutus), cattail (Typha spp.) and brass-buttons (Cotula coronopifolia), and some non-native annual
grasses (Echinochloa crus-gallii, Polypogon monspeliensis).
Also common in diked brackish marshes are baltic rush
(Juncus balticus), saltgrass (Distichlis spicata) and pickleweed (Salicornia virginica). Other species have colonized
these brackish managed wetlands, including goosefoot
(Chenopodium chenopodioides), docks (Rumex crispus, R.
Soil acidity is normally not highly variable in tidal salt
marshes, which are buffered by cations of estuarine water and relatively stable reduction-oxidation conditions
established by groundwater surface position in the marsh
soil profile (Adam 1990). In diked conditions, extreme
seasonal fluctuations in the soil saturation levels may
occur, causing release of sulfides and free metals in marsh
soils with high sulfur contents. Some depressions in
diked wetlands develop very low pH (pH 4 and occasionally lower) and high concentrations of iron oxide precipitates (Madrone Associates et al. 1983, Madrone Associates 1977). These areas are often barren of vegetation,
or develop sparse, low diversity vegetation. Less extreme
but low pH in diked wetlands may inhibit plant production, but the abundant phytomass of many diked wetlands (e.g., rank growth of pickleweed, cattails, peppercress) suggests that the seasonal drainage and aeration
of diked wetland soils commonly has a stronger overall
effect on vegetation production than low pH. Extremely
low redox potential and sulfite toxicity, which often accompany low pH, are highly significant inhibitors of
plant growth (Russell 1973). Soil acidity is highly variable in diked wetlands and depends on local soil conditions and prevailing hydroperiods.
Disturbance in Diked Wetlands – The disturbance regimes of diked wetlands are influenced primarily by discing and flooding. Discing is performed for agriculture, suppression of weed biomass, and suppression
of mosquito production. Episodes of discing have maintained a significant ruderal (weedy) element to the diked
wetland flora of San Francisco Bay, creating large vegetation gaps suitable for invasion by non-native plants,
particularly annuals. Extreme flooding events which are
possible in non-tidal diked marshes also cause disturbances: deep, prolonged flooding causes mass dieback of
most standing perennial vegetation. Following dieback
events, similar or very dissimilar plant associations may
establish.
Diked Bayland Plant Communities
The plant communities present in the diked Baylands
can resemble those of local tidal salt marshes, tidal brackish marshes, non-tidal perennial freshwater marshes, or
seasonally wet grasslands. Some also have characteristics
similar to components of tidal marshes which are now
regionally scarce or extirpated, such as high marsh pans
and alluvial high marsh ecotones. Diked wetlands usually have lower native species richness than their analogous natural plant communities, and often a larger component of exotic plant species. The typical “ weediness”
of many diked wetlands is probably more a result of past
land uses rather than an intrinsic susceptibility to invasion by exotic vegetation. Some diked wetlands are managed actively to maintain community dominance by
marsh plant species favored by wildlife or game manag-
Chapter 1 —
Plant Communities
37
Plants
pulcher), purslane (Sesuvium verrucosum, a recent invader
native to the Great Basin), celery, (Apium graveolens),
Lepidium latifolium, and Conium maculatum. Some
diked brackish marsh communities are essentially artificial, in contrast with the incidental nature of wetland
communities in diked Baylands which are either derelict
or managed for hay production, pasture, or salt production.
Salinity in Diked Wetlands – The substrate salinity conditions in the diked, drained marshes were
modified by leaching the silty clay Bay muds with precipitation, eliminating leached salts through drainage
ditches and tidegates, and excluding tidal inundation by
dikes. This caused rapid desalinization of the substrate,
enabling glycophytes with relatively low salt tolerance
(compared with the salt marsh flora), such as oats and
agricultural weeds, to dominate the converted tidal
marsh soils (Harvey 1987, Madrone Associates et al.
1983, Meiorin et al. 1991). The desalinized conditions
of the substrate were maintained by drainage through
ditches and tidegates. Subsidence caused (and continues to cause today) decreased efficiency of drainage, and
therefore also decreased flushing of residual or reintroduced salts.
Diked wetlands which have been effectively desalinized for agricultural production do not remain so
unless substantial maintenance efforts are applied to
drainage and dikes. Diked wetlands become resalinized
by partial failure of tidegates and levees (Madrone Associates et al. 1983). Leaking or ruptured tidegates allow
influx of saline tidal waters in drainage ditches. Saline
or brackish ditch water can recharge salts locally in
groundwater, and move into the soil through evapotranspiration and capillary movement. In derelict cattail-lined
ditches of abandoned diked hayfields, late summer ditch
water salinity can reach 15 ppt, due to salt leaching and
evaporation. In addition, seepage through dikes (particularly where Bay mud is silty) introduces salts locally.
Overtopping (cresting) of dikes during storm surges
floods reclaimed salt marsh soils with brackish or saline
water. All these processes recharge soil salinity in diked
wetlands. Overtopping typically occurs in winter, and
is not a rare event, particularly in south San Francisco
Bay (USACE 1988). If poor drainage conditions prevail
following a substantial tidal flooding event in a diked
basin, wetlands rapidly become recolonized by salt-tolerant vegetation. High salinity in diked Baylands is often maintained by episodic tidal flooding events which
are not often observed. Residual salinity tends to decline
very rapidly except where drainage is very poor.
Acidification of Diked Wetlands – Soil acidity
affects plant growth primarily by altering the availability of soil nutrients, or liberating excessive amounts of
otherwise low-solubility ions into the soil solution, creating toxicity problems for roots. Acid-related toxicity
occurs only at very low pH (Reuss and Johnson 1986).
Plants
ers (Mall 1969). Most are either managed for purposes
other than wildlife conservation (hayfields, grazed pasture, flood detention basins, salt evaporation ponds) or
are derelict (i.e., pending conversion to urban development), but may still support significant marsh plant communities.
Plant community composition in diked wetlands
is strongly influenced by the degree of residual soil salinity or salt recharge of soils, the efficacy of artificial
drainage, and the relictual factors of land use history.
These factors vary extremely in diked Baylands: some
exhibit insignificant salinity, maximal drainage and disturbance in some intensively cropped oat hayfields in San
Pablo Bay; others exhibit high salinity, poor drainage and
little disturbance in diked pickleweed marshes in south
San Francisco Bay. Other modifications persisting from
past land uses which affect plant community composition include importation of soils or fill (e.g., former airport landing strips, derelict building pads), abandoned
berms and ponds of gun clubs, residual effects of past
fertilizer applications; industrial waste disposal, and soil
contamination.
Relict Halophytic Vegetation – The majority of
derelict diked wetlands in central and southern San Francisco Bay are dominated by species native to local tidal
salt marshes and brackish marshes, such as Distichlis
spicata and Salicornia virginica (BCDC and Harvey
1983, Madrone Associates et al. 1983). Salt-tolerant
glycophyte species have very low physiological nutritional
requirements for salt, and flourish in non-saline and
subsaline soils (Waisel 1972). They often co-exist with
species with little affinity for saline soil, such as Polypogon
monspeliensis and Lolium multiflorum (Harvey 1987,
Kaufman and Harvey 1987). S. virginica and D. spicata
have a significant competitive advantage over salt-intolerant plant species when substrate salinities are in the
range of halophytes (over 5 ppt soil salinity), and rapidly establish dominance during episodes of high salinity conditions. Some halophytes like S. virginica are efficient colonizers of bare wet mud even when salinity is
low if seed rain intensity is high. Pioneer halophytes do
not necessarily decline in abundance, however, when
substrate salinities decline as a result of progressive leaching and drainage of salts. Many apparent diked “ salt
marshes” are composed of relict vegetation halophyte
vegetation which persists in relatively low salinity conditions. This condition is indicated by the presence of a
minor to subdominant component of species with relatively low salt-tolerance (e.g., ruderal composites, bedstraws, mustards), growing vigorously among halophytes
without indications of salt stress (stunted growth, leaf
tip burn, pale leaves) in diked “ salt marshes.” Examples
are sometimes found in abandoned dredge disposal sites
(Zentner and Zentner 1995, Huffman and Associates
1996). Some mixed halophyte-glycophyte associations
may also occur where stratification of rooting zones oc-
38
Baylands Ecosystem Species and Community Profiles
curs in distinct salinity horizons, caused by near-surface
leaching of salts and accumulation in deeper portions of
the soil profile.
Thus, apparent salt marsh vegetation in diked Baylands may indicate either current high salinity or former
high salinity, and does not necessarily indicate sustained
high residual salinity. It often represents inertia in plant
community structure after relaxation of salinity stress.
The term “ non-tidal salt marsh” in the context of the
San Francisco Bay Estuary should be interpreted narrowly in floristic rather than physiological terms, because
dominance of halophytes in the unstable substrate salinity conditions in diked wetlands is an unreliable indicator of current substrate salinity. Some diked salt
marshes are truly saline and tend to remain so because
of chronically poor drainage or frequent partial dike failures. Others are in gradual succession to other vegetation types. Some diked salt marshes with low residual
substrate salinity are subject to rapid conversion to other
vegetation types following disturbances (e.g., discing or
flooding).
Species Richness and Composition – The species
richness and composition of diked marshes is highly variable among sites, and among different marsh types. High
salinity and hypersalinity in diked marshes tend to promote low species diversity, selecting for a few tolerant
species. Other extreme soil conditions, such as strong
acid production and mass release of free iron (often associated with prolonged inundation followed by summer
drought) minimize plant species diversity. Truly hypersaline seasonal wetlands in the Bay usually support only
sparse Salicornia virginica, Distichlis spicata, and Salsola
soda, with a minor component of Frankenia salina. Hypersaline seasonal wetlands are now scarce in the Bay
Area, mostly scattered around South Bay salt ponds and
adjacent lands. A few occur in the North Bay (e.g., parts
of Gallinas Creek diked salt marshes, peripheral portions
of the Napa salt ponds). Many former hypersaline diked
wetlands have been altered by water management for
mosquito abatement and wildlife habitat enhancement,
and are now muted tidal marshes (e.g., New Chicago
Marsh in Alviso, Oro Loma Marsh in Hayward).
Species diversity in nontidal diked brackish and salt
marshes is generally much higher than in hypersaline
basins, but this does not reflect relatively greater overall
diversity of native plant species. Diked brackish to saline nontidal wetlands support a number of common
native tidal brackish and salt marsh species (Salicornia
virginica, Distichlis spicata, Frankenia salina, Cuscuta
salina, Atriplex triangularis) and sometimes support relatively infrequent native species typical of the natural high
tidal marsh and upland ecotone (Iva axillaris, Leymus
triticoides, Baccharis douglasii). The native perennial grass
Leymus triticoides, historically a dominant species of the
upper transition zone of tidal salt and brackish marshes,
is infrequently found in some diked brackish marshes,
back after prolonged deep flooding. Other common herbaceous non-native plant species of diked brackish and
saline marshes include Lepidium latifolium, Bassia hyssopifolia, Beta vulgaris, Salsola kali, and Salsola soda. Lepidium latifolium is especially invasive in brackish diked
marshes, particularly where the soil has been disturbed,
but also in areas of marsh with thin or discontinuous
vegetative cover.
Diked subsaline and nonsaline Baylands are very
seldom the subject of careful floristic surveys (e.g., Rugt
1991, Madrone Associates 1977); vegetation descriptions
usually focus on visually dominant ruderal species, often based on summer survey dates when native annual
species are not identifiable (Jones and Stokes 1977,
Hudson 1980, Werminski 1973). Consequently, the floristic diversity and affinities of diked subsaline to fresh
seasonal wetlands has probably in many (perhaps most)
cases been underestimated. Diked subsaline and nonsaline wetlands are mixtures of exotic species of ruderal
seasonal wetlands and native species typical of vernal
pools and swales. Diked wetlands which are mostly subsaline to nonsaline after years of agricultural drainage
support a range of marsh plant associations. These are
most common around San Pablo Bay and Suisun Bay,
where grazing pasture and oat hayfields have been maintained for many decades in diked former tidal brackish
marshes. They may have inclusions of relatively brackish indicator species where soil salinity and acidity are
locally elevated (e.g., mixtures of Atriplex triangularis,
Polypogon monspeliensis, Distichlis spicata) but are dominated by glycophytic wetland plant species, both native
and non-native. Composition of the fresh/subsaline
diked wetland flora is influenced by disturbance. Annually disked hayfields support wetland “ weeds” which are
a mixture of native annuals (e.g., Plagiobothrys spp., esp.
P. stipitatus, P. leptocladus), Juncus bufonius, Lilaea
scilloides, Callitriche marginata, C. spp. Cicendia quadrangularis, Elatine brachysperma, Eryngium spp., Cressa
truxillensis; locally, Downingia spp.; non-native annuals
(Lythrum hyssopifolium, Cotula coronopifolia, Polygonum
aviculare, Hordeum marinum ssp. gussoneanum, Polypogon
monspeliensis, ) and non-native perennials (Lotus corniculatus, Agrostis avenacea) Grazed pasture land in diked
Baylands in San Pablo Bay may also support native annuals found in diked disked hayfields, as well as native
perennials (Eleocharis machrostachya, Glyceria spp. Juncus
effusus, J. patens) and naturalized non-native perennials
(Rumex crispus, R. pulcher, Cirsium arvense, Lolium multiflorum). The relative abundance of these species in
diked pasture and hayfield wetlands is variable and unstable. Some diked wetlands, after relaxation of intensive agricultural manipulation, develop seasonal wetlands
with plant species composition highly similar to that of
regional vernal pools and swales (locally dominated by
Downingia spp., Eryngium spp. Eleocharis macrostachya,
Callitriche spp. Lilaea scilloides, Plagiobothrys spp., etc.)
Chapter 1 —
Plant Communities
39
Plants
particularly where disturbance has been infrequent. A
relatively rare historic component of subsaline tidal
marsh ecotones, Centaurium muehlenbergii is found in
diked subsaline wetlands at Cullinan Ranch. It is currently reported known from only one tidal marsh/upland
ecotone (China Camp). The sedge Scirpus maritimus, a
dominant native component of tidal brackish marshes,
is often abundant or dominant in brackish to saline
ditches or deep, wet depressions in diked marshes ((Madrone Associates et. al. 1983, Mall 1969).
Conversely, diked salt and brackish marshes generally fail to support some important species of corresponding tidal marsh communities; Spartina foliosa is excluded from nontidal conditions, and Jaumea carnosa,
Plantago maritima, Triglochin spp. and Limonium californicum are absent or very infrequent in nontidal salt
marsh; Grindelia stricta is generally less abundant in nontidal salt marsh than tidal marsh. Diked salt marshes
typically lack rare tidal marsh species (e.g., Cordylanthus
spp., Castilleja ambigua, Lasthenia glabrata, Lilaeopsis
masonii, Cirsium hydrophilum, Aster lentus), and also
usually lack most infrequent tidal marsh species (e.g.,
Pluchea odorata, Senecio hydrophilus, Glaux maritima).
The failure of these tidal marsh species in diked conditions is probably due to the relatively greater competition by robust “ generalist” species with broad ecological amplitude, and physiological intolerance of extremes
of inundation and dryness in diked wetlands. Diked salt
and brackish marshes in some cases, however, provide
refugia for tidal marsh plants of the high tidal marsh
which have become (or in some cases have always been)
regionally rare or infrequent in the modern tidal marsh
ecosystem, such as Suaeda moquinii, Hemizonia
pungens ssp. maritima, Salicornia subterminalis, Downingia pulchella, Juncus mexicanus). As such, they may
serve to maintain genetically differentiated salt-tolerant populations of species displaced from modern tidal
marshes.
Diked brackish and salt marshes are subject to invasion by many non-native species and species which are
not typical of tidal marshes, or are typically restricted to
marginal conditions in tidal marshes. Non-native pasture grasses with moderate salinity tolerance, such as
Lolium multiflorum (and hybrids), Polypogon monspeliensis, Lotus corniculatus and Hordeum marinum ssp.
gussoneanum, and even Rumex crispus are also major components of diked salt and brackish marshes, often locally
dominating either depressions (Polypogon, Hordeum) or
mounds (Lolium). Exotic halophytic grasses Parapholis
incurva and Hainardia cylindrica are also locally common
in diked salt or brackish marsh. Cotula coronopifolia is
usually only a minor component of tidal salt and brackish marsh, colonizing depressions and marsh pan edges,
but is often a major component of diked brackish
marshes, particularly in disturbed or winter-ponded
brackish depressions where other vegetation has died
Plants
Reference Sites
Reference sites for different types of diked wetlands
would generally not be long-lived because of the prevalence of unstable vegetation conditions in diked Baylands. Droughts, wet years, changes in drainage and
pumping, disturbances from agricultural practices, and
succession can cause profound changes in vegetation in
short periods of time. The following reference sites reflect conditions observed in the mid-late 1990s.
1. Diked non-tidal salt marsh (dominant Salicornia
virginica)
• Fremont Airport (King and Lyons site; proposed
for phased tidal restoration), Alameda Co.
• Gallinas Creek diked wetlands, Marin Co.
• Western Marsh and Central Lowlands, Bahia
Site, Novato, Marin Co.
• Dredge pond 3E, Mare Island, Solano Co.
• Area H, Redwood Shores, San Mateo Co.
2. Diked non-tidal brackish marsh
• Cullinan Ranch, Solano Co.
• Suisun Marsh managed marshes, Solano Co.
• Huichica Unit, CDFG Napa-Sonoma Marsh,
Sonoma Co.
3. Diked subsaline to nonsaline seasonal wetlands
• Black Point/Renaissance Faire site, Novato,
Marin Co. (extirpated 1999)
• Twin House Ranch Site, Lower Petaluma River,
Sonoma Co.
• Leonard Ranch, North Point, Dixon parcels,
Sonoma Co., along Hwy 37
Francisco Peninsula, where sand beach ridges occurred.
Seasonal freshwater wetlands (vernal pools and
swales, springs) occurred within grasslands peripheral to
the Bay, particularly in the Petaluma River valley, on alluvial terraces near Fremont, portions of Richmond and
Berkeley, and along much of the Suisun Marsh area.
Their distribution and abundance, as suggested by soil
surveys, were probably not limited to areas mapped as
poorly drained; seasonal freshwater wetlands often occur as local inclusions within soil series in which wetlands are not indicated as prevalent. This is indicated by
records of vernal pool endemics in locations like San
Francisco, where “ vernal pool” soil types are not mapped,
but winter pools with typical endemic annuals were
found.
The historical abundance and distribution of these
wetland types is extremely difficult to quantify in terms
of area. Quantitative estimates of historic abundance of
seasonal wetlands displaced by urbanization depends
heavily on interpretation and assumptions about early
soil surveys (which were not intended to function as
maps of actual or potential native vegetation), historical
accounts, and fragmentary information on species occurrences in old floras. The qualitative differences in natural non-tidal wetland types and their diked Bayland analogues further obscures the relevance of quantitative
comparisons between historic losses of natural seasonal
wetland plant communities and their partial replacement
with wetlands of diked Baylands.
Conservation Issues
Historic and Modern Distribution
Wetlands of diked Baylands are relatively recent historic
artifacts. The plant associations they support are analogous to, but distinct from, wetlands along the margins
of historic tidal marshes. Brackish non-tidal marshes
somewhat similar to diked brackish marshes probably occurred within alluvial deposits at mouths of small streams
which discharged into tidal marshes with locally poor
drainage, such as near Ignacio (Novato), where riparian
areas converged with dense marsh ponds and few or no
tidal creeks. Analogous examples of brackish or subsaline marshes with marginal tidal flooding are found today along Drakes Estero and Tomales Bay, particularly
near shallow backbarrier lagoons. Salt marsh with restricted tidal influence probably occurred along portions
of the Bay where local sand beach ridges were likely to
obstruct tidal flows. One modern example exists at Pinole
Point (Whittell Marsh), where the proximal end of a
sand spit episodically dams small tidal channels, causing seasonal ponding in a small salt marsh cut off from
regular tidal flows. Prehistoric examples of “ pocket”
nontidal salt marsh probably occurred in the vicinity of
Richardson Bay, Alameda, Oakland, and the San
40
Baylands Ecosystem Species and Community Profiles
Plant conservation needs for diked wetlands are dependent on larger-scale wetland management and restoration plans. Diked wetlands usually support less native
plant species diversity than mature tidal marshes at
equivalent locations, but may in some cases still provide
important plant conservation functions. For example, in
San Pablo Bay, agriculture and development have eliminated most historic natural seasonal wetlands in supratidal grasslands peripheral to the Bay. The original vernal pool flora which occurred in subsaline to alkaline
depressions around the historic edge of the Bay (as in
parts of northeastern Suisun Marsh today) has been
largely extirpated in its original location, but persists in
artificial equivalent topography and edaphic conditions
in some diked seasonal wetlands. These populations
maintained in subsaline conditions may provide important founder populations for opportunities to restore
vernal pool and swale systems in the original soil types
and topography along the margins of the Bay, in coordination with tidal restoration. Similarly, one diked salt
marsh in the South Bay (former Fremont Airport) provides refugia for Suaeda moquinii, otherwise found
around the Bay only in remnant alkali vernal pools ad-
Conclusions and Recommendations
Diked wetlands considered for conversion to other marsh
types, such as tidal wetlands, should be studied individually for site-specific floristic values, particularly for potential functions as refugia for species displaced from historic seasonal wetlands and tidal marsh ecotones. Diked
wetlands should not be assumed to have uniformly low
native wetland plant species diversity or “ ruderal” status. In areas where restoration of seasonal fresh wetland
systems (e.g., vernal pools, alkali basins, alluvial Juncus/
Scirpus marsh, etc.) is precluded by development, some
diked wetlands should be considered for modification
and management to maintain regionally scarce plant
communities. Generally, however, priority should be
assigned to restore peripheral estuarine plant communities in their proper original soils and topographic position. Where diked wetlands support regionally rare plant
populations, they should be given interim conservation
priority until suitable population restoration sites are established in more natural or restored habitats. Existing
diked marshes should be managed to minimize impacts
of exotic invasive plants on adjacent managed or natural tidal marshes. Dike maintenance should include best
management practices which favor recolonization of disturbed dike surfaces by native vegetation and suppress
re-invasion by exotic species.
References
Adam, P. 1990. Saltmarsh ecology. Cambridge Univ.
Press.
BCDC (San Francisco Bay Conservation and Development Commission) and H.T. Harvey. 1983.
Aquatic and wildlife resources of Richardson Bay.
Unpublished report prepared for the Richardson
Bay Special Area Plan Study.
Doane, S. N. 1999. Shoreline changes in San Pablo Bay,
California. M.Sc. thesis, Vanderbilt Univ. 116 pp.
Eicher, A. L. 1988. Soil-vegetation correlations in wetlands and adjacent uplands of the San Francisco
Bay estuary, California. U.S. Fish and Wildlife Service Biological Report 88 (21), contract no. 1416-0009-85-001. August 1988
Granholm, S.L. 1989. Endangered habitat: a report on
the status of seasonal wetlands in San Francisco
Bay. Report sponsored by National Audubon Society, San Francisco Bay Area Audubon Society
chapters, Save San Francisco Bay Association, Sierra Club Bay Chapter.
Harvey, T.E. 1987. Fish, wildlife, and habitat management plan for Naval Security Group Activity Skaggs Island, California. U.S. Fish and Wildlife
Service, San Francisco Bay National Wildlife Refuge, Newark, Calif.
Hudson, J. 1980. Bird census of diked-marshland habitat. In: D.S. Sloan (ed). San Pablo Bay: an environmental perspective. Environmental Studies
Group, Univ. of Calif., Berkeley.
Huffman and Associates 1996. Proposed Section 404
wetland delineation for the Bahia Master Plan,
Novato, California. U.S. Army Corps of Engineers,
San Francisco District, file no. 148831.
Jaffee, B., R.E. Smith and L.Z. Torressan. Sedimentation changes in San Pablo Bay. U.S. Geological
Survey open-file report 98-759.
Chapter 1 —
Plant Communities
41
Plants
jacent to the Bay at one site (Zentner and Zentner 1996).
Partial vernal pool floras have also been generated spontaneously after cessation or relaxation of agricultural
manipulation at Montezuma Wetlands (Solano County),
Sears Point (Sonoma County), and a construction site
in Alviso (Santa Clara County). Most diked wetlands are
poorly surveyed, and may act as refugia for many populations of plants of conservation significance.
Diked wetlands are also conservation threats to
plant species diversity when they provide outposts, reservoirs, or dispersal corridors for invasive wetland weeds,
such as Lepidium latifolium and Salsola soda. By increasing seed rain pressures on adjacent tidal marshes, or adjacent marsh restoration sites, diked wetlands may also
cause degradation of tidal marshes.
Sea level rise makes long-term conservation of
diked wetlands problematic. In addition to inherent tendencies of diked systems to suffer levee subsidence and
erosion, sea level rise imposes increasing risks of levee
failure and tidal flooding. Breached diked wetlands spontaneously revert to tidal wetlands, but usually only as low
mudflat or marsh to lower middle marsh after even two
decades (e.g., White Slough, Vallejo, Solano County) In
addition, some high-sulfur diked marsh soils undergo
long-term changes in soil chemistry which make them
unsustainable for any valuable natural or artificial vegetation.
Dike maintenance and repair may cause degradation to diked and tidal marsh plant communities by favoring spread and dominance of exotic invasive marsh
plant species. Dike maintenance practices currently lack
any elements which facilitate recolonization by native
species.
Restoration of diked marshes is somewhat self-contradictory, since true restoration would entail conversion
to the original tidal marsh condition. However, diked
wetlands can be significantly enhanced as non-tidal
marshes by reducing or eliminating adverse land use
practices. Reduction of intensive drainage efforts and
elimination of high-frequency disking can enable diked
fresh/subsaline wetland plant communities to mature
and accumulate greater native species. Pasture management that tolerates some winter inundation in depressions, for example, is more compatible with native wetland plant species diversity than oat crop management.
Plants
Jones and Stokes. 1976. Suisun Marsh Protection Plan.
Calif. Dept. Fish and Game, Sacramento, Calif.
Kaufman, S. and H.T. Harvey. 1987. Plant list: Army
Corps of Engineers; Leslie Salt Site. File no. 30901. Harvey and Associates, Alviso, Calif.
Madrone Associates. 1977. The natural resources of
Napa Marsh. Report prepared for the Calif. Dept.
Fish and Game, Coastal Wetland Series # 19. Sacramento, Calif.
Madrone Associates, Philip William and Associates, J.R.
Cherniss and N. Wakeman. 1983. Ecological values of diked historic baylands. A technical report
prepared for the San Francisco Bay Conservation
and Development Commission, April 1992 (revised 1983).
Mall, R.E. 1969. Soil-water-salt relationships of waterfowl food plants in Suisun Marsh of California.
State of Calif. Dept. Fish and Game Wildlife Bulletin No. 1. Sacramento, Calif.
Meiorin, E.C., M.N. Josselyn, R. Crawford, J. Calloway,
K. Miller, R. Pratt, T. Richardson and R. Leidy
1991. San Francisco Estuary Project status and
trends report on wetlands and related habitats
in the San Francisco Estuary. Public Report prepared under cooperative agreement #815406-010 with the U.S. Environmental Protection
Agency by the Association of Bay Area Governments, Oakland California; Romberg Tiburon
Centers of San Francisco State Univ., and the
U.S. Fish and Wildlife Service, Sacramento Calif.
Moffatt and Nichol and Wetland Research Associates
1988. Future sea level rise: predictions and implications for San Francisco Bay. Report prepared for
the San Francisco Bay Conservation and Development Commission, San Francisco, Calif. Revised
edition.
42
Baylands Ecosystem Species and Community Profiles
Reuss J.O. and D.W. Johnson. 1986. Acid Deposition
and Acidification of Soils and Waters. SpringerVerlag, New York.
Rugyt, J. 1991. Checklist of vascular plant species at
the Montezuma Wetlands Project site. Appendix to
Montezuma Wetlands Project Technical Report,
Levine-Fricke Restoration Corp., Emeryville, Calif.
Russell, E.W. 1973. Soil conditions and plant growth.
10th ed., Longman, London and New York.
Takekawa, J., M. Eagan and R. Laird. 1999. Monitoring tidal wetland restoration projects in the San
Pablo Bay National Wildlife Refuge. Progress Report, May 1999. U.S. Geological Survey Biological Resources Division, Mare Island Field Office,
Calif.
U.S. Army Corps of Engineers (USACE). 1988. San Francisco Bay Shoreline Study: Southern Alameda and
Santa Clara Counties, Interim Office Report. U.S.
Army Corps of Engineers, San Francisco District.
Ver Planck, W.E. 1958. Salt in California. State of Calif. Dept. Natural Resources, Division of Mines,
Bulletin 175. 168 pp.
Wermunski, J. 1973. Ecological attributes of the Hayward Area Shoreline: a habitat survey with recommendations. Unpublished background technical
report prepared for the Hayward Area Shoreline
Planning Agency, Hayward, Calif.
Waisel, Y. 1972. The biology of halophytes. New York:
Academic Press.
Waterways Experimental Station, U.S. Army Corps of
Engineers (WES). 1987. Corps of Engineers Wetlands Delineation Manual. U.S. Army Corps of
Engineers Wetlands Research Program Technical
Report Y-87-1, January 1987.
Zentner and Zentner. 1995. Section 404 wetland delineation for the Bayside Business Park Phase II, Fremont, California. U.S. Army Corps of Engineers,
San Francisco District.
Peter R. Baye
Introduction
The term “ salt pond,” as treated in this discussion, includes both natural and artificial large-scale persistent hypersaline ponds that are intermittently flooded with Bay
water, and which occur within tidal salt marsh systems
of San Francisco Bay and San Pablo Bay. Historic natural salt ponds were characterized by persistent thick accumulation of salt inundated with concentrated seawater brines. They were restricted to a relatively narrow
reach of San Francisco Bay near San Lorenzo Creek.
They are distinguished here from related salt marsh features such as pans and which occur at smaller spatial
scales, have distinctive physiographic traits, and lack
strong persistent (perennial) brines and precipitated
crystaline salt deposits. Artificial salt ponds (solar
salterns) are diked salt marshes which are managed for
the production of concentrated brine and fractional crystallization of sea salts. Natural and artificial salt ponds
are presumed to share the same narrowly adapted hypersaline biota.
Information on modern artificially engineered salt
pond systems is derived principally from the biological
literature on solar salterns and hypersaline environments
(Javor 1989, and references within), historic documentation on the salt industry in California from the State
Division of Mines (Ver Planck 1958, 1951; Dobkin and
Anderson 1994) and regional documentation produced
by the local salt industry and government regulatory
agencies (Corps of Engineers, San Francisco District,
Regulatory Branch permit and compliance files; Office
of Counsel files, and references within). Information on
historic salt pond systems is limited to descriptive historic accounts and descriptions, detailed topographic
maps of natural salt ponds prior to extensive dike construction (U.S. Coast Survey T-charts, 1956), and field
investigations by the author comparing modern salt pans,
marsh ponds, and artificial salt ponds.
Environmental Setting
Salt ponds are large, shallow, hypersaline impoundments
or depressions in tidal salt marsh systems which undergo
a sequence of infrequent flooding with saline or brackish
Bay water, evaporative concentration, and formation of
strong hypersaline brines and deposits of gypsum, calcium
carbonate, and crystalline salt (halite; sodium chloride).
Historic salt ponds were mapped with a high degree of resolution in the 1856 U.S. Coast Survey. They
were nested within particular portions of the salt marshes
Chapter 1 —
Plant Communities
43
Plants
Plants of San Francisco Bay
Salt Ponds
along the Alameda shoreline in the vicinity of San
Lorenzo Creek and Mount Eden Slough. This reach of
salt marsh was distinguished by a relatively straight-edge
erosional marsh shoreline, little tidal drainage at the edge
of the mudflats, and evidence of drowned marsh topography (mapped as emergent sinuous tidal creek levees).
The upland edge was an extensive alluvial lowland, presumably with significant subsurface groundwater discharge. No major freshwater creeks were directly associated with the salt ponds. Atwater et al. (1979) suggested
that natural estuarine beach ridges along outer marsh
edge were responsible for the impoundments of salt
marsh that created salt ponds near San Lorenzo. Some
salt ponds at the northern end of the local San Lorenzo
distribution were certainly associated with well-defined
barrier sand spits (U.S. Coast Survey T-charts, 1850s),
which were probably nourished by sand eroded from
submerged Merritt sand deposits (Pleistocene marine
beach and dune). Less well-defined transgressive berms
of sand and coarse organic detritus may have been deposited on top of the erosional marsh edge south of the
sand spits themselves. Similar transgressive beach-marsh
berms today act as dams enclosing freshwater to brackish ponds and marshes in Drake’s Estero, Point Reyes
and at one location in San Francisco Bay (Whittell
Marsh, Point Pinole, Contra Costa County). U.S. Coast
Survey T-charts also indicate numerous sandy barrier
beaches which dammed (either permanently or intermittently) lagoons. The impoundment of Crystal Salt Pond
by a wave-constructed swash bar or beach ridge would
distinguish it morphologically, hydrologically, and topographically from more common salt marsh ponds (pans)
which occurred as depressions, sometimes extensive,
between tidal creeks. These were widely distributed in
salt marshes in the South Bay. Extensive, elongate pans
also occurred near and below the upland borders of salt
marshes; these have been termed “ transitional” pans, although their position and form do not necessarily indicate a gradual ecotonal relationship with alluvial or upland habitats.
Salt ponds today (solar salterns) are artificially managed and engineered diked Baylands converted from tidal
salt marsh. The first artificial salt ponds began as extensions and improvements of natural salt ponds which occurred near Hayward (Crystal Salt Pond), but most of
the contemporary man-made salt pond system is established in former tidal marsh that included few or no
perennial hypersaline ponds. Artificial salt ponds have
entirely displaced their natural forerunners; no natural
true salt-crystallizing ponds remain in San Francisco Bay
today, although related smaller salt pans and marsh
ponds containing weak brines in summer and fall do
occur.
Classification of Salt Ponds – Javor (1989) placed
marine-derived hypersaline aquatic environments in four
ecological salinity classes:
Plants
The first salinity class (ca. 60 - 100 ppt) contains
a highly diverse, productive biota dominated by marine
species. This class would correspond to “ low salinity”
ponds (a misnomer, since salinity exceeds seawater concentration), from intake ponds to the next one or two
stages that support abundant macroalgae and fish.
The second class (ca. 100 - 140 ppt) is dominated
by specially adapted halophilic species which are related
to freshwater taxa, not marine taxa. The organisms include abundant cyanobacteria, unicellular green algae,
brine shrimp, and various halobacteria.
The third class (ca. 140 - 300 ppt) is distinguished
by marked reduction of species diversity (loss of cyanobacteria, most invertebrates other than brine shrimp),
and dominance of Dunaliella and brine shrimp.
The fourth class (300 ppt to salt saturation, near
360 ppt) contains only Dunaliella and bacteria at low
productivity.
The first class predominates in modern marsh
ponds. The historic natural salt pond complex probably
varied seasonally between Javor’s second to fourth hypersaline classes. Other natural marsh pans were most
likely predominantly in the first class only, becoming seasonally hypersaline, and supporting relatively weak brines
and macroalgal cover. Natural historic salt ponds were
distinguished from other types of inundated depressions
in salt marshes by the persistent thick halite deposits, indicating perennial hypersaline conditions, and their large
lake-like size. In these aspects, they differ from shallow
marsh ponds and marsh pans, which are regularly
flooded during higher spring tides, and either remain
persistently ponded or develop thick algal mats which
desiccate in summer (bleaching white in the sun, resembling salt deposits in aerial photographs), or only develop
thin, temporary salt films on unvegetated mud and peat.
Various marsh pan features are represented in U.S.
Coast Survey maps of the mid-19th century, but only a
few have persisted in modern rare remnant tidal marshes,
such as Petaluma Marsh, Rush Ranch and Hill Slough
(Solano County). Elongate marsh ponds are evident
along the upland edge of historic marshes, particularly
in eastern and southern parts of San Francisco Bay. Some
of these may have been influenced by surface runoff and
groundwater seepage from adjacent alluvial uplands, and
could have been less saline than other marsh depressions
most of the year. Some historic elongate marsh edge pans
may also have been the unvegetated upper intertidal surface of alluvial fans and terraces, consistent with small
modern “ transitional pans” observed at Hill Slough,
Solano County. These also lack brine and halite development. Modern elongate marsh pans have formed in
recently (100 year) prograded marshes adjacent to Mare
Island dredge ponds. These ponds are about 0.3 m deep
in winter and spring, and range from brackish (nearly
fresh) in winter to hypersaline when ponded areas are
highly reduced in summer, but no significant halite pre-
44
Baylands Ecosystem Species and Community Profiles
cipitation is evident in them. These and similar pans may
appear white with sun-bleached dried algal mats, which
resemble salt flats. High densities of true natural marsh
ponds, also termed “ drainage divide ponds” (owing to
their position in poorly drained marsh areas between
tidal creeks), also occur in the Petaluma Marsh. Marsh
ponds are a variation of salt pans which are topographic
depressions flooded by spring tides, and support
submergent vegetation, typically macroalgae (such as
Enteromorpha spp.) and beds of widgeon-grass (Ruppia
maritima), indicating brackish to near-marine salinity.
The beds of marsh ponds are usually a soft organic oillike black muck composed of decayed, waterlogged organic matter.
In contrast with salt ponds in estuaries with strong
marine influence, such as San Diego Bay, San Francisco
Bay salt ponds are relatively nutrient-rich and sustain
high primary productivity (Javor 1989). Nutrient-poor
salt pond conditions promote microbial mats, while
planktonic microalgae tend to dominate nutrient-rich
salt pond systems (Javor 1989). Most salt ponds in San
Francisco Bay support richly pigmented and somewhat
turbid organic “ soups” of Dunaliella, halobacteria, cyanobacteria, dissolved organics and organic particulates
and, often in ponds between approximately 120 200 ppt salinity, large “ blooms” of brine shrimp which
graze primarily on Dunaliella.
Historic natural salt ponds were unlike modern
artificial salt ponds in that they were not differentiated
geographically into stable hypersaline classes, but varied
only seasonally in salinity. Natural salt ponds went
through a seasonal “ intake” phase during extreme high
spring tides (December-January and June-July), when
Bay water flooded them and diluted them with brackish to saline Bay water, seldom exceeding 20 ppt, and
typically between 2 - 10 ppt in winter. During summerfall evaporation periods, brines formed in situ, ranging
in salinity over time up to crystallization (saturation) near
360 ppt. In contrast, the modern engineered salt pond
system is based on timed transfers of brines between
ponds, resulting in spatial separation of brines at different stages of concentration, and fractional crystallization
of various seawater salts (other than sodium chloride,
halite), such as magnesium and potassium salts (bitterns), gypsum (calcium sulfate) and lime (calcium chloride) in different ponds. In this system, crystallization
is restricted to relatively few ponds engineered to facilitate harvest of halite deposits, and relatively stable hypersalinity regimes are established for individual evaporator ponds in the system (Ver Planck 1958).
The sequential and spatial separation of brines in
artificial salt pond systems also produces salt pond
“ types” which are not fully analogous to natural systems.
The late stages of brine production near sodium chloride crystallization produce strong non-sodium brines
called “ bittern.” Bittern brines (or bittern) are a concen-
Salt Pond Plant Community
Salt ponds support a distinctive and highly specialized
halotolerant to halophilic biota consisting of microalgae,
photosynthetic bacteria, and invertebrates, but no vascular plants (except along the edges of artificial salt pond
levees). The dominant photosynthetic organisms of most
hypersaline San Francisco Bay salt ponds are a singlecelled green algal species, Dunaliella salina (Chlorophycophyta) and numerous species of blue-green bacteria
(Cyanobacteria), halobacteria, and purple sulfur-reducing bacteria. The proportions of these organisms vary
with salinity. Artificial eutrophic salt ponds with salinities closer to marine concentrations (near 35 ppt; “ intake ponds” ) are dominated by marine macroalgae such
as sea-lettuce (Ulva spp.), Enteromorpha spp., Cladophora
spp., and also sometimes support Fucus spp. and Codium
spp. where substrate is stable and firm. They also include
marine diatoms, dinoflagellates, and cryptomonads.
There are no detailed studies of the species diversity, distribution or geographic variation of the halophilic microflora communities of San Francisco Bay.
Managed and engineered contemporary salt
ponds are ecologically similar in many respects to their
natural precursor salt ponds, and presumably share the
same algal and bacterial microflora.
Indicator Species – There are no detailed classifications or analytic studies of salt pond algal communities. Following Javor’s (1989) classification of hypersaline environments (see Classification of Salt Ponds,
above), two broad hypersaline algal communities may be
identified: communities dominated by free-floating marine macroalgae typical of upper tidepools near marine
salinities to low-hypersaline conditions, corresponding
to intake ponds and young brines in a saltern series (e.g.,
Ulva spp., Enteromorpha spp., Cladophora spp.; also bottom-mat forming cyanobacterial colonies); and communities dominated by motile unicellular halophilic phyto-
Bob Walker
Plants
trated solution of sodium chloride, magnesium chloride
and sulfate, and potassium chloride and sulfate. The
ionic balance of highly concentrated bittern is toxic even
to bacteria, and saturated bittern is considered sterile
(Javor 1989). During winter rains, dilute bittern stratifies on top of the concentrated bittern, and brine shrimp
may appear seasonally, indicating algal production (Jim
Swanson, Rick Coleman, pers. comm.). Natural salt
pond brines did include bittern salts; in fact, the “ low
quality” of early California solar salt was due to bittern,
and the modern solar saltern system is principally devised
as a method to fractionate sodium and bittern salts. Crystallizer ponds, which are used to precipitate halite, are
also maintained near the limits of halotolerance of Dunaliella (which can nonetheless fix carbon up to salt saturation; Javor 1989), but undergo seasonal dilution during winter rains.
Modern salt ponds are artificially managed and
engineered diked baylands converted from tidal salt
marsh. (South San Francisco Bay)
plankton (principally Dunaliella salina), which characterize moderate to high hypersaline conditions. Macroalgal salt pond communities also correspond with fishdominated animal communities, while phytoplanktondominated brines are associated with brine shrimp abundance.
Dunaliella spp. is ubiquitous in salt ponds in San
Francisco Bay. It is reported to survive, and can be photosynthetically active, in brines which are close to saturated (near 350 ppt), but may be absent in some extremely concentrated brines and bittern (potash-phase,
or potassium-magnesium) brines (Javor 1989, Brock
1975). Its optimum salinity for growth is near 120 ppt,
about four times the concentration of seawater. Dunaliella salina concentrates carotenoid and other pigments
in response to various forms of physiological stress, including salinity. It can be used as a crude color-indicator of brine salinity: cells growing in 50-100 ppt are
greenish, and turn yellowish-green in 150 ppt brine.
Reddish hues occur in brines 200-250 ppt (Javor 1989).
Purplish-red hues in brines over 200 ppt may be contributed by halophilic bacteria. A conspicuous mosaic of
salt pond hues are readily visible from aerial views of San
Francisco Bay, particularly in summer and fall. Dunaliella osmoregulates in hypersaline brines by concentrating glycerol as a compatible osmotic solute in its cytoplasm (Javor 1989).
Reference sites
There are currently no reference sites in the San Francisco Bay Estuary for true natural salt ponds (ponds
which periodically or chronically produce crystalline salt
deposits). The historic salt pond system near San
Lorenzo Creek in Alameda was eliminated by diking in
the 1850s and 1860s. All modern salt pans and marsh
ponds in the Bay Area differ from these historic salt
ponds. Most existing marsh ponds are only slightly hypersaline, or briefly hypersaline in late summer, and support algal mats rather than brines and halite beds. Most
Chapter 1 —
Plant Communities
45
Plants
existing salt pans within small modern Bay Area salt
marshes are comparatively small and produce sparse and
thin (few mm) salt crusts in summer and fall. In contrast, reference sites for artificial salt ponds are abundant.
Examples of (relatively) low salinity intake ponds, which
are saline or slightly hypersaline, are found at Pond B1/
B2 in Mountain View, Pond 1 near Mowry Slough, and
Pond A9 in Alviso. Examples of intermediate hypersaline ponds (known as concentrators or evaporators) are
found in ponds A10-14 in Alviso, ponds 2-8 near Coyote Hills, and ponds 2-6 between Mowry Slough and
Coyote Creek. High hypersaline ponds (strong brines approaching or reaching salt saturation, “ pickle” ) are found
in extensive crystallizer beds near Newark and Redwood
City, ponds 10 and 26 near Newark, and periodically in
drained evaporators before they are re-filled.
Modern salt marsh (and brackish marsh) pans may
be found in few remnant pre-historic tidal marshes at
Petaluma Marsh (abundant), China Camp (scarce) and
Point Pinole (Whittell Marsh; scarce). Pans vary in topography. Some upper marsh pans are similar to patches
of salt flats, while pans in middle marsh zone depressions
are normally shallow ponds 10-20 cm deep. Pans which
become ponded, either because of depressional topography or marsh surface drainage barriers, develop algae
or widgeon-grass. Salt marsh pans also occur in historically accreted marshes at Mowry Marsh. Elongate marsh
pans fringing uplands (“ transitional” pans) have also
formed in the relatively young (20th century) salt marsh
at Emeryville Crescent and adjacent to Mare Island
dredge ponds. Elongate but diffuse shore-parallel marsh
pans, perhaps best regarded as incipient pans, are found
along the east end of the fringing salt marsh at Highway 37. Small but well-differentiated semi-circular to
semi-linear salt marsh pans occur in peaty coastal salt
marshes at Limatour Spit, Point Reyes; Bolinas Lagoon;
Morro Bay; Elkhorn Slough; and along Tomales Bay.
Morro Bay, Bodega Bay, and Bolinas Lagoon also have
elongate shallow salt marsh pans fringing alluvial deposits. Most of these salt marsh pans are brackish in winter
and spring, but become moderately hypersaline (usually
40-60 ppt, rarely > 90 ppt) in summer (Baye, unpub.
data) when inundated.
Historic and Modern Distribution
The historic (pre-1860) location of natural salt ponds
within San Francisco Bay was probably restricted to the
Alameda shoreline in the vicinity of San Lorenzo Creek
(between the historic Thompsons’s Landing and Union
City Creek). This area included an extensive complex of
both connected and isolated large ponds in a matrix of
salt marsh. The complex was labelled as “ Crystal Salt
Pond” on the 1856 U.S. Coast Survey T-chart of the
area. The San Francisco Estuary Institute estimates the
acreage of Crystal Salt Pond to be approximately 1660
46
Baylands Ecosystem Species and Community Profiles
acres, based on the precise pond outline represented on
the 1856 T-chart (R. Grossinger, personal communication). If, however, the pond size fluctuated seasonally (as
expected from winter rainfall and tidal flooding), the
ponded area may have been several thousand acres from
late fall to spring. Two smaller ponds with similar configuration occurred north of San Lorenzo Creek, and
were clearly associated with sandy barrier beach deposits at the bayward edge of the marsh. (It is not clear
whether these northern satellite ponds produced high
concentration brine and halite, or were merely intermittently hypersaline lagoons). Crystal salt pond was used
as a salt source by aboriginal inhabitants of the Alameda
shoreline, and was exploited by early Mexican, Spanish
and U.S. settlers (Ver Planck 1951, 1958). Early descriptions of Crystal Salt Pond indicate that it contained a persistent crust of crystalline salt up to eight
inches thick, and the brines and salt contained “ impurities” of concentrated non-sodium salts (“ bittern”
salts, principally magnesium chloride and sulfate; Ver
Planck 1958).
The natural halite deposits of Crystal Salt Pond
were exhausted rapidly by the infant salt collecting industry; by 1860 they were largely depleted. Artificial enhancement of solar evaporation of brines was initiated
around 1853, when salt harvesters (farmers who used salt
for tanning leather and curing meats, and expanded into
the salt industry) began manual construction of low
berms around natural salt ponds to enhance their capacity to retain saline floodwaters and capture and precipitate their salt loads. These artificially enhanced natural
salt ponds became the nucleus of the solar salt industry.
By the end of the 19th century, the salt ponds of
San Francisco Bay were still confined to the northern
portion of the Alameda shoreline, from San Leandro
Creek to Alvarado (Union City). They did not comprise
a salt pond “ system,” but were an aggregation of many
independently owned and operated enterprises. Extensive conversion of salt marsh to salt ponds in south San
Francisco Bay did not occur until the 20th century. This
was facilitated by the consolidation of almost all the independent salt operations to a few (dominated by Leslie
Salt Company) in the 1930s. Permit requests to the
Corps of Engineers to dam numerous sloughs and
marshes in the South Bay were not filed until the early
1920s. Actual levee construction would have taken at
least several years, and new ponds take about 5 - 7 years
to “ seal” (become impermeable after gypsum and carbonate precipitation; Ver Planck 1958, Dobkin and Anderson 1994); therefore, the 1920s ponds were probably not
fully functional salterns until around 1930. The last
extensive marshes in the Alviso and Sunnyvale areas were
not diked for conversion to salt ponds until the early
1950s (Pacific Aerial Photo archives). Bair Island was not
converted to salt pond until the 1950s, although it had
previously been diked for agricultural use. The modern
Conservation Issues
Exotic Species – Salt pond microbial taxa are widespread geographically, but narrowly distributed ecologically. They are probably subject to dispersal by waterfowl and marine transport. There are no currently
recognized exotic species “ threats” to salt ponds as there
are with vascular plants in salt marshes.
Restoration – The crude technology for creating
artificial salt ponds (levee construction, wind-driven
pumps, tidegates) has been well developed for over a century. There is little doubt that complete artificial salt
pond systems can be created and maintained at a wide
range of sizes, from as little as 20 - 50 acre historic “ family size” or one-man operations (Ver Planck 1958), to
the modern systems in the tens of thousands of acres.
Low-salinity “ intake” ponds can also be maintained independently, in the absence of a salt-producing system,
by balancing influx of Bay water, residence time and redischarge at near-marine salinity. No new salt ponds have
been constructed since the 1950s, although ponds have
been interconverted from one type to another since then
(evaporator ponds to bittern disposal/“ storage” ). Small
and autonomous salt pond systems could be modified
to be less “ productive” of salt, and more biologically “ productive,” by reducing the efficiency of brine and salt production. This could be achieved by increasing the flux
in intake ponds, and reducing the residence time of
brines in each pond transfer. In winter, when brines are
diluted by rainwater, they could also be re-mixed with
intake Bay water and redischarged to the Bay at nearmarine salinities.
There have been recent tidal marsh restoration designs for artificial but naturalistic ponds and pans, but
no marsh restoration designs have included equivalents
of salt ponds. In principle, naturalistic salt ponds could
be artificially created and naturally maintained by replicating the hypothetical historic conditions of Crystal Salt
Pond (as inferred by Atwater 1979). This would entail
deposition of coarse sediments (sand or shell hash) at the
edge of a high-energy marsh shoreline, to be reworked
as beach ridges which restrict marsh drainage. In theory,
beach ridges would maintain form and size as they retreat with the eroding marsh edge, given ample sediment
supply and overwash processes. Under less natural geomorphic settings for salt ponds, artificial naturalistic salt
ponds could be created by constructing low, broad berms
made of bay mud or sand that would be set at elevations
enabling highest spring tides to overtop them. Low, wide
berms would be less prone to gullying and breaching
than steep levees, but would require some degree of
maintenance. Maintenance would be minimized by setting salt pond levees within restored marshes which
would shelter them from wave erosion of the open Bay.
Restored naturalistic salt ponds would undergo extreme
variation of salinity within and between years, depending on rainfall variation, evaporation conditions, and
storm surges.
Sea Level Change and Levee Maintenance – The
modern salt pond levee system requires periodic maintenance, and levees bordering the open Bay (not sheltered by fringing salt marsh) require frequent maintenance, armoring, or both. The need for levee maintenance
(topping with fresh dredged sediment) is likely to become
more frequent if storm frequency increases or sea level
rises, as would be expected with global warming (Moffatt
and Nichol and WRA 1988). Borrow pits along the interior side of salt pond levees become depleted over time,
and some old borrow ditches have been widened so
much that dredges need to re-handle material to bring
it within reach of levees. Dredging tidal marshes as an
alternative source of sediment is unlikely, since it
causes conflicts with endangered species habitat.
Therefore, sea level rise is likely to cause long-term
increases in costs and risk of levee failure of the existing salt pond system. Sea level rise could also make
naturalistic salt pond restoration more difficult, since
beach ridges or low levees are more likely to breach
and allow excessive (though restricted) tidal exchange
to impounded areas.
Chapter 1 —
Plant Communities
47
Plants
extent of salt ponds in the southern reaches of South Bay,
therefore, is relatively recent compared with the northern Alameda salt ponds. The Napa salt ponds are even
more recent: the diked Baylands of the Napa marshes
were converted from derelict agriculture (seasonal subsaline to brackish wetlands) to salt ponds between 1953
and 1959. Salt production ceased there in the mid1990s, but most of the system remains hypersaline.
Relative change – The minimum acreage of true
natural salt pond in San Francisco Bay was less than
2,000 acres (SFEI 1998); the maximum acreage (assuming seasonal expansion of Crystal Salt Pond by flooding,
and assuming that northern satellite ponds were brine/
halite ponds) could have been on the order of 3,000 4,000 acres. Other marsh pan habitats were not likely
to support persistent hypersaline algal communities and
were presumably dominated by marine-related macroalgae or Ruppia, as are most salt marsh pans today. However, if a significant proportion of the historic extensive
elongate lake-size marsh ponds fringing uplands (Redwood City to Palo Alto, and in the Newark vicinity) were
seasonally or perennially hypersaline, the total acreage of salt pond habitat could have been on the order
of 5,000 - 10,000 acres. There is very weak indication that elongate upland-fringing salt marsh pans ever
contained persistent strong brines supporting the narrow hypersaline algal/bacterial community, however.
Today, approximately 9,500 acres of derelict salt
ponds remain in San Pablo Bay, and over 29,000 acres
of artificial salt pond are actively maintained in San
Francisco Bay.
Plants
Conclusions and Recommendations
The commercial salt pond operations of San Francisco
Bay are unlikely to continue indefinitely because of economic changes in the Bay region and in the salt industry, and due to physical changes in the levee and borrow ditch system. Salt ponds are not likely to regenerate
spontaneously as a result of natural geomorphic processes
when salt marshes are restored. Other more common
types of pans and ponds are unlikely to establish in young
salt marshes; they are mature marsh features, associated
with well-differentiated marsh topography. The environmental setting associated with salt ponds has been radically altered; the combination of steep and relatively
high-energy Bay shorelines, coarse sediment supply, and
extensive high salt marsh with impeded tidal drainage
no longer exists. It is also likely that the feasibility of
maintaining the erosion-prone levee system of the artificial salt ponds will decrease over time, as borrow ditches
(sources of mud for levee repair) are depleted. Therefore,
new and artificial measures will be required to conserve
at least historic amounts of salt pond habitats within the
Estuary in the long term. The highest priority setting
for salt pond restoration of some type would be on the
Alameda County shoreline, from approximately San
Leandro to the Dumbarton Bridge, where the Bay shoreline profile and wave fetch may be conducive for formation of beach ridges (marsh berms), given appropriate
sediment size and supply.
There is no minimal ecosystem size for salt ponds.
The basic grazer food chain between Dunaliella and
Artemia can be maintained in extremely small systems.
However, the full microbial diversity of San Francisco
Bay salt ponds, which has not been analyzed in detail,
would probably not persist in small ponds. Also, since
the stability of natural salt ponds is inherently low (subject to ordinary natural fluctuations as well as catastrophic changes), microbial diversity would be better
conserved with a large system of semi-independent salt
ponds. Pre-historic salt pond acreage was probably on
the order of 2,000 acres. Aiming at this minimal acreage, in the absence of any experience at restoration of
naturalistic salt ponds or “ alternative” management of
solar salterns, would probably be insufficient to conserve
a diverse halophilic microflora.
We therefore recommend that long-term conservation of salt ponds entail the following actions:
1. Pilot projects should be undertaken that incorporate naturalistic salt pond designs as integral
components of large-scale tidal marsh restoration
on the northern Alameda shoreline;
2. Some existing salt ponds should be divided into
smaller, autonomous units away from the open
48
Baylands Ecosystem Species and Community Profiles
bay, preferably nested in the landward reaches of
restored salt marsh areas, and managed to maintain
intermediate strength brines rather than salt
production;
3. Salt pond restoration and alternative management
should aim for temporally variable as well as
spatially variable salinity and brine depths;
4. Both artificial and naturalistic salt pond restoration
should aim for designs which minimize maintenance requirements; and
5. An initial target acreage for salt ponds should
reflect the uncertainty of restoring sustainable salt
pond environments after commercial salt production ceases. We suggest that an initial target of
approximately 10,000 acres (equivalent to late 19th
century acreage) be stipulated and modified based
on the results of salt pond restoration and alternative pond management.
R efer
ences
eferences
Atwater, B., S.G. Conard, J.N. Dowden, C.W. Hedel,
R. L. MacDonald and W. Savage. 1979. History,
landforms, and vegetation of the Estuary’s tidal
marshes. In: T.J. Conomos (ed). San Francisco Bay:
The Urbanized Estuary. American Association for
the Advancement of Science, Pacific Division, Proc.
of the 58th Annual Meeting.
Brock, T.D. 1975. Salinity and ecology of Dunaliella from
Great Salt Lake. J. General Microbiol. 89:285-282.
Dobkin, M. and R.B. Anderson 1994. Oliver Bros. Salt
Co. Alameda County, California: Historic Resource
Evaluation Report. Contract 04F828-EP, Task Order #1, ALA-2, P.M. R0.0/R6.4, EA # 003050,
prepared for Calif. Dept. Transportation, District
4 (Oakland).
Javor, B. 1989. Hypersaline Environments. SpringerVerlag.
Moffatt and Nichol and Wetlands Research Associates.
1988. Future sea level rise: predictions and implications for San Francisco Bay. Report prepared for
BCDC (San Francisco Bay Conservation and Development Commission).
San Francisco Estuary Institute (SFEI). 1998. Draft spatial analysis of the Baylands ecosystem. San Francisco Estuary Institute, Richmond, Calif.
Ver Planck, W.E. 1951. Salines in the Bay Area. In: Geologic Guidebook of the San Francisco Bay Counties, Bulletin 154, State of Calif. Dept. Natural
Resources, Division of Mines.
______. 1958. Salt In California. Bulletin 175, State of
Calif. Dept. Natural Resources, Division of Mines.
168 pp.
Glen Holstein
Introduction
The San Francisco Bay estuary wetlands ecosystem historically included vegetated and non-vegetated areas.
Dominant among physical factors influencing estuarine
vegetation was the semi-diurnal tidal cycle. As a consequence, vegetation exposed to tides differed dramatically
from plant communities that existed above the tides. For
non-estuarine vegetation diurnal factors were relatively
insignificant; annual climate cycles and non-cyclic geological factors were the dominant influences. Substrates
in vegetated parts of the Estuary consisted almost entirely
of Bay mud (Louderback 1951, Wahrhaftig et al. 1993).
Beyond it they were much more heterogenous.
Environmental Setting
Vegetation increases in structural diversity and species
richness beyond the estuarine ecosystem boundary in a
complex pattern caused by interactions between the
physical factors of climate, geology, and hydrology.
Climate – The San Francisco Bay Area, like all the
California Floristic Province (Hickman 1993), has a climate characterized by wet winters and dry summers.
Such climates are called “ Mediterranean” because similar climatic conditions occur in the Mediterranean Basin, but the San Francisco Bay Area’s Mediterranean climate is more extreme than much of its namesake since
it rarely receives any significant rainfall during the years’s
warmest five months (Wernstedt 1972). Despite ample
water, plant growth is retarded in winter by low temperatures and short days. Growth is maximal in spring when
temperature and day length significantly increase and
reserves of soil water from winter rains are still abundant
(Walter 1979).
The diversity of the San Francisco Bay Area climate
is explained, to a great extent, by variation in two factors; winter precipitation and summer marine air flow.
Both cause local climates to be relatively mesic, but their
maxima rarely coincide and do not identically affect vegetation. High winter precipitation makes abundant soil
moisture reserves available for rapid spring and early
summer plant growth where low temperatures and fog
brought by marine air flow do not limit it. Since rapid
plant growth increases biomass, high biomass vegetation
types like redwood and mixed evergreen forests are frequently dominant in the Bay Area where rainfall is highest. A popular myth contends redwoods (Seqouoia
sempervirens) require summer fog. What they actually
require (and are limited to) are places with high precipi-
Chapter 1 —
Plant Communities
49
Plants
Plant Communities Ecotonal
to the Baylands
tation that are protected from summer marine air flow
and fog. By leaching mineral nutrients from surface soils,
high rainfall also retards growth of herbaceous vegetation that could otherwise compete with forest tree seedlings (Holstein 1984a).
Bay Area mean annual precipitation varies from 13
inches at San Jose and Antioch to 47 inches at Kentfield
in Marin County (Felton 1965). Not surprisingly, relatively undisturbed upland vegetation consists of redwood
and mixed evergreen forests near Kentfield (Shuford
1993) and of grassland near Antioch and San Jose
(Critchfield 1971). The Bay Area receives its precipitation from cyclonic storms with predominantly southwest
winds arriving from the Pacific Ocean. Consequently
stations with large mountains to the southwest lie in rain
shadows with reduced precipitation. Antioch, for example, is in the lee of Mount Diablo, and San Jose is in
the lee of the Santa Cruz Mountains. Kentfield, paradoxically, is also in the lee of a mountain, Mt. Tamalpais,
but is close enough to receive an increase in rainfall
caused by its orographic lifting. In most of the bay area,
however, mean annual rainfall is between 15 and 25
inches (Gilliam 1962). Within this range, vegetation is
controlled more by geologic substrate and slope exposure
than relatively minor local differences in mean annual
rainfall (Critchfield 1971).
In the San Francisco Bay Area, fog and associated
marine air chilled by offshore upwelling reduce summer
evapotranspiration and cause local climates to be mesic,
where summer marine air flow is strongest and fogs most
frequent. Such conditions reduce plant growth and resultant biomass, however, since they limit light and
warmth. High biomass forest vegetation also seldom occurs in areas directly exposed to salt-laden winds associated with marine air flow (Holstein 1984a).
Summer water stress causes incomplete cover and
much bare ground in most Mediterranean climate vegetation. In parts of the San Francisco Bay Area, however,
marine air flow and fog mitigate summer drought sufficiently for occurrence of vegetation types like coastal
scrub and prairie characterized by very complete cover
and little bare ground despite relatively low biomass
(Holstein 1984a).
Summer marine air flow and fog arrive at the Pacific Coast predominantly from the northwest because
of anticyclonic origins, but a shallow semi-permanent
temperature inversion confines their movement into and
through the San Francisco Bay estuary to just a few low
altitude gaps in the Coast Range. By far, the most important of these is the Golden Gate (Gilliam 1962).
Since marine air flow and fog suppress summer
temperatures, mean July temperature is a reliable indicator of their relative presence or absence in the San
Francisco Bay Area. Not surprisingly, San Francisco’s
July mean of 58.8° F is the lowest around the Estuary
because of its location at the Golden Gate. Antioch’s July
Plants
mean of 74.0° F is the highest of any Estuary station
since the low hills of northern Contra Costa County
protect it from marine air flowing into the Central Valley through the Carquinez Strait. Fairfield’s July mean
of 72.1° F indicates more direct exposure to that air flow
despite its more inland location. Mount Diablo State
Park’s entrance station has a July mean of 74.3°F because
of its location above the inversion that limits marine air
to low elevations. Most Estuary stations have July means
in the sixties, but a difference of just a few degrees within
that range can profoundly effect summer climate. Berkeley (61.5° F), Richmond (62.0° F), and Oakland
(62.4° F) have the lowest summer temperatures in the
Estuary next to San Francisco because of their location
directly east of the Golden Gate. Burlingame (62.3° F)
and the San Francisco Airport (62.7° F) are also relatively
low because of their location at the east end of the San
Bruno coast range gap (Gilliam 1962). Kentfield
(65.9° F), in contrast, is relatively warm in July because
Mount Tamalpais protects it from summer fog as well
as inducing its high winter rainfall. Distance from the
Pacific Coast is generally a poor predictor of summer
marine air flow. Redwood City (67.9° F) on the west side
of the Bay, for example, is warmer than Newark (64.9°F)
on the east side since the latter is more directly exposed
to air flow through the Golden Gate (Felton 1965,
Gilliam 1962). Coastal scrub and coastal prairie, the
vegetation types most associated with summer fog, are
common on the outer Pacific Coast but relatively scarce
in the San Francisco Bay Estuary because the parts of it
most exposed to summer fog were also those settled earliest and urbanized most completely (Hoover et al. 1966,
Donley et al. 1979).
Geology – Holocene alluvium characterized by
abundant clay and level topography surrounds slightly
over half the Estuary and is consequently the most abundant geologic substrate beneath its adjacent non-tidal
vegetation (Jennings 1977, Wahrhaftig et al. 1993). Bay
Area uplands underlain by alluvium were farmed early
and are now largely urbanized, but historic accounts and
relict stands indicate open grassland was their overwhelmingly dominant vegetation type before settlement
(Bryant 1848, McKelvey 1955). An exception was a few
oak savannas where widely spaced valley oaks (Quercus
lobata) occurred in a grassland matrix. Such savannas
were most frequent around the northern part of the
Estuary where rainfall was relatively high, but even there
they were most frequent in areas protected from summer marine air flow.
A specialized feature of California Holocene and
older alluvium with level topography is vernal pools,
small closed basins that fill in winter and dry during
spring. They support a characteristic specialized flora rich
in annual forbs (Holland and Jain 1977). Vernal pools
were long thought to result from gopher activity
(Dalquest and Scheffer 1942), but are better explained
50
Baylands Ecosystem Species and Community Profiles
as microtopographic patterns arising from ground shaking during earthquakes (Berg 1990) or interaction of localized soil processes and wind erosion (Abbott 1984).
Non-alluvial uplands around the Estuary consist of
uplifted hills underlain by a variety of pre-Holocene sedimentary and volcanic rocks. These include the Mesozoic
Franciscan formation and Great Valley Beds; Cenozoic
sediments consisting of Paleocene, Eocene, Miocene,
Pliocene, and Pleistocene marine beds and Pliocene nonmarine deposits; and the Pliocene Sonoma volcanic deposits (Jennings 1977, Norris and Webb 1990). The influence of these rocks on vegetation is most frequently
controlled by their clay content. Those with abundant
clay like Paleocene, Eocene, Miocene, and Pliocene sediments weather to deep soils much like those on Holocene
alluvium and predominantly support similar grassland
vegetation. The Mesozoic deposits include areas where
clay is abundant and others where it is scarce. As on other
clay-rich substrates, deep soils and grasslands dominate
the former in contrast to the thin soils and woody vegetation types predominant where clay is scarce. The
Franciscan Formation, a melange of soft clay sediments
and hard metamorphic rocks, has a particularly complex
vegetation pattern since grass dominates the former and
trees the latter. Pleistocene marine beds and the Sonoma
volcanics are relatively clay poor and consequently largely
support woody vegetation types like oak woodland and
mixed evergreen forest (Ellen and Wentworth 1995,
Critchfield 1971).
Grass is dominant on clay soils because they have
a relatively high water holding capacity (Walter 1979).
West of Cordelia in Solano County, for example, DibbleLos Osos and Hambright loams occur on adjacent hills
in the same climate. Dibble-Los Osos soils develop on
clay-rich Eocene marine sediments and consequently
have B2t horizons containing accumulated clay and a
water-holding capacity of 5 to 7 inches. Hambright soils,
in contrast, develop on Sonoma volcanics, lack a B2t horizon, and have a water holding capacity of only 2 to 3.5
inches (Bates 1977). Despite identical precipitation,
Dibble-Los Osos soils support grassland and Hambright
soils support oak woodland dominated by coast live oak
(Quercus agrifolia) because the former’s B2t retains soil
water that can be used by the shallow fibrous root systems of grasses. Since Hambright soils retain much less
water, the excess infiltrates to the fractured rock below
where it can be utilized by deep roots of trees but not
grasses. In May, evidence of the Dibble-Los Osos B2t’s
water retention capacity is plain in the hills above
Cordelia since grass stays green there several weeks longer
on Dibble-Los Osos soils than it does on the Hambright
despite the frequent shade of oaks. This phenomenon
illustrates that two very different vegetation types can be
equally “ mesic” and that oak woodland and grassland are
competitive enough within this region for slight soil differences to shift dominance from one to the other.
types. Numerous other local hydrological features around
the San Francisco Bay Estuary like springs, seeps, and
shallow water tables are associated with distinctive local
vegetation types. The relatively shallow water table under most valley oak savannas is a notable example.
Ecotonal Plant Communities
Plant communities surrounding the Baylands ecosystem
are here classified using the system of Holland and Keil
(1995). At present the most widely used and influential
classifications of California vegetation are derived from
Munz and Keck’s (1959) mixed system, which includes
taxonomic, physiognomic, and ecological information.
Barbour and Major’s extensive (1977) review of California vegetation, for example, was organized around a
slightly modified and expanded version of Munz and
Keck’s system. The units of their classification were
vegetationally ill-defined, however, since the plant species lists provided for each one lacked even estimates of
relative dominance. Some very important plant communities like riparian forests were also missing from both
Munz and Keck’s system and Barbour and Major’s subsequent review. It is doubtful Munz and Keck intended
their brief plant community synopsis to so profoundly
influence California vegetation science, however, since
the primary purpose of their book was clearly floristic.
Its success at remaining California’s floristic standard for
decades undoubtedly strongly contributed to the influence of its community classification.
Sawyer and Keeler-Wolf (1995) have recently tried
to overcome the Munz and Keck system’s problems by
developing a comprehensive alternative that excludes
ecological information from community definitions except in the case of certain specialized habitats like vernal pools. The Sawyer and Keeler-Wolf system presents
its own new problems, however, since it lumps quite different stable and successional communities when they
are dominated by the same species. Excluding most ecological information also causes very different coastal and
alpine communities sharing only a generic relationship
between their dominant species to be lumped into catchall groupings like “ Sedge series.” Many local dominance
types present in California’s complex vegetation are also
missing from their system despite its numerous series and
apparent comprehensiveness.
Holland and Keil avoid these problems by greatly
increasing the comprehensiveness and consistency of
Munz and Keck’s limited but fundamentally sound system. The result is a system outstanding for simplicity,
ease of use, and realistic description and classification of
California vegetation. Beginning with coastal sand dune
vegetation and concluding with freshwater vegetation
and anthropogenic environments, the plant community
descriptions below follow the system developed by Holland and Keil.
Chapter 1 —
Plant Communities
51
Plants
The geologic factor that most influences vegetation
around the San Francisco Bay Estuary is the physical effect of clay on soil water holding capacity, but chemical
effects are also locally important. Serpentinite, associated
with the Franciscan Formation and occurring at the
Estuary’s edge in Marin and San Francisco counties, is
so chemically distinctive because of its high Mg/Ca ratio and frequent heavy metals that it supports unique
vegetation types and many endemic plant species (Kruckeberg 1984). Soils beyond the limits of tides are also
usually much less saline than those under tidal influence,
but salts can locally accumulate to high levels in nontidal areas where drainage is poor. Salt especially accumulates in non-tidal areas where precipitation is low,
relief is subdued, and Cretaceous Great Valley beds provide a salt source (Chapman 1960; Johnson et al. 1993;
Harris 1991). Geology also strongly affects microclimate
wherever hills have been uplifted since their south slopes
receive more sunlight, warmth, and resultant evapotranspiration than their north slopes. Vegetation on Bay Area
hills is consequently relatively xeric on south slopes and
relatively mesic on north slopes (Bakker 1984).
Hydrology – The influence of geology and climate
on soil water is discussed above. Streams also tend to
increase in frequency and flow duration as rainfall increases. Since they provide water to plants in greater
quantities and different seasons than local climates, they
support distinctive riparian vegetation types not found
in upland areas. Not surprisingly, riparian and upland
vegetation become increasingly distinct as rainfall decreases (Holstein 1984b). Streams and their associated
riparian vegetation are usually narrowly linear landscape
features, but they can broaden dramatically when streams
reach base level and form deltas. A broad willow-composite zone now removed by urbanization that reportedly once occurred around the southern edge of San
Francisco Bay (Cooper 1926) undoubtedly represented
covergent deltaic riparian vegetation of several creeks that
flow into the Bay.
Alluvium in streambeds tends to be coarser and
thus better aerated than interfluvial alluvium, and the
running water of streams is also relatively well-aerated.
Streamsides consequently provide suitable environments
for roots of woody riparian vegetation. In freshwater
marshes, however, standing water in poorly drained
interfluvial areas quickly causes anaerobic reducing conditions to develop at such shallow depths that only herbaceous vegetation with shallow, predominantly fibrous
root systems can occur. The herbaceous freshwater
marsh vegetation is consequently quite distinct from
predominantly woody riparian vegetation (Holstein
1984a).
Freshwater marsh vegetation grades into vernal
pool vegetation through a series of transitional seasonal
marsh vegetation types and into moist grassland through
a transitional series of lowland wet meadow and swale
Plants
1. Coastal Sand Dune Vegetation – Sand is a distinctive substrate for plants since water infiltrates it very
rapidly leaving little moisture available for plants with
shallow root systems (Walter 1979). Sand differs from
other substrates like fractured rock which have similarly
high infiltration rates, however, because of sand’s high
subsurface homogeneity and lack of resistance to root
penetration. Large sand deposits are characteristic landscape features of coasts and arid areas. In Holland and
Keil’s (1995) system, followed here, vegetation on sand
deposits of arid areas is classified as desert sand dune vegetation and consequently distinguished from vegetation
on coastal sands. In California, however, some dune
fields are located in areas neither coastal nor truly arid.
Examples occur on the Merced River alluvial fan in
Merced County and at Antioch in Contra Costa County
(Wahrhaftig et al. 1993). The former was produced by
outwash from glacial erosion of granite in the Yosemite
Valley (Wahrhaftig and Birman 1965), but extensive
Eocene to Pliocene sandstone deposits in nearby hills
(Ellen and Wentworth 1995) are a likely source for the
latter. Neither the Merced or Antioch dunes are discussed by Holland and Keil, but both occur in semi-arid
areas and share more floristic features with their desert
sand dune vegetation type than their coastal sand dune
vegetation types.
Pioneer dune vegetation occurs where significant
aeolian movement of sand limits development of stable
soil and vegetation.
Ambrosia chamissonis is its characteristic dominant,
and Abronia latifolia, Achillea millefolium, Atriplex
californica, Atriplex leucophylla, Calystegia soldanella,
Camissonia cheiranthifolia, Lathyrus littoralis, Leymus
mollis, and Lupinus chamissonis are frequent associated
species (Barbour and Johnson 1977).
Dune scrub occurs where stable soil and vegetation
have developed on sand of dunes usually considerably
older than those supporting pioneer dune vegetation.
Ericameria ericoides is the characteristic dominant of
dune scrub, and associated species include Artemisia
californica, Baccharis pilularis, Lotus scoparius, Lupinus
arboreus and Lupinus chamissonis (Barbour and Johnson
1977).
Sand is relatively rare around the San Francisco Bay
estuary, but a significant deposit, the Pleistocene Merritt
sand, is present at Alameda and adjacent parts of Oakland (Radbruch 1957). Since the local climate is marine,
some areas with surface deposits of Merritt sand probably once supported pioneer dune and dune scrub
communities similar to those now occurring along the
outer Pacific Coast. The sandy area at Alameda and Oakland was one of the first places along the Bay to urbanize, however, and any dune vegetation present there was
consequently eradicated before it could be described. A
modern analogue with similar soils and climate is
Elkhorn Slough (Monterey County), which is incised
52
Baylands Ecosystem Species and Community Profiles
into Pleistocene deposits, the Aromas sand. Agricultural
development has removed some natural vegetation
around Elkhorn Slough, but remaining relict stands are
still numerous. Topographic features recorded prior to
development of the port of Oakland and Lake Merritt
resemble those along Elkhorn Slough (Wahrhaftig and
Birman 1965).
At Antioch, a sandy area is also present immediately east of Broad Slough. It is less urban than Alameda,
but most of its dune vegetation was lost to sand mining
prior to urbanization. A small protected remnant of such
vegetation at Antioch supports several state and federally listed rare animal and plant species (Sawyer and
Keeler-Wolf 1995). Antioch is significantly hotter and
drier than the outer coast, and its sand probably originated from nearby sand deposits that extend southward
along the inner Coast Range. The affinity of its distinctive sand dune flora and vegetation is consequently closer
to Holland and Keil’s (1995) desert sand dune community than to either of his coastal dune communities.
Because a rain shadow occurs along the inner Coast
Range, the ranges of several plant and animal species
with desert affinities, including the relatively well-known
San Joaquin kit fox (Vulpes macrotis ssp. mutica) (Zeiner
et al. 1990), extend north along the western San Joaquin
Valley to near Antioch.
2. Coastal Scrub – Coastal scrub refers to communities dominated by small shrubs in non-desert areas of California. Coastal scrub typically develops on soil
and friable sediments rather than conglomerate or fractured hard rock and consists of shrubs with relatively
shallow root systems.
Northern coastal scrub is a dense shrub-dominated
community which most frequently occurs on steep slopes
receiving strong prevailing onshore winds and at least 20
inches of precipitation, but can also occur as an ecotone
between northern oak woodland and southern oak woodland on slopes with less wind. Most typically, however,
it occurs where precipitation and soils are adequate for
development of forests, but tree growth is prevented by
strong onshore winds. Since moisture is not limiting,
cover is typically complete (Heady et al. 1977).
Baccharis pilularis is the characteristic dominant,
but Mimulus aurantiacus, Rhamnus californica, and Toxicodendron diversilobum can also occasionally be locally
dominant. Characteristic understory species include
Achillea millefolium, Anaphalis margaritacea, Eriophyllum
staechadifolium, Gaultheria shallon, Heracleum lanatum,
Polystichum munitum, Pteridium aquilinum, Rubus
ursinus and Scrophularia californica. Northern coastal
scrub is most common along the outer Pacific Coast but
also occurs at suitable sites around the San Francisco Bay
Estuary near the Golden Gate, in the Berkeley Hills, and
in San Mateo County. Baccharis pilularis frequently invades disturbed grasslands and forms communities which
superficially resemble northern coastal scrub but lack
quent on steep slopes but can also occur with relatively
low relief on stone alluvial fans in valleys.
Holland and Keil (1995) subdivide California
chaparral into 11 subclasses, of which six occur in San
Francisco Bay counties. These are not separately treated
here, however, since relatively little chaparral of any kind
occurs in the Estuary’s immediate vicinity.
Chaparral is dominated by shrubs in the genera
Adenostoma, Arctostaphylos, Ceanothus, Cercocarpus, and
Quercus, which form a functional group characterized by
deep root systems adapted for extracting water from deep
cavities in fractured rock. The sclerophyllous leaves of
chaparral shrubs are adapted for maintaining low levels
of evapotranspiration and associated productivity during long growing seasons (Walter 1979, Mooney and
Miller 1985). Discussions of chaparral ecology have long
emphasized its adaptation to fire since its shrubs use a
variety of strategies to rapidly reoccupy burns and an associated functional group of annuals has seeds that remain dormant for decades and only germinate following chaparral fires (Biswell 1974). Extensive research on
Adenostoma-dominated chaparral suggesting a relatively
short fire cycle my not be directly applicable to other
chaparral types, however, since some other kinds of chaparral may have a much longer fire cycle (Keeley and
Keeley 1988).
The nearest extensive chaparral to the Estuary occurs in Marin County on the slopes of Mt. Tamalpais
two miles west of San Francisco Bay (Shuford 1993,
Wieslander and Jensen 1945). While chaparral on alluvial fans is rapidly disappearing but still fairly common
in parts of southern California (Smith 1980), it is virtually unknown in central and northern California. Cooper (1926), however, reported that chaparral that has
since been extirpated formerly occurred near the southern end of San Francisco Bay on Los Gatos Creek’s alluvial fan.
4. Grassland – Vegetation dominated by grasses
and graminoid sedges was widespread along the shores
of the San Francisco Bay Estuary prior to urban development and is still fairly common there (Bryant 1848,
McKelvey 1955). It occurs in non-wetlands wherever
soils with clay horizons thick enough to hold significant
water near the soil surface and to exclude air from deeper
horizons are directly exposed to solar radiation. Clay soils
are particularly favorable for grasses and other graminoids
because the near-surface water they hold is preferentially
available to the dense, relatively shallow fibrous root systems of such plants. In wet climates the most mesic conditions occur on soils with high clay content because of
their high water holding capacity, but in arid areas that
pattern is reversed. In deserts clay holds much water from
scarce precipitation near the soil surface, where solar
radiation quickly evaporates it (Walter 1979). Conditions intermediate between these extremes prevail in the
semiarid climate surrounding most of the Estuary. Clay
Chapter 1 —
Plant Communities
53
Plants
most of its characteristic species. Eventually such recently
invasive B. pilularis stands may develop into stable coastal
scrub or oak woodland communities (Heady et al. 1977).
A protected example of northern coastal scrub occurs
near the estuary at China Camp State Park.
Southern coastal scrub is a relatively open shrubdominated community occurring most frequently on
steep, dry slopes. It is commonest in areas receiving under 20 inches of precipitation but can occasionally occur in wetter areas on sunny south slopes. It typically
occurs where soils otherwise suitable for grassland are excessively drained because of steepness. Because water is
the primary limiting factor in southern coastal scrub, its
dominant shrubs tend to be widely spaced, forming relatively incomplete cover. In spring, when water stress is
briefly relieved, a diverse annual forb flora develops in
interstices between the dominant shrubs (Mooney
1977).
Artemisia californica is the characteristic dominant,
and common associated species include Eriogonum nudum, Eriophyllum lanatum, Lotus scoparius, Lupinus
albifrons, Mimulus aurantiacus and Nassella pulchra.
Small stands of southern coastal sage scrub occur in hills
around the Estuary and are especially frequent east of
South San Fracisco Bay and south of Suisun Bay, where
precipitation is relatively low. An example occurs along
the Estuary shore at Point Richmond.
Sea-bluff coastal scrub occurs where persistent saltladen onshore winds suppress most other plant communities. Such climatic conditions resemble those in northern coastal scrub but are more extreme. In such sites the
only communities are sea-bluff coastal scrub and northern coastal grassland. The former tends to occur on rocky
sites with thin soils and the latter on deeper soils that
tend to be heavier, but both frequently intermix in a
complex mosaic (Holland and Keil 1995).
Eriophyllum stachaedifolium is the characteristic
dominant, and frequently associated species include Artemisia pycnocephala, Baccharis pilularis, Erigeron glaucus,
Eriogonum latifolium and Lessingia filaginifolia. Saltladen winds strong and persistent enough to support this
community enter the Estuary through the Golden Gate
but rapidly lose their intensely marine character as they
move inland. Havlik (1974) described small stands of this
community at Yerba Buena Island, Brooks Island, Red
Rock, Point Richmond, Point Fleming, and Potrero San
Pablo, all places directly exposed to marine winds entering San Francisco Bay through the Golden Gate.
3. Chaparral – Chaparral refers to a widespread
and characteristic California community dominated by
large shrubs with evergreen sclerophyllous leaves. It is
frequent in areas with precipitation between 10 and 20
inches per year and occasional in wetter areas on sunny
south slopes. Chaparral occurs where rocky soils with
little clay permit rapid infiltration of water and air to
relatively great depths. Such conditions are most fre-
Plants
soils are xeric and grass-covered on plains and south
slopes, where they are directly exposed to solar radiation,
but mesic and covered by forest and woodland on north
slopes, where solar radiation is reduced. Grassland is
most prevalent where annual precipitation is between 10
and 20 inches but becomes progressively scarcer as annual rainfall increases. Some grassland is usually present
even in very wet areas, however, wherever clay is directly
exposed to solar radiation. Soils with sufficient clay for
grassland predominate on the recent alluvium that forms
the floors of virtually all San Francisco Bay Area valleys;
they are also common on hillslopes where clay-rich sediments have been uplifted (Ellen and Wentworth 1995).
While direct solar radiation usually keeps grassland free
of woody plants on valley floors and south-facing hillslopes, similar grassland frequently dominates understories beneath the oak woodland that occurs on north
slopes because of less intense radiation (Holstein 1984a).
Native perennial grassland. Frequent relict stands and
clear descriptions by early travelers leave little doubt that
most native grassland near the Estuary on both valley
floors and hillslopes was dominated by a rhyzomatous
and largely sterile hybrid between Leymus triticoides and
L. condensatus (Stebbins and Walters 1949). Hybrids between these species have been called Leymus xmultiflorus,
but since the hybrid dominant around the Estuary is too
small to match descriptions of xmultiflorus (Hickman
1993), it is here included in L. triticoides. Two frequently
associated rhizomatous graminoids were Carex barbarae
and C. praegracilis, the latter being especially frequent
at upland-wetland ecotones. Nassella pulchra, a non-rhizomatous bunchgrass, has received more attention than
any other species as a native grassland dominant. It frequently dominated grassland but mostly did so only near
ecotones with coastal scrub and oak woodland where
heavy clay grassland soils had begun to thin and dissipate or where specific substrates like serpentinite prevented development of typical grassland soils (Bryant
1848, McKelvey 1955).
Native grassland had numerous local variations
ranging from topographic lows where soil water and clay
accumulated to topographic highs where clay was thin
and water scarce. Species indicating topographic lows
(locally called swales) included Juncus balticus, Juncus
xiphioides, Ranunculus californicus, and Sisyrinchium
bellum, while N. pulchra and a variety of forbs indicated
the highs. Along the Estuary shore at ecotones with tidal
marsh, Distichlis spicata, another rhizomatous grass, was
particularly prominent (Heady 1977, Holland and Keil
1995).
A scarce native grassland type especially significant
for its many rare plants occurs on salt-affected soils associated with inland basins and basin rims rather than
coastal tidelands (Faber 1997). These inland alkaline
grasslands share features like the prominence of Distichlis
spicata with the grassland-tidal marsh ecotone but often
54
Baylands Ecosystem Species and Community Profiles
differ from it in the presence of more bare ground and
many species not occurring at the Estuary shore. Cooper (1926) reported Hemizonia congesta and H. pungens
were formerly dominant on similar soils near the southern end of the Estuary that are now completely covered
by urban development. The best presently extant examples of alkaline grassland in the Estuary’s vicinity occur near Livermore in Alameda County and near
Fairfield in Solano County. Other distinctive grassland
types of unusual substrates supporting rare species are
serpentinite grassland and sandy soil grassland (Skinner and Pavlik 1994).
Native annual forbland. Wester (1981) presented
evidence that the southern San Joaquin Valley, an area
traditionally considered former grassland, was dominated
by annual forbs prior to European settlement. California vegetation classification has traditionally called all
upland vegetation dominated by herbs grassland, but
Wester’s work suggests much of the area traditionally
mapped as grassland (Kuchler 1964) was actually native
annual forbland. Since native annual forbland occurs
where rainfall is insufficient for most perennial grasses,
it consequently was most extensive far south of the Estuary in the southern San Joaquin Valley. Numerous
relict taxa suggest, however, that a narrow native annual
forbland corridor extended north from there to near the
Estuary shore in Contra Costa County because of a rain
shadow along the inner Coast Range’s eastern base.
Forbland elements also probably occurred even more
widely wherever local conditions like soil infertility and
trampling by megafauna suppressed otherwise ubiquitous perennial grasses. Even today wildflower displays
(i.e., annual forb dominance) are most spectacular locally
where soil is relatively infertile (i.e., Bear Valley in Colusa
County and Table Mountain in Butte County [Faber
1997]) and most spectacular generally in years, as in
1991, when winter drought suppression of competitive
grasses is followed by forb-promoting heavy spring rains.
Some forbland species like Eremocarpus setigerus have
adapted well to anthropogenic land use changes but others have become rare (Convolvulus simulans, Madia radiata) or extinct (Eschscholzia rhombipetala).
Non-native annual grassland. Introduction of grazing and agriculture during the nineteenth century caused
a dominance shift in almost all of California’s grasslands
from native perennial graminoids to Eurasian non-native annual grasses. Today dominance among such annuals changes spatially in a complex pattern reflecting
soil conditions. On catenas from thick, heavy clay soils
to thinner, lighter ones a typical annual grass dominance
sequence Lolium multiflorum-Bromus hordeaceus-Avena
fatua-Avena barbata replaces a simpler perennial sequence Leymus triticoides-Nassella pulchra still occasionally extant on the same catenas. Another common dominance
sequence
Bromus
hordeaceus-Bromus
diandrus-Hordeum murinum reflects shifts in soil nitro-
mountain ranges. Contrary to an enduring myth, redwoods are negatively rather than positively associated
with summer fog. Consequently, even at sites protected
from onshore winds they are almost completely absent
along the immediate coast wherever summer fog is frequent. Redwoods survive summer drought not because
of fog drip but by storing surplus water from high winter precipitation in their massive trunks, a strategy that
has produced only slightly less dramatic gigantism in
other conifer species where large winter water surpluses
occur with summer drought. As a consequence northern California and southern Chile, both areas with unusually wet winters and dry summers located at the outer
periphery of more typical Mediterranean climate zones,
are the world’s two greatest centers of tree gigantism
(Holstein 1984a, Zinke 1977).
Shade is so intense in the redwood forest understory that only a few plant species survive there. Two that
do, Oxalis oregana and Polystichum munitum, are usually the sole understory dominants.
6. Mixed Evergreen Forest –
Central California mixed evergreen forest. Forests
dominated by a mix of broadleaf and conifer evergreen
trees are frequent in California where precipitation is
relatively high and winter temperatures are mild. In
northern California the trees most frequently dominating such mixed evergreen forests are Arbutus menziesii,
Lithocarpus densiflorus, Pseudotsuga menziesii, and Umbellularia californica. In central California the term mixed
evergreen forest as presently used is somewhat anomalous, however, since it often designates forests solely
dominated by Umbellularia californica, the California
laurel. Such laurel-dominated forests are frequent around
the Estuary where annual rainfall is between 20 and 40
inches. At the dry end of that precipitation range laurel
forests are entirely confined to very shady north slopes
and canyons, but they also occur on somewhat sunnier
slopes as 40 inches is approached. Above that they are
almost entirely replaced by redwood forests (Sawyer et
al. 1977, Wainwright and Barbour 1984).
The most commonly associated tree species in central California’s laurel forests is a non-evergreen, Acer
macrophyllum. Arbutus menziesii is also a frequent associate but is almost entirely confined to the rockiest slopes.
Shade is so intense beneath laurel forest canopies that
completely bare ground is common where drought is an
added stressor, but as 40 inches is approached Polystichum munitum often dominates the understory. Holodiscus discolor, a deciduous species, is commonly dominant in shrubby openings frequent in laurel forests
(Safford 1995).
7. Oak Woodland – Vegetation with an overstory
dominated by oak trees is common throughout California’s
Mediterranean climate zone including the Estuary’s vicinity. Such oak woodlands primarily vary in species and
spacing of their overstory oaks. Vegetation is called sa-
Chapter 1 —
Plant Communities
55
Plants
gen content from low to relatively high. The above species are the most frequently dominant non-native annual
grasses, but others also occasionally participate. Cynosurus
echinatus, for example, frequently dominates annual
grassland where rainfall is relatively high. Several exotic
forbs are also becoming increasingly important components in a vast exotic herbaceous vegetation type that
may only temporarily be called grassland. Vicia villosa
ssp. varia is increasingly planted for forage in the Avena
zone; Picris echioides is important in the Lolium zone;
and Centaurea solstitialis, especially, is a widespread invader of the B. hordeaceus zone, where Erodium botrys is
also important when soil fertility is particularly low
(Heady 1977, Holland and Keil 1995). Grazing is particularly important for maintaining replacement of native perennial grasses with exotic annual species. At
numerous sites around the Estuary, for example, dominance is shifting back from exotic annual grasses to
Leymus triticoides and Carex barbarae where expanding
urbanization has at least temporarily caused the removal
of grazing.
Coastal prairie. Where clay soils are directly exposed to marine air flow, a floristically distinct grassland
occurs that Holland and Keil (1995) call northern coastal
grassland but is widely known in California as coastal
prairie. Coastal prairie is most frequent along the outer
coast, but small amounts also likely occur near the Estuary where marine air flow is particularly direct.
Much of California’s coastal prairie is now dominated by two exotic perennial grasses, Anthoxanthum
odoratum and Holcus lanatus, but many distinctive native perennial grasses like Agrostis pallens, Calamagrostis
nutkaensis, Danthonia californica, Deschampsia cespitosa,
Festuca idahoensis and Festuca rubra can also frequently
be locally dominant. Two other distinctive plant species
indicative of coastal prairie are Iris douglasiana and Juncus
patens (Heady et al. 1977).
5. Coastal Coniferous Forest – Forests dominated
by large coniferous trees occur along the eastern Pacific
Coast in a high rainfall zone extending from central California to Alaska. Holland and Keil (1995) recognize two
subdivisions of coastal coniferous forest, but only one of
these, redwood forest, occurs near the Estuary.
Redwood forest. Extensive forests dominated by
Sequoia sempervirens, the well-known redwood and the
world’s tallest tree species, occur on the southern slopes
of Mt. Tamalpais within 1.75 miles of the Estuary
(Shuford 1993), but individual redwoods occur in mixed
evergreen forest much less than a mile from the shore
of San Pablo Bay at China Camp State Park. Redwoods
are common up to about 2,000 feet in the California
Coast Range wherever annual precipitation is above 40
inches and soil is relatively fertile. Despite sufficient rainfall, sensitivity to cold prevents their occurrence along
the Oregon coast beyond a few miles north of the border, at high elevations in the Coast Range, or on inland
Plants
vannah where oaks are widely spaced and forest where
spacing is so close their canopies are closed. Woodland,
as a term, describes vegetation with intermediate spacing, but tree separation is so locally variable in
California’s oak-tree dominated vegetation it is appropriate to use the traditional term oak woodland to refer
to all of it. That generalized oak woodland can then be
divided into subclasses based on its dominant species
(Griffin 1977).
Since woodland oaks and grassland grasses occur
in similar environmental conditions, they frequently
compete directly for water and other soil resources. Specific aspects of that competition are discussed for each
subclass but a few of its consequences are general. Oaks
only occur where water is present in deep soil horizons,
where it may arrive horizontally through shallow aquifers or vertically when precipitation is abundant enough
to infiltrate past dense but relatively shallow grass root
systems. Grassland grasses, in contrast, only occur where
solar radiation is direct because overstory trees are either
absent or so widely spaced their canopies are not contiguous (Walter 1979).
Coast live oak woodland, which is dominated by
Quercus agrifolia, is distinctive among oak woodland subclasses because it consists almost exclusively of closed
canopy forests. As a consequence it is frequently treated
as a subclass of mixed evergreen forest rather than oak
woodland. It is included here with oak woodland, however, because of the affinities of both its dominant tree
and the majority of its fauna (Griffin 1977).
Coast live oak woodland occurs widely around the
Estuary where annual precipitation is between 15 and
40 inches and continentality is at least partially moderated by marine influences. Marine air flow through
Carquinez Strait even permits occurrence of coast live
oak woodland with two isolated Coast Range-related
mammal populations (Sylvilagus bachmani riparius and
Neotoma fuscipes riparia) on the Central Valley floor near
Lodi (Zeiner et al. 1990).
In hills on clay soils coast live oak woodland is frequently present as an extensive ecotone between grassland and mixed evergreen forest since it occurs on slopes
shadier than the former but sunnier than the latter. On
slopes where rockier substrates and lighter soils permit
infiltration of more water to greater depths, however,
coast live oak is less limited by solar exposure and can
even occur on south slopes. North of Carquinez Strait,
for example, adjacent ridges with identical microclimates
differ only in their substrates. Ridges underlain by sediments of the clay-rich Eocene Markley Formation are
covered by grassland and have coast live oak woodland
only on north slopes and in canyons, while those underlain by hard but fractured rocks of the Pliocene Sonoma
volcanics are covered by coast live oak woodland on all
exposures but north slopes and canyons, where Umbellularia-dominated mixed evergreen forest occurs. The
56
Baylands Ecosystem Species and Community Profiles
great vegetational difference is a result of the way in
which the two substrates respond to precipitation — rain
rapidly infiltrates to deep levels on the fractured volcanics
where it can be utilized by oak roots, whereas it is held
at the surface on the clay-rich Markley where it is more
available to grass roots. Rapid infiltration on the volcanics
causes such xeric conditions in its surface soils that its
few stands of annual grassland cease productivity and dry
two weeks earlier than Markley grasslands dominated by
the same species (Bates 1977, Ellen and Wentworth
1995).
Coast live oak woodland differs from other oak
woodland subclasses in the relative rarity of annual
grasses in its understory. The most frequent dominant
there is Toxicodendron diversilobum, poison oak, but
Rubus ursinus and Symphoricarpus mollis are also often
important (Safford 1995).
Valley oak woodland consists of several structurally diverse communities sharing dominance by Quercus
lobata that include savannah and woodland on clay hillslopes and savannah, woodland, and closed canopy forest on alluvial plains over shallow unconfined aquifers.
Alluvial valley oak woodland often occurs on the outer
edges of riparian forest corridors (see below) on relatively
fine, heavy soils distinct from the coarse alluvial soils
under typical riparian stream bank vegetation. Tree spacing in alluvial valley oak woodland is related to water
stress since canopies closed when subsurface water is
abundant become progressively more open as water stress
increases, resulting first in woodland and then savannah.
Much alluvial valley oak forest was removed because it
coincided with highly desired agricultural soils, but a few
stands are extant in the Central Valley and elsewhere.
Alluvial valley oak woodland was probably always scarce
near the Estuary, however, since it is better adapted to
inland Califonia’s hot summers than to the outer Coast
Range’s relatively marine climate. One of the few examples near the Estuary is located along Green Valley
Creek near Cordelia in Solano County.
Valley oak woodland is most frequent near the
Estuary on clay hillslopes with annual rainfall between
15 and 40 inches, where its range overlaps coast live oak
woodland and foothill woodland. It is less abundant than
either but more tolerant of clay soils than the former and
less resistant to water stress than the latter. The understory of valley oak woodland’s savannah and woodland
phases typically consists of non-native annual and occasionally native perennial grassland. Vegetation beneath
closed canopy valley oak alluvial forest, however, can include both grassland and features shared with riparian
forest or coast live oak forest understories. Valley oak
reproduction, often low because of competition with
annual grass and predation of seeds and seedlings by a
variety of herbivores, can be abundant in alluvial woodland when suppression of grass by flooding coincides
with large acorn crops. Urban fringes are also favorable
Dudleya, the California genus with the most highly
adapted chasmophytes, has a few taxa near the Estuary
but is much more diverse in Southern California.
Plants of rock outcrops are less specialized than
chasmophytes but may be rare since they occur in distinctive microenvironments that consequently are free
from competition with surrounding vegetation. Rock
outcrops and cliffs are most likely to support rare plants
when they are mineralogically different from surrounding landscapes, and one mineral receiving particular attention because of its frequent association with rare
plants is serpentine. Soil development is so retarded and
vegetation so distinctive on serpentine that its occurrences may be viewed as extended rock outcrops even
though they occasionally cover many square miles
(Bakker 1984, Skinner and Pavlik 1994, Fiedler and
Leidy 1987).
9. Riparian Vegetation – Riparian vegetation refers to the distinctive plant communities of streambanks
and ecologically related habitats. Its most salient environmental features are relatively coarse alluvial soils typically associated with streams and root zone water supplies greater than the local climate provides. When
mature, California riparian vegetation is closed canopy
forest, but early successional riparian vegetation can be
shrubby.
Near the Estuary riparian vegetation is overwhelmingly dominated by three species, Acer negundo, Salix
lasiolepis and Salix laevigata, but others may dominate
in specialized habitats. Populus fremontii and Salix gooddingii are important where climate becomes less marine
and more continental near the Central Valley; Salix
exigua is important on sandbars and other habitats where
early successional riparian vegetation is developing; Alnus
rhombifolia and Salix lucida ssp. lasiandra are important
where, as at Niles Canyon, streams with rocky beds flow
perennially; and Platanus occidentalis and Baccharis
viminea dominate where ones with sandy and rocky beds
flow intermittently.
Typical Acer negundo-Salix lasiolepis-Salix laevigata
riparian vegetation also is common where ecological conditions simulate streambank environments, as at lakeshores and a variety of places with shallow water tables.
On the outer coast non-streambank riparian vegetation
is frequent in dune slacks, but around the Estuary it at
least formerly was most frequent in sausals, microdeltas
occurring where stream channels and their subsurface
water tables spread laterally as they entered tidal marsh.
Most sausals have been lost to urbanization of the
Estuary’s periphery, but a small example occurs at China
Camp State Park.
Common riparian understory plants near the Estuary include Baccharis douglasii, Euthamia occidentalis,
Rosa californica, and Rubus ursinus. For a short distance
these can also replace riparian trees as dominants at the
ecotone with tidal marsh where a veneer of coarse stream-
Chapter 1 —
Plant Communities
57
Plants
sites for valley oak reproduction because their low livestock and wildlife populations result in lowered seed and
seedling predation (Holstein 1984b, Holland and Keil
1995).
Foothill woodland is woodland and savannah vegetation wholly or partially dominated by Quercus douglasii, blue oak, that is widespread on hillslopes surrounding the Central Valley. Near the Estuary, foothill
woodland is largely confined to the inner Coast Range.
The foothill woodland environment has a relatively continental climate with cool to cold winters, very hot summers, and annual rainfall from 15 to 40 inches. Winter
cold reduces understory grass growth and consequently
permits infiltration of a large part of the wet season’s
water surplus to deep subsoil where it can be utilized by
blue oaks during spring and summer. In summer high
temperatures and low humidity produce very low water
potentials in blue oak leaves that permit withdrawal of
water tightly held by clay-rich subsoils.
Blue oak is usually the sole foothill woodland dominant on clay hillslopes, but on slopes with more rock and
thinner soils it often shares dominance with Pinus
sabiniana. Blue oaks occur on a wider range of slope exposures than many other oak species, but foothill woodland dominance often shifts to Aesculus californicus on
shaded north slopes. In canyons and around rock outcrops Quercus wislizenii is also often a local dominant.
Because of foothill woodland’s open canopy its
understory is almost universally dominated by non-native annual grassland. Native forbs like Holocarpha
virgata, however, are also usually frequent there. Competition is particularly intense between annual grasses
and blue oak seedlings before they develop roots long
enough to reach subsoil water. Seedling mortality at this
stage is so intense that much foothill woodland consists
almost entirely of mature blue oaks that germinated in
the 1860’s, a decade when severe overgrazing reduced
much presumably native perennial grassland from
California’s rangelands. Subsequent increase of nonnative annual grassland has severely restricted reproduction of foothill woodland developing at that time (Griffin 1977, Holland and Keil 1995).
8. Cliffs and Rock Outcrops -Vegetation of cliffs
and rock outcrops is usually virtually ignored in surveys
of California vegetation including that of Holland and
Keil (1995) because its areal extent is small and it consists largely of non-vegetated surfaces. It is particularly
important, however, as a habitat for rare plant species.
Cliffs are unique environments where soil and competition with other plants is very limited and solar radiation is often abundant. Plants adapted to cliffs (chasmophytes) resemble epiphytes in producing small easily
dispersed seeds in such great numbers that the likelihood
of reaching rare suitable germination habitats is increased. Seeds reaching these light-rich habitats can afford to be small because they require little stored food.
Plants
side alluvium deposited on tidal mud thins as it nears
the Estuary. Like sausals, however, such riparian-tidal
marsh ecotones have almost entirely disappeared around
the Estuary because of urbanization (Holland and Keil
1995, Holstein 1984b).
10. Freshwater Vegetation – Freshwater wetland
vegetation occurs where land surfaces are saturated by
freshwater or shallowly covered by it. Its two main phases
near the Estuary, freshwater marshes and vernal pools,
are very distinctive but also united by intermediate communities.
Freshwater marsh refers to vegetation dominated
by plant species emergent from at least semi-permanent
shallow freshwater. The most frequently dominant freshwater marsh species near the Estuary is Scirpus acutus,
but Scirpus americanus, Scirpus californicus, Typha angustifolia, Typha domingensis, and Typha latifolia can also
be important there as dominants. The Typha spp., in
particular, are often dominant in early successional and
nitrogen-enriched freshwater marshes.
Climate and geology have less influence on the distribution of freshwater marsh than they do on the occurrence of other plant communities. When vegetation
is primarily limited by precipitation, temperature, and
light, its distribution is controlled by climate, and when
limited by mineral nutrition and soil texture, its distribution is controlled by geology. The primary limiting
factor in freshwater marshes, however, is air, which,
while superabundant at the marsh surface, falls to such
low concentrations a short distance beneath it that environments too anoxic, reduced, and toxic for root
growth are frequent there. All freshwater marsh dominants in California are consequently monocotyledons,
which have shallow fibrous root systems readily supplied
with air by aerenchyma-rich stems. Many dicotyledons
including most trees and shrubs, in contrast, have solid
stems and deep, non-fibrous root systems poorly adapted
to anoxic conditions. California consequently lacks
swamps, vegetation in semi-permanent shallow water
dominated by woody plants, since it has no native trees
or shrubs capable of completing life cycles in flooded
environments. Buttonbush, Cephalanthus occidentalis,
and several species of Salix can tolerate extended flooding, however, and frequently occur at the ecotone between freshwater marsh and riparian vegetation (Holland
and Keil 1995, Holstein 1984a,b).
Both tidal and non-tidal freshwater marshes are frequent around the Estuary, but the former are most important in the Sacramento-San Joaquin Delta immediately upstream from the true estuary. The Delta
consisted almost entirely of tidal freshwater marsh before it was largely converted to agricultural land, but a
few remnant tidal freshwater marshes still occur there.
Small non-tidal freshwater marshes often resulting from
human alteration of hydrologic conditions are also widespread around the estuary (Bowcutt 1996).
58
Baylands Ecosystem Species and Community Profiles
Soils beneath freshwater marshes may be mineral
or organic. Despite otherwise similar vegetation freshwater marshes with organic soils are technically fens.
Since organic soils derived from Scirpus acutus rhizomes
were general beneath the Delta’s tidal freshwater
marshes, they once constituted a single vast fen (Atwater
and Belknap 1980). Mineral soils, however, generally
occur beneath the many small freshwater marshes
around the Estuary. Marshes develop most readily on
fine, heavy mineral soils since these exclude air and create the anaerobic conditions suitable for marsh vegetation more readily than the coarse and readily aerated
sediments common on streambanks beneath riparian
forests. Since waterbirds quickly transport propagules
permitting establishment of freshwater marsh plants at
sites with suitable hydrological conditions regardless of
their climatic and geological environments, freshwater
marshes are among the easiest plant communities to
restore (Kusler and Kentula 1990).
Continua exist between semi-perennial marshes
and both moist grassland swales (see above) and vernal
pools (see below). Vegetation of areas with hydrology intermediate between freshwater marshes and vernal pools
pond longer than the latter but shorter than the former.
These are most frequently dominated by Eleocharis
macrostachya with normal winter wet season inundation
but can also be dominated by Cyperus eragrostis when
ponding resulting primarily from agricultural and urban
runoff occurs in the warm season. Vegetation arising
from both kinds of seasonal ponding is properly called
seasonal marsh, but wildlife managers also frequently use
the term to describe non-tidal mudflat environments
extremely important for shorebird foraging. Such nontidal mudflats have little vegetation and once commonly
occurred where flooding temporarily suppressed normal
grassland development on stream terraces. Streamflow
control and terrace urbanization, however, have greatly
reduced traditional episodically flooded shorebird habitat around the Estuary. Most non-tidal seasonal marshes
presently occurring there consequently result from seasonal drawdowns of artificial ponds and floodways (SFEP
1991a,b).
Limnetic vegetation refers to floating or submerged
vegetation occurring in open freshwater too deep or otherwise unsuitable for marsh vegetation. Important native components of submerged limnetic vegetation near
the Estuary include Ceratophyllum demersum, Najas
guadalupensis, Potamogeton pectinatus and Potamogeton
pusillus, while important floating elements are Azolla
filiculoides, Lemna gibba and Lemna minor. Non-native
species like Egeria densa and Mytiophyllum aquaticum are
now also extremely significant and often predominant
elements of submerged limnetic vegetation near the
Estuary, but the floating component consists almost entirely of extremely widespread and readily dispersed native species except in and near the Delta, where non-na-
hydrologically resembling vernal pools but lacking their
characteristic biota because of elevated salinity also occur. They are called playas when their surrounding saline environment is inland (Chapman 1960, Waisel
1972) and pans when it is coastal (Adam 1990,
Chapman 1960, Long and Mason 1983, Waisel 1972).
Both occur in San Francisco Bay Area counties but only
the latter near the Estuary shore (SFEP 1991a). Today
vegetated pans near the Estuary are ubiquitously dominated by Cotula coronopifolia, an exotic annual that may
have replaced a now extinct native annual Plagiobothrys
glaber.
11. Anthropogenic Environments – Anthropogenic environments must be briefly considered because
they collectively now dominate non-tidal uplands around
the Estuary. The anthropogenic typology used here follows Mayer and Laudenslayer, Jr. (1988) rather than the
more complex one of Holland and Keil (1995).
Agricultural environments historically surrounded
much of the Estuary but have become increasingly scarce
because of displacement by urbanization. Structurally
and physiologically different elements like orchards, vineyards, and both irrigated and dry farmed cropland are
included here, but all share low plant and animal diversity. Irrigated nursery crops are most important near the
southern part of the Estuary; and vineyards, irrigated
pastures, and dry farmed oats (Avena sativa) predominate near the northern part.
Urban and suburban environments now overwhelmingly dominate non-tidal uplands around the
Estuary. They often structurally resemble extended and
unusually diverse riparian zones since irrigated non-native trees predominantly in the genera Acacia, Eucalyptus, and Pinus are ubiquitously present above an even
more diverse understory of ornamental shrubs and herbs.
As a consequence, urban-suburban communities are
probably the landscape unit near the Estuary with the
highest plant diversity but the fewest native plants. Some
native bird species have adapted to using urban areas as
habitat and become common, but far fewer terrestrial
species are able to do so.
Rare Plants of Ecotonal Plant Communities
Table 1.4 lists rare plant species found in the nine Bay
Area counties. The table is organized using the same
classification system (Holland and Keil) as was used in
the previous section. It includes, for each species, the
state and federal listing status, as well as the status derived from the California Native Plant Society (CNPS)
inventory (Skinner and Pavlik 1994). The CNPS inventory is more complete than the state or federal listings
and is organized on the following lists:
1a. Presumed extinct
1b. Rare, threatened, or endangered in California
and elsewhere.
Chapter 1 —
Plant Communities
59
Plants
tive Eichornia crassipes is important (Holland and Keil
1995).
Vernal pool vegetation refers to a distinctive plant
community dominated by annual and short-lived perennial forbs that occurs in microtopographic basins flooded
in the wet season and dry the rest of the year. Vernal pool
plants are consequently adapted for beginning their life
cycle like submerged limnetic species but completing it
as terrestrial plants in completely dry environments. The
vernal pool environment has led to adaptive radiation of
numerous species primarily in the genera Downingia,
Eryngium, Lasthenia, Navarretia, Plagiobothrys, and
Psilocarphus, and it is these that dominate its vegetation.
Plagiobothrys bracteatus, in particular, is the most frequent dominant of vernal pool vegetation around the
Estuary. Upland vegetation around vernal pools is almost
invariably non-native annual grassland (Holland 1977).
The origin of vernal pool basins is obscure but may
result from seismic activity or interaction of wind erosion and soil processes (see above). To pond water and
create an environment suitable for vernal pool vegetation, however, an aquaclude or barrier to water infiltration that may be a clay horizon, duripan, or bedrock
must be present immediately beneath the basin. Most
vernal pools and the plants adapted to them occur entirely or almost entirely in California, and few hydrologic
features resembling vernal pools occur outside North
America even in otherwise similar Mediterranean climates (Thorne 1984).
Vernal pools are at risk even in the Central Valley
where they are most common because virtually all human activities except rangeland grazing destroy the microtopography and aquacludes that create the vernal pool
environment. Around the Estuary they are even more
threatened since they are extremely rare near southern
San Francisco Bay and only slightly more frequent north
of San Pablo and Suisun bays. Vernal pools north of Suisun Bay are particularly environmentally significant because they are often partially dominated by Lasthenia
conjugens, a federally listed endangered species extinct
throughout much of its range (Skinner and Pavlik 1994).
Artificially created basins often sufficiently resemble natural vernal pools to be colonized by a few
wide-ranging and extremely tolerant pool species. More
rarely a few rare species may be present in such artificial
sites. The full suite of vernal pool taxa including the
rarest species almost never develops in such environments, however, because soil characteristics of natural
pools can rarely be replicated. As a consequence creation
of artificial vernal pools has been the least successful of
all wetland restoration efforts (Ferren and Gevirtz 1990,
Kusler and Kentula 1990).
Vernal pools typically are freshwater environments
since their primary water source is precipitation (Hanes
et al. 1990). However, salt diffusion from underlying
soils causes some to be slightly brackish. Seasonal pools
Plants
2. Rare, threatened, or endangered in California
but more common elsewhere.
3. Possibly rare, but more information is needed.
4. Distribution limited: a watch list.
Plants with the greatest need for protection are on
list 1b, and 1a (presumed extinct) plants are placed there
if rediscovered. The CNPS inventories rare plants by
county. To prepare Table 1.4, each of the CNPS-identified rare species was assigned to a modal plant community or ecotone based on information provided by
state and local floras. An effort was made to place each
taxon in the plant community it most frequently (but
not necessarily exclusively) occurs(ed) in, however, frequently reference materials regarding a taxon were contradictory. In these cases I sought to develop a consensus view, and weighted local floras and my own field
experience most heavily.
The greatest constraint in preparing this table was
the frequent sparsity of ecological information regarding rare species. Preparation was easiest in areas with
60
Baylands Ecosystem Species and Community Profiles
local floras since these are full of observations by botanists with deep knowledge of their region’s plants and
habitats. Tragically, however, a number of plants near
the Estuary went extinct or became extremely rare in an
older era when little or no ecological information was
provided when plants were collected. We can only speculate regarding the niches of these taxa.
Many species are rare because they occur in rare
ecological niches. Historically, these have not been the
focus of plant community classification, which is most
concerned with the commonest kinds of vegetation.
While there was an effort to include some of the rarer
niches occurring near the Estuary in Table 1.4, it is not
comprehensive, and rare niches distant from the Estuary are not included. Rare plants that occur primarily in
plant communities distant from the Estuary were not included in the narrative community descriptions.
It is hoped this table will generate discussion and
suggestions for its improvement.
Table 1.4 Rare Plant Species* Found in the Nine Counties Adjacent to the San Francisco Bay Estuary,
by Plant Community or Ecotone
A. Pioneer coastal dune vegetation
1b. Abronia umbellata ssp. breviflora - FSC
B. Coastal dune scrub
1b. Agrostis blasdalei - FSC
Chorizanthe cuspidata var. cuspidata - FSC
Chorizanthe cuspidata var. villosa
Chorizanthe robusta var. robusta - FE
Collinsia corymbosa
Erysimum ammophilum - FSC
Horkelia cuneata ssp. sericea - FSC
Horkelia marinensis - FSC
Layia carnosa - FE, SE
Lessingia germanorum - FE, SE
Lupinus tidestromii - FE, SE
+Gilia capitata ssp. chamissonis
+Gilia millefoliata
4. Monardella undulata
C. Inland dune vegetation
1b. Erysimum capitatum ssp. angustatum - FE, SE
Oenothera deltoides ssp. howellii - FE, SE
2. Coastal scrub
A. Northern coastal scrub
1b. Delphinium bakeri - FPE, SR
Lilium maritimum - FC
+Lupinus latifolius var. dudleyi
3. Lupinus eximius - FSC
4. Cirsium andrewsii
Collinsia multicolor
Piperia michaelii
Sanicula hoffmannii
B. Southern coastal scrub
C. Sea-bluff coastal scrub
1b. Cirsium occidentale var. compactum - FSC
Grindelia hirsutula var. maritima - FSC
Phacelia insularis var. continentis - FSC
Silene verecunda ssp. verecunda - FSC
+Agrostis clivicola var. punta-reyesensis
+Gilia capitata ssp. tomentosa
+Piperia elegans ssp. decurtata
4. Arabis blepharophylla
Ceanothus gloriosus var. gloriosus
Erysimum franciscanum - FSC
Hesperevax sparsiflora var. brevifolia
+Agrostis clivicola var. clivicola
3. Chaparral
1b. Arctostaphylos auriculata
Arctostaphylos densiflora - FSC, SE
Arctostaphylos imbricata - FPT, SE
Arctostaphylos manzanita ssp. laevigata
Arctostaphylos montaraensis - FSC
Arctostaphylos pallida - FPT, SE
Arctostaphylos stanfordiana ssp. decumbens
Ceanothus confusus - FSC
Ceanothus divergens - FSC
Ceanothus foliosus var. vineatus - FSC
Ceanothus masonii - FSC, SR
Ceanothus sonomensis - FSC
Malacothamnus hallii
Plagiobothrys uncinatus - FSC
Plants
1. Sand dune vegetation
3. Calyptridium parryi var. hesseae
+Arctostaphylos manzanita ssp. elegans
4. Ceanothus purpureus
Dichondra occidentalis
Lomatium repostum
Malacothamnus arcuatus
Malacothamnus helleri
Orobanche valida ssp. howellii
Plagiobothrys myosotoides
A. Chaparral burns
4. Calandrinia breweri
+Malacothrix phaeocarpa
4. Grassland
A. Native perennial grassland
1b. Amsinckia grandiflora - FE, SE
Astragalus clarianus - FE, ST
Blepharizonia plumosa ssp. plumosa
Fritillaria pluriflora - FSC
Tracyina rostrata
Trifolium amoenum - FE
+Calochortus argillosus
3. Lessingia hololeuca
4. Androsace elongata ssp. acuta
Fritillaria agrestis
+Allium peninsulare var. franciscanum
+Microseris paludosa
Alkaline grassland
1a. Tropidocarpum capparideum - FSC
1b. Astragalus tener var. ferrisiae - FSC
Astragalus tener var. tener
Atriplex cordulata - FSC
Atriplex depressa
Atriplex joaquiniana - FSC
Cordylanthus mollis ssp. hispidus - FSC
Cordylanthus palmatus - FE, SE
Delphinium recurvatum - FSC
Hemizonia parryi ssp. congdonii - FC
Isocoma arguta – FSC
+Trifolium depauperatum var. hydrophilum
3. Hordeum intercedens
4. Atriplex coronata var. coronata
Thelypodum brachycarpum
* Derived from the inventory of the California Native Plant Society (CNPS) (Skinner and Pavlik 1994)
Key to CNPS list codes:
Key to Federal and State List Codes:
1a.
1b.
2.
FE Federally listed as endangered
FT Federally listed as threatened
FC Federal listing candidate
3.
4.
+
Presumed extinct
Rare, threatened, or endangered in California and elsewhere.
Rare, threatened, or endangered in California but more common
elsewhere.
Possibly rare, but more information is needed.
Distribution limited: a watch list
Proposed new addition to the CNPS inventory
SE State listed as endangered
ST State listed as threatened
SR State listed as rare
FPE Proposed for Federal listing as endangered
FPT Proposed for federal listing as threatened
FSC Federal species of special concern
Chapter 1 —
Plant Communities
61
Table 1.4 (continued) Rare Plant Species* Found in the Nine Counties Adjacent to the San Francisco
Bay Estuary, by Plant Community or Ecotone
Plants
Sandy soil grassland
1a. Eriogonum truncatum
4. Cryptantha hooveri
Linanthus grandiflorus
Serpentinite grassland
1b. Acanthomintha duttonii - FE, SE
Calochortus tiburonensis - FT, ST
Castilleja affinis ssp. neglecta - FE, ST
Fritillaria biflora var. ineziana
Lessingia arachnoidea - FSC
Streptanthus niger - FE, SE
4. Piperia candida
Pityopus californicus
+Galium muricatum
6. Mixed evergreen forest
A. Central California mixed evergreen forest
1b. +Quercus parvula var. tamalpaisensis
3. +Viburnum ellipticum
4. Cypripedium montanum
Dirca occidentalis
Ribes victoris
3. Eriogonum luteolum var. caninum
7. Oak woodland
4. Astragalus breweri
Linanthus ambiguus
A. Coast live oak woodland
Moist grassland
1a. Plagiobothrys hystriculus
1b. Pleuropogon hooverianus - FSC, SR
Sidalcea oregana ssp. hydrophila
2. Carex californica
4. Perideridia gairdneri ssp. gairdneri - FSC
B. Native annual forbland
1a. Eschscholzia rhombipetala - FSC
1b. Madia radiata
4. Convolvulus simulans
+Erodium macrophyllum
C. Non-native annual grassland
D. Coastal prairie
1b. Blennosperma nanum var. robustum - FSC, SR
chorizanthe valida - FE, SE
Erigeron supplex - FSC
Fritillaria lanceolata var. tristulis
Fritillaria liliacea - FSC
Holocarpha macradenia - FC, SE
Limnanthes douglasii ssp. sulphurea - FSC, SE
Plagiobothrys diffusus - FSC, SE
Sanicula maritima - FSC, SR
Triphysaria floribunda - FSC
3. Hemizonia congesta ssp. leucocephala
Plagiobothrys chorisianus var. chorisianus
5. Coastal coniferous forest
A. Redwood forest
4. Elymus californicus
B. Closed-cone coniferous forest
1b. Ceanothus gloriosus var. porrectus - FSC
Cupressus goveniana ssp. pigmaea - FSC
Pinus radiata - FSC
C. North coast coniferous forest
2. Boschniakia hookeri
1b. Clarkia concinna var. automixa
4. Amsinckia lunaris
Isocoma menziesii var. diabolica
B. Valley oak woodland
C. Foothill oak woodland
8. Cliffs and rock outcrops
A. Cliffs
3. +Streptanthus tortuosus var. suffrutescens
4. Arabis modesta
B. Rock outcrops
1b. Arctostaphylos virgata
Coreopsis hamiltonii - FSC
Penstemon newberryi var. sonomensis
Phacelia phacelioides - FSC
Sanicula saxatilis - FSC, SR
Streptanthus callistus - FSC
Streptanthus glandulosus var. hoffmanii - FSC
Streptanthus hispidus - FSC
3. Erigeron biolettii
Monardella antonina ssp. antonina
4. Antirrhinum virga
Arabis oregona
Arctostaphylos hispidula
Navarretia subuligera
Serpentinite outcrops
1a. Arctostaphylos hookeri ssp. franciscana - FSC
1b. Allium sharsmithae
Arctostaphylos bakeri ssp. bakeri - SR
Arctostaphylos bakeri ssp. sublaevis
Arctostaphylos hookeri ssp. montana - FSC
Arctostaphylos hookeri ssp. ravenii - FE, SE
Astragalus rattanii var. jepsonianus
Calochortus raichei - FSC
Campanula sharsmithiae - FSC
Ceanothus ferrisae - FE
Chlorogalum pomeridianum var. minus
Clarkia franciscana - FE, SE
* Derived from the inventory of the California Native Plant Society (CNPS) (Skinner and Pavlik 1994)
Key to CNPS list codes:
Key to Federal and State List Codes:
1a.
1b.
2.
FE Federally listed as endangered
FT Federally listed as threatened
FC Federal listing candidate
3.
4.
+
62
Presumed extinct
Rare, threatened, or endangered in California and elsewhere.
Rare, threatened, or endangered in California but more common
elsewhere.
Possibly rare, but more information is needed.
Distribution limited: a watch list
Proposed new addition to the CNPS inventory
Baylands Ecosystem Species and Community Profiles
SE State listed as endangered
ST State listed as threatened
SR State listed as rare
FPE Proposed for Federal listing as endangered
FPT Proposed for federal listing as threatened
FSC Federal species of special concern
Table 1.4 (continued) Rare Plant Species* Found in the Nine Counties Adjacent to the San Francisco
Bay Estuary, by Plant Community or Ecotone
3. Cardamine pachystigma var. dissectifolia
+Streptanthus glandulosus var. sonomensis
4. Acanthomintha lanceolata
Asclepias solanoana
Aspidotis carlotta-halliae
Calamagrostis ophitidis
Calyptridium quadripetalum
Campanula exigua
Clarkia breweri
Collomia diversifolia
Cordylanthus tenuis ssp. brunneus
Eriogonum argillosum
Eriogonum ternatum
Eriogonum tripodum
Fritillaria purdyi
Galium andrewsii ssp. gatense
Lomatium ciliolatum var. hooveri
Navarretia jepsonii
+Ceanothus jepsonii var. albiflorus
+Streptanthus barbiger
Granite and sandstone outcrops
4. Arctostaphylos regismontana
Plants
Cordylanthus nidularius - FC, SR
Cordylanthus tenuis ssp. capillaris - FE, SR
Cryptantha clevelandii var. dissita
Dudleya setchellii - FE
Erigeron angustatus
Erigeron serpentinus
Eriogonum nervulosum - FSC
Fritillaria falcata - FSC
Hesperolinon bicarpellatum - FSC
Hesperolinon breweri - FSC
Hesperolinon congestum - FT, ST
Hesperolinon drymarioides - FSC
Hesperolinon serpentinum
Lessingia micradenia var. glabrata - FSC
Lessingia micradenia var. micradenia - FSC
Madia hallii - FSC
Navarretia rosulata
Sidalcea hickmanii ssp. viridis - FSC
Streptanthus albidus ssp. albidus - FE
Streptanthus albidus ssp. peramoenus - FC
Streptanthus batrachopus - FSC
Streptanthus brachiatus ssp. brachiatus - FC
Streptanthus brachiatus ssp. hoffmanii - FC
Streptanthus glandulosus ssp. pulchellus
Streptanthus morrisonii ssp. elatus - FC
Streptanthus morrisonii ssp. hirtiflorus - FC
Streptanthus morrisonii ssp. kruckebergii - FSC
Streptanthus morrisonii ssp. morrisonii - FSC
+Hoita strobilina
+Streptanthus breweri var. hesperidis
Volcanic outcrops
1b. Eriastrum brandegeae - FSC
4. Madia nutans
9. Riparian vegetation
1b. Juglans californica var. hindsii - FSC
+Triteleia lugens
4. Astragalus rattanii ssp. rattanii
10. Freshwater vegetation
A. Freshwater marsh
1a. Castilleja uliginosa - FSC, SE
1b. Alopecurus aequalis var. sonomensis - FE
Arenaria paludicola - FE, SE
Campanula californica - FSC
Carex albida - FE, SE
Lilium pardalinum ssp. pitkinense - FE, SE
Potentilla hickmanii - FPE, SE
Rhynchospora californica - FSC
Sidalcea calycosa ssp. rhizomata
Sidalcea oregana ssp. valida - FE, SE
2. Calamagrostis crassiglumis - FSC
Carex comosa
Carex leptalea
Rhynchospora globularis var. globularis
3. Equisetum palustre
4. Calamagrostis bolanderi
Rhynchospora alba
+Zigadenus micranthus var. fontanus
B. Limnetic vegetation
2. Potamogeton filiformis
Potamogeton zosteriformis
4. Azolla mexicana
Ranunculus lobbii
C. Vernal pools
1b. Blennosperma bakeri - FE, SE
Gratiola heterosepala - SE
Lasthenia burkei - FE, SE
Lasthenia conjugens - FE
Legenere limosa - FSC
Limnanthes vinculans - FE, SE
Navarretia leucocephala ssp. bakeri
Navarretia leucocephala ssp. pauciflora - FE, ST
Navarretia leucocephala ssp. plieantha - FE, SE
Neostapfia colusana - FT, SE
Tuctoria mucronata - FE, SE
2. Downingia pusilla
3. Myosurus minimus ssp. apus - FSC
Pogogyne douglasii ssp. parviflora
* Derived from the inventory of the California Native Plant Society (CNPS) (Skinner and Pavlik 1994)
Key to CNPS list codes:
Key to Federal and State List Codes:
1a.
1b.
2.
FE Federally listed as endangered
FT Federally listed as threatened
FC Federal listing candidate
3.
4.
+
Presumed extinct
Rare, threatened, or endangered in California and elsewhere.
Rare, threatened, or endangered in California but more common
elsewhere.
Possibly rare, but more information is needed.
Distribution limited: a watch list
Proposed new addition to the CNPS inventory
SE State listed as endangered
ST State listed as threatened
SR State listed as rare
FPE Proposed for Federal listing as endangered
FPT Proposed for federal listing as threatened
FSC Federal species of special concern
Chapter 1 —
Plant Communities
63
Plants
Table 1.4 (continued) Rare Plant Species* Found in the Nine Counties Adjacent to the San Francisco
Bay Estuary, by Plant Community or Ecotone
4. Eryngium aristulatum var. hooveri - FC
Psilocarphus brevissimus var. multiflorus
Psilocarphus tenellus var. globiferus
D. Thermal springs
1b. Dichanthelium lanuginosum var. thermale - FSC, SE
Plagiobothrys strictus - FE, ST
Poa napensis - FP, SE
11. Anthropogenic environments
A. Agricultural
B. Urban-suburban
12. Coastal marsh
A. Brackish marsh
1b. Aster lentus - FSC
Cirsium hydrophilum var. hydrophilum - FE
Cordylanthus mollis ssp. mollis - FE, SR
Lathyrus jepsonii var. jepsonii - FSC
Lilaeopsis masonii - FSC, SR
2. Limosella subulata
B. Saltmarsh
1b. Castilleja ambigua ssp. humboldtiensis - FSC
Cordylanthus maritimus ssp. palustris - FSC
Suaeda californica - FE
3. Polygonum marinense - FSC
4. Eleocharis parvula
Grindelia stricta var. angustifolia
13. Ecotones
A. Grassland-oak woodland
1b. Helianthella castanea - FSC
Monardella villosa ssp. globosa
4. Linanthus acicularis
B. Grassland-rock outcrops
1b. Balsamorhiza macrolepis var. macrolepis
Clarkia concinna ssp. raichei - FSC
Layia septentrionalis
Pentachaeta bellidiflora - FE, SE
Stebbinsoseris decipiens - FSC
4. Micropus amphibolus
C. Mixed evergreen-chaparral
1b. Arctostaphylos andersonii - FSC
Arctostaphylos canescens ssp. sonomensis
Cupressus abramsiana - FE, SE
Eriogonum nudum var. decurrens
Lupinus sericatus
Penstemon rattanii var. kleei
4. Calystegia collina ssp. oxyphylla - FSC
Erythronium helenae
Lilium rubescens
Monardella viridis ssp. viridis
D. Mixed evergreen-serpentinite outcrops
4. Calochortus umbellatus
E. Rock outcrops-riparian
1b. Delphinium californicum ssp. interius - FSC
4. Trichostema rubisepalum
F. Serpentinite outcrops-riparian (including serpentine
seeps)
1b. Cirsium fontinale var. campylon - FSC
Cirsium fontinale var. fontinale - FE, SE
Cirsium hydrophilum var. vaseyi - FSC
4. Astragalus clevelandii
Cypripedium californicum
Delphinium uliginosum
Helianthus exilis
Mimulus nudatus
Senecio clevelandii var. clevelandii
G. Coastal coniferous forest-riparian
1b. Pedicularis dudleyi - FSC, SR
4. Cypripedium fasciculatum - FSC
Pleuropogon refractus
H. Oak woodland-serpentinite outcrops
1b. Eriophyllum latilobum - FE, SE
I. Oak woodland-chaparral
1b. Calochortus pulchellus
4. Eriophyllum jepsonii
J. Alkaline grassland-pans
1a. Plagiobothrys glaber
Plagiobothrys mollis var. vestitus - FSC
K. Coastal coniferous forest-coastal prairie
1b. Sidalcea malachroides
L. Freshwater marsh-riparian
1b. Sagittaria sanfordii - FSC
2. Hibiscus lasiocarpus
M. Grassland-southern coastal scrub
2. Senecio aphanactis
N. Coastal prairie-northern coastal scrub
1b. Delphinium luteum - FPE, SR
O. Grassland-chaparral
1b. Clarkia imbricata - FE, SE
Horkelia tenuiloba
P. Northern coastal scrub-riparian
4. Veratrum fimbriatum
* Derived from the inventory of the California Native Plant Society (CNPS) (Skinner and Pavlik 1994)
Key to CNPS list codes:
Key to Federal and State List Codes:
1a.
1b.
2.
FE Federally listed as endangered
FT Federally listed as threatened
FC Federal listing candidate
3.
4.
+
64
Presumed extinct
Rare, threatened, or endangered in California and elsewhere.
Rare, threatened, or endangered in California but more common
elsewhere.
Possibly rare, but more information is needed.
Distribution limited: a watch list
Proposed new addition to the CNPS inventory
Baylands Ecosystem Species and Community Profiles
SE State listed as endangered
ST State listed as threatened
SR State listed as rare
FPE Proposed for Federal listing as endangered
FPT Proposed for federal listing as threatened
FSC Federal species of special concern
References
Chapter 1 —
Plant Communities
65
Plants
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Adam, P. 1990. Saltmarsh ecology. Cambridge, UK:
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Atwater, B. F. and D. F. Belknap. 1980. Tidal wetland
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Barbour, M. G. and A. F. Johnson. 1977. Beach and
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Berg, A. W. 1990. Formation of Mima mounds: a seismic hypothesis. Geology 18: 281-4.
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Chapter 1 —
Plant Communities
67
Plants
68
Baylands Ecosystem Species and Community Profiles
2
Estuarine Fish and Associated Invertebrates
Fish
Food and Feeding
Opossum Shrimp
Neomysis mercedis
Bruce Herbold
General Information
The opossum shrimp is a native mysid shrimp that is an
important food for many estuarine fish, especially young
striped bass. Since 1994, their role of dominant planktonic shrimp has been overwhelmed by the introduced
species, Acanthomysis (Orsi and Mecum 1996).
The diet of N. mercedis varies with size. At release, young
shrimp eat mostly phytoplankton and rotifers. Adult
diets include phytoplankton and rotifers but the diet
shifts more to copepods, particularly Eurytemora affinis
(Herbold et al. 1992).
Distribution
N. mercedis is found in greatest abundance in Suisun Bay
and the western Delta, although it occurs as far upstream
as Sacramento, the lower reaches of the Mokelumne
River, and in the San Joaquin River to above Stockton.
Reproduction
ARO, M. Roper
The common name of the opossum shrimp derives from
the fact that females carry their eggs and young in a
pouch at the base of the last two pairs of legs. Young are
released at a well-developed stage. Fecundity is related
both to adult size and season (Heubach 1969).
Reproduction is continuous but the rate is strongly
controlled by temperature and food supply. Thus, the
rate is high during spring and summer months and slows
down as temperature and insolation decline. The wintertime population is composed largely of large adults,
whose greater fecundity allows rapid development of
high densities as temperatures and phytoplankton densities rise. The autumn decline in density has been variously attributed to seasonal changes in high temperature,
low dissolved oxygen, predation, and food supply
(Turner and Heubach 1966, Heubach 1969, Siegfried
et al. 1979, Orsi and Knutson 1979).
Population Status and Influencing Factors
During most of the 1980s, the opossum shrimp population varied considerably, but remained at a lower level
of abundance than existed in the early 1970s. Opossum
shrimp abundance fell dramatically after 1986 and remained at very low levels from 1990 to 1993 (CDFG
1994). As a general trend, opossum shrimp populations
have declined substantially in Suisun Bay, yet they have
occasionally rebounded to high levels (BDOC 1993).
Reasons for the system-wide declines of several
zooplankton taxa in the Bay-Delta Estuary are not
known. Although the declines occurred at about the
same time as declines in phytoplankton and various fish
species, no cause-and-effect relationships have been established (CDWR 1992). However, several factors have
been identified which are believed to have some influence on the decline of zooplankton in the Estuary.
Decrease in food supply has been associated with
the decline in abundance of rotifers and the copepod,
E. affinis. The decline of rotifers in the Delta appears to
be strongly associated with declining concentrations of
chlorophyll a, which formerly characterized the areas of
greatest rotifer abundance (Herbold et al. 1992). However, chlorophyll and many zooplankton species have
similar spatial distributions, and correlations between the
two groups can arise through movement of the entrapment zone in the Estuary. Also, while it is commonly
Chapter 2 — Estuarine Fish and Associated Invertebrates
69
70
Baylands Ecosystem Species and Community Profiles
Fish
Amphibians &
Reptiles
a higher rate of loss to resident zooplankton populations
than export pumping.
Pollutants may be another factor in the decline of
zooplankton in the upper Estuary. For example, rice herbicides have been shown to be toxic to opossum shrimp
(CDWR 1992). However, rice herbicides are largely confined to the Sacramento River, not the entire Estuary.
No Estuary-wide decline in planktonic crustaceans have
been associated with the timing of herbicide occurrence
in the river (NHI 1992).
Trophic Levels
The opossum shrimp is a primary and secondary consumer.
Proximal Species
Predators: Striped bass, longfin smelt, splittail.
Prey: Various copepods, various phytoplankton.
Competitors: Potamocorbula amurensis, Acanthomysis spp.
Good Habitat
Good habitat appears to be similar to that of Delta smelt;
a well-dispersed area of open water with salinities in the
range of 2 to 6 ppt for most of the year and clean, nontoxic over-wintering habitat in freshwater through the
winter and early spring. Dead-end sloughs both in Suisun Marsh and upstream apparently serve as important
refuges from predation during the annual period of low
abundance and slow growth. With the advent of newly
introduced competitors in the open waters of the Estuary it is possible that such refugia will become important for the year-round maintenance of opossum shrimp.
References
Bay-Delta Oversight Council (BDOC). 1993. Draft
briefing paper on biological resources of the San
Fran. Bay/Sac.-San Joaquin Delta Estuary. September 1993. 42 pp. plus appendices.
California Department of Fish and Game (CDFG).
1994. Comments on key issues of the State Water
Resources Control Board’s June workshop for review of standards for San Francisco Bay/Sacramento-San Joaquin Delta Estuary. Presented by
Perry L. Herrgesell, Chief of Bay-Delta and Special Water Projects Division, June 14, 1994. 68
pp.
California Department of Water Resources (CDWR).
1992. Bay-Delta fish resources. Ca. Dept. of Water Res. Sacramento, Ca. July 1992. 46 pp.
(WRINT-DWR-30)
Herbold, B., P.B. Moyle and A. Jassby. 1992. Status and
trends report on aquatic resources in the San Fran-
Plants
assumed that chlorophyll is a good measure of food availability for zooplankton, E. affinis can subsist on detrital
matter and requires larger particles than those that make
up total chlorophyll. In addition, small zooplankton
could provide food for many of the larger zooplankton
species (Kimmerer 1992). Consistently low E. affinis
abundance in recent years has been named as a factor
that has probably contributed to the decline of opossum
shrimp (Herbold et al. 1992).
Introduced species have also been named as a potential cause for the decline in zooplankton abundance.
For example, the introduction of Sinocalanus has been
identified as a possible cause of the decline in abundance
of E. affinis (Kimmerer 1992), although the introduced
copepod does not have the same habitat requirements
as the native copepods (NHI 1992). However, based on
the known feeding habits of a related species of Sinocalanus, S. doerrii may prey on native copepods (Herbold et
al. 1992). In addition, predation by the introduced Asian
clam, Potamocorbula amurensis, has been suggested as a
factor in the decline of rotifer (Herbold et al. 1992) and
E. affinis populations. E. affinis abundance in Suisun Bay
decreased substantially when the clam became abundant
there in 1988 (CDWR 1992). Since 1994 Neomysis
abundance has dropped to less than that of an introduced
species of mysid shrimp which has increased in abundance (Orsi 1996). Competition with both the clam and
new shrimp are likely to prevent re-establishment of
Neomysis at the levels of their former abundance.
The decline in the abundance of opossum shrimp
and other zooplankton species (e.g., E. affinis) that are
found in the entrapment zone in relatively high abundances has been correlated with Delta outflow. It is presumed that low outflow reduces opossum shrimp abundance by: (1) restricting the entrapment zone to deeper,
more upstream channels which are less likely to promote
high densities of opossum shrimp; and (2) producing
weaker landward currents along the bottom so that the
ability of opossum shrimp transported downstream to
return to the entrapment zone is reduced. It has also been
presumed that larger numbers of opossum shrimp may
be exported through the Central Valley Project and State
Water Project pumps as a result of the increased proportion of inflow diverted during drought years when the
entrapment zone is upstream in the Estuary. The location of the entrapment zone within the lower river channels during dry years increases the vulnerability of opossum shrimp to such displacement (Herbold et al. 1992).
However, analyses by Kimmerer (1992) suggest that exports by the water projects are not a major source of
losses for opossum shrimp and E. affinis populations, primarily due to the small percentage of entrapment zone
volume (and entrapment zone organisms) diverted. Depending on the timing, location, and quantity of withdrawals, in-Delta water diversions, whose net consumption is on the same order of export flows, may result in
Dungeness Crab
Cancer magister
Robert N. Tasto
General Information
CDFG
Dungeness crab has been the object of an immensely
popular commercial and recreational fishery in the San
Francisco region since 1848. The San Francisco fishery,
which occurs exclusively outside the Golden Gate, was
long a mainstay of statewide commercial landings. However, beginning in the early 1960s, it underwent a severe and longterm decline which persisted until the
mid-1980s. The principal causes of the decline have been
related to changes in ocean climate, increased predation,
and possibly pollution (Wild and Tasto 1983). Landings
in the past decade have rebounded to some extent and
are generally able to accomodate local market demand,
but the northern California fishery (Eureka and Crescent City) continues to be the major provider of Dungeness crabs throughout the rest of California. The value
of the Dungeness crab resource extends beyond the traditional economic return to the fishermen, seafood processors, and retail markets, as it is an important element
in the tourism industry of San Francisco.
California commercial and recreational fishing
regulations pertaining to Dungeness crab have been designed to protect this species from over-harvesting. The
standard commercial fishing gear is a baited 3.5-foot
diameter metal trap, weighing 60 to 120 pounds (Warner
1992). California regulations set a 6.25-inch carapace
width (cw) size limit, prohibit the take of female crabs,
and, like most states, have established a specific fishing
season to protect reproducing and egg-bearing crabs. A
limited recreational fishery allows the take of female crabs
and has a smaller size restriction (5.75 inches cw); a
10-crab bag limit is placed on the sportfishers. Recreational gear consists of a variety of traps, hoops, and nets
of different sizes, shapes, and materials. It is currently
Chapter 2 —
Estuarine Fish and Associated Invertebrates
71
Fish
cisco Estuary. San Francisco Estuary Project. Public Report. March 1992. 257 pp. plus appendices.
Heubach, W. 1969. Neomysis awatchschensis in the Sacramento-San Joaquin Estuary. Linmol. Oceanogr.
14: 533-546.
Kimmerer, W. 1992. An evaluation of existing data in
the entrapment zone of the San Francisco Bay Estuary. Interagency Ecological Studies Program for
the Sac./San Joaquin Estuary. Tech. Rept. 33. September 1992. FS/BIO-IATR/92-93. 49 pp.
Natural Heritage Institute (NHI). 1992. Causes of decline in estuarine fish species. Presented to the State
Wat. Res. Contr. Bd. Water Rights Phase of the
Bay Delta Estuary Proceedings. St. Wat. Res.
Contr. Bd. exhibit WRINT-NHI-9. 29 p plus appendices.
Orsi, J.J. and A.C Knutson. 1979. The role of mysid
shrimp in the Sacramento-San Joaquin Estuary and
factors affecting their abundance and distribution.
In: T.J. Conmos (ed). San Francisco Bay: The urbanized estuary, Pac. Div., Am. Assoc. Adv. Sci.,
San Francisco, Ca. pp. 401-408.
Orsi, J.J. and W.L. Mecum. 1996. Food limitation as
the probable cause of a long-term decline in the
abundance of Neomysis mercedis, the opossum
shrimp, in the Sacramento-San Joaquin estuary.
In: J.T. Hollibaugh (ed). San Francisco Bay: The
ecosystem. Pacific Division, Amer. Assoc. for the
Advancement of Science, San Francisco, Ca. pp.
375-402,
Siegfried, C.A., M.E. Kopache and A.W. Knight. 1979.
The distribution and abundance of Neomysis
mercedis in relation to the entrapment zone in the
western Sacramento-San Joaquin Delta. Trans. Am.
Fish. Soc. 108: 262-270.
Turner, J.L. and W. Heubach. 1966. Distribution and
concentration of Neomysis awatchschensis in the
Sacramento-San Joaquin Delta. In: D.W. Kelley
(ed). Ecological studies of the Sac.-San Joaquin
Delta. Ca. Dept. Fish and Game, Fish Bull. No.
133: 105-112.
illegal to catch Dungeness crab of any size in San Francisco Bay.
Fish
Reproduction
Mating occurs in nearshore coastal waters, from March
through May, between hard-shelled males and recently
molted, soft-shelled females. Fertilized eggs are extruded
in the fall and lay protected beneath the female’s abdominal flap in a sponge-like mass until hatching occurs from
late December to mid-January (Wild and Tasto 1983).
Fecundity ranges from 500,000 to 2,000,000 eggs, depending upon the size of the female (Warner 1992). C.
magister is capable of about four broods over its reproductive life span (Hines 1991).
Dungeness crab life stages include the egg, larval, juvenile, and adult. Dungeness crab eggs range in diameter
from 0.016 to 0.024 inches (Warner 1992). There are a
total of six larval stages (five zoeae and one megalopa)
which spend about 3 to 4 months in both nearshore and
offshore coastal waters; larval timing is believed to coincide with peak plankton production (Hines 1991).
Late-stage megalopae, which have returned to the coast,
bays, and estuaries via ocean currents and other mechanisms, settle onto relatively open sandy areas (Oresanz
and Gallucci 1988) and subsequently metamorphose to
the first bottom-dwelling instar stage generally between
April and June. It is at this stage that the young crabs
enter San Francisco Bay in large numbers, relative to
year-class strength, seemingly aided by strong bottom
currents (Tasto 1983). San Francisco Bay-reared crabs
molt more frequently than those juveniles found in the
near coastal environment and reach sexual maturity (approximately 4 inches wide) after nearly one year (Wild
and Tasto 1983). This rate of growth is substantially
greater than that found in open areas along the Pacific
coast and may be due to increased availability of food
and/or overall warmer temperatures of estuaries (Tasto
1983, Gunderson et al. 1990, Wainwright and Armstrong 1993). It is believed that the large number of
molts necessary to reach sexual maturity in an estuarine
environment is due, in large part, to the demands of osmoregulation (Oresanz and Gallucci 1988).
Food and Feeding
Larval Dungeness crab in the water column are planktivorous, whereas the juvenile and adult crabs are opportunistic foragers on larger bottom-dwelling organisms.
In the San Francisco Estuary, juvenile crabs have been
shown to feed on clams, crustaceans, and small fishes
(Tasto 1983). In Grays Harbor, Washington, juvenile
crab diets consisted primarily of Crangon shrimp, juve-
72
Baylands Ecosystem Species and Community Profiles
Distribution
Dungeness crab range from the Aleutian Islands to Santa
Barbara, but are rare south of Point Conception (Warner
1992). The pelagic larval forms are found distributed
widely in both nearshore and offshore waters, but return
to the coast, bays, and estuaries where the juvenile and
adult stages are mostly found from the intertidal zone
to approximately 300 feet (Hatfield 1983, Reilly 1983b,
Warner 1992). San Francisco Bay, as is the case with
other coastal estuaries, is an important nursery area for
the offshore stock. The vast majority of individuals in
the Bay are juveniles of a single year-class, having entered
in the spring of one year and exited approximately 1 year
later (Tasto 1983, McCabe et al. 1988). Juveniles are
often found in tidal and navigational channels early in
summer, but spread out over mudflats and into protected
shoreline areas as they develop over the year (Figures 2.1
and 2.2).
Population Status and Influencing Factors
Few population estimates have been made on individual
Dungeness crab stocks along the Pacific coast because
there is significant variation in year-class strength, purportedly due to environmental conditions and density-dependent factors (Botsford and Hobbs 1995). However, commercial crab landings, monitored annually
by state and, in some instances, federal resource agencies, appear to be a reliable indicator of relative abundance.
The most important factors affecting overall population numbers in the San Francisco area (i.e., Half
Moon Bay to Bodega Bay) include ocean temperatures
(hatching success), ocean currents (larval drift), predation, commercial fishing, and, possibly, pollution of
nursery habitat (Wild and Tasto 1983). Although labo-
Plants
Amphibians &
Reptiles
Growth and Development
nile fish, and bivalves (Gunderson et al. 1990). By comparison to other cancrid crabs, the small chelae of C.
magister are better suited for soft-bodied, mobile prey
found on sandy bottoms (Oresanz and Gallucci 1988).
One study has suggested that size-specific feeding on
clams in the laboratory was due to an attempt to minimize handling time of the prey in a competitive situation (Palacios and Armstrong 1990).
The most common predators on juvenile crabs
within the San Francisco Estuary include bottom-feeding
fishes such as starry flounder, English sole, Pacific tomcod, Pacific staghorn sculpin, white croaker, pile perch,
sturgeon, and several elasmobranchs (sharks, skates, and
rays) (Reilly 1983a). The principal predator on young-of-the-year Dungeness crab in Gray’s Harbor Estuary
was found to be the Pacific staghorn sculpin (Fernandez
et al. 1993a). In addition, cannibalism is reported to occur among all age groups (Warner 1992).
Fish
Figure 2.1 Seasonal Distribution of Juvenile Dungeness Crab Within San Francisco Bay (Tasto 1983)
ratory results show that cannibalism may be an important determinant in the abundance and structure of some
populations (Fernandez et al. 1993b), year-class strength
and recruitment to the fishery do not appear to be dependent upon success of any particular “ critical” stage
(McConnaughey and Armstrong 1990). Within the San
Francisco Estuary, juvenile abundance varies considerably from year to year, but is often highest in San
Pablo Bay and lowest in south Bay (Tasto 1983,
CDFG 1987).
Chapter 2 —
Estuarine Fish and Associated Invertebrates
73
Fish
Amphibians &
Reptiles
Plants
Figure 2.2 Annual Distribution of Juvenile Dungeness Crab Within the San Francisco Bay – Caught by
Otter Trawl, May-December (CDFG 1987)
Trophic Levels
Proximal Species
Larvae are planktivores making them primary consumers (phytoplankton) and secondary consumers (zooplankton). Juveniles and adults are higher order consumers.
Predators: Chinook and coho salmon* (prey on late larval stages); Carcinonemertes errans* (predator worm on
egg masses); Dungeness crab (cannibalism by larger instars, principally females, on small juveniles), starry
74
Baylands Ecosystem Species and Community Profiles
flounder, English sole, Pacific tomcod, Pacific staghorn
sculpin, white croaker, brown smoothhound shark, and
skate (prey on juveniles); and humans (commercial and
recreational fishing for adults*).
* Generally takes place outside of San Francisco Bay.
Prey: Crustaceans, bivalves (clams), small fishes.
Good Habitat
References
Botsford, L.W. and R.C. Hobbs. 1995. Recent advances
in the understanding of cyclic behavior of Dungeness crab (Cancer magister) populations. ICES Mar.
Sci. Symp., 199: 157-166.
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-85. Exhibit 60. State Wat. Res. Ctrl.
Bd., Wat. Qual./Wat. Rights Proc. on San Fran.
Bay/Sac.- San Joaquin Delta. 337 pp.
Eggleston, D.B. and D.A. Armstrong. 1995. Pre- and
post-settlement determinants of estuarine Dunge-
Chapter 2 —
Estuarine Fish and Associated Invertebrates
75
Fish
Juvenile crabs appear to prefer sandy or sandy-mud substrate, but can be found on almost any bottom type (e.g.,
shell debris). Structurally complex habitats that provide
protection from predation (e.g., high relief shell, eel
grass, drift macroalgae, etc.) are favored over bare mud
or open sand (Fernandez et al. 1993a, Iribarne et al.
1995, Eggleston and Armstrong 1995, McMillan et al.
1995).
Chemical and physical characteristics of the water
column and sediment are also important habitat features.
Juvenile Dungeness crab in the San Francisco Estuary
seem to be somewhat intolerant of salinities lower than
10 ppt (Tasto 1983, CDFG 1987). Maximum growth
appears to occur at 15° C or above (Kondzela and Shirley
1993, McMillan et al. 1995); and studies in Washington State have shown that juvenile crab have stable metabolic rates at elevated estuarine temperatures (e.g., 14
to 16° C), whereas older crabs were more stable at colder
temperatures (Gutermuth and Armstrong 1989). This
is consistent with the tendancy for juvenile crabs to
emigrate out of estuaries into colder coastal waters as they
approach sexual maturity.
Although no single pollutant, or suite of pollutants,
has been shown to significantly affect Dungeness crab,
various studies on different life stages have shown sensitivity to oiled sediments, dissolved oxygen levels below
5 ppm, low ammonia concentrations, pesticides, and
chlorinated wastewater (Wild and Tasto 1983, Emmett
et al. 1991). Juvenile crab abundance in the Bay has been
shown to be negatively correlated to Delta outflow
(CDFG 1987).
ness crab recruitment. Ecological Monographs, 65
(2): 193-216.
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Volume
II: species life history summaries. ELMR Rep. No.
8 NOAA/NOS Strategic Envir. Ass. Div.,
Rockville, MD. 329 pp.
Fernandez, M.E., O. Iribarne and D.A. Armstrong.
1993a. Habitat selection by young-of-the-year
Dungeness crab Cancer magister Dana and predation risk in intertidal habitats. Mar. Ecol. Prog.
Ser., Vol. 92: 171-177.
Fernandez, M.E., D.A. Armstrong and O. Irbarne.
1993b. First cohort of young-of-the-year Dungeness crab (Cancer magister) reduces abundance of
subsequent cohorts in intertidal shell habitats. Can.
J. Fish. Aquat. Sci. 50: 2100-2105.
Gunderson, D.R., D.A. Armstrong, Y.-B. Shi and R.A.
McConnaughey. 1990. Patterns of estuarine use
by juvenile English sole (Parophrys vetulus) and
Dungeness crab (Cancer magister). Estuaries 13(1):
59-71.
Gutermuth, F. Brandt and D.A. Armstrong. 1989.
Temperature-dependent metabolic response of juvenile Dungeness crab (Cancer magister Dana): ecological implications for estuarine and coastal populations. J. Exp. Mar. Biol. Ecol. 126: 135-144.
Hatfield, S.E. 1983. Intermolt staging and distribution
of Dungeness crab, Cancer magister, megalopae.
In: Wild and Tasto (eds). Life history, environment,
and mariculture studies of the Dungeness crab,
Cancer magister, with emphasis on the central California fishery resource. Ca. Dept. Fish and Game,
Fish Bull. (172): 85-96.
Hines, A.H. 1991. Fecundity and reproductive output
in nine species of Cancer crabs (Crustacea,
Brachyura, Cancridae). Can. J. Fish. Aquat. Sci.
48: 267-275.
Iribarne, O., D. Armstrong and M. Fernandez. 1995.
Environmental impact of intertidal juvenile
Dungeness crab enhancement: effects on bivalves
and crab foraging rate. J. Exp. Mar. Biol. Ecol.
192: 173-194.
Kondzela, C.M. and T.C. Shirley. 1993. Survival, feeding, and growth of juvenile Dungeness crabs from
southeastern Alaska reared at different temperatures. Jour. of Crust. Biol., 13 (1): p. 25-35.
McCabe, G.T. Jr., R.L. Emmett, T.C. Coley and R.J.
McConnell. 1988. Distribution, density, and
size-class structure of Dungeness crabs in the
river-dominated Columbia River Estuary. Northwest Science 62(5): 254-262.
McConnaughey, R.A. and D.A. Armstrong. 1990. A juvenile critical stage in the Dungeness crab (Cancer
magister) life history. Abstracts of the 1990 annual
76
Baylands Ecosystem Species and Community Profiles
Fish
Amphibians &
Reptiles
Rock Crabs
Cancer antennarius and Cancer productus
Robert N. Tasto
General Information
The brown rock crab (Cancer antennarius) is found along
the west coast of North America from Washington State
to Baja California; the red rock crab (Cancer productus)
has a slightly more northerly distribution, i.e., Alaska to
San Diego (Carroll and Winn 1989). A small recreational
fishery exists for brown and red rock crabs in central San
Francisco Bay and parts of south Bay and San Pablo Bay.
Most rock crabs in this fishery are caught from piers and
jetties by a variety of baited hoop nets and traps. A modest commercial fishery also occurs throughout California waters, with the vast majority of the catch taking
place from Morro Bay southward (Parker 1992). Ex-vessel value for the commercial fishery approached $2 million in the mid-1980s (Carroll and Winn 1989) and
appears to be unchanged since then. Unlike their close
relative, the Dungeness crab, which has a significant
amount of muscle tissue in the body, rock crabs, generally, have been sought after for their claws only. In recent years, however, live whole crabs have become a
larger part of the retail market. California Department
of Fish and Game regulations prohibit the commercial
take of crabs less than 4.25 inches carapace width (cw),
require that sport-caught crabs must be 4.0 inches cw
or greater, and impose a bag limit of 35 crabs per day.
Reproduction
Mating takes place between a soft-shelled (recently
molted) female and hard-shelled male. Male brown rock
crabs have been reported to outnumber females by a ratio of 1.6/1 (San Mateo County coast) during all seasons
(Breen 1988), although studies by Carroll (1982) at
Diablo Cove showed that females were more abundant
in the fall, with no other seasonal trends for either sex.
Unfertilized eggs remain within the female for approximately three months, following mating, and then are
fertilized by the stored sperm as they are released (Parker
1992). The fertilized eggs are then carried until hatching (6 to 8 weeks) in a sponge-like mass beneath the
female’s abdominal flap (Parker 1992). Female body size
is the principal determinant of reproductive output and
fecundity, with red rock crab having 172,600 to 597,100
eggs per brood and brown rock crab having 156,400 to
5,372,000 eggs per brood (Hines 1991). Like the
Dungeness crab, ovigerous female rock crabs have been
observed buried in the sand at the base of rocks in shallow waters protecting their eggs (Reilly 1987). Also, some
red rock crab females have been detected emigrating out
Plants
meeting of the National Shellfisheries Assoc.
Williamsburg, VA. p. 133-134.
McMillan, R.O., D.A. Armstrong and P.A. Dinnel.
1995. Comparison of intertidal habitat use and
growth rates of two northern Puget Sound cohorts
of 0+ age Dungeness crab, Cancer magister. Estuaries 18(2): 390-398.
Oresanz, J.M. and V.F. Gallucci. 1988. Comparative
study of postlarval life-history schedules in four
sympatric species of Cancer (Decapoda: Brachyura:
Cancridae). Jour. Crust. Biol. 8(2): 187-220.
Palacios, R. and D.A. Armstrong. 1990. Predation of
juvenile soft-shell clam (Mya arenaria) by juvenile
Dungeness crab. Abstracts of the 1990 annual
meeting of the National Shellfisheries Assoc.
Williamsburg, VA. p. 445-446.
Reilly, P.N. 1983a. Predation on Dungeness crabs, Cancer
magister, in central Califirnia. In: Wild and Tasto
(eds). Life history, environment, and mariculture
studies of the Dungeness crab, Cancer magister,
with emphasis on the central California fishery resource. Ca. Dept. Fish and Game, Fish Bull. (172):
155-164.
______. 1983b. Dynamics of Dungeness crab, Cancer
magister, larvae off central and northern California. In: Wild and Tasto (eds). Life history, environment, and mariculture studies of the Dungeness crab, Cancer magister, with emphasis on the
central California fishery resource. Ca. Dept. Fish
and Game, Fish Bull. (172): 57-84.
Tasto, R.N. 1983. Juvenile Dungeness crab, Cancer magister, studies in the San Francisco Bay area. In: Wild
and Tasto (eds). Life history, environment, and
mariculture studies of the Dungeness crab, Cancer magister, with emphasis on the central California fishery resource. Ca. Dept. Fish and Game,
Fish Bull. (172): 135-154.
Wainwright, T.C. and D.A. Armstrong. 1993. Growth
patterns in the Dungeness crab (Cancer magister
Dana): synthesis of data and comparison of models. Jour. Crust. Biol. 13(1): 36-50.
Warner, R.W. 1992. Dungeness crab, p. 15-18. In: W.S.
Leet, C.M. Dewees and C.W. Haugen (eds).
California’s living marine resources. Ca. Sea Grant
Publ. UCSGEP-92-12, 257 pp.
Wild, P.N. and R.N. Tasto. 1983. Life history, environment, and mariculture studies of the Dungeness
crab, Cancer magister, with emphasis on the central California fishery resource. Ca. Dept. Fish and
Game, Fish Bull. (172): 352 pp.
Rock crabs are both nocturnal predators and scavangers
and have been shown to feed upon hard-shelled organisms such as clams, snails, and barnacles (Parker 1992).
The large chelae of these crabs is well-suited to forage
on the hard shells of more sedentary prey of their rocky
habitats (Oresanz and Gallucci 1988). Red rock crab feed
upon intertidal mussels and barnacles (Robles et al.
1989). Juvenile rock crabs are preyed upon by other macroinvertebrates and demersal fishes, whereas adults are
prey items for marine mammals (Carroll 1982). Very
little is known about the specific food habits of, or predators upon, these two species of rock crabs within San Francisco Bay; however, the sportfishery within the Bay accounts
for the loss of an indeterminate number of adult crabs.
CDFG
Brown rock crabs are known to go through 10 to 12
molts before reaching sexual maturity at about 3 inches
cw, and will likely molt one to two times per year thereafter (Parker 1992). The average number of red rock crab
instars is 13 over the total life span (Oresanz and Gallucci
1988). Studies in Humboldt Bay (O’Toole 1985) found
Food and Feeding
Brown Rock Crab, Cancer antennarius.
Top and bottom: Views of 5 in. male crab.
CDFG
Growth and Development
ovigerous red rock crab as small as 3.7 inches cw. Brown
rock crabs have reached a maximum 6.5 inches cw and
red rock crabs, the larger of the two species, at 8 inches
cw (Carroll and Winn 1989, Parker 1992). Maximum
life span of the brown rock crab has been estimated at
5-6 years (Carroll 1982).
Red Rock Crab, Cancer productus.
Top: top surface of 6.5 in. male. Bottom: under
surface of 5.75 in. female.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
77
Fish
of estuaries prior to spawning to avoid osmotic stress
(Oresanz and Gallucci 1988).
Hatching takes place in spring and early summer
in central California (Carroll 1982). The planktonic larvae then settle to the bottom before beginning the juvenile stage. Juvenile abundance is highest in San Francisco Bay during the summer months (CDFG 1987).
Much like other cancrids, larval release in spring coincides with peak plankton production, and settlement in
the summer is optimal for growth (Hines 1991). The reproductive life span for the red rock crab is approximately
four years with four broods, and for the brown rock crab
it is approximately seven years with up to 10 broods
(Hines 1991).
(Carroll 1982, Breen 1988). Small, local populations of
rock crab can be overfished, although there is no evidence suggesting that overfishing occurs in the Bay. Data
from the Interagency Ecological Study Program indicate
that there is a negative relationship between abundance
of both rock crab species and outflow from the Delta
(CDFG 1987).
Both rock crab species inhabit the low intertidal zone
to depths of 300 feet or more (Parker 1992) and, although their microhabitat utilization patterns are similar, they appear to be different in how they utilize estuaries (Oresanz and Gallucci 1988). The brown rock crab
is principally a marine species and does not osmoregulate
well in brackish waters, whereas the red rock crab can
successfully inhabit brackish areas. All stages of the red
and brown rock crab have been collected in San Francisco Bay, including larvae and ovigerous females (Tables
2.1 and 2.2). Areas of peak abundance appear to be in
Central Bay, the northern portion of South Bay, and the
southern portion of San Pablo Bay, with the red rock
crab having a somewhat greater distribution than the
brown rock crab (CDFG 1987). In general, rock crab
movement is local (Breen 1988, Carroll and Winn
1989). At Fitzgerald Marine Reserve along the San
Mateo County coast, studies demonstrated that juvenile
brown rock crab are most abundant in July, although no
seasonal trend in the settlement of early instars was evident (Breen 1988). In Santa Barbara County, Reilly
(1987) found all stages of rock crabs to be most abundant in the fall.
Trophic Levels
Rockcrab larvae are planktivores and, as such, are both
primary consumers (phytoplankton) and secondary consumers (zooplankton). Juveniles and adults are higher
order consumers.
Proximal Species
Predators: Marine mammals, humans (recreational fishery).
Prey: Bay mussels, barnacles.
Good Habitat
Not surprisingly, both species have been shown to prefer rocky shore, subtidal reef, or coarse gravel and sand
substrate (Carroll and Winn 1989). Opportunity for concealment appears to be an important feature of red rock
crab habitat in British Columbia studies (Robles et al.
1989). Juvenile brown rock crab, when settling from the
last larval stage, appear to accept both sand and rock as
suitable substrate (Carroll and Winn 1989), and red rock
crab also tend to settle out onto structurally complex
substrates (Oresanz and Gallucci 1988).
Population Status and Influencing Factors
There are no known estimates of the overall population
size or knowledge of recruitment mechanisms for San
Francisco Bay rock crabs. Most studies have shown that
population densities of rock crabs were well below 1/m2
Table 2.1 Annual Abundance of Rock Crabs Caught by Otter Trawl (crabs/tow) in the San Francisco
Estuary (CDFG 1987)
Species and Size Class
1980
1981
1982
1983
1984
1985
1986
C. antennarius (all sizes)
0.101
0.047
0.010
0.015
0.071
0.033
0.007
C. antennarius (<50mm)
0.098
0.037
0.005
0.015
0.067
0.024
0.007
C. gracilis (all sizes)
0.035
0.103
0.044
0.182
0.333
0.240
0.174
C. gracilis (<20mm)
0.003
0.005
0.034
0.080
0.079
0.064
0.095
C. productus (all sizes)
0.014
0.032
0.005
0.010
0.055
0.071
0.088
C. productus (<50mm)
0.014
0.027
0.002
0.005
0.040
0.050
0.081
Table 2.2 Annual Abundance of Rock Crabs Caught by Ring Net (crabs/tow) in the San Francisco
Estuary (CDFG 1987)
Species and Size Class
1980
1981
1982*
1983
1984
1985
1986
C. antennarius (all sizes)
-
-
0.113
0.095
0.296
0.491
0.407
C. antennarius (<50mm)
-
-
0
0.009
0.028
0.009
0.176
C. gracilis (all sizes)
-
-
0.014
0.019
0.037
0.009
0.130
C. productus (all sizes)
-
-
0.155
0.067
2.509
4.315
0.806
C. productus (<50mm)
-
-
0
0
0.185
0.148
0.157
* Ring net survey started in May 1982
78
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Fish
Distribution
Sulkin, S.D. and G. McKeen. 1994. Influence of temperature on larval development of four co-occurring species of the brachyuran genus Cancer. Marine Biology. 118: 593-600.
Bat Ray
Myliobatus californica
Kurt F. Kline
References
General Information
The bat ray is a member of the family Myliobatidae
(eagle rays). The family is found worldwide in tropical
and temperate shallow seas. Bat rays are very common
and are found in sandy and muddy bays and sloughs, as
well as in rocky areas and kelp beds. In shallow bays they
can be found feeding in the intertidal zone during high tide.
Reproduction
Mating occurs during the summer months followed by
an estimated gestation period of nine to 12 months (Martin and Cailliet 1988). The young are born alive at 220
to 356 mm wing width and weigh about 0.9 kg (Baxter
1980, Martin and Cailliet 1988). Males are mature at
450 to 622 mm wing width and two to three years, while
50% of the females are mature at 881 mm wing width
and five years.
Growth and Development
The growth of juvenile bat rays is not well documented,
but is likely at least 100 mm per year. They can grow to
CDFG
Breen, R.T. 1988. Sizes and seasonal abundance of rock
crabs in intertidal channels at James V. Fitzgerald
Marine Reserve, California. Bull. Southern Ca.
Acad. Sci. 87(2): 84-87.
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-85. Exhibit 60. State Wat. Res. Ctrl.
Bd., Wat. Qual./Wat. Rights Proc. on San Fran.
Bay/Sac.-San Joaquin Delta. 337 pp.
Carroll, J.C. 1982. Seasonal abundance, size composition, and
growth of rock crab, Cancer antennarius Stimpson, off
central California. J. Crust. Biol. 2: 549-561.
Carroll, J.C. and R.N. Winn. 1989. Species profiles:
life histories and environmental requirements of
coastal fishes and invertebrates (Pacific Southwest)—brown rock crab, red rock crab, and yellow crab. U.S. Fish Wildl. Serv. Biol. Rep.
82(11.117). U.S. Army Corps of Engineers, TR
EL-82-4. 16 pp.
Hines, A.H. 1991. Fecundity and reproductive output
in nine species of Cancer crabs (Crustacea,
Brachyura, Cancridae). Can. J. Fish. Aquat. Sci.
48: 267-275.
Oresanz, J.M. and V.F. Gallucci. 1988. Comparative
study of postlarval life-history schedules in four
sympatric species of Cancer (Decapoda: Brachyura:
Cancridae). Jour. Crust. Biol. 8(2): 187-220.
O’Toole, C. 1985. Rock crab survey of Humboldt Bay.
U.C. Coop. Ext., Sea Grant Mar. Adv. Prog., Eureka, Ca., Interim Rep. 14 pp.
Parker, D. 1992. Rock Crabs. In: W.S. Leet, C.M.
Dewees and C.W. Haugen (eds). California’s living marine resources. Ca. Sea Grant Publ.
UCSGEP-92-12, 257 pp.
Reilly, P.N. 1987. Population studies of rock crab, Cancer antennarius, yellow crab, Cancer anthonyi, and
Kellet’s whelk, Kelletia kelletii, in the vicinity of
little Coho Bay, Santa Barbara County, California.
Ca. Dept. Fish and Game. 73: 88-98.
Robles, C., D.A. Sweetnam and D. Dittman. 1989. Diel variation of intertidal foraging by Cancer productus L. in British Columbia. Jour. Nat. Hist. 23: 1041-1049.
Fish
Rock crabs appear to be influenced by both temperature and salinity. In various laboratory studies, both
brown and red rock crab were adversely affected by exposure to water temperatures above 20° C (Carroll and
Winn 1989, Sulkin and McKeen 1994). The brown rock
crab is considered primarily a marine species, whereas
red rock crabs can osmoregulate in more brackish water; although the latter have been shown to be adversely
affected by salinities below 13 ppt (Oresanz and Gallucci
1988, Carroll and Winn 1989).
Chapter 2 —
Estuarine Fish and Associated Invertebrates
79
a wingspan of six feet (>2000 mm) though individuals
this large are uncommon. The largest bat ray reported
was a 95 kg female from Newport Bay (Baxter 1980).
Good Habitat
Sandy to muddy shallow bottoms with abundant mollusk and polychaete populations.
Fish
Food and Feeding
Bat rays are opportunistic bottom feeders, feeding primarily upon benthic and epibenthic invertebrates. In
Elkhorn Slough, bat rays feed primarily on clams and the
echiuroid worm, Urechis caupo; in La Jolla kelp beds, they
feed on shellfish including abalone and snails; and in
Tomales Bay, they feed on polychaete worms, large clams
and echiuroid worms (Karl and Obrebski 1976, Karl
1979, Talent 1982). Studies done along the southern
California coast (Van Blaricom 1982) found that pits dug
by feeding bat rays were an important controlling factor of
infaunal community organization, opening areas for infauna
recolonization and uncovering food items for other fish.
Distribution
Population Status and Influencing Factors
The current status of the bat ray in San Francisco Bay
is unknown. Its distribution is likely influenced by salinity; it has occaisionally been collected in San Pablo Bay
at salinities lower 20 ppt (Flemming 1999).
Trophic Levels
Bat rays are primary consumers, feeding primarily on
benthic invertebrates. They are taken by fishermen using cut fish as bait, however natural feeding on fishes
has not been documented.
Proximal Species
Prey: Benthic mollusks, polychaetes, crustaceans,
Urechis caupo.
80
Baylands Ecosystem Species and Community Profiles
Baxter, J.L. 1980. Inshore fishes of California. Ca. Dept.
Fish and Game. 72 pp.
Flemming, R. 1999. Elasmobranchs. In: J. Orsi (ed).
Report on the 1980-1995 fish, shrimp, and crab
sampling in the San Francisco Estuary, California.
IEP Technical Report 63.
Karl, S.R. 1979. Fish feeding habit studies from Tomales Bay, California. M.S. Thesis, Univ. of Pacific,
Stockton, Ca., 44 pp.
Karl, S.R. and S. Obrebski. 1976. The feeding biology
of the bat ray, Myliobatis californica, in Tomales
Bay, California. In: C.A. Simsenstad and S.J.
Lipovsk (eds). Fish food habits studies, 1st Pacific
Northwest Technical Workshop, Workshop Proceedings, October 13- 15. Washington Sea Grant,
Div. Mar. Res., Univ. of Washington HG :30, Seattle, pp. 181-186.
Martin, L.K. and G.M. Cailliet 1988. Age and growth
determination of the bat ray, Myliobatis californica
Gell, in Central California. Copeia 3: 763 773.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Dept. Fish and
Game Fish Bull. 157, 249 pp.
Talent, L.G. 1982. Food habits of the gray smoothhound, Mustelus californicus, the brown smoothhound, Mustelus henlei, the shovelnose guitarfish,
Rhinobatos productus, and the bat ray, Myliobatis
californica, in Elkhorn Slough, California. Ca.
Dept. Fish and Game 68(4): 224-234.
Van Blaricom, G.R. 1982. Experimental analyses of
structural regulation in a marine sand community
exposed to oceanic swell. Ecological Monographs
52: 283-305.
Plants
Amphibians &
Reptiles
The bat ray ranges from the Gulf of California to Oregon, and is found from shallow subtidal water to 46 m.
It is common in bays and shallow sandy areas along the
coast (Miller and Lea 1976).
References
Leopard Shark
Triakis semifasciata
Michael F. McGowan
General Information
Reproduction
The leopard shark is a live bearer with internal fertilization, but no yolk-sac placenta. Mating occurs in the
spring, primarily during April and May soon after the
females give birth to from 4-29 pups (Compagno 1984).
Pupping can occur from March through August with a
peak in April or May (Ackerman 1971). In San Francisco Bay leopard sharks pup almost exclusively in South
Bay (CDFG Bay Trawl data). The center of abundance
of pups <300 mm long is south of, and just north of the
Dumbarton Bridge.
Growth and Development
CDFG
Embryonic development is direct and internal and takes
10-12 months. At birth pups are 18-20 cm long. Females
mature when 12-14 years old at a length of 110-129 cm.
Food and Feeding
Primary foods of the leopard shark are benthic and
epibenthic crustaceans, clam siphons, echinuroid worms,
and small fishes.
Fish
The leopard shark (Family: Elasmobranchs) is one of the
most common sharks in California bays and estuaries
(Talent 1976). It is the most abundant shark in San Francisco Bay (Ebert 1986) being found especially around
piers and jetties (Emmett et al. 1991). The leopard shark
is an important recreational species in San Francisco Bay
and a limited commercial long-line fishery has targeted
it in the bay (Smith and Kato 1979). Juveniles and adults
are demersal and sometimes rest on the bottom (Feder
et al. 1974). Although other elasmobranchs occur in
euhaline bays and estuaries of the U. S. Pacific coast, the
leopard shark was the only shark or ray included among
47 fish and invertebrate species in the life history summaries of west coast estuarine species prepared by the
National Oceanic and Atmospheric Administration’s
Estuarine Living Marine Resources (ELMR) program
(Emmett et al. 1991). These species were selected on the
basis of commercial value, recreational value, indicator
species of environmental stress, and ecological importance. That the leopard shark was selected is an indication of its importance in estuaries in general and in San
Francisco Bay where it is the most abundant shark.
Males mature earlier and at smaller sizes than females.
Growth rates are slow. In San Francisco Bay tagged leopard sharks grew 1.4 cm/yr (Smith and Abramson 1990).
Distribution
The leopard shark is found from Mazatlan, Mexico including the Sea of Cortez to Oregon (Miller and Lea
1976). In California it is most common in estuaries and
bays south of Tomales Bay (Monaco et al. 1990). Leopard sharks are apparently resident in San Francisco Bay,
although some move out in fall and winter (Smith and
Abramson 1990) and several size classes appear in the
California Department of Fish and Game length data.
Population Status and Influencing Factors
The leopard shark probably has no predators except
larger sharks and humans. Its broad dietary range should
protect it from food limitation. Heavy fishing mortality
poses a threat to the leopard shark, as it does to all sharks,
because of its slow growth, long time to maturity, and
low fecundity. The minimum size limit recommended
by Smith and Abramson (1990) for sustainable fishing
in San Francisco Bay was 100 cm (40 in). Areas of high
freshwater input causing low salinity are largely avoided
by leopard sharks.
Trophic Levels
Juveniles and adults are secondary and higher carnivores.
Proximal Species
Predators: Larger sharks, humans.
Prey: Yellow shore crab, Urechis caupo, ghost shrimp,
rock crabs, octopus, shiner perch, arrow goby, Pacific
herring, northern anchovy, topsmelt.
Cohabitors: Smoothhound sharks form mixed schools
with leopard sharks.
Good Habitat
Leopard sharks are primarily a marine species which occupies bays and estuaries unless freshwater flows lower salinity excessively. Sandy and muddy bottom areas are preferred, although they may be found near rocky areas and
kelp beds along the coast. Estuaries are used as pupping and
rearing areas for young sharks. Shallow mud and sand flats
are used for foraging during high tide (Compagno 1984).
Chapter 2 —
Estuarine Fish and Associated Invertebrates
81
Pacific Herring
Clupea pallasi
Robert N. Tasto
General Information
The Pacific herring (Family: Clupeidae) resource in the
San Francisco Estuary is widely recognized for its commercial, recreational, and ecological values. The commercial fishery concentrates on ripe females for their roe
(eggs) which are then exported to Japan, although there
is some limited effort for the fresh fish market and for
live bait by recreational salmon trollers (Spratt 1981,
Lassuy 1989). Fishermen traditionally catch herring in
nearshore areas of the Bay with gillnets or in deeper waters with round-haul nets, and there also is a relatively
new roe-on-kelp fishery operated from rafts (Spratt 1981,
CDFG 1992). The economic value of the fishery based
upon ex-vessel prices paid to the fishermen in 1995-96 was
approximately 16.5 million dollars (CDFG, unpub. data).
Reproduction
Adult herring congregate outside of San Francisco Bay
before entering and generally spend about 2 weeks in the
Bay before spawning (CDFG 1987). Spawning takes
place from early November through March, with peak
activity in January (Spratt 1981, CDFG 1992). The timing of spawning is believed to coincide with increased
levels of plankton production as a food source for larvae
(Lassuy 1989), as well as the presence of freshwater flows
(Emmett et al. 1991). Pacific herring spawn primarily
on vegetation, rock rip-rap, pier pilings, and other hard
substrates in intertidal and shallow subtidal waters
(Spratt 1981, Lassuy 1989, Emmett et al. 1991). Spawning occurs in waves of 1 to 3 days, occasionally up to a
week in length, and often at night in conjunction with
high tides (Spratt 1981). Waves are separated by one to
several weeks over the length of the season with larger
fish tending to spawn first (Lassuy 1989). The number
and size of the waves is related to the distribution of the
dominant year classes (CDFG 1992).
Egg-deposition is thought to be facilitated by the
brushing of the female’s vent up against the substrate,
and, while there is no pairing of the sexes, the spawning area will be white with milt from the males so that
the rate of fertilization is usually high (Hart 1973). Pa-
CDFG
Ackerman, L.T. 1971. Contributions to the biology of
the leopard shark, Triakis semifasciata (Girard) in
Elkhorn Slough, Monterey Bay, California. M.A.
Thesis, Sacramento State College, Sacramento, CA,
54 pp.
Compagno, L.J. 1984. FAO species catalogue. Vol. 4.
Sharks of the world. An annotated and illustrated
catalogue of shark species known to date. Part 2.
Carcharhiniformes. FAO Fish. Synop. 125(4): 433434.
Ebert, D.A. 1986. Observations on the elasmobranch
assemblage of San Francisco Bay. Ca. Dept. Fish
and Game 72(4): 244-249.
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Volume
II: species life history summaries. ELMR Rep. N.
8. NOAA/NOS Strategic Environmental Assessments Div., Rockville, MD, 329 pp.
Feder, H.M., C.H. Turner and C. Limbaugh. 1974.
Observations on fishes associated with kelp beds
in southern California. Ca. Dept. Fish Game, Fish
Bull. 160, 144 pp.
Miller, D.J. and R.N. Lea. 1976. Guide to the coastal
marine fishes of California. Fish Bull. 157, Ca.
Dept. Fish and Game , Sacramento, Ca.. Sea Grant
reprint of 1972 edition with addendum added
1976.
Monaco, M.E., R.L. Emmett, S.A. Hinton and D.M.
Nelson. 1990. Distribution and abundance of
fishes and invertebrates in West Coast estuaries,
volume I: data summaries. ELMR Report No. 4.
Strategic Ass. Div., NOS/NOAA, Rockville, Maryland. 240 pp.
Smith, S.E. and N. J. Abramson. 1990. Leopard shark
Triakis semifasciata distribution, mortality rate,
yield, and stock replenishment estimates based on
a tagging study in San Francisco Bay. Fish. Bull
88: 371-381.
Smith, S.E. and S. Kato. 1979. The fisheries of San
Francisco Bay: past, present and future. In: T.J.
Conmos (ed). San Francisco Bay: The urbanized
estuary, Pac. Div., Am. Assoc. Adv. Sci., San Francisco, Ca. pp. 445-468.
Talent, L.G. 1976. Food habits of the leopard shark,
Triakis semifasciata, in Elkhorn Slough, Monterey
Bay, California. Ca. Fish Game 62(4): `286-298.
82
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Fish
References
vertebrates and various fishes, while juveniles and adults
are consumed by a variety of fishes (e.g., spiny dogfish
shark, Chinook salmon, Pacific staghorn sculpin, and
striped bass), seabirds (e.g., Brandts cormorants, brown
pelicans, and western gulls), and marine mammals, such
as harbor seals (Hart 1973, Lassuy 1989, Emmett et al.
1991). Predation is considered to be the greatest source
of natural mortality for juvenile and adult Pacific herring (CDFG 1992).
Fish
cific herring eggs adhere to the substrate in amounts
ranging from a few eggs to as many as eight layers thick
(Spratt 1981). The fecundity of herring is approximately
4,000 to 134,000 eggs per female, depending upon its
distribution and size (Hart 1973, Emmett et al. 1991).
As with spawning, most hatching takes place at night,
and will occur in 10 to 15 days under 8.5° to 10.7° C
temperatures; longer if the water is colder (Emmett et
al. 1991). The average in San Francisco Bay is 10.5 days
at 10.0° C (CDFG 1992).
Distribution
Growth and Development
Pacific herring eggs are approximately 1.0 mm in diameter, and 1.2 to 1.5 mm after fertilization (Hart 1973).
A newly hatched larva, with yolk sac, is about 6 to 8 mm
total length (TL) and will develop swimming powers at
about 20 mm TL (CDFG 1992). Metamorphosis to the
juvenile stage occurs from 25 to 35 mm TL and takes
place over two to three months (Emmett et al. 1991).
They are free-swimming at this stage and begin to form
shoreline-oriented schools (CDFG 1992). Juveniles are
35 to 150 mm TL depending upon regional growth
rates, which in turn are affected by population size and
environmental conditions (Emmett et al. 1991). In the
Bay Area, there are no apparent differences in the growth
rates of males and females (Spratt 1981). Adults range
in size from 130 to 260 mm TL, and locally it takes two
to three years to reach maturity (Spratt 1981, Emmett
et al. 1991). The San Francisco Bay population ranges
from 110 to 250 mm TL (CDFG 1992; Ken Ota, Pers.
Comm.) It is possible that some Pacific herring in more
northern climates may exceed 15 years in age, but few
have been noted to live longer than nine years (Emmett
et al. 1991).
Food and Feeding
Pacific herring larvae, juveniles, and adults are selective
pelagic planktonic feeders and move toward the water’s
surface to feed at dusk and dawn (Emmett et al. 1991).
Generally, prey items will change with growth and geographic distribution. Larvae feed on diatoms, invertebrate and fish eggs, crustacean and mollusc larvae,
bryzoans, rotifers, and copepods (Hart 1973). Juveniles
consume a variety of crustaceans, as well as mollusc and
fish larvae; while adults eat mostly planktonic crustaceans
and fish larvae (Hart 1973, Emmett et al. 1991). In
winter, there is an overall reduction in adult Pacific herring feeding as stored energy is used for ripening reproductive products and, during their spawning migration
and inshore “ holding” period, herring may severely limit
or stop feeding entirely (Lassuy 1989).
Herring eggs are eaten by various species of fish
(e.g., sturgeon), ducks (e.g., surf scoter), and gulls
(CDFG 1992). Larvae are often prey for large pelagic in-
Major populations exist in the eastern Pacific between
San Francisco Bay and central Alaska (Hart 1973).
Within San Francisco Bay, the principal spawning areas are found along the Marin County coastline (i.e.,
Sausalito, Tiburon Penninsula, and Angel Island), at the
San Francisco waterfront and Treasure Island, on the east
side of the Bay from the Port of Richmond to the Naval
Air Station at Alameda, and on beds of vegetation in
Richardson Bay and South Bay (Figure 2.3) (Spratt
1981, CDFG 1992). After hatching, the larvae are
clumped and controlled largely by tidal factors, and following disappearance of the yolk sac and the onset of
feeding, their distribution becomes patchy (CDFG
1992). Larvae and young juveniles are found in the Bay
between November and April and their greatest densities are in the shallow waters of upper South Bay, Central Bay, and San Pablo Bay. Juveniles are found in the
deeper areas of the Bay (peak in Central Bay) between
April and August, and, for the most part, have left the
Bay by late June at sizes that approach 80 mm TL
(CDFG 1987). They eventually move to offshore or
nearshore areas and do not return to the Bay until they
are mature and ready for spawning. There is conflicting
evidence of a strong correlation between juvenile abundance, as measured by young-of-the-year surveys, and recruitment to the adult spawning population two years
later (Herbold et al. 1992)
Population Status and Influencing Factors
San Francisco Bay population levels fluctuate widely and
have ranged between approximately 6,000 tons and
100,000 tons spawning biomass, as measured by spawn
deposition surveys and hydroacoustic monitoring of fish
schools (CDFG 1992). 1995-96 season estimates were
approximately 99,000 tons, second highest on record
(CDFG, unpub. data). Year-class strength is often determined in the first six months of life (Hart 1973,
Lassuy 1989, Emmett et al. 1991). Egg mortalities can
result from tidal exposure and dessication, abrubt or severe temperature or salinity changes, low oxygen levels,
wave action, suffocation by high egg densities or siltation, pollution, and predation (Lassuy 1989, Emmett
et al. 1991). Factors related to natural mortality of larvae
Chapter 2 —
Estuarine Fish and Associated Invertebrates
83
Plants
Amphibians &
Reptiles
Fish
Figure 2.3 Traditional Pacific
Herring Spawning Areas in
Central San Francisco Bay
in the Bay include competition and other density dependent mechanisms, as well as starvation during their initial
feeding period and changes in dispersal patterns. Juveniles
and adult survival is affected by competition, predation, disease, spawning stress, and fishing (Emmett et al. 1991).
Predation appears to be the single most important
factor affecting population levels (Lassuy 1989). In addition to commercial and recreational fishing, humans
influence herring survival by impacting water and habitat quality. Spawning habitat quantity and Delta outflows are not thought currently to be limiting factors in
determining the Bay’s herring population size (CDFG
1987 and 1992).
Trophic Levels
Larvae are planktivores (primary and secondary consumers). Juveniles and adults are primary and higher order
consumers.
Proximal Species
Egg Predators: Gulls, diving ducks, white sturgeon,
atherinids (topsmelt and jacksmelt), surf perches, rock crabs.
84
Baylands Ecosystem Species and Community Profiles
Larvae predators: Young salmonids, pelagic invertabrates.
Juvenile Predators: California halibut, young salmonids, harbor seals, harbor porpoise.
Adult Predators: California halibut, California sea lion,
harbor seals, harbor porpoise.
Habitat: Eel grass (spawning substrate).
Prey: Striped bass, copepods.
Good Habitat
It is frequently stated that herring prefer sea grasses (e.g.,
Zostera marina) or algae (e.g., Gracilaria sp.) as spawning substrate (Lassuy 1989, Emmett et al. 1991); however, a variety of seemingly less attractive surfaces have
proven to be very successful in the Estuary. Rigidity,
smooth texture, and the absence of sediment appear to
be important components of suitable substrates (Lassuy
1989). Larvae and juveniles need quiescent and productive shallow subtidal areas as rearing habitat.
In northern waters, the optimal salinity range for
spawning is reported to be 8 to 22 ppt and 13 to 19 ppt
for eggs and larval survival (CDFG 1987). Also in these
areas, temperatures in the range of 5.5 to 8.7° C have
References
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-85. Exhibit 60. State Wat. Res. Ctrl.
Bd., Wat. Qual./Wat. Rights Proc. on San Fran.
Bay/Sac.-San Joaquin Delta. 337 pp.
______. 1992. Pacific herring commercial fishing regulations. Draft environmental document. 183 pp.
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in West Coast estuaries, Volume
II: species life history summaries. ELMR Rep. No.
8 NOAA/NOS Strategic Environmental Ass. Div.,
Rockville, MD, 329 pp.
Hart, J.L. 1973. Pacific herring. In: Pacific fishes of
Canada. Fisheries Research Bd. of Canada, Bull.
180: 96-100.
Herbold, B., A.D. Jassby and P.B. Moyle. 1992. Status
and trends report on aquatic resources in the San
Francisco Estuary. U.S. Env. Prot. Agency, San
Fran. Est. Proj. 368 pp.
Lassuy, D.R. 1989. Species profiles: life histories and
environmental requirements of coastal fishes and
invertebrates (Pacific Northwest)—Pacific herring.
U.S. Fish and Wildl. Serv. Biol. Rep. 82 (11.126).
U.S. Army Corps of Engrs., TR-EL-82-4. 18 pp.
Spratt, J.D. 1981. Status of the Pacific herring, Clupea
harengus pallasii, resource in California, 1972 to
1980. Ca. Dept. Fish and Game, Fish Bull. (171).
107 pp.
Northern Anchovy
Engraulis mordax
Michael F. McGowan
General Information
The northern anchovy (Family: Engraudidae) has the
largest biomass and is the most abundant fish in San
Francisco Bay (Aplin 1967). It is an important forage
species for larger predators and consumes substantial
amounts of phytoplankton and zooplankton (McGowan
1986). There is a bait fishery for northern anchovy at
the mouth of the Bay. Most of the stock occurs outside
the Bay in the California Current. Although northern anchovy can be found inside the bay throughout
the year, their seasonal peak is generally April to October. The spring influx may be associated with the
onset of coastal upwelling (P. Adams, pers. comm.).
Their exodus in the autumn may be linked to cooling water temperatures inside the bay (McGowan
1986).
Reproduction
Northern anchovy spawn oval, pelagic eggs approximately 1.5 x 0.75 mm in size. Peak spawning is
thought to occur at night at about 10 pm. Females
can produce up to 130,000 eggs per year in batches
of about 6,000. The eggs hatch in approximately 48
hours depending on temperature. Larvae were collected in Richardson Bay within San Francisco Bay by
Eldridge (1977). Spawning was documented in San
Francisco Bay in 1978 by collections of eggs and larvae from south of the Dumbarton Bridge to San Pablo
Bay (McGowan 1986). Based on differential distributions of eggs and larvae, spawning occurs in the channels while larvae seek out the productive shallows.
Although the biomass of northern anchovy within the
bays is small relative to that in the California Current,
the bay is a favorable habitat for reproduction because
of ample food for adults to produce eggs, abundant
zooplankton prey for larvae, and protection of eggs
and larvae from offshore transport to less productive
areas by coastal upwelling.
Personal Communications
CDFG
Ken Ota. Ca. Dept. of Fish and Game, Pacific Herring
Research Project, 1996.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
85
Fish
been shown to be best for egg development (Emmett et
al. 1991); however, 10 to 12° C temperatures are about
average for the spawning grounds in San Francisco Bay
(Lassuy 1989). Optimal temperatures for juveniles and
adults appear to be a few degrees higher than for eggs
or larvae (Lassuy 1989). It has been suggested that eggs
need a minimum dissolved oxygen concentration of 2.5
mg/L at the surface and, therefore, eggs elevated from
the bottom on vegetation or other structures avoid siltation and receive better circulation for waste removal and
oxygenation (Lassuy 1989). Water quality is an important factor as eggs are vulnerable to high levels of suspended particulate matter, particularly if the sediments
are laden with contaminants (e.g., dredged material from
urban ports). Additionally, larvae have been shown to
be sensitive to the water-soluble fraction of hydrocarbons
from spilled oil or other sources (Lassuy 1989).
Growth and Development
Fish
Larvae grow from 2.5 to 25 mm in about two months,
at which time they are considered juveniles. Growth is
rapid within the warm productive bay environment.
Based on analysis of length frequencies, some juveniles
that were spawned late in the summer overwinter in the
bay (McGowan 1986). The others apparently depart at
the same time as the adults in autumn.
Food and Feeding
Larvae eat dinoflagellates and zooplankton, while adults
filter-feed in dense patches of large phytoplankton or
small zooplankton, but selectively pick larger zooplankters from the water (O’Connell 1972).
The northern anchovy occurs from Queen Charlotte Islands, Canada to Cabo San Lucas, Baha California and
into the Sea of Cortez. It can be found in all estuaries
within this range. There is a subpopulation which occupies the Columbia River plume, an “ offshore estuary.”
In San Francisco Bay, they occur from Suisun Bay to
South Bay, but are most abundant downstream of the
Carquinez Strait (Herbold et al. 1992). There is a seasonal influx of northern anchovy into the bay in spring
when water temperatures and plankton production begin to rise in the bay and when nearshore upwelling generally begins. Adults exit the bay in autumn, but some
late-spawned juveniles may overwinter within the bay.
Population Status and Influencing Factors
Northern anchovy populations off California range in
the hundreds of thousands of tons. Their biomass increased dramatically following the decline of the sardine
stock, suggesting that competitive interactions might
control population fluctuations. Historical records of fish
scales in sediments suggests that large fluctuations in
both anchovy and sardine populations have occurred in
the past and were not strongly correlated with each other.
Variable survival of eggs and larvae due to environmental factors probably influences population size more
than predation or fishing. Active research into the causes
of northern anchovy population dynamics has contributed immensely to our understanding but without resolving whether starvation, predation, advection, or other
cause is the key limiting factor.
Proximal Species
Predators: California halibut, Chinook and coho
salmon; rockfishes, yellowtail, tunas, sharks, and almost
all California current fish; harbor seal; northern fur seal;
sea lions; common murre; brown pelican; sooty shearwater; cormorant spp.
Potential Competitors: Sardine. Jacksmelt, topsmelt,
and other schooling planktivores are potential competitors and predators on young life stages.
Good Habitat
Northern anchovy occupy near surface waters where the
water temperature should be between 10° and 25° C.
Eggs tend to be in water with salinities from 32-35 ppt,
but juveniles and adults are abundant in fresher bays and
estuaries as well as marine waters. Spawning in San Francisco Bay occurs at higher temperatures and lower salinities than spawning in coastal areas. Northern anchovy
are typical species of areas with high production such as
coastal upwelling regions and estuaries.
References
Aplin, J.A. 1967. Biological survey of San Francisco Bay
1963-1966. Report for Ca. Dept. Fish and Game,
MRO, Sacramento, Ca. Ref. 67-4. 131pp.
Eldridge, M.B. 1977. Factors influencing distribution
of fish eggs and larvae over eight 24-hour samplings in Richardson Bay, California. Ca. Dept.
Fish Game 63: 101-106.
Herbold, B., A.D. Jassby and P.B. Moyle. 1992. Status
and trends report on aquatic resources in the San
Francisco estuary. The San Francisco Estuary
Project, Oakland, Calif., 257 pp. plus apps.
McGowan, M.F. 1986. Northern anchovy, Engraulis
mordax, spawning in San Francisco Bay, California 1978-1979, relative to hydrography and zooplankton prey of adults and larvae. Fish. Bull., U.S.
84(4):879-894.
O’Connell, C.P. 1972. The interrelation of biting and
filtering in the feeding activity of the northern anchovy (Engraulis mordax). J. Fish. Res. Board Can.
29:285-293.
Personal Communications
Trophic Levels
First-feeding larvae may eat phytoplankters, larger larvae selectively pick copepods and other zooplankters
from the water, juveniles and adults pick or filter plank-
86
Baylands Ecosystem Species and Community Profiles
P. Adams, National Marine Fisheries Service, Tiburon.
Plants
Amphibians &
Reptiles
Distribution
ton, fish eggs, and fish larvae, depending on food concentrations. Larvae and older stages should be considered as secondary and higher consumers.
Sacramento Splittail
Pogonichthys macrolepidotus
Ted R. Sommer
General Information
Reproduction
Adult splittail generally reach sexual maturity at about
2 years of age (Caywood 1974). Some males mature at
the end of their first year and a few females mature in
their third year. An upstream spawning migration occurs
November through May, with a typical peak from
January-March. Spawning is thought to peak during
February-June, but may extend from January-July.
Although submerged vegetation is thought to be the preferred spawning substrate, egg samples have not yet been
collected on any substrate. Reproductive activity appears
to be related to inundation of floodplain areas, which
provides shallow, submerged vegetation for spawning,
rearing and foraging (Caywood 1974, Sommer et al.
1997). Splittail have high fecundity like most cyprinids.
Reported fecundities range from 5,000 to 266,000
eggs per female, depending on age (Daniels and Moyle
1983). Generally, female splittail will have more than
100,000 eggs each year.
Growth and Development
Ted Sommer
The morphological characteristics of splittail eggs, larvae,
and juveniles have been described and recent culturing
studies (Bailey 1994) are providing preliminary information on early life history requirements and development.
Very little is known about factors that influence splittail
egg and larval development.
Food and Feeding
Feeding studies describe splittail as opportunistic benthic
foragers. Splittail feeding appears highest in the morning and early afternoon. Studies from the Sacramento
River found that their diets were dominated by oligochaetes, cladocerans, and dipterans (Caywood 1974).
Samples from the lower San Joaquin River included
copepods, dipterans, detritus and algae, clams (Corbicula)
and amphipods (Corophium spp.). Copepods were the
dominant food items. These findings were similar to
results of feeding studies from Suisun Marsh (Daniels
and Moyle 1983), where the diet consisted predominantly of detritus in both percent frequency of occurrence (74%) and percent volume (57%). A smaller portion of the stomach contents (41% by volume) consisted
Chapter 2 —
Estuarine Fish and Associated Invertebrates
87
Fish
The Sacramento splittail (Family: Cyprinidae) is one
of California’s largest native minnows and is the only
surviving member of its genus. In 1994 it was proposed
for listing as a Threatened species by U.S. Fish and Wildlife Service based on concerns about reduced abundance
and distribution (Meng and Kanim 1994, Meng and
Moyle 1995). The species supports a small sport fishery
in winter and spring, when it is caught for human consumption and live bait for striped bass angling.
Mature splittail eggs are 1.3 to 1.6 mm in diameter with a smooth, transparent, thick chorion (Wang
1986 cited in CDWR and USBR 1994). The eggs are
adhesive or become adhesive soon after contacting water (Bailey 1994). The eggs appear to be demersal and
it is assumed that they are laid in clumps and attach to
vegetation or other submerged substrates. Under laboratory conditions, fertilized eggs incubated in fresh water at 19° C (±0.5° C) start to hatch after approximately
96 hours. Asynchronous hatching of egg batches from
single females has been observed in preliminary culturing tests.
Early hatched larvae are 6 mm long, have not developed eye pigment, and are physically underdeveloped.
The last larvae to hatch have developed eye pigmentation and are morphologically better developed. Larvae are
7.0 to 8.0 mm total length (TL) when they complete
yolk-sac absorption and become free swimming; postlarvae are up to 20 mm (±4.2 mm) TL. First scale formation appears at lengths of 22 mm standard length (SL)
or 25 mm to 26 mm TL. It is unknown when exogenous
feeding actually begins, but preliminary observations
indicate that newly hatched larvae may have undeveloped
mouths. Well-developed mouths are observed in postlarvae between 8.1 mm and 10.4 mm TL.
Sacramento splittail are a relatively long-lived
minnow, reaching ages of 5, and possibly, up to 7 years.
Studies from Suisun Marsh indicate that young-of-theyear (YOY) grow approximately 20 mm per month (mm/
month) from May through September and then decrease
to < 5 mm/month through February (Daniels and Moyle
1983). In their second season they grow at about 10
mm/month until the fall when somatic growth declined
and gonadal development began. The adult growth rate
ranges from 5 to 7 mm/month. During gonad development, which occurs primarily between September and
February, the growth rate slows to less than 5 mm/
month. The largest recorded splittail measured between
380 mm and 400 mm.
Fish
of animal matter, mostly crustaceans (35% by volume).
Opossum shrimp (Neomysis mercedis) were the dominant
crustacean food item (37% frequency; 59% volume less
detritus) both daily and seasonally for splittail in Suisun
Marsh. Other minor prey items included molluscs, insects, and fish.
Food selection studies from Suisun Marsh suggest
that splittail specifically select Neomysis as their main
prey item in the Estuary (Herbold 1987). Fullness indices data indicate that condition factors of splittail are
linked to Neomysis abundance. Splittail did not switch
to alternate and more prevalent food items, as was observed for other native resident species.
The historical range of splittail included all low gradient portions of all major tributaries to the Sacramento
and San Joaquin rivers, as well as some other freshwater tributaries to San Francisco Bay (Meng and Moyle
1995). A confounding issue is that the collection season and life stage for most of the early observations
are unknown, so the relative importance of each location to different age classes of splittail cannot be established.
Splittail are presently most common in the brackish waters of Suisun Bay, Suisun Marsh, and the Sacramento-San Joaquin Delta. The data suggest that splittail
inhabit much of their historical range and have been located in previously unreported sites (Table 2.3). Much
of the loss of splittail habitat is attributable to migration
barriers, but loss of floodplain and wetlands due to dik-
Population Status and Influencing Factors
Abundance estimates for YOY and adult splittail were
developed recently (Sommer et al. 1997) from several
Interagency Ecological Program surveys. The survey
equipment for the Program includes otter trawls, midwater trawls, beach seines, and townets.
Abundance of YOY declined in the Estuary during the six-year drought, which commenced in 1987
(Figure 2.4). There was a strong resurgence in YOY
in 1995, when abundance estimates were the highest
on record for State Water Project, Central Valley
Project, beach seine, Outflow/Bay otter trawl, and
Outflow/Bay study midwater trawl. The midwater trawl
index was the second highest on record. The response
Table 2.3 Historical and Recent Collections of Splittail(a)
River
Distance (km) from Mouth of River
to Collection Site
Distance (km) to first dam
Rutter (1908)
Caywood (1974)
483
387
331
387 (Red Bluff)
109
(b)
94
109 (Oroville)
49
37
19
37 (Nimbus)
San Joaquin
435(c)
(b)
201
295 (Sack)
Mokelumne
n/a
25
63
63 (Woodbridge)
Sacramento
Feather
American
Sommer et al. (1997)
Napa
n/a
21
10
n/a
Petaluma
n/a
25
8
16(d)
(a) For the purposes of comparing present and historical distribution, we assumed that collection of any life stage of splittail constituted
evidence that a given location was part of the range of the species. The results should be considered as the minimum range only;
there had not been sufficient sampling in sites farther upstream to conclusively show that they were not present. To illustrate the fact
that much of the loss of channel habitat is attributable to migration barriers, the location of the first dam on each river is included.
(b) Records indicate that splittail were collected, but it is unclear where.
(c) Rutter (1908) was cited by FWS (1994) as the source of an observation of splittail at Fort Miller (km 435), near the current site of Friant
Dam on the San Joaquin River . However, Rutter’s distribution was based on Girard (1854), who reported two Pogonichthys species,
P. symetricus and P. inaquilobus in the San Joaquin system. P. symetricus, collected from Fort Miller, is unlikely to have been a splittail
(P. macrolepidotus) because Girard reported the “ lobes of the caudal fin are symmetrical” . Girard’s description of P. inaequilobus
had an asymmetrical tail and other features similar to that of splittail, but the collection location is listed as “ San Joaquin River” without reference to a specific site.
(d) Dam was removed in 1994.
88
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Distribution
ing and draining activities during the past century probably represents the greatest reduction in habitat.
Within the San Francisco Estuary, splittail were
collected from southern San Francisco Bay and at the
mouth of Coyote Creek in Santa Clara County around
the turn of the century. To our knowledge, no other
splittail have been collected in this part of San Francisco
Bay (Aceituno et al. 1976). However, splittail are caught
in San Francisco Bay and San Pablo Bay in wet years.
Adults and young are abundant in two tributaries to San
Pablo Bay, the Napa and Petaluma rivers. The core of
distribution of adult splittail during summer appears
to be the region from Suisun Bay to the west Delta.
Splittail are also present in some of the smaller tributaries and sloughs of Suisun Bay, including Peyton Slough,
Hastings Slough, and Pacheco Creek.
Fish
Figure 2.4 Trends in Age-0 Splittail Abundance for 1975-1995 as Indexed by Eight Independent Surveys.
The first data point in each series is marked with a circle. Dry years are identified with asterisks above
the data points— all other years are wet.
was not as dramatic for the Suisun Marsh, Chipps
Island, or townet surveys, but there was a clear increase in abundance for each relative to the previous
nine years.
There appears to be no consistent decline in adult
abundance for most of the surveys (Figure 2.5). However, both the Suisun Marsh and Chipps Island surveys
show significantly lower abundance in the early to
mid-1980s (Sommer et al. 1997).
Floodplain inundation appears to be a key factor
responsible for strong year classes, based on both statistical and limited observational data (Sommer et al.
1997). Higher flows increase inundation of floodplain
areas, such as the Yolo Bypass, which provides spawning, rearing, and foraging habitat. The species has little
or no stock recruitment relationship. This is best illustrated from data collected in 1995, when exceptionally
large numbers of young splittail were produced by a stock
Chapter 2 —
Estuarine Fish and Associated Invertebrates
89
Fish
Amphibians &
Reptiles
Plants
Figure 2.5 Trends in Adult Splittail Abundance for 1976-1995 as Indexed by Six Independent Surveys. The
first data point in each series is marked with a circle. Dry years are identified with asterisks above the
data points— all other years are wet.
that should have been depleted by drought conditions
in seven of the previous eight years.
Attributes that help splittail respond rapidly to
improved environmental conditions include a relatively
long life span, high reproductive capacity, and broad environmental tolerances (Sommer et al. 1997). Additional
factors that may affect population levels include habitat
loss, recreational fishing, entrainment, and toxic compounds.
Trophic Levels
Splittail are secondary consumers.
Proximal Species
Predators: Striped bass, centrarchids.
90
Baylands Ecosystem Species and Community Profiles
Prey: Oligochaetes, zooplankton (cladocerans and copepods), terrestrial insects, opossum shrimp (Neomysis
mercedis), mollusks.
Good Habitat
Sacramento splittail are one of the few freshwater
cyprinids that are highly tolerant of brackish water. Although they have been collected at salinities as high as
18 ppt, abundance is highest in the 0-10 ppt salinity
range (Sommer et al. 1997). Physiological studies show
that splittail have critical salinity maxima of 20-29 ppt
(Young and Cech 1996). Splittail also tolerate a wide
range (7-33° C) of water temperatures in the laboratory, which fits well with thermal fluctuations associated with its habitat. Depending upon the acclimation temperature (range 12-20° C), critical thermal
Aceituno, M.E., M.L. Caywood, S.J. Nicola and W.I.
Follett. 1976. Occurrence of native fishes in
Alameda and Coyote Creeks, California. Ca. Dept.
Fish and Game 62(3):195-206.
Bailey, H. 1994. Culturing studies on splittail.
IEP Newsletter, Summer 1994. p. 3.
Caywood, M.L. 1974. Contributions to the life history of
the splittail Pogonichthys macrolepidotus (Ayres).
Master’s thesis. Ca. State Univ., Sacramento.
Daniels, R.A. and P.B. Moyle. 1983. Life history of
the splittail (Cyprinidae: Pogonichthys macrolepidotus
(Ayres)) in Sacramento-San Joaquin estuary. U.S.
Natl. Marine Fish. Bull. 81: 647-654.
California Department of Water Resources and U.S. Bureau of Reclamation (CDWR and USBR). 1994.
Effects of the Central Valley Project and State
Water Project on Delta smelt and Sacramento
splittail. Prepared for U.S. Fish and Wildl. Serv.,
Ecol. Services, Sacramento, CA.
Herbold, B. 1987. Patterns of co-occurrence and resource
use in a non-coevolved assemblage of fishes. Ph.D.
dissertation. Univ. of Ca., Davis.
Meng, L. and N. Kanim. 1994. Endangered and threatened wildlife and plants; proposed determination
of threatened status for the Sacramento splittail.
Fed. Reg. 59:004: 862-868.
Meng, L. and P.B. Moyle. 1995. Status of splittail in
the Sacramento-San-Joaquin estuary. Transactions
of the Amer. Fisheries Society 124: 538- 549.
Sommer, T.R., R. Baxter and B. Herbold. 1997. The
resilience of splittail in the Sacramento-San Joaquin
Estuary. Transactions of the Amer. Fisheries Society 126: 961-976.
Young, P.S. and J.J. Cech. 1996. Environmental tolerances and requirements of splittail. Transactions
of the Amer. Fisheries Society 125: 664-678.
Oncorhynchus tshawytscha
Lt. Dante B. Maragni
General Information
The Chinook salmon (Family: Salmonidae) is morphologically distinguished from other Oncorhynchus species
of the northern Pacific Ocean by its large size, small black
spots on both caudal fin lobes, black pigment along the
base of the teeth (McPhail and Lindsey 1970 as cited in
Healey 1991), and varying shades of flesh color from
white through shades of pink and red (Healey 1991).
The Chinook salmon life history (Figure 2.6) is characterized by adult migration from the ocean to natal
freshwater streams to spawn, and juvenile migration
seaward as smolts in their first year of life. During the
smoltification process, juvenile Chinook salmon undergo
physiological, morphological, and behavioral changes
that stimulate emigration and prepare them for life in
the marine environment (Healey 1991).
The Sacramento-San Joaquin Chinook salmon of
California exists as four races—winter, spring, fall, and
late-fall—as defined by the timing of adult spawning migration (Mason 1965, Frey 1971, Moyle 1976, Healey
1991). In 1989, the Sacramento River winter-run Chinook salmon was listed as threatened under the federal
Endangered Species Act by the National Marine Fisheries Service (NMFS) (54 FR 32085). NMFS reclassified the winter-run as endangered in 1994 (59 FR 440)
based on: 1) the continued decline and increased variability of run sizes since its listing as a threatened species in 1989, 2) the expectation of weak returns in certain years as a result of two small year classes (1991 and
1993), and 3) continuing threats to the population. The
State of California listed the winter-run as endangered
under the California Endangered Species Act in 1989.
In 1995, the Oregon Natural Resources Council and R.
Nawa petitioned NMFS to list Chinook salmon along
the entire West Coast, including the States of California, Idaho, Oregon, and Washington, under the federal
Endangered Species Act (54 FR 32085). The State of
California presently includes on its list of species of special concern the late-fall (Class 2– special concern) and
the spring-run (Class 1– qualified as threatened or endangered) Chinook salmon. Spring-run Chinook salmon
Moyle 1976
References
Chinook Salmon
Chapter 2 —
Estuarine Fish and Associated Invertebrates
91
Fish
maxima ranged from 22-33° C. As further evidence
of the general hardiness of the species, splittail appear
to be tolerant of low dissolved oxygen levels and strong
water currents.
Splittail are numerous within small dead-end
sloughs, those fed by freshwater streams, and in the
larger sloughs such as Montezuma and Suisun (Daniels
and Moyle 1983). Juveniles and adults utilize shallow
edgewater areas lined by emergent aquatic vegetation.
Submerged vegetation provides abundant food sources
and cover to escape from predators. Shallow, seasonally
flooded vegetation is also apparently the preferred spawning habitat of adult splittail (Caywood 1974).
Figure 2.6 Life History of
Chinook Salmon (USFWS
have also been given a special category by the state and
are considered a “ monitored” species.
Chinook salmon support commercial, recreational,
and tribal subsistence fisheries. However, due to the state
of Pacific Coast Chinook salmon populations, the U. S.
Department of Commerce declared the U.S. Pacific
Coast salmon commercial fishery, excluding Alaska, a
disaster and has provided emergency relief funding for
displaced fisherman in 1995 and 1996 (59 FR 51419,
60 FR 5908). Also, the federal Central Valley Project
Improvement Act requires restoration actions to double
the Chinook salmon population in the Sacramento-San
Joaquin River system in California by the year 2002 estimated from average population levels from 1967 to
1991 (CDFG 1993).
Reproduction
The Chinook salmon is anadromous; that is, it spends
most of its adult life in the ocean and returns to freshwater streams to spawn. Chinook salmon typically spend
3-6 years maturing in the ocean before returning as
adults to their natal streams to spawn (Moyle 1976,
Eschmeyer et al. 1983). Historically, most SacramentoSan Joaquin Chinook salmon returning to spawn have
been four years of age (Clark 1929). The Chinook
salmon is also semelparous in its reproductive strategy
in that it dies after it spawns. Thus, the life span of
the Chinook salmon is 3-6 years. All adults die after
spawning except some “ jacks” (i.e., precocious males
92
Baylands Ecosystem Species and Community Profiles
that mature early in freshwater) (Miller and Brannon
1982).
Chinook salmon can be grouped into two types
based on variations in their life histories: stream-type and
ocean-type. Stream-type Chinook salmon populations
are most commonly found north of 56° N latitude along
the North American coast and characterized by long
freshwater residence as juveniles (1+ years). Ocean-type
Chinook salmon populations are most commonly found
south of 56° N latitude and characterized by short freshwater residence as juveniles (2-3 months). Chinook
salmon of the Sacramento-San Joaquin River system are
predominantly ocean-type (Healey 1991). Adult upstream migration and juvenile downstream migration of
the Sacramento-San Joaquin Chinook salmon differ
among the four races. Sacramento-San Joaquin Chinook
salmon populations’ migration characteristics are listed
in Table 2.4 (Bryant, pers. comm.).
The Chinook salmon normally spawns in large rivers and tributaries, and typically in deeper water and
larger gravel than other Pacific salmon (Scott and Crossman 1973). In preparation for spawning, a female Chinook salmon digs a shallow depression in the gravel of
the stream bottom in an area of relatively swift water by
performing vigorous swimming movements on her side
near the bottom (Emmett et al. 1991, Healey 1991).
This depression is referred to as a “ redd,” and can be 1.210.7 m (3.9-35.1 ft) in diameter (Chapman 1943). The
female then deposits a group or ” pocket” of eggs in the
redd (Emmett et al. 1991, Healey 1991). From 2,000
Plants
Amphibians &
Reptiles
Fish
1995)
to 14,000 eggs are laid per female, with 5,000 eggs per
female being average (Rounsefell 1957, Moyle 1976, Bell
1984). The eggs are in turn fertilized by one or more
males. During spawning, a female will be attended by
one dominant male and occasionally other subdominant
males. The female then buries the eggs by displacing
gravels upstream of the redd (Emmett et al. 1991, Healey
1991).
Chinook salmon eggs are spherical, non-adhesive, and
the largest of all the salmonids (6.0-8.5 mm (0.24-0.33
in) in diameter) (Rounsefell 1957, Scott and Crossman
1973, Wang 1986). The incubation range is from 4-6
weeks, depending on levels of dissolved oxygen, biochemical oxygen demand, water temperature, substrate,
channel gradient and configuration, water depth, water
velocity and discharge (Reiser and Bjornn 1979, Alaska
Department of Fish and Game 1985).
Larval sizes range from 20-35 mm (0.79-1.38 in)
in length (Wang 1986). Yolk sac fry, termed “ alevins,”
remain in the gravel from 2-3 weeks until the yolk sac is
absorbed (Scott and Crossman 1973, Wydoski and
Whitney 1979), whence they emerge from the gravel as
fry. Fry develop into parr beginning the smoltification
process as they encounter increasing salinities during
their migration from freshwater to the ocean. Parr acquire a silver color as they transform into smolts during
the smoltification process (Healey 1991). Fry and smolts
can stay in freshwater from 1-18 months (Beauchamp
et al. 1983), with residency periods differing with race
(Table 2.4). Outmigration periods vary with outflow
conditions. High outflows will carry fry downstream,
while seasons with low outflow cause fry to rear longer
in upstream areas where they grow much larger. Juvenile Chinook salmon in these two differing scenarios
Food and Feeding
Chinook salmon larvae and alevins feed on their yolk.
Chinook salmon juveniles and adults are carnivorous,
“ opportunistic” feeders, feeding on a variety of terrestrial and aquatic insects, crustaceans, and fish (Emmett
et al. 1991).
Juveniles in freshwater consume primarily terrestrial and aquatic insects, amphipods and other crustaceans, and sometimes fish (Becker 1973, Higley and
Bond 1973, Scott and Crossman 1973, Craddock et al.
1976, Muir and Emmett 1988, Sagar and Glovea 1988).
Table 2.4 Migration Characteristics of Sacramento-San Joaquin Chinook Salmon Runs (Bryant 1997)
Characteristic
Winter
Spring
Fall
Late-fall
ADULT
Immigration Period
December - July
March - July
June - December
October - April
Peak Immigration
March
May -June
September - October
December
Spawning Period
late April early August
late August late October
late September December
January late April
Peak Spawning
early June
mid September
late October
early February
Emergence Period
July - October
November - March
December - March
April - June
Freshwater
Residency
Period
5 - 10 months
July - April
3 - 15 months
November - January
(year 2)
4 - 7 months
December - June
7 - 13 months
April - April (year 2)
Estuarine
Emigration
Period
November - May
March - June &
November - March
March - July
October - May
JUVENILE
Chapter 2 —
Estuarine Fish and Associated Invertebrates
93
Fish
Growth and Development
have substantially different habitat requirements (Kjelson
et al. 1982). The fry to smolt life stages’ size range is 2152 cm (0.6-42.9 in), but is usually less than 91 cm (25.7
in), in length (Wydoski and Whitney 1979).
Juvenile Chinook salmon migration into estuaries
has been reported to occur at night (Seiler et al. 1981)
and during daylight (Dawley et al. 1986). Juveniles may
move quickly through estuaries (Dawley et al. 1986) or
reside there for up to 189 days (Simenstad et al. 1982).
Juvenile Chinook salmon gain significant growth in estuarine habitats as they smolt and prepare for the marine phase of their life (MacDonald et al. 1987). The juveniles of most stocks of Chinook salmon appear to migrate north upon entering the ocean (Wright 1968,
Healey 1991). Chinook salmon produced in streams
from the Rogue River (Oregon) and south appear to rear
in the ocean off northern California-southern Oregon
(Cramer 1987). The stream-type Chinook salmon move
offshore early in their ocean life, whereas ocean-type
Chinook salmon remain in sheltered coastal waters.
Stream-type Chinook salmon maintain a more offshore
distribution throughout their ocean life than do oceantype (Healey 1991). Chinook salmon reach maturity in
3-6 years (Moyle 1976).
Distribution
Chinook salmon eggs and alevins are benthic and infaunal. Fry and parr are benthopelagic. Parr become pelagic
as they enter smoltification. Smolts, ocean-dwelling and
maturing juveniles, and adults are pelagic (Alaska Department of Fish and Game 1985). Adults are bottomoriented in freshwater (Emmett et al. 1991).
Chinook salmon eggs, alevins, fry, and parr occur
in riverine areas from just above the intertidal zone to
altitudes of 2,268 m (7,441 ft) above sea level (Allen et
al. 1991). Smolts are riverine and estuarine. Oceandwelling juveniles are neritic and epipelagic, and found
within 128 m (420 ft) of the surface (Fredin et al. 1977).
Adults may be neritic and estuarine, but are riverine
during their spawning migration and may travel upstream more than 4,700 km (2,920 mi) from the
ocean (Emmett et al. 1991) as flows and passage allow. Most tributaries are now dammed for water supply, which limits the extent of upstream migration
(USFWS 1995).
The Chinook salmon is the least abundant of the
major Pacific salmon species (Emmett et al. 1991,
Healey 1991). However, it is the most abundant salmon
in California (McGinnis 1984). The Chinook salmon is
recorded as far north as the Coppermine River in Arctic
Canada, and south to northeastern Hokkaido, Japan, and
southern California (Ventura River) (Hart 1973, Scott
and Crossman 1973). It is, however, rarely found in
freshwater south of the Sacramento-San Joaquin River
system of California (Eschmeyer et al. 1983).
94
Baylands Ecosystem Species and Community Profiles
Fish
While Chinook salmon are found in all estuaries
north of San Francisco Bay in California, except Tomales Bay (Monaco et al. 1990), California’s largest populations of Chinook salmon originate in the SacramentoSan Joaquin River system (Fry 1973). Spring-run Chinook salmon are extinct in the San Joaquin River and
only remnant runs remain in a few Sacramento River
tributaries. Historically, spring-run Chinook salmon
spawned in small tributaries that have essentially all been
blocked to migration by large dams. Fall and late-fall
Chinook salmon are main stem spawners. Winter-run
Chinook salmon are unique to the Sacramento River and
spawned in coldwater tributaries above Shasta Dam prior
to its construction (Sacramento River Winter-Run Chinook Salmon Recovery Team 1996). While distribution
of outmigrating juvenile Chinook salmon is not well
known in the San Francisco Bay, they have been found
throughout, including the South Bay on high outflow
years.
Population Status and Influencing Factors
Chinook salmon populations have declined substantially,
with winter-run at the point of near extinction and
spring-run at severely depressed population levels (Table
2.5). Whereas spring-run historically outnumbered all
other runs, fall-run comprises the bulk of the present
Chinook salmon population. The remnant “ endangered”
population of winter-run now depend on cold water releases from Shasta Reservoir, and the protection of the
federal Endangered Species Act.
No single impact can be attributed to the decline
of Chinook salmon populations and the important Chinook salmon fishery. High mortality for Chinook salmon
occurs during the early freshwater life stages (eggs, fry,
parr) (Emmett et al. 1991). This mortality is caused by
redd destruction, siltation and destruction of spawning
grounds, extremely high or low water temperatures, low
dissolved oxygen, loss of cover, disease, food availability
and competition, and predation (Reiser and Bjornn
1979). Besides the above factors, human impacts such
as river flow reductions, the construction of dams and
the consequent creation of reservoirs, water diversions,
logging practices, and pollution have affected population
abundances (Raymond 1979, Netboy 1980, Stevens and
Miller 1983). Factors influencing survival of adult Chinook salmon are equally numerous. In the ocean, Chinook salmon are impacted by oceanographic conditions, disease, food availability and competition, predation, and overfishing (Fraidenburg and Lincoln
1985, Emmett et al. 1991). In freshwater, adults are
subject to natural factors such as drought and flood,
and human impacts including fishing, dams, road
construction and other development, flood protection,
dredging, gravel mining, timber harvest, grazing, and
pollution (USFWS 1995).
Plants
Amphibians &
Reptiles
In estuaries, juveniles feed in intertidal and subtidal
habitats of tidal marshes. In these habitats, juveniles prey
upon insects, gammarid amphipods, harpacticoid copepods, musids, chironomids, decapod larvae, and small
(larval and juvenile) fish (Levy and Levings 1978, Levy
et al. 1979, Northcote et al. 1979, Healey 1980a, Levy
and Northcote 1981, Healey 1982, Kjelson et al. 1982,
Simenstad et al. 1982, Simenstad 1983, McCabe et al.
1986). In low flow years when juveniles are larger, their
food source will include crab megalops, squid, and small
fish (e.g., northern anchovy, Pacific herring, rockfish)
(Beauchamp et al. 1983).
Smaller juvenile Chinook salmon having recently
migrated into the marine environment feed on amphipods, euphausiids, and other invertebrates, and small
(larval and juvenile) fish (Healey 1980b, Peterson et
al.1983, Emmett et al. 1986). Larger juvenile and adult
Chinook salmon in the ocean feed primarily on fish (e.g.,
northern anchovy, Pacific herring, and Pacific sandlance), as well as squid, euphausiids, decapod larvae, and
other invertebrates (Silliman 1941, Merkel 1957,
Prakash 1962, Ito 1964, Hart 1973, Fresh et al. 1981).
Immigrating adult Chinook salmon do not actively feed
in freshwater (Emmett et al. 1991).
Juvenile Prey: Terrestrial insects, aquatic insects, chironomids, copepods, amphipods, mysids, euphausiids,
decapod larvae, bay shrimp.
Adult Prey: Euphausiids, decapods, squid, Pacific herring
(Clupea pallasi, northern anchovy (Engraulis mordax),
osmerids, rockfish (Sebastes spp.), Pacific sandlance
(Ammodytes hexapterus).
Trophic Levels
Chinook salmon are primary and secondary consumers
as juveniles and secondary consumers as adults.
Proximal Species
Good Habitat
Chinook salmon eggs develop only in freshwater, but
larvae can tolerate salinities of up to 15 ppt at hatching.
Three months after hatching juvenile Chinook salmon
can tolerate full seawater, with faster growing individuals better able to handle salinity changes (Wagner et al.
1969). Juveniles and adults occur in freshwater to
euhaline waters. Successful egg incubation occurs from
just above freezing to 20.0° C (68.4° F) (Olsen and Fos-
Table 2.5 Estimated Number of Sacramento-San Joaquin Chinook Salmon Returning to Spawn: 19671991 (Mills and Fisher 1994) (Continued on next page.)
Year
2
3
San Joaquin
Fall-run Chinook2
grilse
adult
total
1967
38,410
104,790
1968
18,181
155,859
1969
48,528
1970
1971
Sacramento
Late-fall-run Chinook3
grilse
adult
total
grilse
adult
143,200
1,176
21,359
22,535
5,730
31,478
37,208
174,040
11,211
6,577
17,788
1,910
32,823
34,733
208,289
256,817
1,935
49,662
51,597
1,747
35,431
37,178
30,121
147,279
177,400
8,539
28,550
37,089
1,823
17,367
19,190
35,775
140,691
176,466
2,986
38,580
41,566
2,277
12,046
14,323
1972
43,795
80,622
124,417
2,454
12,321
14,775
2,398
29,155
31,553
1973
40,640
197,193
237,833
674
6,438
7,112
711
21,493
22,204
total
1974
25,364
185,953
211,317
762
3,625
4,387
329
6,116
6,445
1975
29,691
141,884
171,575
968
6,258
7,226
816
15,847
16,663
1976
21,926
155,767
177,693
505
3,894
4,399
581
14,699
15,280
1977
22,831
139,971
162,802
60
990
1,050
873
8,217
9,090
1978
23,635
115,363
138,998
254
2,473
2,727
959
7,921
8,880
1979
46,397
152,982
199,379
456
3,897
4,353
44
8,696
8,740
1980
25,472
110,833
136,305
702
5,600
6,302
566
7,181
7,747
1981
42,575
145,503
188,078
8,022
20,295
28,317
168
1,429
1,597
1982
43,396
129,388
172,784
2,681
14,214
16,895
186
955
1,141
1983
41,714
88,676
130,390
32,312
10,970
43,282
1,221
12,053
13,274
1984
41,030
115,509
156,539
18,335
37,641
55,976
2,357
3,550
5,907
1985
41,563
211,695
253,258
4,311
71,873
76,184
1,670
5,990
7,660
1986
27,356
212,739
240,095
3,117
18,588
21,705
490
6,220
6,710
1987
66,364
150,965
217,329
18,269
6,689
24,958
780
13,663
14,443
1988
26,517
197,841
224,358
1,138
20,798
21,936
2,094
8,589
10,683
1989
24,060
116,726
140,786
282
3,489
3,771
286
9,589
9,875
1990
9,443
83,499
92,942
312
663
975
1,536
5,385
6,921
1991
11,546
87,070
98,616
207
647
854
888
5,643
6,531
33,053
143,083
176,137
4,867
15,844
20,710
1,298
12,861
14,159
AVERAGE
1
Sacramento
Fall-run Chinook1
Escapement data for the Sacramento River and its tributaries north of and including the American River.
Escapement data for the Mokelumne, Cosumnes, Calaveras, Stanislaus, Tuolumne and Merced rivers.
Escapement data for the main stem Sacramento River above Red Bluff Diversion Dam.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
95
Fish
Juvenile Predators: Sacramento squawfish (Ptychocheilus grandis), riffle sculpin (Cottus gulosus), channel catfish (Ictalurus punctatus), steelhead trout (Oncorhynchus
mykiss), striped bass (Morone saxatilis), centrarchids,
rockfish (Sebastes spp.), kingfishers, egrets, herons, terns,
grebes, pelicans.
Adult Predators: Pacific lamprey (Lampetra tridentata),
harbor seal (Phoca vitulina), California sea lion (Callorhinus ursinus), killer whale (Orcinus orca), North American river otter (Lutra canadensis), American black bear
(Ursus americanus), bald eagle (Haliaeetus leucocephalus).
Fish
ter 1955), however, best incubation temperatures are
5.0-14.4° C (41.0-57.9° F) (Bell 1984). The upper lethal
temperature for Chinook salmon is 25.1° C (77.2° F)
(Brett 1952), but may be lower depending on other water quality factors (Ebel et al. 1971). Eggs and alevins
are found in areas with flow of 20-150 cm/sec (0.7-5 ft/
sec) and juveniles where flows are 0.5-60.0 cm/sec (0.022 ft/sec) (at pool edges). Adults can migrate upstream
in flows up to 2.44 m/sec (8 ft/sec) (Thompson 1972).
Successful egg development requires redds to have adequate dissolved oxygen (>5.0 mg/L), water temperatures
(4-14° C [39-57° F]), substrate permeability, sediment
composition (<25% fines, <6.4 mm [0.25 in] in diameter), surface flows and velocities, and low biochemical
oxygen demand (Reiser and Bjornn 1979).
Juveniles in freshwater avoid waters with <4.5 mg/L
dissolved oxygen at 20°C (68°F) (Whitmore et al. 1960).
Migrating adults will pass through water with dissolved
oxygen levels as low as 5 mg/L (Hallock et al. 1970). Excessive silt loads (>4,000 mg/L) may halt Chinook
salmon movements or migrations. Silt can also hinder
fry emergence, and limit benthic invertebrate (food)
production (Reiser and Bjornn 1979). Freshwater inflow
into estuaries is critical for providing adequate water temperatures, food production, and overall beneficial environmental conditions for juvenile outmigration. High
freshwater flows allow for cooler water temperatures,
while also stimulating and sustaining production of food.
High river flows improve juvenile survival and enable
active migration into estuaries and on to the ocean.
In addition to specific hydrologic components,
physical habitat requirements of interrelated instream
gravel, riparian, and tidal marsh habitats comprise the
healthy ecosystem in which Chinook salmon spawn and
rear. Chinook salmon eggs and alevins require clean,
loose gravel and occur in spawning gravel or cobble that
Table 2.5 (continued) Estimated Number of Sacramento-San Joaquin Chinook Salmon Returning to
Spawn: 1967-1991 (Mills and Fisher 1994)
YEAR
Sacramento
Springl-run Chinook4
grilse
adult
total
grilse
adult
total
11,397
12,297
23,694
24,985
32,321
57,306
81,698
202,245
283,943
1968
3,317
11,827
15,144
10,299
74,115
84,414
44,917
281,202
326,119
1969
2,843
24,492
27,335
8,953
108,855
117,808
64,006
426,729
490,735
1970
1,420
6,017
7,437
8,324
32,085
40,409
50,228
231,297
281,525
1971
2,464
6,336
8,800
20,864
32,225
53,089
64,366
229,878
294,244
1972
1,343
7,053
8,396
8,541
28,592
37,133
58,531
157,743
216,274
1973
2,082
9,680
11,762
4,623
19,456
24,079
48,729
254,261
302,990
1974
2,538
5,545
8,083
3,788
18,109
21,897
32,782
219,347
252,129
1975
7,683
15,670
23,353
7,498
15,932
23,430
46,656
195,591
242,247
1976
4,067
22,006
26,073
8,634
26,462
35,096
35,712
222,829
258,541
1977
5,421
8,409
13,830
2,186
15,028
17,214
31,372
172,614
203,986
1978
1,093
7,063
8,156
1,193
23,669
24,862
27,134
156,489
183,623
1979
707
2,203
2,910
113
2,251
2,364
47,717
170,029
217,746
1980
3,734
8,081
11,815
1,072
84
1,156
31,545
131,780
163,325
1981
8,249
13,066
21,315
1,744
18,297
20,041
60,757
198,591
259,348
1982
4,528
21,644
26,172
270
972
1,242
51,061
167,947
219,008
1983
672
3,809
4,481
392
1,439
1,831
76,311
116,947
193,258
1984
4,373
3,988
8,361
1,869
794
2,663
67,965
161,481
229,446
1985
3,792
7,631
11,423
329
3,633
3,962
51,665
300,822
352,487
1986
1,606
17,290
18,896
451
2,013
2,464
33,020
256,850
289,870
1987
4,177
7,330
11,507
236
1,761
1,997
89,826
180,408
270,234
1988
2,132
9,521
11,653
708
1,386
2,094
32,589
238,136
270,725
1989
884
6,304
7,188
53
480
533
25,566
136,587
162,153
1990
948
4,376
5,324
16
425
441
12,256
94,347
106,603
433
1,208
1,641
38
153
191
13,112
94,721
107,833
3,276
9,714
12,990
4,687
18,421
23,109
47,181
199,955
247,136
1991
5
total
1967
AVERAGE
4
adult
Central Valley
Total Chinook Salmon
Escapement data for the main stem Sacramento River above Red Bluff Diversion Dam.
Escapement data for the main stem Sacramento River above Red Bluff Diversion Dam.
96
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
grilse
Sacramento
Winter-run Chinook5
References
Alaska Department of Fish and Game. 1985. Alaska
habitat management guide, southcentral region,
volume I: life histories and habitat requirements
of fish and wildlife. Dept. of Fish and Game, Juneau, Al. 429 pp.
Allen, M.A. and T.J. Hassler. 1986. Species profiles: Life
histories and environmental requirements of coastal
fishes and invertebrates (Pacific Southwest), Chinook salmon. U.S. Fish and Wildl. Serv. Biol. Rept.
82 (11.49). U.S. Army Corps of Engrs., TR EL82-4. 26 pp.
Allen, M.J., R.J. Wolotira, Jr., T.M. Sample, S.F. Noel
and C.R. Iten. 1991. Salmonids: life history descriptions and brief harvest summaries for salmonid species of the northeast Pacific Ocean and eastern Bering Sea. Technical memorandum, NOAA,
NMFS, Northwest and Alaska Fisheries Center,
Seattle, Wash.
Beauchamp, D.A., M.F. Shepard and G.B. Pauley. 1983.
Species profiles: life histories and environmental
requirements of coastal fishes and invertebrates
(Pacific Northwest), Chinook salmon. U.S. Fish
and Wildl. Serv. Biol. Rept. 82(11.6). U.S. Army
Corps of Engrs., TR EL-82-4. 15 pp.
Becker, C.D. 1973. Food and growth parameters of juvenile Chinook salmon, Oncorhynchus tshawytscha, in
central Columbia River. Fish. Bull., U.S. 71:387-400.
Bell, M.C. 1984. Fisheries handbook of engineering requirements and biological criteria. Fish Passage Development and Evaluation Program, U.S. Army Corps
of Engrs., North Pac. Div., Portland, OR. 290 pp.
Brett, J.R. 1952. Temperature tolerance in young Pacific salmon, genus Onchorhynchus. Journal Fish.
Res. Bd. of Canada 9(6):265-323.
California Department of Fish and Game (CDFG).
1993. Restoring Central Valley streams: A plan for
action. Sacramento, California.
Cannon, T.C. 1991. Status of the Sacramento-San
Joaquin Chinook salmon and factors related to their
decline. Report prepared for the National Marine
Fisheries Service, Southwest Region, by Envirosphere
Company, Newport Beach, Ca.. 11pp.
Chapman, W.M. 1943. The spawning of Chinook
salmon in the main Columbia River. Copeia
1943:168-170.
Chapman, W.M. and T.C. Bjornn. 1968. Distribution
of salmonids in streams with special reference to
food and feeding. In: T.G. Northcote (ed). Salmon
and trout in streams, pp. 153-176. H.R. MacMillan Lectures in Fisheries, Univ. of British Columbia, Vancouver, British Columbia, Canada.
Clark, G.H. 1929. Sacramento-San Joaquin salmon
(Oncorhynchus tshawytscha) fishery of California.
Fish Bull. No. 17, Ca. Dept. Fish and Game, Sacramento, Ca.
Craddock, D.R., T.H. Blahm and W.D. Parente. 1976.
Occurrence and utilization of zooplankton by juvenile Chinook salmon in the lower Columbia River.
Transactions of the Amer. Fish. Soc. 105:72-76.
Cramer, S.P. 1987. Oregon studies to increase regional
salmon production. Annual progress report, Marine Resources Region, Oregon Dept. Fish and
Wildl., Portland, Oregon. 15 pp.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
97
Fish
is 1.3-10.2 cm (0.5-10.2 in) in diameter (Reiser and
Bjornn 1979). Juveniles in freshwater are found within
areas of shallow riffles and deep pools over various substrates, ranging from silt bottoms to large boulders
(Chapman and Bjornn 1968). Juveniles in estuaries occur in intertidal and tidal habitats over mud, sand, gravel,
and eelgrass (Zostera spp.) (Healey 1980a). Adults in
marine waters show no sediment preference, but may be
associated with gravel-cobble bottoms in rivers and
streams during upstream migration (Alaska Department
of Fish and Game 1985).
In riverine areas, both submerged cover, such as
boulders, woody debris, and aquatic vegetation, and
overhead cover, such as continuous riparian vegetation
canopies, undercut banks, and turbulent water, provide
shade, food, and protection against predation to juvenile Chinook salmon. Estuaries appear to play a vital role
in Chinook salmon life history as well, and specifically,
tidal marsh habitat is of great importance to juvenile
salmonids (Dorcey et al. 1978, Levy et al. 1979,
Meyer 1979, Levy and Northcote 1981, Healey 1982,
MacDonald et al. 1987, 1988). Juvenile Chinook salmon
forage in the intertidal and shallow subtidal areas of tidal
marsh mudflat, slough, and channel habitats, and open
bay habitats of eelgrass and shallow sand shoal areas.
These productive habitats provide both a rich food supply and protective cover within shallow turbid waters
(McDonald 1960; Dunford 1975, cited from Cannon
1991). The distribution of juvenile Chinook salmon
changes tidally, with fry moving from tidal channels
during flood tides to feed in nearshore marshes.
Tidal marshes are most heavily used by fry, whereas
smolts tend to utilize deeper waters. Fry disperse along
the edges of marshes at the highest points reached by the
tide, then retreat into the tidal channels with the receding tide. Smolts congregate in surface waters of main and
secondary sloughs and move into shallow subtidal areas
to feed (Levy and Northcote 1981, Levings 1982, Allen
and Hassler 1986, Healey 1991).
In addition to good water quality, adequate flows,
and productive spawning and rearing habitat, state-ofthe-art positive barrier screens on water diversions, protection from excessive harvest, and free access to upstream migration or well-designed ladders for adult passage offers promising overall habitat for healthy Chinook
salmon populations.
98
Baylands Ecosystem Species and Community Profiles
Fish
Amphibians &
Reptiles
Frey, H.W. 1971. California’s living marine resources
and their utilization. Ca. Dept. Fish and Game,
Sacramento, Ca. 148 pp.
Fry, D.H. 1973. Anadromous fishes of California. Ca.
Dept. Fish and Game, Sacramento, Ca. 111 pp.
Hallock, R.J., R.F. Elwell and D.H. Fry, Jr. 1970. Migrations of adult king salmon (Oncorhynchus
tshawytsha) in the San Joaquin Delta. Ca. Dept.
Fish and Game Bull. 151:1-92.
Hart, J.L. 1973. Pacific fishes of Canada. Bulletin of
the Fisheries Research Board of Canada, bulletin
no. 180. 740 pp.
Healey, M.C. 1980a. Utilization of the Nanaimo River
Estuary by juvenile Chinook salmon, Oncorhynchus tshawytscha. Fish. Bull., U.S. 77(3):653-668.
______. 1980b. The Ecology of Juvenile Salmon in
Georgia Strait, British Columbia. In: W.J. McNeil
and D.C. Himsworth (eds). Salmonid Ecosystems
of the North Pacific, pp. 203-229. Oregon State
Univ. Press, Corvallis, Oregon.
______. 1982. Juvenile Pacific salmon in estuaries: the
life support system. In: V.S. Kennedy (ed). Estuarine comparison. Academic Press, New York, NY,
pp. 315-341.
______. 1991. Life history of Chinook Salmon (O.
Tshawytscha). In: C. Groot and L. Margolis (ed).
Pacific salmon life histories, pp. 331-391. UBC
Press, Univ. of British Columbia, Vancouver, British Columbia, Canada.
Higley, D.L. and C.E. Bond. 1973. Ecology and production of juvenile spring Chinook salmon, Oncorhynchus tshawytscha, in an entropic reservoir. Fish.
Bull., U.S. 71(3):877-891.
Ito, J. 1964. Food and feeding habits of Pacific salmon
(genus Oncorhynchus) in their ocean life. Bull. of
the Hokkaido Reg. Fish. Res. Lab. 29:85-97. (Fisheries Research Board of Canada Translation Service 1309).
Kjelson, M.A., P.F. Raquel and R.W. Fisher. 1982. Life
history of fall-run Chinook salmon, Oncorhynchus
tshawytscha in the Sacramento-San Joaquin Estuary,
California. In: V.S. Kennedy (ed). Estuarine comparisons. Academic Press, New York, NY, pp. 393-411.
Levings, C.D. 1982. Short term use of a low-tide refugia in a sandflat by juvenile Chinook, (Oncorhynchus tshawytscha), Fraser River Estuary. Can. Tech.
Rpt. Fish and Aq. Sci. 1111. 7 pp.
Levy, D.A. and C.D. Levings. 1978. A description of
the fish community of the Squamish River Estuary, British Columbia: relative abundance, seasonal
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Muir, W.D. and R.L. Emmett. 1988. Food habits of
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Chapter 2 —
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99
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Steelhead
Oncorhynchus mykiss irideus
Robert A. Leidy
General Information
Reproduction
Moyle 1976
Polymorphic salmonids exhibit a high degree of life history variation (Titus et al., in press). Steelhead within the
San Francisco Estuary may be classified as “ ocean-maturing” or “ winter” steelhead that typically begin their
spawning migration in the fall and winter, and spawn
within a few weeks to a few months from when they
enter freshwater (McEwan and Jackson 1996). Ocean
maturing steelhead enter freshwater with well-developed
gonads and spawn shortly after entering a river or stream.
Steelhead begin upstream migration after one to four
growing seasons at sea (Burgner et al. 1992). A small
number of immature fish (i.e., grilse) may also move
upstream after spending only a few months in the ocean.
Growth and Development
Steelhead eggs are spherical to slightly irregular in shape,
non-adhesive, demersal, and range in diameter from 36 mm (Wang 1986). Incubation of eggs is dependent
upon water temperature in the redd. Wales (1941) observed hatching at approximately 19 days at an average
water temperature of 15.5° C and 80 days at about 4.5°
C. For Waddell Creek in coastal San Mateo County,
steelhead hatching time was estimated at 25 to 35 days,
with emergence beginning at 2-3 weeks following hatching (Shapovalov and Taft 1954). Steelhead length at
hatching ranges between 14 to 15.5 mm total length
(TL), with alevins ranging between 23-26 mm TL
(Wang 1986). Alevins emerge from the gravel following
yolk sac absorption as fry or juveniles ready to actively
feed.
Steelhead remain in freshwater for one to four years
(usually two years) before downstream migration as
“ smolts” , at an average size ranging between 13 cm and
25 cm TL (Moyle 1976). Age at emigration is highly
variable, but may occur earlier in warmer, more productive streams where juveniles can reach smolt size at a
younger age (Moyle et al. 1995). Most Sacramento River
juvenile steelhead emigrate as 1-year-old fish during
spring and early summer (Barnhart 1986, Reynolds et
al. 1993), although Shapovalov and Taft (1954) found
that steelhead moved downstream in Waddell Creek
during all months of the year. While steelhead may spend
up to four years in the ocean, most only survive to age
two. In the ocean steelhead may grow at a rate of 1.2
inches per month and reach a length of 23 inches in two
years.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
101
Fish
Steelhead (Family: Salmonidae) are the anadromous (searun) form of resident rainbow trout. Behnke (1992) proposed classification of steelhead on the west coast of the
United States into a coastal subspecies, O. m. irideus, and
an inland subspecies, O. m. gairdneri. California is considered to have only coastal steelhead (Behnke 1992). In
California steelhead may be classified into two races,
summer and winter steelhead, based upon the timing of
upstream migration into freshwater. The San Francisco
Estuary and its tributary streams support winter steelhead. Steelhead are a polymorphic species and as such
populations within a stream may be anadromous, resident, or mixtures of the two forms that presumably interbreed (Titus et al., in press). Steelhead do not support
a commercial fishery within the San Francisco Estuary
and its tributaries. It is illegal for commercial salmon
trollers to possess steelhead (McEwan and Jackson 1996).
There is a inland recreational sportfishery for steelhead
that is dependent largely on hatchery operations to sustain populations. The estimated net annual economic
benefit of doubling steelhead stocks within the Sacramento/San Joaquin river systems is estimated at 8.0
million dollars (Meyer Resources Inc. 1988).
Releases of cold water from several large Central Valley
reservoirs on the Sacramento River system may induce
steelhead to begin to move into upstream tributaries as
early as August and September. This means that upstream migrating steelhead may be observed within San
Francisco Bay and Suisun Marsh/Bay between August
and March. Ocean-maturing steelhead typically spawn
between December and April, with most spawning occurring between January through March.
Steelhead are iteroparous and do not die after
spawning as do other Pacific salmon; therefore, they may
return to the ocean and spawn again the following year.
The frequency of return spawning for a given population is generally unknown. Steelhead spawn in redds constructed by the female over a gravel/cobble substrate.
Eggs are deposited in the redd and then fertilized by the
male. The number of eggs produced is largely a function of the size of the female, and may range from 200
to 12,000 eggs over the geographic range of steelhead
(Scott and Crossman 1973, Moyle 1976). Steelhead
within the Sacramento River drainage average between
1,000 to 4,500 eggs (Mills and Fisher 1994).
Fish
Food and Feeding
Rearing juvenile steelhead are primarily drift feeders utilizing a variety of terrestrial and aquatic insects, including emergent aquatic insects, aquatic insect larvae, snails,
amphipods, opossum shrimp, and various species of
small fish (Moyle 1976). Larger steelhead will feed on
newly emergent steelhead fry. Emigrating adult and juvenile steelhead may forage in the open water of estuarine subtidal and riverine tidal wetland habitats within
the Estuary, although the importance of these areas as
rearing habitat for juveniles is not well documented. Apparently upstream migrating steelhead rarely eat and
therefore exhibit reduced growth (Pauley and Bortz
1986).
Steelhead populations are native to Pacific Ocean coastal
drainages of the Kamchatka Peninsula and scattered
mainland locations of Asia and in the western Pacific
from the Kuskokwim River in Alaska to Malibu Creek
in southern California (Titus et al., in press, McEwan and
Jackson 1996, Moyle 1976). Although the life-history
characteristics of steelhead are generally well known, the
polymorphic nature of the subspecies has resulted in
much confusion over the status and distribution of steelhead in San Francisco Estuary and its tributaries. Historically, the Sacramento-San Joaquin River systems
supported large runs of steelhead (McEwan and Jackson
1996). Presumably, most streams with suitable habitat
within the San Francisco Estuary also supported steelhead, however accurate population estimates for individual streams are not available (Skinner 1962, Leidy
1984).
Currently, small steelhead runs of unknown size are
known to exist in South San Francisco Bay in San
Francisquito Creek, San Mateo County; Guadalupe
River and Coyote and Upper Penitencia creeks, Santa
Clara County; Alameda Creek, Alameda County; and
possibly San Leandro Creek, Alameda County (R. Leidy,
unpub. data). Within Central San Francisco Bay steelhead runs are believed to occur in Corte Madera Creek
and its tributaries, Miller Creek, Novato Creek, and
possibly Arroyo Corte Madera del Presideo Creek, Marin
County (R. Leidy, unpub. data). Within San Pablo Bay,
steelhead make spawning runs in the Napa River and
several of its tributary streams and Huichica Creek, Napa
County; and the Petaluma River and Sonoma Creek and
several of their tributary streams, Sonoma County (R.
Leidy, unpub. data). Tributaries to Suisun Bay and adjacent drainages that support steelhead runs of unknown
size include the Sacramento and San Joaquin rivers;
Green Valley and Suisun creeks, Solano County; and
Walnut Creek and possibly Alhambra, Pinole, Wildcat,
and San Pablo creeks, Contra Costa County (R. Leidy,
102
Baylands Ecosystem Species and Community Profiles
Population Status and Influencing Factors
Nehlsen et al. (1991) identified at least 43 steelhead
stocks at moderate to high risk of extinction, with more
than 23 stocks believed to have been extirpated, on the
west coast of the United States. Steelhead in California
are estimated to number roughly 250,000 adults, which
is one half the adult population of 30 years ago (McEwan
and Jackson 1996). As a result of this precipitous decline,
the National Marine Fisheries is currently reviewing the
status of steelhead to determine if they warrant listing
under the Endangered Species Act. Estimates of the average annual steelhead run size for the Sacramento-San
Joaquin River system, including San Francisco Bay tributaries, range between 10,000 and 40,000 adults (Hallock
et al. 1961, McEwan and Jackson 1996). The California Fish and Wildlife Plan (CDFG 1965) estimated an
annual run size for the Sacramento above the mouth of
the Feather River of approximately 30,000 fish, and a
total for the reminder of the entire Central Valley of
40,000 steelhead, including tributaries to San Francisco
Bay. This likely places the size of steelhead runs in San
Francisco Bay tributaries at well below 10,000 fish, however, the fact remains that reliable estimates for individual streams tributary to San Francisco Estuary do not
exist.
General factors influencing steelhead population
numbers during upstream migration, spawning, and incubation include barriers to passage, diversions, flow
fluctuations, water temperature, and other water quality parameters, such as sedimentation of spawning habitats. Factors affecting juvenile rearing habitat and emigration within the San Francisco Estuary and its tributary streams include low summer flows combined with
high water temperatures. Within Suisun Bay/Marsh the
downstream migrating steelhead are adversely affected
by altered flows, entrainment, and mortality associated
with trapping, loading, and trucking fish at state and
federal pumping facilities. In addition, dredging and
dredged material disposal within the San Francisco Estuary may contribute to degradation of steelhead habitat and interference with migration, foraging, and food
resources (LTMS 1996).
Trophic Levels
Larvae are primary consumers. Juveniles and adults are
primary and higher order consumers.
Plants
Amphibians &
Reptiles
Distribution
unpub. data). Steelhead may also be present in other
tributary streams below migration barriers within the
Estuary, but currently there is little or no data on their
status in many streams. Steelhead adults and smolts may
be found foraging in and migrating through estuarine
subtidal and riverine tidal habitats within all areas of the
San Francisco Estuary.
Proximal Species
Good Habitat
The preferred water depth for steelhead spawning ranges
from six to 24 inches, while fry and parr prefer water
depths of between two to 14 inches and 10 to 20 inches,
respectively (Bovee 1978). Steelhead prefer to spawn in
areas with water velocities of approximately two ft/sec
(range = 1-3.6 ft/sec), although optimal spawning velocity is partially a function of the size of fish; larger fish
can successfully spawn in higher water velocities
(Barnhart 1986). Optimal spawning substrate is reported
to range from 0.2 to 4.0 inches in diameter, but steelhead will utilize various mixtures of sand-gravel and
gravel-cobble (Bovee 1978, Reiser and Bjornn 1979).
Optimal temperature requirements for steelhead vary as
follows: adult migration, 46° to 52° F; spawning, 39° to
52° F; incubation and emergence, 48° to 52° F; fry and
juvenile rearing, 45° to 60° F, and smoltification, < 57°
F (Bovee 1978, Reiser and Bjornn 1979, Bell 1986).
While egg mortality begins to occur at 56° F and fish
are known to have difficulty extracting oxygen from the
water at temperatures exceeding 70° F (Hooper 1973),
steelhead populations are often adapted to local environmental conditions where preferred temperature conditions are regularly exceeded for prolonged time periods
(McEwan and Jackson 1996).
Some other important factors that are critical to
maintaining optimal steelhead habitat include water
quality and quantity, habitat heterogeneity, migration
barriers, and introduced salmonids. Steelhead require
relatively “ good” water quality (e.g., low suspended sediment and contaminant loads and other forms of pollution), as well as sufficient flows for spawning, rearing,
and migration. Diverse stream habitats consisting of shallow riffles for spawning and relatively deep pools, with
well-developed cover, for rearing are important factors.
The importance of estuarine or riverine tidal wetlands
within the San Francisco Estuary for rearing/foraging or
migrating steelhead are not well understood.
References
Barnhart, R.A. 1986. Species profiles: life histories and
environmental requirements of coastal fishes and
Chapter 2 —
Estuarine Fish and Associated Invertebrates
103
Fish
Egg Predators: Freshwater sculpins.
Juvenile and Smolt Predators: Other large freshwater,
estuarine, and marine piscivorous fish.
Juvenile and Adult Predators: Harbor seals and other
pennipeds.
Habitat/Cover: Riparian, emergent, and palustrine wetland vegetation.
Major Prey Items: Aquatic and terrestrial insects, amphipods, snails, mysid shrimp, small fish.
invertebrates (Pacific Southwest)—steelhead. U.S.
Fish. Wildl. Serv. Biol. Rep. 82(11.60). U.S. Army
Corps of Engrs., TR EL-82-4. 21 pp.
Behnke, R.J. 1992. Native trout of western North America.
Amer. Fish. Soc. Monograph no. 6. 275 pp.
Bell, M.C. 1986. Fisheries handbook of engineering requirements and biological criteria. Fish Passage Development and Evaluation Program, U.S. Army
Corps of Engrs., No. Pacific Div., Portland, OR.
290 pp.
Bovee, K.D. 1978. Probability-of-use-criteria for the
family Salmonidae. Instream Flow Information
Paper 4, U.S. Fish Wildl. Serv., FWS/OBS-78/07.
79 pp.
Burgner, R.L., J.Y. Light, L. Margolis, T. Okazaki, A
Tautz and S. Ito. 1992. Distribution and origins
of steelhead trout (Oncorhynchus mykiss) in offshore
waters of the north Pacific Ocean. International
North Pacific Fisheries Commission. Bull No. 51.
California Department of Fish and Game (CDFG).
1965. California fish and wildlife plan. Sacramento,
CA.
Hallock, R.J., W.F. Van Woert and L. Shapovalov. 1961.
An evaluation of stocking hatchery-reared steelhead rainbow trout (Salmo gairdneri gairdneri) in
the Sacramento River system. Ca. Dept. Fish and
Game Fish Bull. No, 114. 74 pp.
Hooper, D.R. 1973. Evaluation of the effects of flows
on trout stream ecology. Dept. of Eng. Res., Pacific Gas and Electric Co., Emeryville, CA. 97 pp.
Leidy, R.A. 1984. Distribution and ecology of stream
fishes in the San Francisco bay drainage. Hilgardia
52(8): 1-175.
Long-term Management Strategy (LTMS). 1996. Draft
long term management strategy for the placement
of dredged material in the San Francisco Bay Region. Prepared by the LTMS Multi-Agency Writing Team for the LTMS Management Committee. Vol II, Appendices.
Meyer Resources Inc. 1988. Benefits from present and
future salmon and steelhead production in California. Report to the California Advisory Committee on Salmon and Steelhead. 78 pp.
McEwan, D. and T.A. Jackson. 1996. Steelhead restoration and management plan for California. Ca.
Dept. Fish and Game, Inland Fisheries Division,
Sacramento, CA. 234 pp.
Mills, T.J. and F. Fisher. 1994. Central Valley anadromous sport fish annual run-size, harvest, and population estimates, 1967 through 1991. Ca. Dept.
Fish and Game. Inland Fisheries Division Tech.
Rept.. Draft. 70 pp.
Moyle, P.B. 1976. Inland Fishes of California. Univ. of
California Press. 405 pp.
Moyle, P.B., R.M. Yoshiyama, J.E. Williams and E.D.
Wikramanayake. 1995. Fish species of special con-
Hypomesus transpacificus
Ted R. Sommer
Bruce Herbold
General Information
The Delta smelt (Family Osmeridae) is a small, shortlived native fish which is found only in the Bay-Delta
Estuary. The species was listed as threatened in 1993 under the Federal Endangered Species Act. Habitat loss is
thought to be one of the most important elements in
causing its decline. New water quality standards adopted
by the state in 1995 are aimed in part at improving habitat conditions (SWRCB 1995).
Reproduction
The Delta smelt has low fecundity and is primarily an
annual species, although a few individuals may survive
a second year (Herbold et al. 1992). The location and
season of Delta smelt spawning varies from year to year.
Spawning, which occurs in shallow freshwater (CDFG
1992b, USFWS 1994), has been known to occur at various sites within the Delta, including the lower Sacramento and San Joaquin rivers and Georgiana Slough,
and in sloughs of the Suisun Marsh (USFWS 1994). In
1996, newly emerged Delta smelt larvae were found in
the Napa River, Cordelia Slough, Montezuma Slough,
and in the San Joaquin River up to Stockton (CDFG
unpub. data). Based on egg and larval trawls in recent
low flow years, it appears that a significant portion of
Delta smelt spawning now takes place in the northern
and western Delta (CDWR 1992).
Spawning may occur from late winter (December)
to early summer (July). In 1989 and 1990, two spawning peaks occurred, one in late-April and another earlyMay (USFWS 1994). Spawning has been reported to
occur at about 45° to 59° F (7-15° C) in tidally influenced rivers and sloughs, including dead-end sloughs and
shallow edge waters of the upper Delta. Most spawning
occurs in fresh water, but some may occur in brackish
water in or near the entrapment zone (Wang 1991). The
demersal, adhesive eggs sink and attach to hard substrates, such as submerged tree branches and roots, gravel
Moyle 1976
Fish
Amphibians &
Reptiles
Delta Smelt
104
Baylands Ecosystem Species and Community Profiles
Plants
cern in California. Ca. Dept. Fish and Game. Inland
Fisheries Division, Rancho Cordova, CA. 272 pp.
Nehlsen, W., J.E. Williams and J.A. Lichatowich. 1991.
Pacific salmon at the crossroads: stocks at risk from
California, Oregon, Idaho, and Washington. Fisheries 16 (2): 4-21.
Pauley, G.B. and B.M. Bortz. 1986. Species profiles:
life histories and environmental requirements of
coastal fishes and invertebrates (Pacific northwest):
steelhead trout. U.S. Fish and Wildl. Serv. Biol.
Rept. 82(11.62). 24 pp.
Reiser, D.W. and T.C. Bjornn. 1979. Habitat requirements of anadromous salmonids. USDA, Forest
Service, Pacific Northwest Forest and Range Experiment Station, Portland, OR, General Tech.
Rept. PNW-96. 54 pp.
Reynolds, F.L., T.J. Mills, R. Benthin and A. Low. 1993.
Restoring Central Valley streams: a plan for action. Ca. Dept. Fish and Game. Inland Fisheries
Division, Rancho Cordova, CA.
Scott, W.B. and E.J. Crossman. 1973. Freshwater fishes of
Canada. Bull. Fish. Res. Brd. Can. 184. 966 pp.
Shapovalov, L. and A.C. Taft. 1954. The life histories of
the steelhead rainbow trout (Salmo gairdneri
gairdneri) and silver salmon (Oncorhynchus kisutch)
with special reference to Waddell Creek, California, and recommendations regarding their management. Ca. Dept. Fish and Game, Fish Bull. 98.
375 pp.
Skinner, J.E. 1962. An historical review of the fish and
wildlife resources of the San Francisco Bay Area.
Ca. Dept. Fish and Game Water Projects Br. Rpt.
No. 1.
Titus, R.G., D.C. Erman and W.M. Snider. In press.
History and status of steelhead in California coastal
drainages south of San Francisco Bay. Ca. Dept.
Fish and Game, Fish Bull.
Wales, J.H. 1941. Development of steelhead trout eggs.
Ca. Dept. Fish and Game 27: 250-260.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: A
guide to the early life histories. Interagency Ecolog. Study Prog. for the Sacramento-San Joaquin
Estuary. Interagency Ecological Workshop,
Asilomar, CA.
(Moyle 1976, Moyle et al. 1992). Downstream distribution is generally limited to western Suisun Bay. During periods of high Delta outflow, Delta smelt populations do occur in San Pablo Bay, although they do not
appear to establish permanent populations there (Herbold et al. 1992). Recent surveys, however, show that
Delta smelt may persist for longer periods in Napa River,
a tributary to San Pablo Bay (IEP, unpub. data).
Rearing and pre-spawning Delta smelt generally
inhabit a salinity range of less than 2 ppt (parts per thousand), although they have been collected at salinities as
high as 10 to 14 ppt (CDFG 1992b). Abundance of prespawning adults typically peaks upstream of the entrapment zone (CDWR and USBR 1994).
Population Status and Influencing Factors
Growth and Development
Newly hatched larvae are planktonic and drift downstream near the surface to the freshwater/saltwater interface in nearshore and channel areas. Maeger (1993)
found that larvae hatched in 10 to 14 days under laboratory conditions and started feeding on phytoplankton
at day four and on zooplankton at day six. Growth is
rapid through summer, and juveniles reach 40 to 50 mm
fork length (FL; the measure to the bottom of the fork
of the tail fin) by early August. Growth slows in fall and
winter, presumably to allow for gonadal development.
Adults range from 55 to 120 mm FL, but most do not
grow larger than 80 mm FL. Delta smelt become sexually mature in the fall at approximately seven to nine months
of age. The majority of adults die after spawning.
Food and Feeding
Newly hatched larvae feed on rotifers and other microzooplankton. Older fish feed almost exclusively on copepods. Prior to 1988, Delta smelt ate almost solely the
native Eurytemora affinis (Herbold 1987). During the
1980s Eurytemora affinis was displaced by the introduced
copepod Pseudodiaptomus forbsii throughout Suisun Bay,
and Delta smelt shifted to a diet of Pseudodiaptomus
forbsii (P. Moyle, pers. comm.).
Distribution
Delta smelt are endemic to the Sacramento-San Joaquin
Estuary. They have been found as far north as the
confluence of the American and Sacramento rivers and
as far south as Mossdale on the San Joaquin River. Their
upstream range is greatest during periods of spawning.
Larvae subsequently move downstream for rearing. Juvenile and adult Delta smelt commonly occur in the surface and shoal waters of the lower reaches of the Sacramento River below Isleton, the San Joaquin River below
Mossdale, through the Delta, and into Suisun Bay
Seven surveys, although not specifically designed to
gather data on Delta smelt populations in the Estuary,
have charted the abundance of Delta smelt. The summer townet survey, which began in 1959 and was primarily designed to measure striped bass abundance, is
considered one of the best measures of Delta smelt abundance because it covers much of the species’ habitat and
represents the longest historical record. Although the
abundance indices vary considerably, they generally remained low between 1983 and 1993. In recent years
moderately wet conditions have produced relatively high
abundances in the summer townet survey. The reduced
population levels during the 1980s appear to have been
consistent throughout the Delta and Suisun Bay, but
declines may have occurred as early as the mid-1970s in
the eastern and southern portions of the Delta (CDWR
and USBR 1993).
The midwater trawl survey provides one of the best
indexes of smelt abundance because it covers most of the
range of Delta smelt (CDWR and USBR 1994). From
1967 through 1975, fall catches were generally greater
than 10 smelt per trawl per month (in 6 of 8 years); from
1976 through 1989, catches were generally less than 10
smelt per trawl per month (in 13 of 14 years). Since
1986, catches have averaged considerably less than one
smelt per trawl per month. The frequency of occurrence
of Delta smelt in the trawls has also declined. Prior to
1983, Delta smelt were found in 30% or more of the
fall trawl catches. In 1983-1985, they occurred in less
than 30% of the catches, and since 1986, they have been
caught in less than 10% of the trawls (Herbold et al.
1992). In 1993, the midwater trawl index was the sixth
highest of the 25 years of record. In 1994, the index
dropped to a 28-year low, but it rebounded again in
1995. Unlike the summer townet survey indices, the
mean catches of Delta smelt have not declined in the
midwater trawl survey. The smelt population is more dispersed in the summer than in the fall. The summer
populations have decreased in average densities while the
Chapter 2 —
Estuarine Fish and Associated Invertebrates
105
Fish
or rocks, and submerged vegetation. Survival of adhesive eggs and larvae is probably significantly influenced
by hydrology at the time of spawning (CDWR and
USBR 1994). Moyle et al. (1992) found no correlation
between female length and fecundity. Females of 5970 mm standard length (SL) ranged in fecundity from
1,247 to 2,590 eggs per fish, with an average of 1,907.
Spawning stock does not appear to have a major
influence on Delta smelt year class success. However, the
low fecundity of this species, combined with planktonic
larvae which likely have high rates of mortality, requires
a large spawning stock if the population is to perpetuate itself. This may not have been an important factor
in the decline of Delta smelt, but it may be important
for its recovery (CDFG 1992b).
106
Baylands Ecosystem Species and Community Profiles
Fish
Amphibians &
Reptiles
from Interagency Ecological Program studies have found
few Delta smelt in agricultural diversions.
Although the effects of the recent high diversions
of fresh water, especially when coupled with drought
conditions from 1987-1992, are the most likely causes
of the decline in the Delta smelt population, other contributing factors may include: the presence of toxic compounds in the water, competition and predation, food
supply, disease, very high outflows, and low spawning
stock.
Toxic contaminants have been identified as a factor that could affect Delta smelt survival (USFWS 1991).
Possible pollutants include heavy metals, pesticides, herbicides, and polycyclic aromatic hydrocarbons. An inverse relationship has been found between copper applications to ricefields and Delta smelt abundance (Herbold, unpub. data), but no toxicity studies have been
conducted to verify the degree to which pollutants in
water and sediments affect Delta smelt.
Research conducted by Bennett (1995) suggests
that competition with inland silversides, a non-native
fish that arrived in the Bay around 1975, working synergistically with low flows, has contributed to Delta smelt
decline. Inland silversides were found to be voracious
predators of larval fish in both field and laboratory experiments. In addition, smelt and silversides may compete for copepods and cladocerans. Hatching and larval
smelt may be extremely vulnerable to schools of foraging silversides, especially in low-outflow years when
Delta smelt are forced into narrower, upstream channels,
where silverside competition and predation may be increased. Evidence suggests that other non-native species,
such as chameleon goby and striped bass, are either direct predators or compete with Delta smelt for food or
habitat (CDWR and USBR 1994). However, it is questionable if striped bass is an important factor when both
striped bass and Delta smelt were abundant in the 1960s,
and the smelt was not a significant prey of the bass
(CDFG 1992b).
Exact food requirements of Delta smelt are not
known, but prey densities in the Estuary appear low relative to other systems in the United States, creating the
potential for food limitation (Miller 1991). Moreover,
there have been several changes in the species composition of zooplankton, with unknown effects on Delta
smelt. The 1988 decline of Eurytemora affinis, a copepod which has been the primary food supply of Delta
smelt, has been identified as a possible factor in the decline of smelt in the Estuary (CDFG 1992b). However,
it may be that declines in E. affinis abundance, due to
the introduction of other copepod species, is not an important factor because the smelt has shifted its diet and
now consumes Pseudodiaptomus forbesi, which was introduced into the Estuary in 1986. The clam, Potamocorbula amurensis, may have an indirect effect on smelt
Plants
fall populations have decreased in numbers of schools
(CDFG 1992b). Data from the Bay Study and the Suisun Marsh study show sharp declines in Delta smelt at
about the same time. The exact timing of the decline is
different in most of the sampling programs, but falls between 1982 and 1985 (Herbold et al. 1992).
As a result of the sharp decline in abundance in the
1980s, the Delta smelt was listed as a federal “ threatened” species by the U.S. Fish and Wildlife Service in
March 1993 and as a State “ threatened” species by the
California Department of Fish and Game in December
1993.
No single factor appears to be the sole cause of the
Delta smelt decline; however declines have been attributed primarily to restricted habitat and increased losses
through entrainment by Delta diversions (CDWR 1992,
Herbold et al. 1992, USFWS 1994). Reduced water flow
may intensify entrainment at pumping facilities as well
as reduce the quantity and quality of nursery habitat.
Outflow also controls the location of the entrapment
zone, an important part of the habitat of Delta smelt. A
weak, positive correlation exists between fall abundance
of Delta smelt and the number of days during spring that
the entrapment zone remained in Suisun Bay (Herbold
1994). The number of days when the entrapment zone
has been in Suisun Bay during the February through
June period is one of only two parameters found so far
that predicts Delta smelt abundance (Herbold 1994).
Reduced suitable habitat and increased entrainment occurs when the entrapment zone moves out of the shallows of Suisun Bay and into the channels of the lower
Sacramento and San Joaquin rivers as a result of low
Delta outflow. The movement of the entrapment zone
to the river channels not only decreases the amount of
area that can be occupied by smelt, but also decreases
food supply (Herbold et al. 1992).
Delta smelt in the western delta are vulnerable to
entrainment by the pumps of the State Water Project
and the Central Valley Project, as well as local agricultural diversions (CDWR 1992, NHI 1992, Herbold et
al. 1992). Diversions in the northern and central Delta,
where smelt are most abundant, are likely the greatest
source of entrainment (USFWS 1994). Larvae and juveniles appear to be particularly vulnerable to pumping
because screens are not effective for these life stages
(CDWR and USBR 1994). Whether entrainment, as estimated by salvage, affects abundance remains to be demonstrated statistically. However, the relative effects of
entrainment are higher in dry years, when the abundance
of Delta smelt is typically lowest and the distribution of
the species shifts closer to the pumps in the interior
Delta. Water diversions such as Contra Costa Canal,
PG&E’s power plants, and in-Delta agricultural diversions, potentially entrain Delta smelt in numbers comparable to or greater than at the Central Valley Project
and State Water Project pumps. However, initial results
Trophic Levels
Delta smelt are secondary consumers.
Proximal Species
Egg and larvae predators: Inland silversides, Menidia
beryllina.
Juvenile and adult predators: Striped bass, Morone
saxatilis (likely).
Prey: Eurytemora affinis, Pseudodiaptomus forbsii, rotifers (e.g., Trichocerca).
Good Habitat
Spawning habitat has been as widely dispersed as the
Napa River to Stockton in 1996. The predominate feature appears to be shallow, freshwater conditions with
some sort of solid substrate for the attachment of eggs.
Spawning has been reported to occur at about 45-59° F
( 7-15° C) in tidally influenced rivers and sloughs including dead-end sloughs and shallow edge waters of the
upper Delta.
Juvenile and adult Delta smelt commonly occur in
the surface and shoal waters of the lower reaches of the
Sacramento River below Mossdale, through the Delta,
and into Suisun Bay (Moyle 1976, Moyle et al. 1992).
Rearing and pre-spawning Delta smelt generally inhabit
a salinity range of less than 2 ppt, although they have
been collected at salinities as high as 10 to 14 ppt
(CDFG 1992a). Analysis of the salinity preferences using midwater trawl data indicate that Delta smelt distribution peaks upstream of the entrapment zone
(Obrebski 1993)1. It should be noted, however, that the
distribution of Delta smelt is fairly broad, particularly
in years when abundance levels are high (CDWR and
USBR 1993). Evidence from the 1993 year class also
demonstrates that salt field position does not necessarily regulate Delta smelt distribution in all years. In late
1993 and early 1994, Delta smelt were found in Suisun
Bay region despite the fact that X22 was located upstream. Samples collected in this area demonstrated that
high levels of the copepod Eurytemora were present, suggesting that food availability may also influence smelt distribution (CDWR and USBR 1994).
Although these results show that the Delta smelt
is not an entrapment zone specialist, there is evidence
that their abundance is correlated with X2. Herbold
(1994) found a significant relationship between the
number of days X2 was in Suisun Bay during February
through June versus midwater trawl abundance. Furthermore, when the entrapment zone is in Suisun Bay and
both deep and shallow water exists, Delta smelt are
caught most frequently in shallow water (Moyle et al.
1992).
Results from the University of California, Davis
provide an indication of environmental tolerances of
Delta smelt (Swanson and Cech 1995). The study found
that although Delta smelt tolerate a wide range of water
temperatures (<8° C to >25° C), warmer temperatures
apparently restrict their distribution more than colder
temperatures.
References
Bennett, W.A. 1995. Potential effects of exotic inland
silversides on Delta smelt. Interagency Program
Newsletter. Winter 1995: 4-6.
California Department of Fish and Game (CDFG).
1992a. A re-examination of factors affecting striped
bass abundance in the Sacramento-San Joaquin Estuary. Entered by the Ca. Dept. of Fish and Game
for the State Wat. Res. Cont. Bd. 1992 Water
Rights Phase of the Bay-Delta Estuary Proceedings. (WRINT-DFG-2) 59 pp.
______. 1992b. Written testimony on Delta smelt. Submitted by the Ca. Dept. of Fish and Game to the
1
The entrapment zone, also referred to by a variety of other
discriptive terms, such as the “ mixing zone,” the “ null
zone,” and and the “ zone of maximum turbidity,” is the
area within an estuary where the freshwater from a stream
meets with the salt water of the ocean. This zone is biologically highly productive, and considered to be of critical
importance to the aquatic food web of the Estuary.
2
“ X2” is the geographic location, measured in kilometers
above the Golden Gate, of the entrapment zone. X2 is
largely a function of outflow, such that when outflow is
high, X2 is closer to the Golden Gate. X2 was used by U.S.
Fish and Wildlife Service in defining Delta Smelt’s critical
habitat under the Endangered Species Act (USFWS 1994).
Chapter 2 —
Estuarine Fish and Associated Invertebrates
107
Fish
populations by reducing its food supply (Herbold et al.
1992).
In some years disease is thought to cause widespread mortality of some fish species in the Estuary, but
mortality of Delta smelt has not been specifically observed (Stevens et al. 1990). Mycobacterium, a genus of
bacteria known to cause chronic infections in fish and
other species, has been the major cause of mortality of
Delta smelt held in the laboratory, and it may cause
deaths among wild fish as well (Hedrick 1995).
The period of the Delta smelt decline includes unusually wet years with exceptionally high outflows. Very
high outflows may be detrimental to the planktonic smelt
larvae, which may be transported out of the Delta and
into San Pablo and San Francisco bays with no way to
get back upstream (CDFG 1992b).
It is possible that the size of the spawning stock
influences population levels. However, there is not a statistically significant stock-recruitment relationship for
Delta smelt, so this factor is not considered a primary
factor regulating abundance (CDWR and USBR 1994).
108
Baylands Ecosystem Species and Community Profiles
Fish
Amphibians &
Reptiles
State Wat. Res. Cont. Bd. June 1992. 35 pp.
(WRINT-NHI-9)
Obrebski, S. 1993. Relationships between Delta smelt
abundance and the entrapment zone position.
Draft report for Dept. of Water Res.. 29 pp.
Stevens, D.E., L.W. Miller and B.C. Bolster. 1990. Report to Fish and Game Commission: A status review of Delta smelt (Hypomesus transpacificus) in
California. Dept. of Fish and Game. Candidate
Species Status Report 90-2.
Swanson, C. and J. Cech. 1995. Environmental tolerances and requirements of the Delta smelt,
Hypomesus transpacificus. Report to Dept. of Water Resources: Contracts B-59449 and B-58959.
State Water Resources Control Board (SWRCB). 1995.
Water quality control plan for the San Fran. Bay/
Sac.-San Joaquin Estuary. 95-1WR, May 1995.
Sacramento, Ca.. 45 pp plus appendices.
Wang, J.C.S. 1991. Early life stages and early life history of Delta smelt, Hypomesus transpacificus, in
the Sacramento-San Joaquin Estuary, with comparison of early life stages of the longfin smelt,
Spirinchus thaleichthys. Interagency Program Tech.
Rept. 28. 52 pp.
U.S. Fish and Wildlife Service (USFWS). 1991. Federal Register: Endangered and threatened wildlife
and plants: proposed threatened status for the Delta
smelt. Dept. of the Interior, Fish and Wildlife Service. 50 CFR Part 17. 56(192): 50075-50083. October 3, 1991.
______. 1994. Biological opinion on the operation of
the Central Valley Project and State Water Project
effects on Delta smelt. February 4, 1994. U.S.
Fish and Wildlife Service, Region 1, Portland,
OR. 34 pp.
Personal Communications
Peter Moyle, University of California, Davis.
Plants
State Wat. Res. Cont. Bd. June 1992. (WRINTDFG-9) 44 pp.
California Department of Water Resources (CDWR).
1992. Bay-Delta fish resources. Sacramento, CA.
July 1992. (WRINT-DWR-30) 46 pp.
California Department of Water Resources and U.S.
Bureau of Reclamation (CDWR and USBR).
1993. Biological Assessment. Effects of the Central Valley Project and State Water Project on
Delta smelt. Prepared by the Ca. Dept. of Water Res. and the U.S. Bureau of Reclamation for
the U.S. Fish and Wildlife Service. October
1993. 134 pp.
______. 1994. Biological Assessment. Effects of the Central Valley Project and State Water Project on Delta
Smelt and Sacramento Splittail.
Hedrick, R.P. 1995. Disease Research in Delta Smelt.
Report to Dept. of Wat. Res.: Contract B-59299.
Herbold, B. 1987. Patterns of co-cccurrence and resource
use in a non-coevolved assemblage of fishes.
Ph.D dissertation. Univ. of California, Davis.
Vii+81 pp.
______. 1994. Habitat requirements of Delta smelt. Interagency Program Newsletter. Winter 1994: 1-3.
Herbold, B., P. Moyle and A. Jassby 1992. Status and
trends report on aquatic resources in the San Francisco Estuary. San Fran. Est. Proj. Public Rept.
March 1992. 257 pp. plus appendices.
Miller, L. 1991. 1990 Working Papers of the Food Chain
Group. Interagency Program Working Papers 1-6.
FCG-1990. 71 pp.
Moyle, P. 1976. Inland Fishes of California. Univ. of
California Press, Berkeley. 405 pp.
Moyle P., B. Herbold, D. Stevens and L. Miller. 1992.
Life history and status of Delta smelt in the Sacramento-San Joaquin Estuary, California. Transactions of the Am. Fisheries Soc. 121:67-77.
Natural Heritage Institute (NHI). 1992. Causes of decline in estuarine fish species. Testimony presented
by Dr. Peter Moyle, Univ. of Ca., Davis to the
Longfin Smelt
Spirinchus thaleichthys
Frank G. Wernette
General Information
CDFG
Reproduction
Maturation of longfin smelt begins late in the second
summer of their life in August and September. As they
mature, the smelt begin migrating upstream from San
Francisco and San Pablo bays toward Suisun Bay and the
Delta. Longfin smelt spawn in fresh water, primarily in
the upper end of Suisun Bay and in the lower and middle
Delta. In the Delta, they spawn mostly in the Sacramento River channel and adjacent sloughs (Wang 1991).
During the recent drought, when saline water intruded
into the Delta, larval longfin smelt were found near the
Central Valley Project and State Water Project export
facilities in the southern Delta (Wang 1991). Ripe
adults, larvae, and juveniles are salvaged at the export
facilities in every below normal or drier water year
(Baxter, pers. comm.). The eggs are adhesive and are
probably deposited on rocks or aquatic plants. Longfin
smelt eggs hatch in 37-47 days at 45° F.
Growth and Development
Shortly after hatching, longfin smelt larvae develop a gas
bladder that allows them to remain near the water surface (Wang 1991). The larvae do not vertically migrate,
but instead remain near the surface on both the flood
and ebb tides (CDFG 1992). Larvae are swept downstream into nursery areas in the western Delta and Suisun and San Pablo bays with larval dispersal farther
downstream in years of high outflow than in years of low
outflow (CDFG 1992; Baxter, pers. comm.). Early development of gas bladders by longfin smelt causes the
larvae to remain near the surface much longer than Delta
smelt larvae. That factor and earlier spawning period help
explain why the longfin smelt larvae are dispersed much
farther downstream in the Estuary than are Delta smelt
larvae (Baxter, pers. comm.). Larval development occurs
primarily in the February through May period and peaks
during February-April (CDFG 1992).
Metamorphosis of longfin smelt from the larval to
juvenile form begins 30-60 days after hatching, depending on temperature. Most longfin smelt growth occurs
during the first summer, when length typically reaches
Chapter 2 —
Estuarine Fish and Associated Invertebrates
109
Fish
The longfin smelt (Family: Osmeridae) is a three to
seven-inch long silvery fish (Moyle 1976). Longfin smelt
were the most abundant smelt species in the Bay-Delta
Estuary prior to 1984 and have been commercially harvested (Wang 1986). In 1993, the U.S. Fish Wildlife
Service (USFWS) was petitioned to list the longfin smelt
under the federal Endangered Species Act. In January
1994, however, USFWS determined that the longfin
smelt did not warrant listing because other longfin smelt
populations exist along the Pacific Coast, the Bay-Delta
Estuary population does not appear to be biologically significant to the species as a whole, and the Bay-Delta
Estuary population may not be sufficiently reproductively isolated (Federal Register Vol. 59 No. 869, January 6, 1994). Still, longfin smelt are typically addressed
in Biological Assessments because of the decline in their
abundance after 1982 and the relatively small increase
in abundance following a wet year in 1993. The species
may also be considered in the future for listing under
the California Endangered Species Act.
The longfin smelt is an euryhaline species with a
2-year life cycle. Spawning occurs in fresh water over
sandy-gravel substrates, rocks, or aquatic plants. Spawning may take place as early as November and extend into
June, although the peak spawning period is from January to April. After hatching, larvae move up into surface
water and are transported downstream into brackishwater nursery areas. Delta outflow into Suisun and San
Pablo bays has been positively correlated with longfin
smelt recruitment because higher outflow increases larval dispersal and the area available for rearing. The
longfin smelt diet consists of mysids, although copepods and other crustaceans also are eaten. Longfin smelt
are preyed upon by fishes, birds, and marine mammals
(Federal Register Vol. 59 No. 4, January 3, 1994).
In the Bay-Delta Estuary, the decline in longfin
smelt abundance is associated with freshwater diversion
from the Delta. Longfin smelt may be particularly sensitive to adverse habitat alterations because their 2-year
life cycle increases their likelihood of extinction after con-
secutive periods of reproductive failure due to drought
or other factors. Relatively brief periods of reproductive
failure could lead to extirpations (Federal Register Vol.
59 No. 4, January 3, 1994).
Although the southernmost populations of longfin
smelt are declining, little or no population trend data are
available for estuaries in Oregon and Washington.
Longfin populations may not be isolated since there is
little genetic variation between northern and southern
populations. Under prolonged drought conditions however, only the Colombia River and San Francisco Bay
stocks may survive.
6 to 7 cm. During their second summer, smelt reach 9
to 11 cm in length (NHI 1992). Most longfin smelt
spawn and die at two years of age (CDFG 1992).
Fish
Food and Feeding
The main prey of adult longfin smelt is the opossum
shrimp, Neomysis mercedis (NHI 1992). There is little
information on food habitats of longfin smelt larvae, but
fish larvae of most species, including Delta smelt, are
known to feed on phytoplankton and small zooplankton,
such as rotifers and copepod nauplii (Hunter 1981,
USBR 1993). Juvenile longfin smelt feed on copepods,
cladocerans, and mysids. The mysid Neomysis mercedis
is the most important prey of larger juveniles.
Longfin smelt are widely distributed in estuaries on the
Pacific Coast. They have been collected from numerous
river estuaries from San Francisco to Prince William
Sound in Alaska (Moyle 1976).
Longfin smelt are euryhaline meaning they are
adapted to a wide salinity range. They are also anadromous. Spawning adults are found seasonally as far upstream in the Delta as Hood, Medford Island, and the
Central Valley Project and State Water Project fish collection facilities in the southern Delta. Historically, before construction of Shasta Dam in 1944, saline water
intruded in dry months as far upstream in the Delta as
Sacramento, so longfin smelt may have periodically
ranged farther upstream than they do currently (Herbold
et al. 1992).
Except when spawning, longfin smelt are most
abundant in Suisun and San Pablo bays, where salinity
generally ranges between 2 ppt and 20 ppt (NHI 1992).
Pre-spawning adults and yearling juveniles are generally
most abundant in San Pablo Bay and downstream areas
as far as the South Bay and in the open ocean.
Population Status and Influencing Factors
Abundance estimates were developed from otter trawl
and midwater trawl sampling conducted by the Outflow/
Bay study as part of the Interagency Ecological Program.
Fall midwater trawl surveys provide the longest index of
longfin smelt abundance.
Results of the fall midwater trawl surveys indicate
that, like Sacramento splittail abundance, longfin smelt
abundance has been highly variable from year to year,
with peaks and declines coinciding with wet and dry
periods. Longfin smelt abundance has steadily declined
since 1982. Abundance continued to be suppressed during the drought years beginning in 1987. Longfin abundance was very low from 1987 to 1992, with 1992 having the lowest index on record. Abundance increased
110
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Distribution
moderately in 1993 following the drought during a period of improved Delta outflow.
Year-class abundance of longfin smelt appears to
depend on the environmental conditions experienced by
the eggs and young fish. Generally, year-class abundance
is positively related to Delta outflow (i.e., high abundance follows high outflow during winter and spring).
Factors possibly contributing to the recent decline in
longfin smelt abundance are reduced Delta outflow, entrainment in diversions, introductions of exotic species,
loss of habitat, and the recent drought.
Delta Outflow – Higher outflows result in higher
longfin smelt survival. An index of survival computed as
the ratio of the index of abundance from fall midwater
trawl surveys to an index of larval abundance in previous springs was strongly correlated (r=0.95) with December-August outflow. Delta outflow or factors associated
with outflow affect survival of larvae and early juveniles.
Delta outflow may be the single most important factor
controlling longfin smelt abundance. High outflows
increase dispersion downstream, available habitat, and
possibly, food availability. High outflow may also reduce
predation and the effects of other adverse factors (i.e.,
toxin concentrations). Low outflow conditions reduce
downstream dispersion and increase vulnerability to entrainment in Delta diversions.
Longfin smelt abundance (according to the fall
midwater trawl survey index) is positively related to Delta
outflow (Stevens and Miller 1983, CDFG 1992). Regression analysis indicated that 79% of variability in the
midwater trawl survey index is explained by changes in
January and February Delta outflow. The significant relationship between the index of abundance from the fall
midwater trawl surveys and Delta outflow may reflect the
effect of outflow on survival of larvae and early juveniles.
Year-class strength may be largely determined by survival
of the early life stages.
High Delta outflow may increase the amount of
suitable brackish water rearing habitat; reduce salinity
in the Estuary, reducing competition and predation by
marine organisms; reduce predation because young smelt
are more dispersed and turbidity is higher; increase phytoplankton and zooplankton production; and increase
transport of larvae out of the Delta and away from diversions (CDFG 1992; Stevens and Miller 1983; Baxter,
pers. comm.). Any of these mechanisms may be responsible for the observed relationship between Delta outflow and longfin smelt abundance.
The position of the entrapment zone1, location of
2
X2 , and volume of critical nursery habitat are determined
by Delta outflow. In addition to the relationship with outflow, the fall midwater trawl survey index has a positive
relationship with the location of X2 and the volume of critical nursery habitat (Jassby 1993, Herrgesell 1993).
Delta smelt abundance tends to be highest when
X2 has an intermediate value (i.e., X2 is located in up-
1
The entrapment zone, also referred to by a variety of other
discriptive terms, such as the “ mixing zone,” the “ null
zone,” and and the “ zone of maximum turbidity,” is the
area within an estuary where the freshwater from a stream
meets with the salt water of the ocean.
2
“ X2” is the geographic location, measured in kilometers
above the Golden Gate, of the entrapment zone.
tection facilities. Based on the high salvage rates of
young-of-year juveniles in some years, it can be assumed
that many thousands of longfin smelt larvae were also
entrained, especially during February through April.
During years of high flows, most longfin smelt
adults spawn in the western Delta, and their larvae are
generally transported out of the Delta and therefore are
unlikely to be entrained in Delta diversions in large numbers. During the 1987-1992 drought and other low flow
years, however, outflows were low and exports were high.
Adults, larvae, and juveniles remained in the Delta, as
indicated by salvage at the Central Valley Project and
State Water Project fish protection facilities. Most juveniles were entrained during April-June and averaged
30-45mm long, with length correlated with the month
of entrainment. Thus, longfin smelt suffer not only loss
of larval dispersal and rearing habitat in a drought, but
also from higher rates of entrainment.
Adult, juvenile, and larvae longfin smelt are vulnerable to entrainment in diversions other than exports
at the Central Valley Project and State Water Project
pumps, including diversions to PG&E’s power generating plants, industrial diversions, agricultural diversions,
and others. However, entrainment of longfin smelt in
these diversions has not been extensively evaluated.
Other Factors – Other factors that may affect survival of longfin smelt include food limitation and presence of toxic materials and introduced species. Abundance of Neomysis and other zooplankton prey (e.g.,
rotifers) of longfin smelt have declined in recent years
(Obrebski et al. 1992). It is not known what effect the
decline in prey abundance has had on longfin smelt;
however, food limitation may be important because yearclass strength of many fish populations, particularly species with planktonic larvae, may be strongly influenced
by feeding conditions during the larval life stage (Lasker
1981).
Agricultural chemicals (including pesticides and
herbicides), heavy metals, petroleum-based products,
and other waste materials toxic to aquatic organisms
enter the Estuary through nonpoint runoff, agricultural
drainage, and municipal and industrial discharges. The
effects of toxic substances have not been tested on longfin
smelt, but recent bioassays indicate that water in the Sacramento River is periodically toxic to larvae of the fathead
minnow, a standard EPA test organism (Stevens et al.
1990). The short life span of longfin smelt and relatively
low position in the food chain probably reduce the accumulation of toxic materials in their tissues and make
them less susceptible to injury than species that live
longer (NHI 1992).
Many exotic species have invaded the Estuary in
recent years. These species may compete with or prey
on longfin smelt. No single invasion of exotic species parallels the decline the longfin smelt closely enough to suggest that competition from or predation by the species
Chapter 2 —
Estuarine Fish and Associated Invertebrates
111
Fish
per Suisun Bay). The location of X2 is also a good predictor of longfin smelt abundance. Since X2 and the
volume of critical nursery habitat are largely determined
by Delta outflow, the relationship between longfin smelt
abundance and the location of X2 or volume of critical
habitat may simply reflect effects of outflow or other
correlates of outflow on longfin smelt abundance.
Lower San Joaquin River – Reverse flow in the
lower San Joaquin River usually transports relatively
fresh water drawn from the Sacramento River and may
increase upstream migration of adults to the southern
Delta. Reverse flow may also transport larvae to the
southern Delta. In the southern Delta, adults, larvae, and
juveniles are vulnerable to entrainment, predation, and
other sources of mortality.
Entrainment – Entrainment of longfin smelt by
Delta diversions affects spawning adults, larvae, and early
juveniles. Older juveniles and prespawning adults generally inhabit areas downstream of the Delta. Salvage at
both the Central Valley Project and State Water Project
fish protection facilities has varied greatly between years.
Salvage represents entrainment, but the number of fish
salvaged is often much lower than total number entrained because fish, particularly those smaller than
about 20-30 mm, pass through the fish screens at the
salvage facilities and, therefore, are not salvaged.
With the exception of 1986, a wet year, the annual
salvage of longfin smelt at the Central Valley Project and
State Water Project pumps was much higher during
1984-1990 than during 1979-1983. The decline in
abundance in 1984 may be attributable to increased entrainment by the Central Valley Project and State Water Project pumps and other diversions, but reduced
Delta outflow, discussed previously, may be a more important factor affecting abundance.
Entrainment of adult longfin smelt has a potentially
greater adverse effect on the population than entrainment of larvae and young juveniles because unless the
adults have already spawned, their reproductive value is
much greater than that of younger fish. Adult smelt are
entrained at the State Water Project and Central Valley
Project pumping facilities primarily during NovemberFebruary. The number of adults entrained is low relative to the number of juveniles entrained.
Longfin smelt larvae have been captured in the
southern Delta near the Central Valley Project and State
Water Project export facilities (Spaar 1990, 1993; Wang
1991). Larval smelt are too small to be salvaged at the
State Water Project and Central Valley Project fish pro-
was a primary cause of the longfin smelt’s recent decline.
The effects of multiple-species invasion, which have occurred in the Estuary, are extremely difficult to assess.
The effects of exotic species invasions on longfin smelt
is likely not large since Delta outflow explains over 60%
of the variation in abundance (Baxter, pers. comm.).
Fish
Trophic Levels
Longfin smelt are secondary consumers.
Proximal Species
Predators: Brown pelican, river otter, striped bass,
centrarchids.
Prey: Zooplankton (cladocerans), opossum shrimp
(Neomysis mercedis), crustaceans (copepods).
Longfin smelt are typically pelagic and use the larger
sloughs and rivers of the Delta and Bay. The optimal
salinity habitat for non-spawning adults is 2 to 20 ppt.
Optimal salinity habitat for spawning adults is 0 to 2 ppt.
Optimum habitat for spawning includes submergent
vegetation that can be used as a substrate for the adhesive eggs. High quality habitat is also defined as having
low levels of exposure to entrainment into water export
facilities and agricultural or managed wetland diversions.
Adjacent runoff of agricultural pesticides is minimal or
does not occur in good habitat areas.
Juvenile longfin use the open water, shallow shoal
areas of San Pablo and Suisun bays after being transported downstream from spawning areas in the Delta.
An average X2 location in upper Suisun Bay defines good
habitat conditions for longfin smelt. Adjacent tidal wetlands are important to supporting the nutrient cycling
and carbon input functions which in turn support the
prey species upon which longfin feed.
References
California Department of Fish and Game (CDFG).
1992. Estuary dependent species. (Exhibit 6.)
Entered by the Ca. Dept. of Fish and Game for
the State Wat. Res. Cont. Bd. 1992 Wat. Qual./
Wat. Rights Proceedings on the San Fran. Bay/
Sac.-San Joaquin Delta. Sacramento, CA.
Herbold, B., A.D. Jassby and P.B. Moyle. 1992. Status
and trends report on aquatic resources in the San
Francisco Estuary. San Francisco Estuary Project,
U.S. Env. Protection Agency. Oakland, CA.
Herrgesell, P.L. 1993. 1991 annual report. Interagency
Ecological Studies Program for the SacramentoSan Joaquin Estuary. Ca. Dept. Fish and Game.
Stockton, CA.
112
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Good Habitat
Hunter, J.R. 1981. Feeding ecology and predation of
marine fish larvae. In: R. Lasker (ed). Marine fish
larvae. Univ. of Washington Press. Seattle, WA.
Jassby, A.D. 1993. Isohaline position as a habitat indicator for estuarine resources: San Francisco estuary. In: SFEP 1993. Managing freshwater discharge
to the San Francisco Bay/Sacramento-San Joaquin
Delta Estuary: the scientific basis for an estuarine
standard, Appendix 3.
Lasker, R. 1981. The role of a stable ocean in larval fish
survival and subsequent recruitment. In: R. Lasker
(ed). Marine fish larvae. pp. 80-87. Univ. of Washington Press. Seattle, WA.
Moyle, P.B. 1976. Inland Fishes of California. Univ. of
California Press. Berkeley, CA.
Natural Heritage Institute (NHI). 1992. Petition for listing under the Endangered Species Act, longfin
smelt and Sacramento splittail. San Francisco, CA.
Obrebski, S., J.J. Orsi and W.J. Kimmerer. 1992. Longterm trends in zooplankton distribution and abundance in the Sacramento-San Joaquin estuary in
California. (FS/BIO-IATR/92-93, Tech. Rept. 32.)
Ca. Dept. of Water Res.. Sacramento, CA. Prepared for Interagency Ecological Studies Program
for the Sacramento-San Joaquin Estuary, Stockton, CA.
Spaar, S.A. 1990. Results of 1988 striped bass egg and
larva study near the State Water Project and Central Valley Project facilities in the Sacramento-San
Joaquin Delta. A cooperative study by Ca. Dept.
of Water Res., State Wat. Res. Cont. Bd., U.S.
Fish and Wildl. Serv., Ca. Dept. of Fish and Game,
U.S. Bur. of Rec., and U.S. Geol. Survey. Prepared
for Interagency Ecological Study Program for the
Sacramento-San Joaquin Estuary, Stockton, CA.
______. 1993. 1992 entrainment of eggs and larvae to
the State Water Project and Central Valley Project
intakes in the Sacramento-San Joaquin Delta. January. Ca. Dept. of Water Res. Sacramento, CA.
Stevens, D.E. and L.W. Miller. 1983. Effects of river
flow on abundance of young Chinook salmon,
American shad, longfin smelt, and Delta smelt in
the Sacramento-San Joaquin River system. No.
Amer. J. Fish. Mgmt. 3:425-437.
Stevens, D.E., L.W. Miller and B.C. Bolster. 1990. Report to the Fish and Game Commission: a status
review of the Delta smelt (Hypomesis transpacificus)
in California. (Candidate Species Status Report 902.) Ca. Dept. Fish and Game. Stockton, CA.
U.S. Bureau of Reclamation (USBR). 1993. Effect of
the Central Valley Project and State Water Project
on Delta smelt. Prepared by Ca. Dept. of Water
Res. and U.S. Bureau of Reclamation, Mid-Pacific
Region. Prepared for U.S. Fish and Wildl. Service,
Ecological Services, Sacramento Field Office, Sacramento, CA.
Personal Communications
Randall Baxter, Assoc. Fisheries Biologist, California
Department of Fish and Game.
survive and spawn beyond one or multiple spawning seasons). Spawning occurs several times during a spawning
season (Clark 1929). The eggs are demersal and adhesive, and can often be found on vegetation in shallow
nearshore coastal habitats and in estuaries and bays
(Clark 1929, Wang 1986).
Adults move inshore and into bays and estuaries
to spawn during late winter and early spring (Clark 1929,
Wang 1986). In San Francisco Bay, spawning occurs
from October to early August (Wang 1986). Spawning
in San Pablo Bay reportedly occurs from September to
April (Ganssle 1966). Eggs are laid on substrates/vegetation (e.g., Zostera spp., Gracilaria spp., hydroids) in
which they become entangled (Frey 1971, Wang 1986).
Embryonic development is indirect and external, and if
given a suitable environment, the yellowish-orange eggs
hatch within seven days (Wang 1986). The fecundity of
jacksmelt is not yet documented, but probably exceeds
2,000 eggs per female (Emmett et al. 1991). Unfertilized jacksmelt eggs are spherical in shape and 0.92.2 mm in diameter (Clark 1929); fertilized eggs are 1.92.5 mm in diameter (Wang 1986).
Growth and Development
Jacksmelt
Atherinopsis californiensis
Michael K. Saiki
General Information
Although jacksmelt (Family: Atherinidae) is not an important commercial fish, it nevertheless constitutes the
largest portion of “ smelt” captures in California (Emmett
et al. 1991). This species is also commonly caught by
recreational anglers fishing from piers (Frey 1971). In
an ecological sense, jacksmelt occupy an important niche
in trophic pathways of nearshore coastal, bay, and estuarine ecosystems (Clark 1929, Allen and DeMartini 1983,
CDFG 1987).
After hatching, larvae remain on the bottom for a moment and then actively swim near the surface (Wang
1986). Larvae vary in size from 7.5 to 8.6 mm immediately after hatching, to about 25 mm long prior to the
juvenile transformation (Clark 1929, Wang 1986). At
eight days posthatch, they average 10.5-11.7 mm in
length whereas at 24 days posthatch, they average 17.620.3 mm in length (Middaugh et al. 1990). Juveniles can
attain 110 mm during their first year, and 180-190 mm
after two years (Clark 1929). All individuals mature by
their third year, but some may grow quickly and mature
in their second year (Clark 1929). Adult jacksmelt
have been reported to attain a length of 780 mm and
an age of 11 years (Miller and Lea 1972, Frey 1971)
but, more typically, the maximum size is 200 mm total
length, and the maximum age is 9-10 years (Clark
1929).
Reproduction
CDFG
Emmett et al. (1991) describes the sexual and reproductive characteristics of jacksmelt as gonochoristic (its gender is determined by developmental rather than hereditary mechanisms) and iteroparous (it has the capacity to
Food and Feeding
The jacksmelt is omnivorous (Bane and Bane 1971,
Ruagh 1976). Larvae live on their yolk-sac for about 48
hours after hatching when it is fully absorbed (Middaugh
et al. 1990). Major food items for jacksmelt include algae (Ulothrix spp., Melosira moniliformis, Enteromorpha
spp., and other filamentous algae), benthic diatoms, crustaceans (mysids, copepods, decapod larvae), and detritus (Bane and Bane 1971, Ruagh 1976). In addition,
stomach analyses of juvenile jacksmelt show that amphipods are a common food item, indicating that juveniles
may feed on the bottom (Wang 1986).
Chapter 2 —
Estuarine Fish and Associated Invertebrates
113
Fish
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and adjacent waters, California:
A guide to the early life histories. (FS/10-4ATR869). Ca. Dept. of Water Res. Sacramento, CA. Prepared for Interagency Ecological Study Program
for the Sacramento-San Joaquin Estuary, Sacramento, CA.
______. 1991. Early life stages and early life history of
the Delta smelt, Hypomesus transpacificus, in the
Sacramento-San Joaquin Estuary, with comparison of early life stages of the longfin smelt,
Spirinchus thaleichthys. (FS/BIO-IATR/91-28.
Tech. Rept. 28). Ca. Dept. of Water Res. Sacramento, CA. Prepared for Interagency Ecological
Studies Program for the Sacramento-San Joaquin
Estuary, Stockton, CA.
Fish
Proximal Species
Jacksmelt occur from Santa Maria Bay, Baja California,
northward to Yaquina Bay, Oregon (Miller and Lea 1972,
Eschmeyer et al. 1983). However, this species is uncommon north of Coos Bay, Oregon (Emmett et al. 1991).
Prior to or after the spawning season, adult jacksmelt typically occur in coastal waters near shore (Baxter
1960). Ruagh (1976) mentioned that jacksmelt are usually caught within 5 km of shore where they often school
with topsmelt (Atherinops affinis).
Locally, jacksmelt have been reported to spawn in
San Francisco Bay (Wang 1986) and San Pablo Bay
(Ganssle 1966, Wang 1986). Juveniles are also present
in San Francisco Bay (Baxter 1960, Aplin 1967), San
Pablo Bay (Ganssle 1966), Carquinez Strait (Messersmith 1966), and occasionally in Suisun Bay (Wang
1986, Herbold et al. 1992, Jones and Stokes Assoc.
1979) and Napa marsh (Jones and Stokes Assoc. 1979).
The amount of freshwater inflow seemingly affects the
local distribution of jacksmelt. During years of low freshwater inflow, jacksmelt occur as far upstream as Carquinez
Strait and San Pablo Bay, but during high-flow years they
are seemingly restricted to Central San Francisco Bay and
South San Francisco Bay (CDFG 1987).
Predators: Yellowtail (Seriola lalandei), sharks and other piscivorous fishes, piscivorous birds (e.g., brown pelicans and gulls).
Prey: Small crustaceans, algae.
Habitat: Kelp (cover for juveniles and adults); algae,
hudroids, and eelgrass (spawning substrate).
Parasites: Nematodes sometimes found living in flesh.
Population Status and Influencing Factors
Presently, jacksmelt are particularly abundant in Tomales, Central San Francisco, South San Francisco, and San
Pablo bays (Emmett et al. 1991). Midwater trawl samples
performed in South San Francisco Bay between 19801988 showed that jacksmelt were the second most common species caught, behind northern anchovy (Herbold
et al. 1992). Furthermore, jacksmelt were more abundant and occurred more frequently than topsmelt in the
South Bay (Herbold et al. 1992). In San Pablo and Central San Francisco bays, Herbold et al. (1992) reported
that jacksmelt were the third most common species
caught. Midwater trawl samples performed in the
Carquinez Strait between 1961-1962 found over 9% of
the total catch consisted of jacksmelt (Messersmith
1966). Herbold et al. (1992) noted that during 19801988 jacksmelt numbers seemed to vary widely in the
Central Bay and are seemingly unpredictable from year
to year, whereas numbers in the South Bay show little
variation from year to year.
Although specific studies relating fish abundance
to environmental variables were not found during our
search of the literature, jacksmelt may be vulnerable to
pollution and habitat modifications because they depend
on embayments and estuaries for spawning.
Trophic Levels
Omnivorous (primary and higher order consumers).
114
Baylands Ecosystem Species and Community Profiles
Good Habitat
Bays and estuaries provide important spawning habitat
for jacksmelt. In general, the preferred spawning areas
are situated in shallow nearshore habitats containing submerged vegetation (Wang 1986). Water quality variables
suitable for embryo development are as follows: temperature, 10-12° C; and salinity, polyhaline and as low as 5
ppt (Wang 1986). Schools of larvae occur near the water surface over a variety of substrates, but mostly sandy
and muddy bottoms and in the kelp canopy (Frey 1971).
Optimum larval and juvenile survival and growth appears
to be at salinities of 10-20 ppt, indicating that larvae may
prefer mesohaline environments (Middaugh and Shenker
1988, Middaugh et al. 1990). Juveniles and adults prefer sandy bottoms in murky water at depths of 1.5-15
m below the surface (Feder et al. 1974). Furthermore,
they seem to use open waters in San Francisco Bay and
sloughs in and near Suisun Marsh and Napa Marsh
(Jones and Stokes Assoc. 1979). Jacksmelt are apparently
more sensitive than topsmelt to fluctuations in salinity
and temperature (Emmett et al. 1991).
References
Allen, L.G. and E.E. Demartini. 1983. Temporal and
spatial patterns of near shore distribution and abundance of the pelagic fishes off San Onofre-Oceanside,
California. Fish Bull., U.S. 81(3):569-586.
Aplin, J.A. 1967. Biological survey of San Francisco Bay,
1963-1966. Ca. Dept. Fish and Game, Marine
Resources Operations. MRO Ref. 67-4, 131 pp.
Bane, G.W. and A.W. Bane. 1971. Bay fishes of northern
California with emphasis on the Bodega Tomales Bay
area. Mariscos Publ., Hampton Bays, NY, 143 pp.
Baxter, J.L. 1960. Inshore fishes of California. Ca. Dept.
Fish and Game, Sacramento, CA, 80 pp.
California Department of Fish and Game (CDFG). 1987.
Delta outflow effects on the abundance and distribution of San Francisco Bay fish and invertebrates,
1980-1985. Exhibit 60, entered at the State Wat. Res.
Cont. Bd. 1987 Wat. Qual./Wat. Rights Proceeding
on the San Fran. Bay/Sac.-San Joaquin Delta. Ca.
Dept. Fish and Game, Stockton, CA, 345 pp.
Clark, F.N. 1929. The life history of the California jack
smelt, Atherinopsis californiensis. Ca. Fish and
Game, Fish Bull. 16, 22pp.
Plants
Amphibians &
Reptiles
Distribution
Interagency Ecological Study Program for the Sacramento-San Joaquin Estuary. Ca. Dept. Water Res.,
Ca. Dept. Fish and Game, U.S. Bureau Reclam., U.S.
Fish Wildl. Serv., various paginations.
Topsmelt
Fish
Atherinops affinis
Michael K. Saiki
General Information
On the West Coast, topsmelt (Family: Atherinidae) are
represented by five recognized subspecies of which only
one, the San Francisco topsmelt (Atherinops affinis
affinis), inhabits San Francisco Bay (Wang 1986).
Topsmelt are a small but tasty food fish taken from piers
by recreational anglers (Emmett et al. 1991). However,
commercial fishing for topsmelt is limited, with the species comprising only about 15-25% of the total “ smelt”
catch (Bane and Bane 1971, Frey 1971). Ecologically,
topsmelt are an important prey item for many piscivorous birds and fishes (Feder et al. 1974).
Reproduction
According to Emmett et al. (1991), the topsmelt is
gonochoristic (its gender is determined by developmental
rather than hereditary mechanisms) and iteroparous (it
has the capacity to survive and spawn beyond one or
multiple spawning seasons). Adults move into shallow
sloughs and mud flats in late spring and summer to
spawn (Wang 1986). In San Francisco Bay, spawning
occurs from April to October, with peaks in May and
June (Wang 1986). Although eggs are deposited singly,
the thick chorion bearing 2-8 filaments becomes entangled in aquatic vegetation, resulting in the formation
of large clusters of eggs (Wang 1986). Topsmelt seemingly spawn in batches, laying eggs more than once during a spawning season (Fronk 1969, Wang 1986). The
fecundity of topsmelt ranges from 200 eggs/fish for females measuring 110-120 mm in length to about 1,000
eggs/fish for females measuring 160 mm or more in
length (Fronk 1969). Hatching time varies from 35 days
at 13° C to less than 9 days at 27° C (Hubbs 1969).
Chapter 2 —
CDFG
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Vol. II:
species life history summaries. ELMR Rep. No. 8.
NOAA/NOS Strategic Environ. Assessments Div.,
Rockville, MD, pp. 190-193.
Eschmeyer, W.N., W.S. Herald and H. Hammann. 1983.
A field guide to Pacific coast fishes of North America.
Houghton Mifflin Co., Boston, MA, 336 pp.
Feder, H.M., C.H. Turner and C. Limbaugh. 1974.
Observations on fishes associated with kelp beds
in southern California. Ca. Fish and Game, Fish
Bull. 160, 144 pp.
Frey, H.W. 1971. California’s living marine resources
and their utilization. Ca. Dept. Fish and Game,
Sacramento, CA, 148 pp.
Ganssle, D. 1966. Fishes and decapods of San Pablo
and Suisun Bays. In: D.W. Kelley (comp). Ecological studies of the Sacramento-San Joaquin Estuary. Ca. Fish and Game, Fish Bull. 133:64-94.
Herbold, B.A., A.D. Jassby and P.B. Moyle. 1992. Status and trends report on aquatic resources in the
San Francisco Estuary. U.S. EPA Publ. Report, San
Francisco, CA. pp. 163 & 172.
Jones and Stokes Assoc., Inc. 1979. Protection and restoration of San Francisco bay fish and wildlife habitat. Vol. 1 & 2. U.S. Fish and Wildl. Serv. and Ca.
Fish and Game. Sacramento, CA.
Klingbeil, R.A., R.D. Sandell, and A.W. Wells. 1974.
An annotated checklist of the elasmobrachs and
teleosts of Anaheim Bay. In: E.D. Lane and C.W.
Hill (eds). The marine resources of Anaheim Bay.
Ca. Fish and Game, Fish Bull. 165:79-90.
Messersmith, J.D. 1966. Fishes collected in Carquinez
Strait in 61-62, In: D.W. Kelley, (comp). Ecological Studies of the Sacramento-San Joaquin Estuary, Part I. pp. 57-63. Ca. Dept. Fish and Game,
Fish Bull 133.
Middaugh, D.P. and J.M. Shenker. 1988. Salinity tolerance of young topsmelt, Atherinops affinis, cultured
in the laboratory. Ca. Fish and Game 74(4):232-235.
Middaugh, D.P., M.J. Hemmer, J.M. Shenker and T.
Takita. 1990. Laboratory culture of jacksmelt,
Atherinopsis californiensis, and topsmelt, Atherinops
affinis (Pisces: Atherinidae), with a description of
larvae. Ca. Fish and Game 76(1):4-43.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Dept. Fish and
Game, Fish Bull. 157, 235 pp.
Ruagh, A.A. 1976. Feeding habits of silversides (Family
Atherinidae) in Elkhorn Slough, Monterey Bay,
California. M.S. Thesis, Ca. State Univ. Fresno,
CA, 60 pp.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: A
guide to the early life histories. Tech. Rept. No. 9.
Estuarine Fish and Associated Invertebrates
115
Fish
Growth and Development
Topsmelt eggs are spherical in shape and approximately
1.5-1.7 mm in diameter (Wang 1986). Between nine
and 35 days after fertilization, eggs hatch into planktonic
larvae that measure 4.3-4.9 mm total length (TL) and
0.0011 grams wet weight (Emmett et al. 1991) or 5.15.4 mm standard length (SL) (Middaugh et al. 1990).
Larvae measure 9.5-10.0 mm after the yolk-sac is absorbed, and begin to develop juvenile characteristics
when approximately 18.5 mm long (Wang 1986). Juveniles may vary in length from 18.5 to 120.0 mm
(Schultz 1933, Fronk 1969). Topsmelt mature in their
second or third year, depending on subspecies, and may
live six to nine years (Schultz 1933, Feder et al. 1974).
Adults can attain as much as 120 mm in length for the
southernmost subspecies (A. affinis littoralis) and as
much as 370 mm in length for the northernmost subspecies (A. affinis oregonia) (Schultz 1933, Fronk 1969,
Eschmeyer et al. 1983). In general, northern varieties
grow larger than southern subspecies (Schultz 1933).
The topsmelt is characterized by an omnivorous diet
(Quast 1968, Horn and Allen 1985). Topsmelt from bay
and estuarine habitats consume mostly plant material
(diatoms, filamentous algae, and detritus), whereas those
from ocean habitats feed mainly on planktonic crustaceans (gammarid and caprellid amphipods, mysids, ostracods, copepods, and crustacean larvae) (Moyle 1976,
Quast 1968, Fronk 1969). Juveniles and adults forage
mostly during daylight near the surface in deep water or
on the bottom in shallow water (Hobson et al. 1981).
Distribution
Topsmelt can occur from the Gulf of California northward to Vancouver Island, but are usually rare north of
Tillamook Bay, Oregon (Miller and Lea 1972, Hart
1973, Eschmeyer et al. 1983). The five subspecies are
A. affinis oregonia (occurs from Oregon to Humboldt
Bay, California), A. affinis affinis (occurs in San Francisco Bay and surrounding waters to Monterey, California), A. affinis littoralis (occurs from Monterey to San
Diego Bay, California), A. affinis cedroscensis (the kelp
topsmelt), and A. affinis insularium (the “ island topsmelt,”
occurs around the Santa Barbara Islands, California)
(Schultz 1933, Feder et al. 1974).
In San Francisco Bay, spawning has been observed
in the South Bay near the Aquatic Park in Berkeley and
at the Dumbarton Bridge (Wang 1986). Small schools
of larvae often occur near the surface of both shallow
water and open water, and are particularly abundant in
tidal basins (e.g., Aquatic Park in Berkeley; Lake Merritt
in Oakland) and the sluggish waters of the South Bay
116
Baylands Ecosystem Species and Community Profiles
Population Status and Influencing Factors
Field studies indicate that topsmelt are among the most
abundant fish species occurring in shallow-water sloughs
of South San Francisco Bay (Jones and Stokes Assoc.
1979, Woods 1981, Herbold et al. 1992). Herald and
Simpson (1955) reported that topsmelt were commonly caught in a fixed fish-collecting device located at the
Pacific Gas and Electric Company power plant in South
San Francisco Bay. Furthermore, Wild (1969) reported
that topsmelt was the most abundant species of fish
sampled at the mouth of Plummer Creek (located in
South San Francisco Bay). Midwater trawls fished at several locations in South San Francisco Bay during 19801988 also yielded numerous topsmelt (Herbold et al. 1992).
South Bay topsmelt increased in abundance during two of
the recent drought years, but otherwise did not show consistent year-to-year patterns (Herbold et al. 1992).
Several factors may directly influence the abundance of topsmelt: salinity, water temperature, freshwater inflows, entrainment on intake screens at power
plants and water diversions, and availability of spawning substrate. In Newport Bay, California, topsmelt
abundance was significantly correlated with water
temperature and salinity (Allen 1982). By comparison, no
relationship was found between abundance indices and river
flow in San Francisco Bay (CDFG 1987). Although this
species is commonly impinged on intake screens of power
plants and water diversions, this source of mortality may
not be significant for bay populations (San Diego Gas
and Electric 1980). In the Tijuana Estuary of southern
California, abundance of topsmelt eggs and larvae was
positively correlated with algal mats (Nordby 1982). In
other words, topsmelt eggs and larvae were seemingly more
abundant in areas with dense algal growth. Because this
species uses algal mats and shallow-water eelgrass beds for
spawning, destruction or removal of these types of vegetation may adversely affect topsmelt abundance.
Trophic Levels
Topsmelt are omnivorous (primary and higher order
consumers).
Proximal Species
Predators: Many piscivorous birds and fishes.
Plants
Amphibians &
Reptiles
Food and Feeding
(e.g., Robert Crown Memorial Park; Hunters Point; San
Mateo Bridge; Dumbarton Bridge) (Wang 1986). Juvenile topsmelt generally move into open waters of the bay
or into coastal kelp beds. Some juveniles may occur in
Suisun Bay during summer and early fall as the salt
wedge moves to the upper reaches of the Estuary (Wang
1986). In general, topsmelt seem to be much less common outside of the South Bay.
Prey: Diatoms (major); diatoms, chironomid midge larvae, and amphipods (minor).
Habitat: Eel grass and micro algae (spawning substrate);
kelp beds (adult and juvenile cover).
Cohabitors: Schools with shiner perch and jacksmelt.
Good Habitat
References
Allen, L.G. 1982. Seasonal abundance, composition, and
productivity of the littoral fish assemblage in upper Newport Bay, California. Fish Bull., U.S.
80(4):769-790.
Anderson, B.S., D.P. Middaugh, J.W. Hunt and S.L.
Turpen. 1991. Copper toxicity to sperm, embryos
and larvae of topsmelt, Atherinops affinis, with notes
on induced spawning. Mar. Environ. Res. 31:17-35.
Bane, G.W. and A.W. Bane. 1971. Bay fishes of northern California with emphasis on the Bodega Tomales Bay area. Mariscos Publ., Hampton Bays, NY,
143 pp.
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-1985. Exhibit 60, entered for the State
Wat. Res. Cont. Bd. 1987 Wat. Quality/Wat.
Rights Proceeding on the San Fran. Bay/Sac.-San
Chapter 2 —
Estuarine Fish and Associated Invertebrates
117
Fish
In general, topsmelt can tolerate a relatively broad range
of environmental conditions during the time that they
inhabit San Francisco Bay. However, for successful
spawning to occur, they require submerged vegetation
for egg attachment, water temperatures of 10-25° C, and
salinities of less than 72 ppt (Schultz 1933, Carpelan
1955, Fronk 1969). By comparison, larvae must be able
to school near the surface in shallow open-water areas,
particularly tidal basins (Wang 1986). Young-of-the-year
topsmelt are common in middle to low salinity portions
of the Estuary (Wang 1986). Although juveniles can tolerate salinities varying from 2 ppt to 80 ppt, growth and
survival are reduced at salinities above 30 ppt (Middaugh
and Shenker 1988). In addition, juveniles and adults are
seemingly eurythermal, but temperatures of 26-27° C or
higher may cause stress (Carpelan 1955, Ehrlich et al.
1979). Within San Francisco Bay, topsmelt utilize mudflats for breeding, spawning, and as nursery areas for
young. Subtidal areas with sandy bottoms are relied on
heavily as nursery and foraging areas. Intertidal streambeds are major foraging areas (Jones and Stokes Assoc.
1979). Recent studies indicate that embryonic and larval stages of topsmelt are sensitive to the effects of pollution (Singer et al. 1990, Anderson et al. 1991,
Goodman et al. 1991, Hemmer et al. 1991). Thus, habitats used by topsmelt for spawning and rearing must not
be exposed to appreciable amounts of pollution.
Joaquin Delta. Ca. Dept. Fish and Game, Stockton, CA, 345 p.
Carpelan, L.H. 1955. Tolerance of the San Francisco
topsmelt, Atherinopsis affinis affinis, to conditions
in salt-producing ponds bordering San Francisco
Bay. Ca. Dept. Fish and Game 41(4):279-284.
Ehrlich, K.F., J.M. Hood, G. Muszynski and G.E.
McGowen. 1979. Thermal behavioral responses of
selected California littoral fishes. Fish bull., U.S.
76(4):837-849.
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in West Coast estuaries, Volume
II: Species life history summaries. ELMR Rep. No.
8. NOAA/NOS Strategic Envir. Assess. Div.,
Rockville, MD, 329 pp.
Eschmeyer, W.N., W.S. Herald and H. Hammann. 1983.
A field guide to Pacific Coast fishes of North America.
Houghton Mifflin Co., Boston, MA, 336 pp.
Feder, H.M., C.H. Turner, and C. Limbaugh. 1974.
Observations of the fishes associated with kelp beds
in Southern California. Ca. Dept. Fish and Game,
Fish Bull. 160, 138 pp.
Frey, H.W. 1971. California’s living marine resources
and their utilization. Ca. Dept. Fish and Game,
Sacramento, CA, 148 pp.
Fronk, R.H. 1969. Biology of Atherinops affinis littoralis
Hubbs in Newport Bay. M.S. Thesis, Univ. Ca.,
Irvine, CA, 106 pp.
Goodman, L.R., M.J. Hemmer, D.P. Middaugh, and
J.C. Morre. 1991. Effects of fenvalerate on the early
life stages of topsmelt (Atherinops affinis). Environ.
Toxicol. Chem. 11:409-414.
Hart J.L. 1973. Pacific fishes of Canada. Fish. Res. Board
Can., Bull. No. 180, 740 pp.
Hemmer, M.J., D.P. Middaugh, and V. Comparetta.
1991. Comparative acute sensitivity of larval
topsmelt, Atherinops affinis, and inland silversides,
Menidia beryllina, to 11 chemicals. Environ.
Toxicol. Chem. 11:401-408.
Herald, E.S. and D.A. Simpson. 1955. Fluctuations in
abundance of certain fishes in South San Francisco Bay as indicated by sampling at a trash screen.
Ca. Dept. Fish and Game 41(4):271-278.
Herbold, B.A., A.D. Jassby and P.B. Moyle. 1992. Status and trends report on aquatic resources in the
San Francisco Estuary, U.S. EPA publ. Rept., San
Francisco, CA.
Hobson, E., W.N. McFarland and J.R. Chess. 1981.
Crepuscular and nocturnal activities of Californian
near shore fishes, with consideration of their scotopic visual pigments and the photic environment.
Fish Bull., U.S. 79(1):1-30.
Horn, M.H. and L.G. Allen. 1985. Fish community
ecology in southern California bays and estuaries.
Chapter 8. In: A. Yanez-Arancibia (ed). Fish com-
118
Baylands Ecosystem Species and Community Profiles
Threespine Stickleback
Gasterosteus aculeatus
Robert A. Leidy
General Information
The threespine stickleback (Family: Gasterosteidae) is a
small laterally-compressed fish with three spines on the
dorsum and from 1 to 35 bony plates on the sides (Moyle
1976). Largely as a matter of taxonomic convenience,
Miller and Hubbs (1969) suggested that there are two
forms: G. a. aculeatus for the fully-plated, anadromous
form; and G. a. microcephalus for the partially-plated
freshwater/resident form. The threespine stickleback is
a polymorphic species and as such, populations within
the San Francisco Estuary and its tributary streams support resident/freshwater and anadromous/saltwater
forms, as well as mixtures of the two forms that presumably interbreed (Moyle, pers. comm.). The threespine
stickleback has no commercial value, but has important
scientific value, especially to evolutionary biologists.
Reproduction, Growth and Development
The following discussion is taken largely from Moyle
(1976) unless otherwise referenced. Threespine sticklebacks typically complete their life cycle within one year
although some individuals may live two to three years.
Individuals from freshwater populations typically do not
exceed 60 mm total length (TL), while anadromous
forms may exceed 80 mm TL. Adult females are usually
larger than adult males.
Anadromous forms migrate into freshwater breeding areas as water temperatures increase during April
through July, although some stickleback populations
may remain in estuarine environments to spawn if suitable habitat is present (Moyle 1976, Wang 1986).
CDFG
Fish
Amphibians &
Reptiles
Woods, E. 1981. Fish utilization. In: T. Niesen and M.
Josselyn (eds). The Hayward Regional Shoreline
marsh restoration: Biological succession during the
first year following dike removal. pp 35-46. Rep.
1. Tiburon center for environmental studies,
Tiburon, Ca., 178 pp.
Plants
munity ecology in estuaries and coastal lagoons:
towards an ecosystem integration, p. 169-190 DR
(R) UN AM Press, Mexico.
Hubbs, C. 1969. Developmental temperature tolerance
and rates of four Southern California fishes, Fundulus parvipinnis, Atherinops affinis, Leuresthes
tenuis, and Hypsoblennius sp. Ca. Dept. Fish and
Game 51(2):113-122.
Jones and Stokes Assoc., Inc. 1979. Protection and restoration of San Francisco bay fish and wildlife habitat. Vol. 1 & 2. U.S. Fish and Wildl. Serv. and Ca.
Dept. Fish and Game. Sacramento, CA.
Middaugh, D.P. 1990. Laboratory culture of jacksmelt,
Atherinopsis californiensis, and topsmelt, Atherinops
affinis (Pisces: Atherinidae), with a description of
larvae. Ca. Dept. Fish and Game 76(1):4-43.
Middaugh, D.P., M.J. Hemmer, J.M. Shenker and T.
Takita. 1990. Laboratory culture of jacksmelt,
Atherinopsis californiensis, and topsmelt, Atherinops
affinis (Pisces: Atherinidae), with a description of
larvae. Ca. Dept. Fish and Game 76(1):4-43.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Dept. Fish and
Game, Fish Bull. 157, 235 pp.
Moyle, P.B. 1976. Inland Fishes of California. Univ. Ca.
Press, Berkeley, CA. 405 pp.
Nordby, C.S. 1982. The comparative ecology of
ichthyoplankton within Tijuana Estuary and its
adjacent nearshore waters. M.S. Thesis, San Diego State Univ., San Diego, CA. 101 pp.
Quast, J.C. 1968. Observations on the food of the kelpbed fishes. Ca. Dept. Fish and Game, Fish Bull.
139:109-142.
San Diego Gas and Electric. 1980. Silvergate power plant
cooling water intake system demonstration (in accordance with section 316(b) Federal Water Pollution
Control Act Amendment of 1972). San Diego Gas and
Electric, San Diego, CA, various pagination.
Schultz, L.P. 1933. The age and growth of Atherinops
affinis oregonia. Jordan and Snyder and other subspecies of bay smelt along the Pacific coast of the
United States. Wash. State Univ. Publ. Biol.
2(3):45-102.
Singer, M.M., D.L. Smalheer, R.S. Tjeerdema, and M.
Martin. 1990. Toxicity of an oil dispersant to the
early life stages of four California marine species.
Environ. Toxicol. Chem. 9:1387-1395.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: A
guide to the early life histories. Tech. Rep. No. 9.
Interagency ecological study program for the Sacramento-San Joaquin Estuary. Ca. Dept. of Water
Res., Ca. Dept. Fish and Game, U.S. Bureau
Reclam., U.S. Fish Wildl. Serv.
Wild, P. 1969. Marine species present in Plummer Creek, San
Francisco Bay. M.S. Thesis, San Jose State Univ., Ca.
Food and Feeding
Threespine sticklebacks are visual feeders primarily on
small benthic organisms or organisms living on submerged, rooted, or floating macrophytes such as insect
larvae, chironomid midge larva, and ostracods (Hynes
1950, Beukema 1963, Hagen 1967). Anadromous forms
feed mostly on free-swimming crustaceans (Barraclough
and Fulton 1967, 1978; Barraclough et al. 1968). In a
study of threespine stickleback diet in San Pablo Creek,
a tributary to San Pablo Bay, Snyder (1984) found the
diet consisted of approximately 42% insects (mainly chironomid larvae), 28% crustacea (mainly ostracods), and
10% earthworms (Lumbricidae). Fish eggs and plant
material accounted for approximately 9% of the diet
(Snyder 1984).
Distribution
Threespine stickleback are native to the coastal waters
of Mediterranean Europe, north to Russia, and east to
Japan and Korea (Moyle 1976). In North America, threespine stickleback populations occur on the East coast
south to Chesapeake Bay, and on the West coast south
from Alaska to Baja California. In California, populations
are found below barriers such as dams and falls in coastal
streams, including the San Francisco Estuary and its tributary streams, and in the Central Valley (Moyle 1976).
Within the San Francisco Estuary, threespine stickleback are widely distributed and often locally abundant
in fresh-, brackish-, and saltwater intertidal upper marsh
and riverine tidal marsh habitats (Leidy 1984; Leidy,
unpub. data; Cathy Hieb, unpub. data). Leidy (1984)
recorded threespine stickleback in 43% of 457 samples
of Estuary streams between elevation 0 to 123 m.
Threespine stickleback are also abundant in large
areas of formerly tidal salt and brackish marsh that have
been converted to salt ponds in the South Bay and San
Pablo Bay (Lonzarich 1989, Herbold et al. 1992).
Carpelan (1957) recorded threespine stickleback as one
of the most numerous fish in the Alviso salt ponds in
the South Bay. Apparently, threespine stickleback persist in these ponds, particularly near the mouth of the
Napa River, until salinities become too high (i.e., salinities between 40 to 50 ppt) (Herbold et al. 1992). There
are approximately 9,059 acres of salt ponds in the NapaSolono area of the North Bay and 27,497 acres in the
South Bay that may be considered available for use by
threespine stickleback on a seasonal basis (Meiorin et al.
1991).
Population Status and Influencing Factors
The current status of threespine stickleback within the
San Francisco Estuary may be regarded as secure. Threespine stickleback populations currently are widespread
and locally abundant in suitable habitats within the San
Francisco Estuary. Because sticklebacks can readily disperse through estuarine and marine environments they
are able to regularly recolonize habitats from which they
may been extirpated. Important factors negatively influencing population numbers likely include excess siltation
and turbidity, increased water temperatures by the removal of riparian vegetation through stream channelization, pollution, loss of nesting, feeding, and cover habitat by the removal of aquatic macrophytes, the construction of barriers such as dams or drop structures, and the
introduction of exotic piscivorous fish.
Trophic Levels
Larvae are primary consumers. Juveniles and adults are
primary and higher order consumers.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
119
Fish
Anadromous forms typically spawn earlier than freshwater populations. Spawning typically occurs at 15° to 18°C
(Vrat 1949, Wang 1986). Males begin to display bright
green and red breeding coloration as they move away from
schools to set up breeding territories and construct nests.
Nests are excavated in the substrate as shallow pits.
The pits are then covered with algae or other plant fragments and formed into a tunnel that is held together by
a sticky renal secretion (Greenbank and Nelson 1959).
Females are then courted by males into the nest where
the female may lay between 50 and 300 eggs in several
spawnings. Eggs are spherical and average 1.5-1.7 mm
in diameter (Wang 1986). A pair can spawn up to six
times within a 10-15 day period (Wang 1986). Following egg laying, the male drives away the female, fertilizes the eggs, and then begins to incubate the eggs while
defending the nest from other sticklebacks and predators. The male is known to circulate water over the eggs
by fanning his pectoral fins and to clean the eggs with
his mouth. Immediately prior to hatching the male tears
apart the nest and breaks apart the egg clusters which is
thought to increase the survival of hatching young
(Wang 1986). Length at hatching is between 4.2 and 5.5
mm TL (Vrat 1949, Kuntz and Radcliffe 1917).
Stickleback eggs hatch in six to eight days at temperatures of between 18° to 20° C (Breder and Rosen
1966). The fry remain in the nest for several days where
they continue to be guarded by the male. Fry eventually form schools of similar-size sticklebacks or other species, usually in shallow water habitats containing dense
vegetation (Wang 1986).
Juveniles are most abundant in late summer, followed by drastic declines in abundance in the fall and
winter (Wang 1986). It is unknown whether populations
of juveniles within the San Francisco Estuary make extensive migrations into open water/subtidal habitats
within the Estuary. Moyle (1976) states that freshwater
and anadromous populations range from complete ecological separation to complete interbreeding.
Fish
Proximal Species
Major Predators: Kingfisher, egrets, herons, and other
wading birds.
Other Predators: Adult salmonids and other large freshwater, estuarine, and marine piscivorous fish terrestrial
and aquatic snakes.
Major Prey: Aquatic insects and crustacea, earthworms,
fish eggs and vegetation.
Habitat/cover: Riparian, submerged, floating, and
emergent wetland and aquatic vegetation.
Good Habitat
References
Barraclough, W.E. and J.D. Fulton. 1967. Data record.
Number, size composition and food of larval and
juvenile fish caught with a two-boat surface trawl
in the Strait of Georgia. July 4-8, 1966. Fish. Res.
Board Can. Rep. Ser. 940. 82 pp.
______. 1978. Data record. Food of larval and juvenile
fish caught with a surface trawl in Saanich Inlet
during June and July 1966. Fish. Res. Board Can.
Rep. Ser. 1003. 78 pp.
Barraclough, W.E., D.G. Robinson and J.D. Fulton.
1968. Data record. Number, size composition,
weight and food of larval and juvenile fish caught
with a two-boat surface trawl in Saanich Inlet, April
23 -July 21, 1968. Fish. Res. Board Can. Rep. Ser.
1004. 305 pp.
Beukema, J. 1963. Experiments on the effects of the
hunger state on the risk of prey of the three-spined
stickleback. Arch. Nees. Zool. 15: 358-361.
Breder, C.M. and D.E. Rosen. 1966. Modes of reproduction in fishes. Am. Mus. Nat. Hist., New York.
941 pp.
Carpelan, L.H. 1957. Hydrobiology of the Alviso salt
ponds. Ecology 38(3): 375-390.
Greenbank, J. and P.R. Nelson. 1959. Life history of
the threespine stickleback, Gasterosteus aculeatus
Linnaeus in Karluk Lake and Bare Lake, Kodiak
120
Baylands Ecosystem Species and Community Profiles
Personal Communications
Peter Moyle, University of California, Davis
Plants
Amphibians &
Reptiles
Freshwater populations of threespine stickleback prefer
clear, cool backwater and pool habitats containing submerged, floating, or emergent vegetation, with sand or
small-sized gravel substrates (Moyle 1976, Leidy 1984).
This species is typically uncommon in silted pools with
moderate to high turbidities (Leidy 1984). Marine and
estuarine populations are pelagic, although they tend to
remain to close to the shore (Moyle 1976). Threespine
stickleback is uncommon where water temperatures
regularly exceed 24° C (Moyle 1976).
Island, Alaska. U.S. Fish Wild. Serv. Fish Bull.
59(153): 537-559.
Hagen, D.W. 1967. Isolating mechanisms in three-spine
sticklebacks (Gasterosteus). J. Fish. Res. Bd. Canada
24(8): 1637-1692.
Herbold, B., P. Moyle and A. Jassby 1992. Status and
trends report on aquatic resources in the San Francisco Estuary. San Fran. Est. Proj. Public Rept.
March 1992. 257 pp. plus appendices.
Hynes, H.B.N. 1950. The food of freshwater sticklebacks (Gasterosteus aculeatus and Pygosteus
pungitius), with a review of methods used in studies of the food of fishes. J. An. Ecol. 19(1): 36-58.
Kuntz, A. and R. Radcliffe. 1917. Notes on the embryology and larval development of twelve teleostean
fishes. Bull. U.S. Bur Fish. 35: 87-134.
Miller, R.R. and C. L. Hubbs. 1969. Systematics of
Gasterosteus aculeatus with particular reference to
intergradation and introgression along the Pacific
Coast of North America: a commentary on a recent contribution. Copeia 1: 52-69.
Leidy, R.A. 1984. Distribution and ecology of stream
fishes in the San Francisco Bay drainage. Hilgardia
52(8): 1-175
Lonzarich, D. 1989. Life history and patterns of distribution in salt pond fishes: a community level study.
M.S. Thesis, San Jose State University, Ca..
Meiorin, E.C., M.N. Josselyn, R. Crawford, J. Calloway,
K. Miller, R. Pratt, T. Richardson, and R. Leidy.
1991. Status and trends report on wetlands and
related habitats in the San Francisco Estuary. U.S.
Environmental Protection Agency, San Francisco
Estuary Project. 209 pp. + appendices.
Moyle, P.B. 1976. Inland Fishes of California. Univ. of
California Press. 405 pp.
Snyder, R.J. 1984. Seasonal variation in the diet of the
threespine stickleback, Gasterosteus aculeatus, in
Contra Costa County, California. Ca. Fish and
Game 70(3): 167-172.
Vrat, V. 1949. Reproductive behavior and development
of eggs of the threespine stickleback (Gasterosteus
aculeatus) of California. Copeia 4: 252-260.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: a
guide to the early life histories. Tech. Rept. 9. Prepared for the Interagency Ecological Study program for the Sacramento-San Joaquin Estuary. Interagency Ecological Workshop, Asilomar, CA.
1990.
from December-January and May-July in Central California (Wyllie-Echeverria 1987).
Although gravid brown rockfish have been collected in San Francisco Bay, most parturition is believed
to occur in coastal waters (Kendall and Lenarz 1986,
Wang 1986). In San Francisco Bay, mature females were
observed in winter and spring and larvae have been collected in winter and spring (Wang 1986).
Brown Rockfish
Sebastes auriculatus
Kurt F. Kline
General Information
Reproduction
All rockfishes, including the brown rockfish, are viviparous. Fertilization is internal and the larvae develop in
the egg capsule within the ovarian cavity. The larvae
hatch within the ovary and are released with little yolk
remaining and ready to feed. The embryos develop in
40-50 days after fertilization and the larvae hatch about
1 week before extrusion (Kendall and Lenarz 1986).
Brown rockfish larvae are 4.7-6.7 mm at hatching
(Delacy et al. 1964) and pelagic for several months. Although brown rockfish fecundity is not known, Sebastes
females typically produce 100,000 to 1,000,000 eggs per
brood (Kendall and Lenarz 1986). Brown rockfish may
have multiple broods within one year, with parturition
Growth and Development
Brown rockfish juveniles are pelagic until 20-30 mm,
whereas older juveniles settle out of the water column
and are strongly association with some type of physical
structure (Turner et al. 1969, Kendall and Lenarz 1986).
Pelagic juveniles have been collected in nearshore coastal
waters from April through June, while benthic juveniles
are common in nearshore coastal waters and the Bay
(Kendall and Lenarz 1986). In San Francisco Bay, age0 juveniles were usually first collected from April to July
and were common through summer and fall (Wang
1986, Baxter 1999).
Juvenile brown rockfish apparently spend several
years in a very restricted home range in the Bay and
gradually move to deeper waters and nearshore. Juvenile
brown rockfish tagged in the Bay have been recaptured
more than 80 km away in nearshore coastal waters
(Kendall and Lenarz 1986).
Both male and female brown rockfish reach maturity as early as age 3 (260 mm TL); half reach maturity at age 5 (310 mm TL); and all are mature at age 10
(380 mm TL) (Wyllie-Echeverria 1987). Both sexes grow
at similar rates and reach a maximum size of about 550
mm TL (Miller and Lea 1972). In southern California,
the oldest male was 18 years, the oldest female 20 years
(Love and Johnson 1998).
Food and Feeding
In San Francisco Bay, smaller juvenile brown rockfish
(<130 mm TL) prey primarily upon small crustaceans,
including amphipods, copepods, caridean shrimp, and
Cancer crabs. Larger fish (130-310 mm TL) prey upon
larger crustaceans (caridean shrimp, Cancer crabs,
Upogebia) and fish (Ryan 1986).
CDFG
Distribution
The brown rockfish ranges from Hipolito Bay, Baja California, to southeast Alaska (Miller and Lea 1972). It most
often solitary, but may be found in small aggregations
(Love and Johnson 1998). In the ocean, it is most common in shallow rocky reefs (5-20 m), but also found over
sand flats near eelgrass and in kelp beds while in bays
and estuaries it is found near piers and over rubble (Feder
et al. 1974, Matthews 1990, Love and Johnson 1998).
Chapter 2 —
Estuarine Fish and Associated Invertebrates
121
Fish
The brown rockfish (Sebastes auriculatus) is a member
of the family Scorpaenidae, one of the largest fish families in the western Pacific. The family is dominated by
the rockfishes (Sebastes spp.), a genus which is represented by over 50 species on the northwest Pacific coast.
The brown rockfish is the most common rockfish
in San Francisco Bay (Alpin 1967, Wang 1986), and the
Bay appears to be an important nursery area for juveniles
(Kendall and Lenarz 1986, Baxter 1999). Brown rockfish are the most common rockfish caught by sport anglers in the Bay (W. Van Buskirk, pers. comm) and the
third most frequently caught rockfish in the San Francisco region (Karpov et al. 1995). Most brown rockfish
are caught by anglers fishing from partyboats, skiffs,
piers, and the shoreline (Miller and Gotshall 1965,
Karpov et al. 1995). It is also a minor, but important,
component of the nearshore commercial fishery; in the
San Francisco area, the majority of brown rockfish are
caught by hook and line for the live or whole fresh fish
markets. Since the early 1990s, the brown rockfish has
been the most common species sold in the live in San
Francisco markets (C. Ryan, pers. comm.).
Fish
In San Francisco Bay the brown rockfish is found primarily in Central San Francisco Bay, to a lesser degree in
South San Francisco and San Pablo bays, and occasionally in Carquinez Strait and western Suisun Bay (Ganssle
1966, Messersmith 1966, Wang 1986, Baxter 1999).
Suitable habitat and salinity are the primary factors influencing distribution of brown rockfish in the
Bay. Benthic juveniles and adults are strongly associated
with structure, including rocky reefs, piers and jetties,
breakwaters, and riprap. In the Bay, most brown rockfish were collected at salinities > 20l (median 28.3l , 90th
percentile 31.8l, 10 th percentile 21.5l, Baxter 1999,
CDFG, unpubl. data).
There is a modest brown rockfish population in the San
Francisco Bay region. San Francisco Bay is a nursery area
for brown rockfish, and most juveniles immigrate to the
Bay from the nearshore coastal area soon after settlement.
It is not clear if resident adult brown rockfish spawn successfully in the Bay. Juveniles rear in the Bay for several
years, and the population is comprised of several year
classes. But there is no reliable index or measure of year
class strength in the Bay, as brown rockfish are strongly
associated with structure, and are undoubtedly
undersampled by trawls or other towed nets typically
used by research studies.
Trophic Levels
Secondary carnivore. Feeds primarily on crustaceans and
fishes.
Proximal Species
Prey: Crustaceans (caridean shrimp, Cancer crabs,
Upogebia, amphipods, copepods), polychaetes, fishes,
herring eggs.
Predators: Larger predatory fishes, including striped bass.
Good Habitat
Structure, including piers and rocky shores, in the higher
salinity regions of the Bay
Acknowledgments
Some of the material in this report was summarized from
the brown rockfish chapter in IEP Technical Report 63,
which is referenced below (Baxter 1999).
References
Aplin, J.A. 1967. Biological survey of San Francisco
Bay, 1963-1966. Ca. Dept. Fish and Game,
122
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Population Status and Influencing Factors
Marine Resources Operations. MRO Ref. 67-4,
131 pp.
Baxter, R. 1999. Brown rockfish. In: J. Orsi, ed. Report
on the 1980-1995 fish, shrimp, and crab sampling
in the San Francisco Estuary, California. Interagency Ecological Program for the Sacramento-San
Joaquin Estuary Tech. Rept. No. 63.
Delacy, A.C., C.R. Hitz, and R.L. Dryfoos. 1964. Maturation, gestation, and birth of rockfish (Sebastodes)
from Washington and adjacent waters. Washington Dept. of Fisheries, Fishery Research Paper
2(2):51-67.
Feder, H.M., C.H. Turner and C. Limbaugh. 1974.
Observations of the fishes associated with kelp beds
in southern California. Ca. Dept. of Fish and Game
Fish Bull. 160, 144 pp.
Ganssle, D. 1966. Fishes and decapods of San Pablo
and Suisun Bays. In: D.W. Kelly (ed). Ecological
studies of the Sacramento San Joaquin Estuary, Part
1, Ca. Dept. of Fish and Game Fish Bull. 133: 64-94.
Karpov, K., P. Albin and W. Buskirk. 1995. The marine
recreational fishery in northern and cetral California. Ca. Dept. Fish and Game Fish Bull. 176.
Kendall Jr., A.W. and W.H. Lenarz. 1986. Status of early
life history studies of northeast Pacific rockfishes.
Proceedings of the International Rockfish Symposium. October 1986. Anchorage, Alaska.
Love, M.S. and K. Johnson. 1998. Aspects of the life
histories of grass rockfish, Sebastes rastrelliger, and
brown rockfish, S. auriculatus, from southern California. US Fish. Bull. 87:100-109.
Matthews, K.R. 1990. An experimental study of the
habitat preferences and movement patterns of copper, quillback, and brown rockfish (Sebastes spp.).
Env. Biol. Fishes 29:161-178.
Messersmith, J. 1966. Fishes collected in the Carquinez
Strait in 1961-1962. In: D.W. Kelly (ed). Ecological Studies of the Sacramento-San Joaquin Estuary, Part 1. Ca. Dept. of Fish and Game Fish Bull.
133:57-63.
Miller, D.J. and D. Gotshall. 1965. Ocean sportfish
catch and effort from Oregon to Pt. Arguello, July
1957 to June 1961. Ca. Dept of Fish and Game
Fish Bull. 130, 135 pp
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Dept. of Fish and
Game Fish Bull. 157, 249 pp.
Ryan, C.J. 1986. Feeding habits of brown rockfish,
Sebastes auriculatus, associated with a dock in San
Francisco Bay, California. M. A. Thesis, San Francisco State University, 88 pp.
Turner, C.H., E.E. Ebert and R.R. Given. 1969. Manmade reef ecology. Ca. Dept. Fish and Game, Fish.
Bull. 146: 221p.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and adjacent waters, California:
A guide to the early life histories. Interagency Ecological Study Program for the Sac.-San Joaquin
Estuary, Tech. Rept. No. 9.
Wyllie-Echeverria, T. 1987. Thirty-four species of California rockfishes: maturity and seasonality of reproduction. US Fishery Bulletin 85(2):229-250.
Personal Communications
Pacific Staghorn Sculpin
Leptocottus armatus armatus
Robert N. Tasto
General Information
The Pacific staghorn sculpin (Family: Cottidae) is found
from Kodiak Island, Alaska to San Quintin Bay, Baja,
California (Miller and Lea 1972). It is the only true euryhaline species among the California cottids (CDFG
1987), and appears to move freely between fresh and saltwater environments (Moyle 1976). It is regarded as a
nuisance species by many sportfishermen, but has shown
some limited value as bait for gamefish (particularly
striped bass) in the Estuary. Bolin (1944) recorded its
maximum depth of capture offshore coastal California
at 300 feet. It is a target species of the National Status
and Trends Program (Emmett et al. 1991), as it is considered an indicator of stress in the estuarine environment, and may spend its entire life in Pacific coast estuaries.
Reproduction
CDFG
Pacific staghorn sculpin may reach sexual maturity in
their first year, and sex ratios within a population appear to favor females slightly (Boothe 1967, Tasto 1975).
In northern California, spawning begins in October
(Tomales Bay) or November (San Francisco Bay), peaks
in January-February, and ends in March (Jones 1962,
Growth and Development
At hatching, Pacific staghorn sculpin larvae range from
3.9 to 4.8 mm total length (TL) (Jones 1962). Metamorphosis to the juvenile begins after about 2 months, when
the larvae are 15 to 20 mm standard length (SL) (Emmett et al. 1991). The juvenile size range is approximately 20 to 120 mm TL (Jones 1962), and there appears to be considerable overlap in the length distribution of 0+ and 1+ fish, particularly in the summer and
fall (CDFG 1987). The staghorn sculpin reaches maturity at about 120 mm TL its first year, and can grow to
over 200 mm TL (3 years old) in California (Jones 1962).
In southern California, growth was determined to be
curvilinear (Tasto 1975). The largest specimen recorded
was about 30 cm (Barnhart 1936).
Food and Feeding
Pacific staghorn sculpin larvae are planktivorous (Emmett et al. 1991). The juvenile and adult forms are,
however, demersal predators, particularly over intertidal
and shallow subtidal mudflats, and have been shown to
feed on a variety of non-burrowing benthic organisms
(Jones 1962, Boothe 1967, Tasto 1975). Feeding behavior of the staghorn sculpin is thought to be continuous,
although there appears to be a preference for feeding at
night (Tasto 1975). The principal food items for staghorn sculpin within San Francisco Bay were found to
be bay shrimp (Crangon spp.), bay goby (Lepidogobius
lepidus), mud crab (Hemigrapsus oregonensis), callianassid
shrimp (i.e., Upogebia), and a variety of amphipods, isopods, and polychaetes (Boothe 1967). Elkhorn Slough
studies showed predation on epifaunal crustaceans and
infaunal and epifaunal worms (Barry et al. 1996). In
Anaheim Bay, major food items were similar to Elkhorn
Slough and San Francisco Bay, including callianasiid
shrimp (i.e., Callinassa sp.), mud crab, and arrow goby
(Clevelandia ios) (Tasto 1975). Jones (1962) found that
in Tomales Bay, staghorn sculpin fed heavily upon
Upogebia and Crangon shrimp. In Grays Harbor, Washington, the staghorn sculpin’s diet consisted of amphipods, crangonid shrimp, small fish, Upogebia sp., juvenile Dungeness crab, and polychaetes (Armstrong et al.
1995). Several studies indicate that the staghorn sculpin
Chapter 2 —
Estuarine Fish and Associated Invertebrates
123
Fish
Wade Van Buskirk. Recreational Fisheries Information
Network Database, Pacific States Marine Recreational Fisheries Monitoring
Connie Ryan. California Department of Fish and Game,
Ocean Fisheries Research Unit - Menlo Park
Boothe 1967). In southern California (Anaheim Bay),
spawning does not begin until December, but also peaks
in January-February and ends around mid-March (Tasto
1975). Fertilization is external. Staghorn sculpin eggs are
adhesive and laid in shallow subtidal and intertidal waters. Fecundity averages 5,000 eggs per female (Jones
1962), and ranges from 2,000 to 11,000 eggs per female
(Moyle 1976). Eggs range from 1.36 to 1.50 mm in diameter and hatch in 9 to 14 days at 15.5° C (Emmett et
al. 1991).
Fish
Surveys
Plants
Amphibians &
Reptiles
Figure 2.5 Spatial and Temporal Distribution of Young-of-the-Year Pacific Staghorn Sculpin (CDFG 1987)
Surveys
Figure 2.6 Spatial and Temporal Distribution of Adult Pacific Staghorn Sculpin (CDFG 1987)
is an important prey item for aquatic birds, particularly
the great blue heron (Tasto 1975, Bayer 1985, Emmett
et al. 1991).
Distribution
Pacific staghorn sculpin have been collected in all four
subregions of the Bay. Larval abundance was determined
to be highest from December through March, peaking
in February, in various parts of the Estuary south of the
Carquinez Bridge (CDFG 1987). Small juveniles are
often found intertidally; catch patterns suggest that, during their first year, these early post larval forms move
gradually from shallow inshore areas to deeper Bay waters (CDFG 1987, Emmett et al. 1991). In studies conducted in Yaquina Bay, Oregon, young-of-the-year first
appeared in December, and were collected through April
124
Baylands Ecosystem Species and Community Profiles
(Bayer 1985). Juveniles and adults are most frequently
captured in central Bay and San Pablo Bay, and are more
abundant in the channels in winter, and on the shoals
in spring and summer (Figures 2.5 and 2.6). Adults experience their widest distribution during high Delta outflow, and it appears that a portion of the adult population moves out of the Estuary by late spring of their second year (CDFG 1987). In Elkhorn Slough (Monterey
County), staghorn sculpin were highest in abundance,
and frequently the dominant species, at sampling stations
furthest inland, near sources of fresh water (Yoklavich
et al. 1991). A tidal marsh population studied in Anaheim Bay, a relatively small embayment in southern California with little freshwater input, was composed almost
entirely of juveniles (Tasto 1975). Pacific staghorn sculpin
can also be found a mile or two up coastal streams in association with exclusively freshwater species (Moyle 1976).
References
Multiple gear catch statistics from 1980-85 showed that
Pacific staghorn sculpin was the most abundant of all the
sculpins caught in the Estuary, and approximately 4%
of all fishes caught by otter trawl and beach seine (CDFG
1987). The highest abundance of larvae noted in this
study occurred during years of low Delta outflows, yet
juvenile and adult numbers showed no quantifiable relationship to magnitude of flows (CDFG 1987). Larval
success is thought to be the determining factor in overall recruitment to local populations (Emmett et al. 1991).
Armstrong, J.L., D.A. Armstrong and S.B. Mathews.
1995. Food habits of estuarine staghorn sculpin,
Leptocottus armatus, with focus on consumption
of juvenile Dungeness crab, Cancer maagister. Fishery Bulletin. Vol. 93: 456-470.
Barnhart, P.S. 1936. The marine fishes of southern California. Univ. Ca. Press. Berkeley. 209 pp.
Barry, J.P., M.M. Yoklavich, G.M. Cailliet, D.A.
Ambrose and B.S. Antrum. 1996. Trophic ecology of the dominant fishes in Elkhorn Slough, California, 1974-1980. Estuaries., Vol. 19 (1): 115-138.
Bayer, R.D. 1981. Shallow water ichthyofauna of the
Yaquina Estuary, Oregon. Northwest Science,
55(3): 182-193.
______. 1985. Shiner perch and Pacific staghorn
sculpins in Yaquina Estuary, Oregon. Northwest
Science, 59(3): 230-240.
Bolin, R.L. 1944. A review of the marine cottid fishes
of California. Stanford Ichthyol. Bull., 3: 1-135.
Boothe, P. 1967. The food and feeding habits of four
species of San Francisco Bay fish. Ca. Dept. Fish and
Game, Mar. Res. Oper. Reference (67-13). 155 pp.
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-85. Exhibit 60. Ca. Dept. Fish and
Game. State Wat. Res. Ctrl. Bd., Wat. Qual./Wat.
Rights Proc. on San Fran. Bay/Sac.-San Joaquin
Delta. 337 pp.
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Volume
11: species life history summaries. ELMR Rep. No.
8. NOAA/NOS Strategic Environmental Assessments Div., Rockville, MD. 329 pp.
Jones, A.C. 1962. The biology of the euryhaline fish
Leptocottus armatus armatus Girard (Cottidae).
Univ. of Ca. Publ. in Zool., 67 (4): 321-368.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Fish Bull (157).
Coop. Ext. Univ. Ca. Div. of Ag. and Nat. Res.
Publ. 4065. 249 pp.
Morris, R.W. 1960. Temperature, salinity, and southern limits of three species of Pacific cottid fishes.
Limnology and Oceanography, 5(2): 175-179.
Moyle, P.B. 1976. Inland Fishes of California. Univ. Ca.
Press. Berkeley. 405 pp.
Tasto, R.N. 1975. Aspects of the biology of Pacific staghorn sculpin, Leptocottus armatus Girard, in Anaheim Bay. In: E. David Lane and Cliff W. Hill (eds).
The marine resources of Anaheim Bay. Ca. Dept.
Fish and Game, Fish Bull. (165): 123-135
Yoklavich, M.M., G.M. Cailliet, J.P. Barry, D.A.
Ambrose and B.S. Antrum. 1991. Temporal and
Trophic Levels
Larvae are first and second order consumers (Emmett et al.
1991). Adults and juveniles are higher order consumers.
Proximal Species
Predators: Diving ducks, great blue heron, western
grebe, Caspian tern, loons, cormorants, gulls, marine
mammals.
Prey: Crangon shrimp (principal prey item), bay goby
(prey of large adults), mud crab, callianassid shrimp, amphipods (juvenile prey item, dominant in fresh water).
Competitor: Starry flounder.
Good Habitat
Success of local staghorn sculpin populations depends
upon the quality and quantity of suitable habitat. Newly
settled juveniles use intertidal and shallow subtidal mudflats for protection and feeding (Tasto 1975), although
older juveniles and adults are said to prefer more sandy
substrates and somewhat deeper waters (Bayer 1981,
Emmett et al. 1991). Pacific staghorn sculpin are known
to bury themselves in soft substrates, and have been
found buried in mudflats after the tide has retreated
(Tasto 1975, Bayer 1985). Staghorn sculpin have also
been found associated with eelgrass (Bayer 1981).
Water quality factors are equally important for successful populations. Demersal eggs hatch most successfully at 26 ppt and larvae survive best at 10 to 17 ppt
(Jones 1962). Greatest catches of larvae were in surface
salinities of 18 to 30 ppt (CDFG 1987). The juvenile
stage appears to be the most euryhaline, with the maturing and adult forms most likely to be found in the
higher salinity waters (CDFG 1987, Emmett et al.
1991). Laboratory experiments have shown that adult L.
armatus can survive 67.5 ppt at 12° C, but gradually lose
their tolerance of high salinities as temperatures rise to
25° C (Morris 1960). Since larval development is planktonic, it does not appear that, under normal conditions
in the San Francisco Estuary, either temperature or salinity are very limiting to distribution.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
125
Fish
Population Status and Influencing Factors
spatial patterns in abundane and diversity of fish
assemblages in Elkhorn Slough, California. Estuarine Research Federation, Vol. 14, No. 4: 465-480.
Prickly Sculpin
Fish
Cottus asper
Bruce Herbold
Sculpins (Family: Cottidae) are specialized for living on
the bottom, generally hiding in the nooks and crannies
among rocks or rooted vegetation. Their large, flattened
heads and proportionally small bodies, their fan-shaped
pectoral fins, and their lack of an air bladder allow
sculpins to hold their position even in wave-swept coasts
or high-velocity mountain streams. The use of such
habitats, combined with their secretive habits and cryptic
coloration, make sculpins difficult to see by predators,
prey, or inquisitive fish biologists. The large mouth relative to body size permits sculpins to consume prey almost as large as themselves. Sculpins are found in the
northern Pacific Ocean and New Zealand. Most members of the family are marine but a number of species
(most in the genus Cottus) occur in the fresh waters of
North America.
Reproduction
Sculpins generally spawn in the late winter or early
spring, although some upstream populations seem to
delay spawning into the early summer (Wang 1986).
Male sculpins prepare for spawning by moving downstream and establishing a nest site where they clean off
some kind of overhanging structure such as a flat rock,
tule root, or beer can (Kresja 1965, Moyle 1976). Females then enter the spawning area and, after a nocturnal courtship, attach their eggs to the prepared overhanging structure. Females produce between 280 and 11,000
eggs (Patten 1971), but one male may court many females and end up with a nest containing up to 30,000
eggs (Kresja 1965). Males stay in the nest protecting the
eggs and circulating water around them until they hatch.
Hatching rates appear to improve in saltier water
(Millikan 1968). After hatching, the larvae become
Fry at hatching average six mm total length. Newly
emerged fry swim soon after hatching and appear to drift
downstream as plankton for three to five weeks. This
early developmental pattern leads to high concentrations
in the slower waters of the Delta (Turner 1966). Young
fish assume a bottom-feeding existence at sizes of 20 to
30 mm, at which time they appear to begin moving upstream (McLarney 1968).
Food and Feeding
Sculpins have a reputation amongst anglers as predators
on salmonid eggs and fry (Munro and Clements 1937,
Shapovalov and Taft 1954, Reed 1967) which is probably undeserved (Moyle 1976, 1977). Diet studies generally show that sculpins prey principally on invertebrates, with younger prickly sculpins eating planktonic
crustaceans and older fish eating larger, benthic animals
and small fish (Moyle 1976). In Suisun Marsh their diet
is predominately benthic amphipods of the genus Gammarus (Herbold 1987).
Distribution
Prickly sculpins are found in fresh to brackish water from
the Kenai Peninsula in Alaska to the Ventura River in
southern California. In California’s Central Valley, they
can be found in the lower reaches of most foothill
streams. Prickly sculpins often overlap in distribution
with the similar riffle sculpin (Cottus gulosus) which is
Moyle 1976
126
Growth and Development
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
General Information
planktonic and are carried further downstream. Young
sculpins (15-30 mm SL, Broadway and Moyle 1978)
settle to the bottom and begin a general upstream movement (McLarney 1968, Mason and Machidori 1976).
The amount of movement associated with spawning appears to vary tremendously among sculpin populations (Wang 1986). Earlier observers suggested that
substantial downstream movements were only found in
coastal populations, not in the Central Valley (Kresja
1967). However, very high densities of newly hatched
prickly sculpins have been reported from the Delta and
Suisun Bay (Turner 1966, Wang 1986), as well as in
upstream sites (Wang 1986) which has led to the conclusion that the Central Valley contains both ‘migratory’
and ‘non-migratory’ populations. Recent studies suggest
that the same may be true in coastal streams, such as the
Eel River, where young prickly sculpins were found 100
km above the river mouth (Brown et al. 1995). Regardless of the degree to which they move for spawning,
mainstem rivers appear to be an important habitat for
most prickly sculpin populations. Young prickly sculpins
are often found in saline water at the tributary mouths
in spring months (Leidy pers. comm.).
found more in upper elevations. Neither is found in the
upper Pit River drainage. Their range includes tidal habitats of brackish salinity, such as Suisun Marsh. Prickly
sculpins are found from headwaters to the mouths of
many of the small tributaries that flow into San Francisco Bay, (including Alameda Creek, Walnut Creek,
Corte Madera Creek, Coyote Creek and the Petaluma
River; Leidy 1984).
Like freshwater sculpins generally, prickly sculpins use
very diverse habitats from small headwater streams to
coastal estuaries, and are widely distributed from Alaska
to southern California (Moyle 1976). Whatever the habitat, prickly sculpins usually are found under some sort
of cover: rocks in streams, vegetation in pools and
marshes, or simply at depth in lakes and reservoirs
(Moyle 1976, Brown et al. 1995).
Population Status and Influencing Factors
Many of the most recent, successful invading species of
the Estuary have the potential to affect prickly sculpins.
In 1986, the Asiatic clam (Potamocorbula amurensis) began a rapid and thorough domination of the benthic
community. Although the decline in abundance of other
benthic species has been well-documented, there is no
information on the impact of these changes on the diet,
distribution, or abundance of prickly sculpin. Also in the
mid-1980s, the Estuary was invaded by the shimofuri
goby (Tridentiger bifasciatus) which lives in the same
kinds of habitats and microhabitats as prickly sculpin.
However, the very small mouth of the goby reduces the
likelihood of interspecific competition. Since 1996,
mitten crabs (Eriochier sinensis) have become extremely
abundant and are voracious and indiscriminate predators on benthic organisms. Mitten crabs undergo an
annual upstream migration to spawn that results in a
large overlap with the range of prickly sculpins. In the
Eel River of northern California, it appears likely that
the introduction of predatory pikeminnows (Ptychocheilus grandis) has resulted in a substantial change in sculpin
behavior when compared to the similar Smith River
(Brown and Moyle 1991, Brown et al. 1995, White and
Harvey in press). In the tributary creeks of the San Francisco Bay drainage, prickly sculpins are often associated
with native species and are usually absent in areas where
large non-native predatory fish are found (Leidy 1984).
No work has been done to document interactions of
prickly sculpin with the vastly changed benthic community of the Central Valley.
Habitat changes and degradations of water quality are associated with a restricted range of prickly
sculpins in the San Joaquin River watershed (Brown
1998). Sculpins are part of an assemblage of native spe-
Trophic Levels
Prickly sculpins are secondary and tertiary consumers.
Proximal Species
Predators: Centrarchids and pikeminnows.
Prey: Planktonic crustacea (for young); benthic invertebrates, particularly gammarid amphipods; neomysis; juvenile fish.
Habitat: Emergent aquatic vegetation (root masses).
Good Habitat
In contrast to staghorn sculpins (Leptocottus armatus),
prickly sculpins larger than 20 mm are usually found in
association with some kind of complex, physical cover.
In upstream sites, cover consists of interstices in cobble,
root wads and woody debris and even discarded soda cans
and tires. In downstream sites, cover usually consists of
root wads of emergent aquatic vegetation. Although
Chapter 2 —
Estuarine Fish and Associated Invertebrates
127
Fish
Habitats
cies that are characteristic of smaller San Joaquin tributaries that have suffered little change in habitat structure or water quality. Unfortunately, the close associations of land use practices, habitat alteration and water
quality degradation in the rest of the watershed make it
impossible to identify the effects of individual environmental variables on sculpin biology.
As in the San Joaquin River, prickly sculpins in
Suisun Marsh tend to be found most often in association with other native fishes and in less disturbed habitats (Herbold 1987). However, the actual physical parameters of low dissolved solids and high gradient that
characterize usual sculpin sites in the San Joaquin River,
are absent in Suisun Marsh. This suggests that the impacts of land use and disturbance on the distribution and
abundance of prickly sculpins are not simple and that
the parameters that reflect disturbance in one area may
not be causally connected to the parameter of importance
to sculpins in that area.
California’s immense water projects appear to have
had little effect on prickly sculpins. Construction of dams
has isolated populations and prevented the downstream
movements exhibited elsewhere, but populations have
remained large in the warmwater reservoir behind Friant
Dam. Prickly sculpins are also found in stream habitats
upstream of impassable dams on a number of other Central Valley streams. Water export from the Delta has resulted in the establishment of new populations of prickly
sculpins within the facilities of the state and federal
projects, as well as within aquatic habitats in southern
California outside the historic range of prickly sculpin
(Wang 1986). The impacts of these introductions on the
native species in southern California streams have been
little studied.
more tolerant of salinity than most other California freshwater fish, sculpins are seldom found in salinities greater
than 10 ppt.
Broadway, J.E. and P.B. Moyle. 1978. Aspects of the
ecology of the prickly sculpin, Cottus asper Richardson, a persistent native species in Clear Lake,
Lake County, California. Environ. Biol. Fishes
3(4):337-343.
Brown, L.R. and P.B. Moyle. 1991. Changes in habitat
and microhabitat partitioning within an assemblage of stream fishes in response to predation by
Sacramento squawfish (Ptychocheilus grandis). Can.
J. Fish. Aquat. Sci. 48:849– 856.
Brown, L.R., S.A. Matern and P.B. Moyle. 1995. Comparative ecology of prickly sculpin, Cottus asper,
and coastrange sculpin, C. aleuticus, in the Eel
River, California. Env. Biol. Fish. 42:329– 343.
Herbold, B. 1987. Patterns of co-cccurrence and resource
use in a non-coevolved assemblage of fishes. Ph.D
dissertation. Univ. of California, Davis. Vii+81 pp.
Krejsa, R.J. 1965. The systematics of the prickly sculpin,
Cottus asper: an investigation of genetic and nongenetic variation within a polytypic species. Ph.D.
dissertation, Univ. of British Columbia.
Kresja, R. 1967. The systematics of prickly sculpin
(Cottus asper Richardson) a polytypic species: part
II. Studies on the life history, with especial reference to migration. Pac. Sci. 21:414-422.
Leidy, R.A. 1984. Distribution and ecology of stream
fishes in the San Francisco Bay drainage. Hilgardia
52(8):1-173.
Mason, J.C. and S. Machidori. 1976. Populations of
sympatric sculpins, Cottus aleuticus and Cottus
asper, in four adjacent salmon-producing coastal
streams on Vancouver Island, B. C. Fish. Bull.
74:131– 141.
McLarney, W.O. 1968. Spawning habits and morphological variation in the coastrange sculpin, Cottus
Personal Communications
Robert Leidy, U.S. Environmental Protection Agency
128
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Fish
References
aleuticus, and the prickly sculpin, Cottus asper.
Trans. Amer. Fish. Soc. 97:46-48.
Millikan, A.E. 1968. The life history and ecology of
Cottus asper Richardson and Cottus gulosus (Girard)
in Conner Creek, Washington. M.S. Thesis. Univ.
Wash. 81 pp.
Moyle, P.B. 1976. Inland Fishes of California. Univ. of
California Press. 405 pp.
______. 1977. In defense of sculpins. Fisheries 2:20-23.
Munro, J.A. and W.A. Clemens. 1937. The American merganser in British Columbia and its relation to the
fish population.. Biol. Bd. Canada Bull. 6(2):1-50
Patten, B.G. 1971. Spawning and fecundity of seven
species of northwest American Cottus. Amer. Mid.
Nat. 85(2): 493-506.
Reed, R.J. 1967. Observations of fishes associated with
spawning salmon. Trans. Amer. Fish. Soc. 96(1)
62-66.
Shapovolov, L. and A.C. Taft. 1954. The life histories
of the steelhead trout (Salmo gairdneri gairdneri)
and silver salmon (Oncorhynchus kisutch) Ca. Dept.
Fish and Game, Fish Bull. 98:1-375.
Turner, J.L. 1966. Distribution of threadfin shad,
Dorosoma petenense, tule perch, Hysterocarpus
traskii, and crayfish spp. in the Sacramento-San
Joaquin Delta. In: J.L. Turner and D.W. Kelley
(eds). Ecological studies of the Sacramento-San
Joaquin Delta, Part II. pp 160-168. Ca. Dept. Fish
and Game, Fish Bull. 136.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and Adjacent Waters: a Guide to
the Early Life Histories. Interagency Ecological
Study Program for the Sacramento-San Joaquin
Estuary, Tech. Rep. No 9. Sacramento, Ca.
White, J.L. and B.C. Harvey. In press. Habitat separation of prickly sculpin, Cottus asper, and Coast
range sculpin, C. aleuticus, in the mainstem Smith
River, northwestern California. Copeia.
the measure to the bottom of the fork of the tail fin) on
their third year, and 48 to 50 cm in their fourth year.
Growth of older adults is 1 to 3 cm annually. Most females mature at four to six years, but many are mature
by the end of their third year. Males typically mature
at two to three years old. Although striped bass apparently have the potential to live in excess of 30 years, most
adults are three to seven years old.
Striped Bass
Morone saxatilis
Ted R. Sommer
General Information
Reproduction
Striped bass are present in the San Francisco Estuary
throughout the year (Moyle 1976). They generally congregate in San Pablo and Suisun Bays in autumn and
move into the Delta and Sacramento River system on
their spawning migration during winter and early spring.
The timing and location of spawning depends on temperature, flow and salinity, but typically peaks in May
and early June. The annual spawning distribution appears to shift between the Sacramento and San Joaquin
rivers and the Delta.
Striped bass spawn in freshwater, with optimum
spawning at salinities of less than 1 ppt (Moyle 1976).
The species has exceptionally high fecundity—females
commonly broadcast from 0.5 to 4.5 million semi-buoyant eggs into the water column. The drifting eggs hatch
in the current in about 2 days. Eggs and newly-hatched
larvae are carried downstream to the Delta and Suisun
Bay. Larvae show peak abundance at the upstream edge
of the entrapment zone, located at a salinity of approximately 2 ppt.
Growth and Development
Striped bass grow to about 38 mm by late July or
August (Moyle 1976). They typically reach 23 to 35 cm
FL by their second year, 38 to 39 cm fork length (FL;
Food and Feeding
Striped bass are gregarious pelagic predators (Moyle
1976). They begin feeding at a length of 5-6 mm on several invertebrates including cladocerans and copepods.
Copepods generally dominate the diet of 7 to 11 mm
larvae, but the opposum shrimp, Neomysis, become a
more important food source in larger individuals. Youngof-the-year feed mostly on opossum shrimp, but amphipods, copepods, and threadfin shad are important alternative prey items. Fish gradually become a more important food source in juvenile bass (13 to 35 cm FL). Subadult and adult bass (age 2+) are primarily piscivorous,
although they are highly opportunistic depending on
prey availability.
Distribution
In contrast to the coastal Atlantic populations of striped
bass, most of the local population spend their lives in the
San Francisco Estuary. However, recent tagging studies
suggest that striped bass are spending more time in Suisun Bay, the Delta, and surrounding freshwater areas
(Sweetnam 1990). The current distribution of the species includes San Francisco Bay, San Pablo Bay, Suisun
Bay, the Delta, tributaries of the Sacramento River and
the Pacific Ocean (Herbold et al. 1992).
Population Status and Influencing Factors
Adult abundance has declined over the past 30 years,
from over 1.5 million in the late 1960s to about 0.5 million in recent years (CDFG 1992). The decline was most
dramatic between the beginning and the end of the
1970s, prompting the initiation of a hatchery stocking
program to supplement natural production (Harris and
Kohlhorst 1996). Stocking was conducted from 1981
through 1991—hatchery fish presently comprise a
substantial percentage (e.g., 35% of the 1990 year class)
of the adult population.
Year class abundance is assumed to depend on
the environmental conditions experienced by the eggs
and young fish (CDFG 1987, 1992). However, a steady
decline in the survival rate of yearlings stocked into the
Estuary suggests that habitat conditions for older fish
also play an important role (Harris and Kohlhorst 1996).
Abundance of young bass is strongly correlated with
Chapter 2 —
Estuarine Fish and Associated Invertebrates
129
Fish
Striped bass (Family: Percichthyidae) were introduced
into the Estuary in 1879, leading to a successful commercial fishery within 10 years (Herbold et al. 1992).
The commercial fishery for striped bass was banned in
1935 following a substantial decline in abundance which
appears to have begun at the turn of the century. The
species are presently the principal sport fish caught in San
Francisco Bay and is estimated to bring approximately
$45 million per year into local economies in the Estuary.
Herbold, B., A.D. Jassby and P.B. Moyle. 1992. Status
and trends report of aquatic resources in the San
Francisco estuary. San Francisco Estuary Project,
USEPA, Oakland, CA.
Moyle, P.B. 1976. Inland Fishes of California. Univ.
of California Press.405 pp.
Sweetnam, D. 1990. Recent changes in striped bass
migratory patterns in the Sacramento-San Joaquin
estuary. Interagency Ecological Workshop,
Asolimar, CA., 1990.
White Croaker
Genyonemus lineatus
Kurt F. Kline
Striped bass are secondary and higher order consumers.
General Information
Proximal Species
The white croaker (Family: Sciaenidae) is found in small
schools (Skogsberg 1939) and ranges from Magdalena
Bay, Baja California, to Mayne Bay, Vancouver Island,
British Columbia (Baxter, 1980, Hart 1973, Miller and
Lea 1972). It is abundant in San Francisco Bay, and supports both commercial and sport fisheries in nearshore
coastal waters, and a sport fishery in the Bay.
Major Prey Items: Zooplankton (cladocerans and copepods), terrestrial insects, opossum shrimp (Neomysis
mercedis), splittail, salmon, threadfin shad, American shad.
Good Habitat
Striped bass are able to tolerate a wide range of environmental conditions, illustrated by their ability to move
regularly between salt- and fresh-water (Moyle 1976).
Optimal temperatures for spawning appear to be from
15.6° to 20.0° C. Low oxygen (4 ppm) and high turbidity are also tolerated. Large rivers or tidal channels with
moderate water velocities are required to keep the eggs
and larvae suspended in the water column. Young-ofthe-year striped bass show highest abundance in the
entrapment zone, the region where fresh- and saltwater mix.
Reproduction
Approximately 50% of all white croakers are mature after their first year and all are mature by their fourth year
(Love et al. 1984). Along the coast, spawning appears
to take place in water from eight to 36 meters deep (Love
et al. 1984). In San Francisco Bay spawning occurs from
September through May (Wang 1986), with most
yolk-sac larvae (YSL) collected from November through
March (CDFG, unpub. data). Females batch spawn
18-24 times per season, with a batch consisting of 80037,200 eggs (Love et al. 1984).
References
California Department of Fish and Game (CDFG).
1987. Factors affecting striped bass abundance in
the Sacramento-San Joaquin river system. CDFG
Exhibit 25, State Water Resources Control Board
Bay Delta Hearings, Sacramento, CA.
______. 1992. A re-examination of factors affecting
striped bass abundance in the Sacramento-San
Joaquin estuary. State Water Resources Control
Board Bay Delta Hearings, Sacramento, CA.
Harris, M.D. and D.W. Kohlhorst. 1996. Survival and
contribution of artificially-reared striped bass in the
Sacramento-San Joaquin Estuary. IEP Newsletter,
Spring 1996.
130
Baylands Ecosystem Species and Community Profiles
Growth and Development
White croaker eggs are pelagic, spherical and transparent. Under laboratory conditions (~20° C), eggs hatched
in 52 hours. The newly hatched YSL are poorly devel-
Plants
Amphibians &
Reptiles
Trophic Levels
CDFG
Fish
Delta outflow and entrapment zone position, although
in recent years this relationship has deteriorated. For
example, in 1995 striped bass production was exceptionally poor despite wet conditions that increased the abundance of several other outflow-dependent species. Entrainment at diversions is known to be substantial, and
there is statistical evidence that these losses affect abundance. Nonetheless, losses at the projects during the
1980s were at least partially mitigated using hatchery
fish, yet the population decline has continued. The reduction in several invertebrate prey species has also been dramatic, particularly since the introduction of the Asian clam
Potamocorbula. The decline in survival of stocked fish
strongly suggests that competition for food has had an effect on the population. Other potentially important factors
include toxic substances, exotic species and illegal fishing.
oped, but by the sixth day the yolk-sac is absorbed, the
swim bladder is inflated and feeding begins (Watson
1982).
Throughout their life, white croaker growth is
fairly constant (Love et al. 1984). They may live to 12
years (Love et al. 1984) and reach a total length (TL) of
41.1 cm (Miller and Lea 1972).
Distribution
Fish
Along the coast, the greatest densities of larvae are found
near the bottom between 15 and 20 meters. The smallest juveniles are common from 3 to 6 meters, and move
to deeper water as they grow. Most adults are found in
waters less than 30 meters, although white croakers have
been recorded to 183 meters (Love et al. 1984).
Within San Francisco Bay, most of the pelagic eggs
and YSL are found in Central Bay (Wang 1986; CDFG,
unpub. data). As the larvae develop to the post yolk-sac
stage, they move toward the bottom. Tidal currents
probably transported white croaker larvae to South and
San Pablo bays. High outflow events during the winter,
which increases the gravitational currents, may increase
the transport of larvae to San Pablo Bay (Fleming, pers.
comm.). By September, most of the young-of-the-year
(YOY) migrate to Central Bay and by winter, emigrate
from the Bay (Fleming 1999)
Within the Bay, YOY white croaker are found at
lower salinities and higher temperatures than the one
year and older fish (1+), reflecting the broader distribution of YOY. The movements of older YOY and 1+ white
croaker out of the Bay during the late fall and winter may
be temperature related.
classes tend to dominate subsequent years’ 1+ catch and
the monthly catch per unit effort (CPUE) shows seasonal migration patterns within and out of the Bay. From
these data, one could draw the following conclusions: 1)
the white croaker “ population” within the Bay is an extension of the nearshore coastal population; 2) factors
that influence the nearshore population of white croaker
are independent of the Bay; and 3) factors that influence
the Bay “ population” appear to be the salinity, temperature and, perhaps most importantly, the size and distribution of the nearshore population.
Population Status and Influencing Factors
Prey: Northern anchovies, Cancer spp., shrimp spp.,
polychaetes.
The California Department of Fish and Game’s Bay
Study has generated annual abundance indices for white
croaker since 1980. The abundance of YOY white
croaker has fluctuated greatly over the past 19 years (Figure 2.7). Highest abundance indices of YOY were in
1980, 1986, 1992, 1993, and 1994. White croaker 1+
indices peaked between 1988 and 1991.
From 1981 to 1986, white croaker 1+ catches were
dominated by the 1980 year class and from 1987 to
1993, they were dominated by the large 1986 year class.
However, the relative size of a year class as YOY is not
indicative of the future abundance of 1+ fish in the Bay.
For example, the 1986 year class apparently contributed
to the subsequent 1+ indices more than either the 1980
or the 1993 year classes. The drought from 1987-1992
may have caused greater use of the Bay by the 1986 year
class than either 1980 or 1993 year classes.
Examination of the annual indices shows no relationship between the number of mature fish and YOY,
while the length frequency data shows that single year
Figure 2.7 Annual Abundance Indices of White
Croaker (Hieb 1999)
Trophic Levels
Secondary consumers.
Proximal Species
Good Habitat
White croaker are associated with soft substrates (Love
et al. 1984). In the Bay, white croaker are primarily
found in areas with the most marine-like (salinity and
temperature) conditions.
Acknowledgments
Some of the materials in this report are summarized from
the white croaker chapter of IEP Technical Report 63
(Flemming 1999).
References
Baxter, R.L. 1980. White croaker. Genyonemus lineatus.
Inshore fishes of California. Calif. Office of State
Printing, Sacramento, Ca.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
131
Kevin Fleming, California Department of Fish and
Game, Bay-Delta and Special Water Projects Division, Stockton, CA.
Cymatogaster aggregata
Michael F. McGowan
General Information
The shiner perch (Family: Embiotocidae) is a small but
abundant species common to the intertidal and subtidal
zones of bays, estuaries, and the nearshore regions of
California. They are commonly caught by anglers around
rocks, and pilings, from shore and docks, and just about
any fishing area. They are also used as live bait in the
San Francisco fisheries for striped bass and California
halibut.
Reproduction
The shiner perch, like other members of the family
Embiotocidae, is a live-bearer. Mating is accompanied
by elaborate courtship behavior and occurs primarily in
the spring and summer in California (Shaw 1971). Females give birth during April and May (Odenweller
1975) in California. Fecundity ranges from 5-36 young
per female, depending on size (Emmett et al. 1991).
Growth and Development
At birth, the fully developed young are 34.0-43.7 mm
long (Wang 1986). Juveniles become adults at 5 cm in
length. Growth is rapid the first year but slows subsequently (Odenweller 1975). Most females mature their
first year. They may live 8 years and reach 20 cm long.
Males mature soon after birth and rarely grow beyond
13 cm.
Food and Feeding
Embryos receive nutrition and gas exchange through
ovarian placenta tissues and fluids. Juveniles and adults
feed on plankton and benthos depending on availability. Prey items include copepods, isopods, amphipods,
mussels, barnacle appendages, mysids, crab larvae, and
other small invertebrates or protruding parts of invertebrates (Emmett et al. 1991).
CDFG
Fish
Amphibians &
Reptiles
Personal Communications
Shiner Perch
132
Baylands Ecosystem Species and Community Profiles
Plants
Fleming, K. 1999. White croaker. In J. Orsi (ed). Report on the 1980-1995 fish, shrimp, and crab sampling in the San Francisco Estuary, California. IEP
Tech. Rept. No. 63.
Hart, J.L. (ed). 1973. Pacific fishes of Canada. Fish.
Res. Board Can., Bull. No. 180. 740 pp.
Hieb, K. 1999. San Francisco Bay species abundance
(1980-98). IEP Newsletter, Vol. 12(2): 30-34.
Love, M.S., G.E. McGowen, W. Westphal, R.J.
Lavenberg and L. Martin. 1984. Aspects of the
life history and fishery of the white croaker,
Genyonemus lineatus (Sciaenidae), off California.
Fishery Bull. 82(1): 179-198.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Fish Game, Fish
Bull. 157, 235p.
Skogsberg, T. 1939. The fishes of the family Sciaenidae
(croakers) of California. Ca. Fish Game, Fish Bull.
54, 62p.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and adjacent waters, California:
A guide to the early life histories. IEP Tech. Rep.
No. 9. Ca. Dept. Water Res., Ca. Dept. Fish Game,
U.S. Bureau Reclam. U.S. Fish Wildl. Serv.
Watson, W. 1982. Development of eggs and larvae of
the white croaker, Genyonetemus lineatus ayres
(Pices: Sciaenidae), of the southern California
coast. Fish. Bull. 80:403-417.
Distribution
Population Status and Influencing Factors
The availability and quality of estuarine areas for giving
birth and rearing young may limit populations. Key factors are water temperature, not excessively hot (Odenweller 1975), and seagrass beds for shelter and feeding.
San Francisco Bay shiner perch catches in trawl surveys
declined in 1983, perhaps due to high outflow (and resulting low salinity) that year (Herbold et al. 1992). Because it uses nearshore areas, the shiner perch may have
high body burdens of pesticides and other compounds
(Earnest and Benville 1971), but population effects of
chronic pollution have not been documented.
Trophic Levels
Shiner perch are secondary and higher level consumers.
Plant matter found in some stomach analysis studies may
be due to feeding on invertebrates that occur on the
aquatic vegetation.
Proximal Species
Predators: Sturgeon spp., salmon spp., striped bass,
California halibut, cormorant spp., great blue heron,
bald eagle.
Prey: Copepods, isopods, amphipods, mussels, barnacle
appendages, mysids, crab larvae, and other small invertebrates or protruding parts of invertebrates.
Good Habitat
The shiner perch appears to favor aquatic vegetation if
present, but is also found over shallow sand and mud
bottoms. They prefer salinities greater than 8-10 ppt and
were reported in water temperatures ranging from 4 to
21° C (Emmett et al. 1991). In San Francisco Bay, they
are widespread but are most abundant downstream of
References
Bayer, R.D. 1979. Intertidal shallow-water fishes and
selected macroinvertebrates in the Yaquina estuary, Oregon. Unpubl. Rep., 134 pp. Oregon State
Univ. Marine Sci. Cent. Library, Newport, OR.
Not seen, cited in Emmett et al. 1991.
Earnest, R.D. and P.E. Benville, Jr. 1971. Correlation
of DDT and lipid levels for certain San Francisco
Bay fish. Pest. Monitor. Journal 5(3):235-241.
Emmett, R.L., S.L. Stone, S.A. Hinton and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Vol.
II: species life history summaries. ELMR Rept. No.
8. NOAA/NOS Strategic Environmental Assessments Division, Rockville, MD, 329 p.
Hart, J.L. 1973. Pacific fishes of Canada. Fish. Res.
Board Can., Bull. No. 180, 740 p.
Herbold, B., A.D. Jassby and P.B. Moyle. 1992. Status
and trends report on aquatic resources in the San
Francisco estuary. San Francisco Estuary Project,
Oakland, CA, 257 pp. + App.
Moyle, P.B. 1976. Inland Fishes of California. Univ. Ca.
Press, Berkeley, CA, 405 pp.
Odenweller, D.B. 1975. The life history of the shiner
surfperch, Cymatogaster aggregata Gibbons, in Anaheim Bay, California. In: E. D. Lane and C.W.
Hill (eds). The marine resources of Anaheim Bay.
Ca. Fish Game, Fish Bull. 165:107-115.
Shaw, E. 1971. Evidence of sexual maturation in young
adult shiner perch, Cymatogaster aggregata Gibbons
(Perciformes, Embiotocidae). Am. Mus. Nov.
2479:1-10.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: A
guide to the early life histories. Tech. Rep. 9. Interagency ecological study program for the Sacramento-San Joaquin estuary. Ca. Dept. Water Res.,
Ca. Dept. Fish Game, U.S. Bureau Reclam. U.S.
Fish Wildl. Serv., various pagination.
Yoklavich, M.M., G.M. Cailliet, J.P. Barry, D.A.
Ambrose and B. S. Antrim. 1991. Temporal and
spatial patterns in abundance and diversity of fish
assemblages in Elkhorn Slough, California. Estuaries 14(4):465-480.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
133
Fish
The shiner perch occurs near shore and in bays and estuaries from Baja California to Alaska commonly associated with aquatic vegetation. Juveniles prefer intertidal
and shallow subtidal habitats in bays and estuaries
(Moyle 1976). In winter they may move out of estuaries to nearshore areas and have been found as deep as
70 m (Hart 1973). In Elkhorn Slough, where they are a
numerically dominant component of the fish fauna, they
were classified as partial residents (Yoklavich et al. 1991).
the Carquinez Strait. Herbold et al. (1992) considered
them a euryhaline species. Eelgrass beds may be important feeding areas because shiner perch use them more
at night than during the day (Bayer 1979).
Tule Perch
Hysterocarpus traskii
Fish
Robert A. Leidy
General Information
Food and Feeding
Tule perch are the only viviparous freshwater fish native to California and the only freshwater member of the
surfperch family (Embiotocidae)(Baltz and Moyle 1982).
They are deep-bodied, spiny-rayed fish found in lakes,
rivers, streams, and estuaries in habitats characterized by
complex cover, especially well developed beds of aquatic
macrophytes (Moyle 1976). There are three recognized
subspecies of tule perch, H. t. pomo from the Russian
River drainage, H. t. lagunae from Clear lake, and H. t.
traskii from the Sacramento-San Joaquin drainage,
which includes populations found in the San Francisco
Estuary (Hopkirk 1973, Baltz and Moyle 1981 and
1982). Because of their small size, tule perch have no
commercial and limited sport value.
Within the Sacramento-San Joaquin Delta and upper Estuary, tule perch feed primarily on mysid shrimp, small
amphipods, midge larvae (Chironomidae), and clams
(Cook 1964, Turner 1966). Hopkirk (1962) recorded
that tule perch collected in brackish water habitats near
the mouth of the Napa River fed mostly on small-sized
brachyuran crabs, while juvenile fish feed predominantly
on midge larvae and pupae. Tule perch are also known
to feed on zooplankton, aquatic insects, and a variety of
benthic and plant-dwelling invertebrates in lakes and rivers (Moyle 1976, Wang 1986).
Reproduction
Tule perch breed during July through September, but
fertilization of the eggs is delayed within the female until January (Bundy 1970, Bryant 1977). Embryos develop
within the females ovarian compartments and are born
as juveniles in May or June, at a length of between 3040 mm standard length (SL) (Bryant 1977). The number of fish produced per female is positively correlated
with the size of the female fish and ranges between 22
and 93 (Bundy 1970, Bryant 1977).
Growth and Development
Moyle 1976
Juveniles begin schooling immediately following birth
within aquatic vegetation, submerged logs, or boulders
(Wang 1986). It is not known whether juveniles move
into tributaries following birth, but it is interesting to
note that several streams feeding into Suisun Marsh and
San Pablo Bay contain large numbers of juvenile tule
perch (Leidy, pers. observ.). Juveniles grow rapidly and
individuals in the Sacramento-San Joaquin Delta may
reach 80 to 100 mm SL following the first year of growth
134
Baylands Ecosystem Species and Community Profiles
Distribution
Tule perch are native to low-elevation valley waters of
the Central Valley, the Sacramento-San Joaquin Delta,
including Suisun Marsh and several streams tributary to
the San Francisco Estuary, Clear Lake, and the Russian,
Salinas, and Pajaro Rivers (Moyle 1976). Within the San
Francisco Estuary tule perch have been recorded from
Suisun Marsh (Herbold et al. 1992), including Montezuma Slough, Suisun Bay (Ganssle 1966), Carquinez
Strait (Messersmith 1966), the Napa River and its
marshes (Moyle 1976; Leidy 1984; Leidy, unpub. data),
and Sonoma, Alameda, and Coyote creeks (Leidy 1984).
Tule perch may be considered locally abundant in lower
estuarine and riverine intertidal marsh and pelagic habitats of Suisun Marsh and several of its tributary streams,
the Napa and Sonoma Creek marshes, and portions of
San Pablo Bay (Leidy, unpub. data). Tule perch no
longer occur in the Pajaro and Salinas rivers, and are rare
in Alameda and Coyote creeks (Leidy, unpub. data).
Population Status and Influencing Factors
While the historical range of tule perch within the San
Francisco Estuary has been reduced, tule perch are still
locally abundant in Suisun Marsh and the Napa River
and Sonoma Creek and its tidal marshes. Important factors negatively influencing population numbers likely
include excess siltation and turbidity, reduced freshwater flows, pollution, removal of riparian vegetation and
aquatic macrophytes through stream channelization and
other flood control measures, and the resultant loss of
nesting, feeding, and cover habitat, and possibly the introduction of exotic centrarchids (Moyle et al.1995).
Moyle et al. (1995) identified introduced fish predators,
such as smallmouth bass (Micropterus dolomieui), pond
Plants
Amphibians &
Reptiles
(Moyle 1976). Maximum size for tule perch is approximately 160 mm SL, although a single individual measuring 175 mm SL was collected in Napa Slough, Napa
County (Leidy, unpub. data). Tule perch rarely live
longer than five years (Moyle 1976).
and dam construction, and reduced flows and poor water quality as threats to the Russian River subspecies of
the tule perch. These are likely threats to the other two
subspecies of tule perch as well. Interestingly, otter trawl
data collected in Suisun Marsh shows a significant decline in tule perch numbers during 1983-84, a year of
extremely high outflow (Herbold et al. 1992).
Trophic Levels
Proximal Species
Juvenile predators: Other large freshwater and estuarine piscivorous fish, egrets, herons and other wading birds.
Prey: Aquatic and terrestrial insects, zooplankton, mysid
shrimp, amphipods, clams, brachyuran crabs, midge larvae and pupae.
Good Habitat
Tule perch may be found in a variety of habitats from
the slow-moving, turbid channels of the Delta, marshes
between the mouths of Sonoma Creek and the Napa
River, to relatively clear, fast-flowing rivers and streams
(Moyle 1976; Leidy, unpub. data). In tidal riverine
marshes, tule perch prefer slow-moving backwater and
slough habitats with structurally-complex beds of floating or emergent aquatic macrophytes, overhanging banks
and/or submerged woody debris. These areas serve as
important feeding and breeding habitats, as well as protective rearing areas (Moyle 1976). Structurally-complex
cover appears to be essential for near-term females and
juveniles as refugia from predators (Moyle et al. 1995).
Although Moyle (1976) states that tule perch seldom venture into brackish water, they are present in the
pelagic zone of tidal riverine and intertidal estuarine environments, such as the Napa River marshes and Suisun Marsh (Leidy, unpub. data). This suggests that some
populations of tule perch may be able to tolerate brackish water conditions, or at least utilize these areas when
freshwater outflows dilute surface water. In Suisun
Marsh tule perch are most frequently collected in the
small, heavily vegetated, dead-end sloughs where introduced centrarchids are uncommon (Moyle et al. 1985).
References
Baltz, D.M. and P.B. Moyle. 1981. Morphometric analysis of the tule perch (Hysterocarpus traski) populations
in three isolated drainages. Copeia 1981: 305-311.
______. 1982. Life history characteristics of tule perch
(Hysterocarpus traski) populations in contrasting
environments. Environ. Biol. of Fishes 7:229-242.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
135
Fish
Juveniles and adults are primary and higher order consumers.
Bryant, G.L. 1977. Fecundity and growth of the tule perch,
Hysterocarpus traski, in the lower Sacramento-San
Joaquin Delta. Ca. Fish, Game 63(3): 140-156.
Bundy, D.S. 1970. Reproduction and growth of the tule
perch, Hysterocarpus traskii (Gibbons), with notes
on its ecology. M.S. Thesis, Univ. of Pacific, Stockton, Ca.. 52 pp.
Cook, S.F., Jr. 1964. The potential of two native California fish in the biological control of chironomid
midges (Diptera: Chironomidae). Mosquito News
24(3): 332-333.
Ganssle, D. 1966. Fishes and decapods of San Pablo
and Suisun Bays. In: J. L. Turner and D. W. Kelley
(eds). Ecological studies of the Sacramento-San
Joaquin Estuary, Part I. Ca. Dept. of Fish and
Game Bull. 133: 57-63.
Herbold, B., A.D. Jassby and P.B. Moyle, 1992. Status
and trends report on aquatic resources in the San
Francisco Estuary. U.S. EPA, San Francisco Estuary Project. 257 pp.
Hopkirk, J.D. 1962. Morphological variation in the
freshwater embiotocid Hysterocarpus traskii Gibbons.
M.A. Thesis, Univ. of California, Berkeley. 159 pp.
______. 1973. Endemism in fishes of the Clear Lake
region of central California. Univ. of Ca. Publ.
Zool. 96: 160 pp.
Leidy, R.A. 1984. Distribution and ecology of stream
fishes in the San Francisco Bay drainage. Hilgardia
52(8): 1-175.
Messersmith, J.D. 1966. Fishes collected in Carquinez
Strait in 1961-1962. In: J. L. Turner and D. W.
Kelley, eds., Ecological studies of the SacramentoSan Joaquin Estuary, Part I. pp. 57-63. Ca. Dept.
Fish and Game Bull. 133.
Moyle, P.B. 1976. Inland Fishes of California. Univ. of
California Press. 405 pp.
Moyle, P.B., R.A. Daniels, B. Herbold and D.M. Baltz.
1985. Patterns indistribution and abundance of a
noncoevolved assemblage of estuarine fishes in Ca..
Fish. Bull. 84: 105-117.
Moyle, P.B., R.M. Yoshiyama, J.E. Williams, and E.D.
Wikramanayake. 1995. Fish species of special concern in California. Prepared for the Resources
Agency, Rancho Cordova, Ca.. 272 pp.
Turner, J.L. 1966. Distribution of threadfin shad,
Dorosoma petenense, tule perch, Hysterocarpus
traskii, and crayfish spp. in the Sacramento-San
Joaquin Delta. In: J. L. Turner and D. W. Kelley
(eds). Ecological Studies of the Sacramento-San
Joaquin Delta, Part II. Ca. Dept. Fish and Game
Bull. 136:160-168.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and adjacent waters, California:
A guide to the early life histories. Tech. Rept. 9.
Prepared for the Interagency Ecological Study program for the Sacramento-San Joaquin Estuary.
Arrow Goby
Clevelandia ios
Kathryn A. Hieb
Fish
General Information
The arrow goby (Family: Gobiidae) is probably the most
abundant native goby in San Francisco Bay. It ranges
from the Gulf of California to Vancouver Island, British Columbia (Miller and Lea 1972) and is common to
intertidal mudflats and shallow subtidal areas of bays, estuaries, and coastal lagoons. It is often commensal with
burrowing invertebrates. The arrow goby grows to a
maximum size of 45 to 50 mm total length (TL). This
small fish is an important component of the intertidal
food web, as it is a common prey item for a variety of
birds and fishes. It has no sport or commercial value.
In Elkhorn Slough, ripe females were collected from
December through August, but were most common
from March through June (Prasad 1948). The reproductive period occurs approximately one to two months
earlier in southern California—in Mission Bay, ripe females were collected from September through June, with
peak abundance from November through April (Brothers 1975), while in Anaheim Bay, ripe females were collected from December through September, with peak
abundance from February through June (Macdonald
1972). Ovary development is asynchronous, as ovaries
are found in various stages of maturation during the
spawning season (Macdonald 1972, Brothers 1975).
This indicates that each female may spawn several times
during the spawning season. Fecundity ranges from 800
to 1,200 eggs per female, with clutch size ranging from
150 to 350 eggs (Brothers 1975) or from 750 to 1,000
eggs (Prasad 1948).
Some disagreement exists in the literature on the
deposition of the eggs and parental care. The eggs are
either deposited on surfaces with no additional parental
investment (Prasad 1948, Macdonald 1972) or deposited on the wall of burrows constructed by the male and
guarded by the male until hatching (Brothers 1975). In
Mission Bay, all males collected in January and February were brooding clutches of eggs in burrows. Typical
of most gobies, the fertilized eggs are club-shaped, with
Newly hatched larvae range from 2.75-3.25 mm TL
(Prasad 1948). Juvenile arrow gobies settle from the
plankton at approximately 8 mm standard length (SL)
and are found in burrows when they are 10-14 mm SL
(Macdonald 1972). The arrow goby matures at one year
and a length of 30 to 40 mm SL in Anaheim and Mission bays (Macdonald 1972, Brothers 1975); in Elkhorn
Slough females begin to mature at 29 mm SL and all are
mature at 34 mm SL (Prasad 1948). In southern California, most arrow gobies die after spawning, with a few
living to two years (Macdonald 1972, Brothers 1975).
In Elkhorn Slough, arrow gobies commonly live two to
three years (Prasad 1948). Fish from Elkhorn Slough apparently spawn later, grow slower, mature later, and
reach a larger size than fish from southern California
populations (Brothers 1975).
Food and Feeding
The arrow goby preys on a variety of small invertebrates.
In Mission Bay, the major prey items (percent occurrence) of juveniles and adults are harpacticoid copepods
(88%), ostracods (58%), tanacians (32%), gammarid
amphipods (19%), mollusc siphon tips (11%), caprellids
(8%), nematodes (7%), and polychaetes (7%) (Brothers
1975). In Anaheim Bay, the most important prey items
are harpacticoid copepods, nematodes, oligochaetes, ostracods, and cylcopoid copepods (Macdonald 1972).
Larvae prey primarily upon the calanoid copepod Acartia
tonsa (Macdonald 1975).
The arrow goby is preyed upon by a variety of demersal fishes, including Pacific staghorn sculpin (MacGinitie and MacGinitie 1949, Brothers 1975, Tasto 1975),
California halibut (Haaker 1975, Drawbridge 1990), and
diamond turbot (Lane 1975). MacGinitie and MacGinitie (1949) presumed probing shorebirds, including
willets, godwits, and curlews would capture arrow gobies while exploring burrows at low tides. Arrow gobies
have been found in the stomachs of greater yellowlegs
and dowitchers (Reeder 1951).
Distribution
The arrow goby is common on mudflats inhabited by
its invertebrate commensal hosts (Brothers 1975), with
densities up to 20/m 2 in Anaheim Bay (Macdonald
1972). It apparently utilizes invertebrate burrows as a
refuge from predators and as a temporary shelter during low tides. The arrow goby primarily inhabits burrows of the ghost shrimp (Callianassa californiensis), the
CDFG
136
Growth and Development
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Reproduction
an attachment thread at one pole. Hatching occurs in
10 to 12 days and the newly hatched larvae are pelagic
(Prasad 1948, Brothers 1975).
Population Status and Influencing Factors
Because the arrow goby is most common in intertidal
and shallow subtidal habitats, it is more effectively
sampled by seines than trawls. In a beach seine survey
of San Francisco Bay conducted by California Department of Fish and Game in the 1980s, the arrow goby
comprised approximately 4% of the catch, ranking eigth
of all fishes collected. In contrast, it comprised only
0.04% of the fishes collected by the otter trawl (Orsi
1999). As the beach seine survey has been discontinued,
there is no long-term monitoring program in the Bay that
effectively samples the arrow goby, and its current status is difficult to assess. From 1981 to 1986, the arrow
goby beach seine annual “ abundance index” varied almost 10-fold, with the highest indices in 1981 and 1986
(Figure 2.8).
Brothers (1975) hypothesized that arrow goby
abundance and distribution could be controlled by the
abundance and distribution of the commensal invertebrates, especially the ghost shrimp. Because the arrow
goby is an annual species, devoting a large proportion
of its resources to reproduction (“ r-strategist” ), it would
be expected to undergo large population fluctuations.
Trophic Level
Arrow goby larvae, juveniles, and adults are secondary
consumers, preying primarily on small benthic and
epibenthic invertebrates.
Proximal Species
Predators: Pacific staghorn sculpin, California halibut,
diamond turbot.
Prey: Harpacticoid copepods, ostracods, tanacians,
gammarid amphipods, mollusk siphon tips, nematodes,
oligochaetes.
Commensal Hosts: Burrowing invertebrates. Bat rays
and leopard sharks impact the abundance and distribution of burrowing invertebrates.
Figure 2.8 Annual Abundance Indices of Arrow
Goby from San Francisco Bay, Beach Seine
(CDFG, unpublished data)
Chapter 2 —
Estuarine Fish and Associated Invertebrates
137
Fish
fat innkeeper worm (Urechis caupo), the mud shrimp
(Upogebia spp.), and various bivalves (Prasad 1948,
Brothers 1975). Males also construct burrows for reproduction (Brothers 1975). At low tides the arrow goby is
also common in remnant pools of water on the mudflats
(Prasad 1948).
In San Francisco Bay, larval arrow gobies are most
abundant in South and San Pablo bays, with few collected upstream of Carquinez Strait in years with low
freshwater outflow (Wang 1986, CDFG 1987). Juveniles
and adults are common in shallow subtidal and intertidal areas of South, Central, and San Pablo bays and
have occasionally been collected in Suisun Bay (CDFG
1987). The arrow goby is also common in some tidal
marsh habitats from South Bay to lower San Pablo Bay.
It was the second most common species collected in otter trawl samples from Hayward Regional Shoreline
Marsh channel sites (Woods 1981). The arrow goby was
common in weir samples collected in Plummer Creek
(South Bay near Newark), although gobies were not
speciated in this study, so their relative abundance is
unknown (Wild 1969). In a survey of Castro Creek,
Corte Madera Creek, and Gallinas Creek marshes, the
arrow goby was relatively common in otter trawl samples
from creek channels and mudflats adjacent to the
marshes, but rare in gill nets and not collected by minnow traps set in the marsh channels (CH2M Hill 1982).
A few arrow gobies were collected in Petaluma River
marshes, Napa-Sonoma Marsh, but none in Suisun
Marsh (CDFG, unpub. data; ANATEC Laboratories
1981; CH2M Hill 1996; Matern et al. 1996).
Arrow goby larvae have been collected year-round
in San Francisco Bay, with peak larval abundance from
April through July (CDFG 1987). Peak abundance in
beach seine samples from the Bay is from March though
August; these catches include recently settled juveniles
and adults (CDFG 1987). In southern California, most
juveniles settle in the spring (February through May in
Mission Bay, February through June in Anaheim Bay),
although juveniles have been collected all but one or two
months in the fall (Macdonald 1972, Brothers 1975).
Juvenile and adult arrow gobies are euryhaline and
have been reported to tolerate salinities ranging from
freshwater to greater than seawater (Carter 1965, as cited
in Emmett et al. 1991). In San Francisco Bay, arrow
goby juveniles and adults have been collected from a wide
range of salinities (0.9-33.9‰ ), with 90% collected from
11.7 to 32.4‰ (5th and 95th percentiles, respectively,
CDFG 1987 and unpub. data). The arrow goby is also
reported to be eurythermal; in aquaria, gobies withstood
temperatures from 4-26° C, but were “ distressed” at temperatures above 22° C (Prasad 1948). In San Francisco
Bay, arrow gobies were collected from 7.5 to 30.5° C,
with 90% collected between 16.9 and 24.3° C (5th and
95th percentiles, respectively, CDFG 1987 and unpub.
data).
Good Habitat
References
ANATEC Laboratories, Inc. 1981. Infaunal, fish, and
macroinvertebrate survey in the Hudeman SloughSonoma Creek watercourses. Prepared for Sonoma
Valley County Sanitation District. Draft Final Report, November 1981.
Brothers, E.B. 1975. The comparative ecology and behavior of three sympatric California gobies. Ph.D.
Thesis, Univ. of California, San Diego, 365 pp.
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-1985. Exhibit 60, entered for the
SWRCB 1987 Water Quality/Water Rights Proceeding on the San Francisco Bay and SacramentoSan Joaquin Delta, 345 pp.
CH2M Hill. 1982. Equivalent protection study intensive investigation. Prepared for Chevron USA. Final Report, April 1982.
______. 1996. Fish Sampling and Water Quality Monitoring Results. In: S. Miner (ed). Annual Monitoring Report, Sonoma Baylands Wetlands Demonstration Project, U.S. Army Corps of Engineers,
San Francisco District and Ca. State Coastal Conservancy.
Drawbridge, M.A. 1990. Feeding relationships, feeding activity, and substrate preferences of juvenile
California halibut (Paralichthys californicus) in
coastal and bay habitats. M.S. Thesis, San Diego
State University, 214 pp.
Emmett, R.L., S.A. Hinton, S.L. Stone, and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Volume
II: species life history summaries. ELMR Report
No. 8, NOAA/NOS Strategic Environmental Assessments Division, Rockville, MD, 329 pp.
Haaker, P.L. 1975. The biology of the California halibut, Paralichthys californicus (Ayres), in Anaheim
Bay, California. In: E.D. Lane and C.W. Hill (eds).
The marine resources of Anaheim Bay. Ca. Dept.
of Fish and Game Fish Bull. 165:137-151.
Lane, E.D. 1975. Quantitative aspects of the life history of the diamond turbot, Hypsopsetta guttulata
138
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Fish
Good habitat for the arrow goby is shallow subtidal and
intertidal mudflats inhabited by the commensal invertebrate hosts. All habitats in tidal marshes may not suitable, as the arrow goby has been collected from larger
channels and adjacent mudflats, but not from smaller
order channels.
(Girard), in Anaheim Bay. In: E.D. Lane and C.W.
Hill (eds). The marine resources of Anaheim Bay.
Ca. Dept. of Fish and Game Fish Bull. 165:153173.
Macdonald, C.K. 1972. Aspects of the life history of
the arrow goby, Clevelandia ios (Jordan and Gilbert), in Anaheim Bay, California, with comments
on the cephalic-lateralis system in the fish family
Gobiidae. M.A. Thesis, California State University, Long Beach, 137 pp.
______. 1975. Notes on the family Gobiidae from Anaheim Bay. In: E.D. Lane and C.W. Hill (eds). The
marine resources of Anaheim Bay. Ca. Dept. of
Fish and Game Fish Bull. 165:117-121.
MacGinitie, G.E. and N. MacGinitie. 1949. Natural
history of marine animals. McGraw-Hill, New
York, N.Y. 473 pp.
Matern, S.A., L. Meng, and P.B. Moyle. 1996. Trends
in fish populations of Suisun Marsh. January 1995December 1995. Annual Report for Contract B59998. Ca. Dept. of Water Res., Sacramento, Ca.,
42 pp.
Miller, D.J. and R.N. Lea. 1976. Guide to the coastal
marine fishes of California. Ca. Dept. Fish and
Game Fish Bull. 157, 249 pp. SeaGrant reprint of
1972 edition with addendum added 1976.
Orsi, J. (ed). 1999 Report on the 1980-1995 fish,
shrimp, and crab sampling in the San Francisco
Estuary, California. IEP Tech. Rept. No. 63.
Prasad, R.R. 1948. The life history of Clevelandia ios
(Jordan and Gilbert). Ph.D Thesis, Stanford Univ.,
141 pp.
Reeder, W.G. 1951. Stomach analysis of a group of shorebirds. Condor 53:43-45.
Tasto, R.N. 1975. Aspects of the biology of the Pacific
staghorn sculpin, Leptocottus armatus, in Anaheim
Bay. In: E.D. Lane and C.W. Hill (eds). The marine resources of Anaheim Bay. Ca. Dept. of Fish
and Game Fish Bull. 165:123-135.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: A
guide to the early life histories. Interagency Ecological Studies Program for the Sacramento-San
Joaquin Estuary. Tech. Rept. No. 9.
Wild, P.W. 1969. Macrofauna of Plummer Creek of San
Francisco Bay collected by a specially designed trap.
M.A. Thesis, San Jose State University, San Jose,
Ca.. 85 pp.
Woods, E. 1981. Fish utilization. In: T. Niesen and M.
Josselyn (eds). The Hayward Regional Shoreline
marsh restorations: Biological succession during the
first year following dike removal. Tiburon Center
for Env. Studies, Tech. Rept. 1:35-46.
Growth and Development
Bay Goby
Lepidogobius lepidus
Kathryn A. Hieb
General Information
Reproduction
CDFG
Females with yolk filled eggs were collected from September through March, with the peak of reproductive
activity from January through March in Morro Bay
(Grossman 1979b). Gonadal development is asynchronous, typical of species that spawn several times a season and have a protracted spawning period. As for
many other species of gobies from temperate waters,
it is assumed the eggs are laid in burrows constructed
by either the males or commensal invertebrate hosts
and are guarded by the male until hatching (Wang
1986). Eggs are club shaped with an adhesive thread
at one pole for attachment to the burrow wall or substrate.
In San Francisco Bay, larvae were collected
throughout the year, with peak abundance from June to
October (CDFG 1987). The period of peak abundance
is similar in other Pacific Coast estuaries—peak larval
abundance is from April to September in Yaquina Bay,
Oregon (Pearcy and Myers 1974) and larvae were collected from April to September in Humboldt Bay
(Eldridge and Bryan 1972). In San Francisco Bay, most
larvae were collected in Central Bay and northern South
Bay, with relatively few collected upstream of San Pablo
Bay (CDFG 1987).
Food and Feeding
The bay goby is an opportunistic predator and major
prey items include polychaetes, harpacticoid copepods,
gammarid amphipods, and bivalves (Grossman et al.
1980). Although larger fish (>50 mm SL) and smaller
fish (<50 mm SL) consume similar prey items, larger fish
include more mollusks, polychaetes, and other larger
prey items in their diet.
Predators of the bay goby include the California
halibut (Drawbridge 1990) and the Pacific staghorn
sculpin (Boothe 1967). It is assumed that other demersal piscivorous fish prey upon bay gobies.
Distribution
In San Francisco Bay, the bay goby is common from
South to San Pablo bays, and is occasionally collected
in Carquinez Strait and lower Suisun Bay. Densities of
young-of-the-year (YOY) bay gobies are usually highest
in South or San Pablo bays while densities of older fish are
usually highest in Central Bay (CDFG 1987, Fleming
1999). From 1980 to 1995, the bay goby was the most
common goby and the second most common fish collected by an otter trawl survey of San Francisco Bay, comprising 14.3% of all fishes collected (Orsi 1999). Although mean densities of YOY fish were higher at shoal
stations than channel stations all months, older fish appear to move from the shoals to the channels in the late
summer and fall (CDFG 1987 and unpub. data).
Surprisingly, the bay goby was not common in a
beach seine survey conducted by CDFG in San Francisco Bay from 1980-1987; it was the fourth most common goby and comprised only 0.06% of all fishes collected by this net (Orsi 1999). These data indicate that
the bay goby may not be common in the very shallow
subtidal and intertidal areas of San Francisco Bay, although Grossman (1979a) concluded it to be one of the
Chapter 2 —
Estuarine Fish and Associated Invertebrates
139
Fish
The bay goby (Family: Gobiidae) ranges from Baja California to Vancouver Island, British Columbia (Miller and
Lea 1976). It is common to bays and estuaries and often commensal with burrowing invertebrates on intertidal mudflats (Grossman 1979a). Because it often occupies burrows, the bay goby is not effectively sampled
by trawls and seines and its relative abundance is undoubtedly greater than indicated by most surveys. It is
the most abundant native goby in larval surveys of San
Francisco, Humboldt, and Yaquina bays. The bay goby
grows to approximately 100 mm total length (TL) and
has no commercial or sport value.
Bay goby larvae are approximately 2.5-3.0 mm TL at
hatching (Wang 1986). The larvae are planktonic for
three to four months (Grossman 1979b) and settle to the
bottom as juveniles at approximately 25 mm TL (Wang
1986). Although the bay goby is reported to grow to
about 87 mm TL (Miller and Lea 1976), specimens as
large as 108 mm TL have been collected in San Francisco Bay (CDFG, unpub. data). Some bay gobies reach
sexual maturity by the end of their first year and by the
end of their second year all are mature (Grossman
1979b). Bay gobies reportedly live up to 7+ years
(Grossman 1979b), although based upon length frequency data from San Francisco Bay (CDFG 1987,
Fleming 1999), their life span may be as short as one to
two years.
Fish
Amphibians &
Reptiles
tribution of YOY somewhat further upstream than older
fish and by the peak abundance of YOY in the winter
and spring and older fish in summer and fall.
Population Status and Influencing Factors
Although trawls are usually considered ineffective for gobies, the bay goby is a very common fish in San Francisco Bay otter trawl surveys. As such, the abundance
indices derived from trawl data may be good indicators
of population trends. California Department of Fish and
Game otter trawl data from 1980-1998 is the longest
data set available for the Bay. The indices from 1988 to
1997 were generally higher than the pre-1988 indices
(Figure 2.9). The relatively stable salinities year-round
during the 1987-92 drought may have resulted in increased nursery habitat for this species (Hieb and Baxter
1993). The multiple cohorts of YOY collected these
years, which indicate successful recruitment over a period of several months, in part support this hypothesis.
Additionally, high winter outflow events may carry larvae or pelagic juveniles from the Bay.
Abundance of predators, as California halibut and
Pacific staghorn sculpin, could influence the bay goby
population. Additionally, factors controlling the abundance of the commensal burrowing invertebrate hosts
may effect the bay goby population. This would include
the abundance and distribution of intertidal and subtidal
mudflat invertebrate predators, such as the bat ray and
leopard shark.
Trophic Level
Secondary consumer.
Proximal Species
Predators: California halibut, Pacific staghorn sculpin.
Prey: Polychaetes, gammarid amphipods, harpacticoid
copepods, bivalves.
Table 2.6 Bay Goby Salinity and Temperature
Statistics: 1980-92 (CDFG unpublished data)
Mean
5th
percentile
Median
95th
percentile
YOY
27.3
14.9
29.2
31.7
1+ and older
28.1
17.1
29.7
32.4
Age Class
Salinity (ppt):
Temperature (°C):
YOY
15.4
11.3
15.2
18.8
1+ and older
16.0
12.4
16.3
18.9
140
Baylands Ecosystem Species and Community Profiles
Figure 2.9 Annual Abundance Indices of All Sizes of
Bay Goby, Otter Trawl (CDFG unpublished data)
Plants
numerically dominant fish species of Morro Bay lower
intertidal mudflats. The bay goby inhabits burrows of
the blue mud shrimp (Upogebia pugettensis) and the innkeeper worm (Urechis caupo) and siphon holes of the
geoduck clam (Panope generosa) in Morro Bay (Grossman
1979a). As for several other species of gobies common
to San Francisco Bay, including arrow goby and longjaw
mudsucker, the bay goby probably utilizes burrows as a
refuge from predators and to avoid desiccation at low
tides.
Few bay gobies have been collected in San Francisco Bay tidal marshes. One bay goby was reported from
Gallinas Marsh and one from Corte Madera Marsh
(CH2M Hill 1982). Both fish were collected by gill nets,
which were used to sample the larger channels. In contrast, the bay goby was the most common species collected in otter trawl samples from Corte Madera Creek
channel, adjacent to Corte Madera Marsh. No bay gobies have been collected by other San Francisco Bay tidal
marsh studies (Wild 1969; Woods 1981; ANATEC
Laboratories1981; CH2 MHill 1996; CDFG, unpub.
data) or by a study of fishes of Elkhorn Slough tidal
marshes (Barry 1983).
Bay goby YOY are most abundant in otter trawl
samples from February through June, which is a one or
two months after peak abundance period for smaller juveniles from the ichthyoplankton net (CDFG 1987 and
unpub. data). In several years, multiple cohorts of YOY
fish have been collected; this was especially noticeable
in four of the six years of the 1987-1992 drought
(CDFG, unpub. data). Peak abundance of older fish is
usually from May through September, which corresponds with the peak period of larval abundance in San
Francisco Bay.
The bay goby has been collected primarily from
polyhaline salinities in San Francisco Bay, with YOY fish
collected at lower salinities than older fish (Table 2.6).
YOY were also collected at slightly lower temperatures
than older fish (Table 2.6). These differences in salinity and temperature by age class are reflected by the dis-
Commensal Hosts: Blue mud shrimp, inn-keeper
worm, geoduck clam. Bat ray and leopard shark impact
the abundance of commensal hosts.
Good Habitat
References
ANATEC Laboratories, Inc. 1981. Infaunal, fish, and
macroinvertebrate survey in the Hudeman SloughSonoma Creek watercourses. Prepared for Sonoma
Valley County Sanitation District. Draft Final Report, November 1981.
Barry, J.P. 1983. Utilization of shallow marsh habitats
by fishes in Elkhorn Slough, California. MA Thesis, San Jose State Univ., San Jose, Ca. 91 pp.
Boothe, P. 1967. The food and feeding habits of four
species of San Francisco Bay fish. Ca. Dept. Fish
and Game MRO Reference No. 67-13, 155 pp.
California Department of Fish and Game (CDFG).
1987. Delta outflow effects on the abundance and
distribution of San Francisco Bay fish and invertebrates, 1980-1985. Exhibit 60, entered for the
SWRCB 1987 Water Quality/Water Rights Proceeding on the San Francisco Bay and SacramentoSan Joaquin Delta, 345 pp.
CH2M Hill. 1982. Equivalent protection study intensive investigation. Prepared for Chevron USA. Final Report, April 1982.
______. 1996. Fish Sampling and Water Quality Monitoring Results. In: Annual Monitoring Report,
Sonoma Baylands Wetlands Demonstration
Project, S. Miner, ed. U.S. Army Corps of Engineers, San Francisco District and Ca. State Coastal
Conservancy.
Drawbridge, M.A. 1990. Feeding relationships, feeding activity and substrate preferences of juvenile
California halibut (Paralichthys californicus) in
coastal and bay habitats. M.S. Thesis, San Diego
State University, San Diego, Ca. 214 pp.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
141
Fish
Good habitat for the bay goby is shallow subtidal areas
with mud or a mud/sand mixture and possibly intertidal
mudflats. The presence of burrowing invertebrates,
which may serve as commensal hosts, would be beneficial. There is no evidence that this species utilizes tidal
marshes in San Francisco Bay or elsewhere in its range.
Eldridge, M.B. and C.F. Bryan. 1972. Larval fish survey of Humboldt Bay, California. NOAA Tech.
Rept. NMFS SSRF-665, 8 pp.
Flemming, K. 1999. Gobidae. In: J.Orsi (ed). Report
on 1980-1995 fish, shrimp, and crab sampling in
the San Francisco Estuary. IEP Tech. Rept. 63.
Grossman, G.D. 1979a. Symbiotic burrow-occupying
behavior in the bay goby, Lepidogobius lepidus. Ca.
Fish and Game 65(2):122-124.
______. 1979b. Demographic characteristics of an intertidal bay goby (Lepidogobius lepidus). Environmental Biology of Fishes 4(3): 207-218.
Grossman, G.D., R. Coffin and P.B. Moyle. 1980. Feeding ecology of the bay goby (Pisces:Gobiidae). Effects of behavioral, ontogenetic, and temporal variation on diet. J. Exp. Mar. Biol. Ecol. 44:47-59.
Hieb, K. 1999. San Francisco Bay species abundance
(1980-1998). IEP Mewsletter, Vol. 12(2): 30-34.
Hieb, K. and R. Baxter. 1993. Delta Outflow/San Francisco Bay Study. In: 1991 Annual Report, Interagency Ecological Studies Program for the Sacramento-San Joaquin Estuary, pp. 101-116
Pearcy, W.G. and S.S. Myers. 1974. Larval fishes of
Yaquina Bay, Oregon: A nursery ground for marine fishes? Fishery Bulletin 72(1):201-213.
Miller, D.J. and R.N. Lea. 1976. Guide to the coastal
marine fishes of California. Ca. Dept. Fish and
Game Fish Bull. 157, 249 pp.
Orsi, J. (ed). 1999 Report on the 1980-1995 fish,
shrimp, and crab sampling in the San Francisco
Estuary, California. IEP Tech. Rept. No. 63.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin estuary and adjacent waters, California: A
guide to the early life histories. Interagency Ecological Studies Program for the Sacramento-San
Joaquin Estuary Tech. Rept. 9.
Wild, P.W. 1969. Macrofauna of Plummer Creek of San
Francisco Bay collected by a specially designed trap.
M.A. Thesis, San Jose State University, San Jose,
Ca.. 85 pp.
Woods, E. 1981. Fish Utilization. In: T. Niesen and M.
Josselyn (eds). The Hayward Regional Shoreline
marsh restorations: Biological succession during the
first year following dike removal. Tiburon Center
for Env. Studies, Tech. Rept. 1, pp 35-46.
Longjaw Mudsucker
Gillichthys mirabilis
Kathryn A. Hieb
The longjaw mudsucker (Family: Gobiidae) is the largest goby native to San Francisco Bay, reaching a size of
200 mm total length (TL). It ranges from Baja California to Tomales Bay (Miller and Lea 1972) and was successfully introduced to the Salton Sea in 1930 (Walker
et al. 1961). The longjaw mudsucker is a common resident of mudflats and sloughs in estuaries and coastal
streams. It is also common in salt ponds, as it can tolerate a wide range of salinities. As the tide ebbs, the
longjaw mudsucker retreats to burrows or buries in the
mud rather than migrate to deeper water. Due to their
ability to live out of water and in freshwater for several
days, mudsuckers or “ mud puppies” are a sought after
bait-fish; however, in recent years, the San Francisco Bay
area bait fishery has targeted the yellowfin goby, a large
introduced species that is very common in many shallow water habitats.
Reproduction
CDFG
Male longjaw mudsuckers construct burrows for breeding, which they aggressively guard until the eggs hatch.
A single female lays 4,000 to 9,000 eggs, depending on
size (Weisel 1947). In southern California, spawning
occurs from January through July, with peak activity
apparently from February through April (Weisel 1947).
In South Bay salt ponds, the spawning period is also protracted, occurring from November through June, with
peak activity in February and March (Lonzarich 1989).
Gonadal regression occurs from July to September, when
temperatures in the salt ponds reach their maximum (de
Vlaming 1972). Females were reported to spawn more
than once per season in South Bay salt ponds (de
Vlaming 1972) and two and possibly three times per
season in the Salton Sea (Walker et al. 1961), with an
interval of 40 to 50 days between spawnings (Barlow
1963). Ovarian development and spawning are asynchronous, which is typical of species that spawn more than
once per season and have a protracted spawning season
(de Vlaming 1972).
142
Baylands Ecosystem Species and Community Profiles
Growth and Development
In South Bay salt ponds, longjaw mudsuckers grow to
80-100 mm standard length (SL) by the end of year one
and 120-140 mm SL by the end of year two (Lonzarich
1989). Few live more than one year and none more than
two years; both sexes mature at age one (Barlow 1963,
Lonzarich 1989). In the Salton Sea, longjaw mudsuckers hatched in the early spring reach 60-80 mm SL by
fall and 80-120 mm SL by the next spring (Walker et
al. 1961).
Food and Feeding
In Elkhorn Slough, California, the longjaw mudsucker
preys primarily on gammarid amphipods, especially
Orchestia traskiana, Eogammarus confervicolus, Corophium
spp., and polychaetes (Barry 1983). Dipterans, harpacticoid copepods, and grapsid crabs (primarily Hemigrapsus oregonensis) are also important food items. In South
San Francisco Bay salt ponds, longjaw mudsucker diet
varies by salinity—in the lower salinity (20-40‰ ) ponds,
they consume primarily polychaetes and amphipods
while in the higher salinity (to 84‰ ) ponds they consume primarily brine shrimp and waterboatmen (Lonzarich 1989). Copepods are an important prey item in
the winter, when brine shrimp are unavailable.
Distribution
In San Francisco Bay, the longjaw mudsucker has been
collected in South, Central, San Pablo, and Suisun bays,
although it is not common upstream of Carquinez Strait.
It is the least common goby collected in trawl surveys of
open water habitats and larger channels, but usually the
most common goby collected in smaller marsh channels.
For example, it was not collected in trawls near Castro
Creek, Corte Madera Creek, and Gallinas Creek
marshes, but was the most abundant goby and third
most abundant species collected in minnow traps set in
the marsh channels (CH2M Hill 1982). Similarly, in a
study of a restored marsh near Hayward, it was not common in trawls of the larger channels, but the only goby
and most common species collected in minnow traps set
on the mudflats (Woods 1981). It was also the second
most common species collected in first and second order channels of tidal marshes in lower Petaluma River
(CDFG, unpub. data). This distribution has also been
Plants
Amphibians &
Reptiles
Fish
General Information
The eggs are club shaped, 2.8-3.4 mm long, with
an adhesive thread at one pole that attaches to the burrow wall. Hatching occurs in 10 to 12 days at 18° C
(Weisel 1947). Larvae have been collected year-round in
the Bay, with peak abundance in May and June (CDFG,
unpub. data); in South Bay salt ponds, larvae were collected at salinities up to 70‰ (Lonzarich 1989).
Population Status and Influencing Factors
There is no survey which routinely samples the longjaw
mudsucker or its preferred habitat in San Francisco Bay,
so the current status of the population cannot be assessed. With the introduction and establishment of the
yellowfin goby in the 1960s, the longjaw mudsucker is
no longer as sought after for bait. However, the introduction of the yellowfin goby may have had a negative
impact on the longjaw mudsucker, as there is substantial overlap in the habitats of the two species.
Trophic Levels
The longjaw mudsucker is a secondary consumer.
Proximal Species
Predators: Bait fishers and possibly great blue herons,
egrets, and larger shorebirds.
Prey: Gammarid amphipods, polychaetes, dipterans,
copepods, Hemigrapsus oregonensis, waterboatmen, brine
shrimp.
Good Habitat
The intertidal area of tidal marsh channels is the typical
habitat of the longjaw mudsucker. Because this species
can tolerate a wide range of environmental conditions,
“ good habitat” is probably defined by the complexity of
these sloughs. More complex channels, with undercut
banks and pools of water at low tide, would offer more
protection from predators than sloughs with little incision and ponded water. These more complex channels
are typical of mature marshes vs. recently “ restored”
marshes.
References
Barlow, G.W. 1963. Species structure of the gobiid fish
Gillichthys mirabilis from coastal sloughs of the eastern Pacific. Pacific Science 17:47-72.
Barry, J.P. 1983. Utilization of shallow marsh habitats
by fishes in Elkhorn Slough, California. M.A. Thesis, San Jose State Univ., San Jose, Ca., 91pp.
Carpelan, L.H. 1957. Hydrobiology of the Alviso salt
ponds. Ecology 38(3): 375-390.
CH2M Hill. 1982. Equivalent protection study intensive investigation. Prepared for Chevron USA. Final Report, April 1982.
Courtois, L.A. 1973. The effects of temperature, availability of oxygen, and salinity upon the metabolism of the longjaw mudsucker, Gillichthys mirabilis.
M.A. Thesis, Ca. State Univ., Hayward, Ca., 32 pp.
de Vlaming, V.L. 1971. Thermal selection behavior in
the estuarine goby, Gillichthys mirabilis Cooper.
J. Fish Biology 1971(3):277-286.
______. 1972. Reproduction cycling in the estuarine
gobiid fish, Gillichthys mirabilis. Copeia 1972(2):
278-291.
Lonzarich, D.G. 1989. Temporal and spatial variations
in salt pond environments and implications for fish
and invertebrates. M.A. Thesis, San Jose State
Univ., San Jose, Ca. 81 pp.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Dept. Fish and
Game, Fish Bull. 157, 249 pp.
Todd, E.S. and W. Ebeling. 1966. Aerial respiration in
the longjaw mudsucker Gillichthys mirabilis
(Telostei:Gobiidae). Biology Laboratory, Woods
Hole 130:265-288.
Walker, B.W., R.R. Whitney and G.W. Barlow. 1961.
The fishes of the Salton Sea. In: B.W. Walker (ed).
The Ecology of the Salton Sea, California, in relation to the sportfishery. Ca. Dept. Fish and Game,
Fish Bull. 133:77-91.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
143
Fish
reported from Elkhorn Slough, where the longjaw mudsucker was not an important component of the otter
trawl samples from deeper (>1.5 m) channels, but was
the third most abundant species and most common goby
in beach seine and channel net samples from shallower
(<1.5 m) channels (Barry 1983).
The longjaw mudsucker is also common in salt
ponds in San Francisco Bay. It was the most common
goby and the second most common fish collected in
South Bay salt ponds (Carpelan 1957, Lonzarich 1989).
Lonzarich (1989) reported highest catches in the summer and fall.
Although longjaw mudsucker can tolerate a wide
range of salinities, they are usually absent from fresh or
slightly brackish water (Barlow 1963). They have been
collected from salinities as high as 82.5‰ in the upper
Gulf of California (Barlow 1963), and as high as 84‰
in South Bay salt ponds (Lonzarich 1989).
Although longjaw mudsuckers have been collected
at temperatures as high as 33° C (Carpelan 1957), in
laboratory thermal selection studies, they preferred temperatures from 9-23° C and strongly avoided temperatures greater than 23° C (de Vlaming 1971). In another
laboratory study, Courtois (1973) concluded that the
longjaw mudsucker was best adapted to temperatures
between 20 and 30° C.
In intertidal areas, the longjaw mudsucker often
remains in the mud or burrows at low tide and is subject to fluctuating oxygen concentrations. The jaw membranes are richly vascularized and serve as an accessory
respiratory apparatus (Weisel 1947). Additionally, the
longjaw mudsucker will respire aerially at low (<2.0 mg/
L) oxygen concentrations; they gulp air at the water surface and hold the bubbles in their large buccopharyngeal
cavity (Todd and Ebeling 1966).
Fish
Weisel, G.F.J. 1947. Breeding behavior and early development of the mudsucker, a gobiid fish of California. Copeia 2:77-85.
Woods, E. 1981. Fish Utilization. In: T. Niesen and M.
Josselyn (eds). The Hayward Regional Shoreline
marsh restorations: Biological succession during the
first year following dike removal. Tiburon Center
for Env. Studies, Tech. Rept. 1: 35-46.
California Halibut
Paralichthys californicus
Michael K. Saiki
The California halibut (Family: Bothidae) is a large marine flatfish that is sought after in the market place because of its large size and excellent taste (Frey 1971).
Commercial fishing for California halibut was historically centered in the Baja California-Los Angeles area,
but has recently shifted northward to the Santa Barbara
region (Barsky 1990). It is harvested by gill net, trammel net, and trawl nets (Schultze 1986). California commercial fishermen landed an average of 534 tons per year
from 1983 to 1987, and received $0.64-$1.59/kg in
1987 (CDFG 1988). California halibut is also highly
prized by recreational anglers and is caught primarily
from piers and boats using hook, line, and live bait. Over
916,000 California halibut were caught by anglers in
1985 (USDC 1986).
Reproduction
CDFG
Emmett et al. (1991) described the California halibut
as being gonochoristic (its gender is determined by developmental rather than hereditary mechanisms) and
iteroparous (it has the capacity to survive and spawn beyond one or multiple spawning seasons). It is a broadcast spawner whose eggs are fertilized externally (Emmett
et al. 1991).
The eggs of California halibut are pelagic (Allen
1988). In a laboratory tank with water depth of 2-3 m,
144
Baylands Ecosystem Species and Community Profiles
Growth and Development
California halibut eggs are spherical in shape and 0.740.84 mm in diameter (Ahlstrom et al. 1984). Eggs hatch
approximately two days after fertilization at 16° C (Emmett et al. 1991). Newly hatched larvae of California
halibut measure about 2.0 mm total length (TL) (Ahlstrom
and Moser 1975, Ahlstrom et al. 1984). The larval yolk
sac is depleted about six days after hatching (Gadomski
and Petersen 1988).
Metamorphosis occurs at a length of 7.5-9.4 mm
(Ahlstrom et al. 1984) when the pelagic, bilaterally symmetrical larvae become benthic, asymmetrical juveniles.
Along with other physical changes, the most visible part
of this process is a change in pigmentation patterns and
the migration of one eye across the top of the head to
its final resting place close to the other eye (Moyle and
Cech 1988).
Temperature has a major effect on survival of eggs
and larvae of the California halibut. Successful hatching occurred at 12° , 16° , and 20° C, but death occurred
prior to embryo formation at 8° and 24° C (Gadomski
and Caddell 1991). At 17 days posthatch, all larvae died
at 12° C, whereas survival varied from 23% to 46% at
16° , 20° , and 24° C. The survival of older larval stages
of California halibut progressively increased as incubation temperatures rose from 16° C to 28° C. Temperature also affected the settlement rate of juveniles that had
just completed metamorphosis.
Although juveniles are reported to vary in length
from 8 mm to 430 mm (Emmett et al. 1991), males can
mature at 200-300 mm standard length (SL) when 2-3
years old whereas females can mature at 380-430 mm
SL when 4-5 years old (Fitch 1965, Fitch and Lavenberg
Plants
Amphibians &
Reptiles
General Information
California halibut spawned while swimming near the
water surface (Allen 1990). Adults typically move into
shallow (6-20 m deep) coastal waters in early spring and
usually spawn over sandy substrates (Ginsburg 1952,
Frey 1971, Feder et al. 1974, Haaker 1975). Spawning
occurs from February through August, peaking in May
with a great number of mature fish (Frey 1971, Feder
et al. 1974, Wang 1986). Spawning most often occurs
when water temperatures are 15.0-16.5° C, and day
lengths are greater than or equal to 10.5 hours (Caddell
et al. 1990). However, abundant eggs and larvae have
also been reported from nearshore coastal waters during
winter-spring when surface temperatures are 13-15° C,
and even during summer when surface waters occasionally reach 22° C (Lavenberg et al. 1986, Petersen et al.
1986).
During the spawning season, small (55.9-61.0 cm
long) California halibut produce approximately 300,000
eggs every 7 days, whereas large (>114.3 cm long) halibut produce about 1 million eggs per day (Emmett et
al. 1991).
1971, Haaker 1975). California halibut may reach a
maximum length of 1,520 mm and a maximum weight
of 33 kg (Eschmeyer et al. 1983), with certain individuals
living for as long as 30 years (Frey 1971).
Food and Feeding
Distribution
The geographic distribution of California halibut extends
from the Quillayute River, Washington, southward to
Magdalena Bay, Baja California (Ginsburg 1952, Miller
and Lea 1972, Eschmeyer et al. 1983). However, it is
common only in bays and estuaries south of Tomales
Bay, California, and reaches peak abundance in estuaries south of Point Conception (Emmett et al. 1991).
Recently, large numbers of mostly female California
halibut were caught by recreational anglers in Humboldt
Bay, with some caught as far north as Crescent City and
southern Oregon (R. Baxter, pers. comm.). A survey of
carcasses suggested that the females had not developed
mature eggs.
Larvae of California halibut occur primarily in the
upper 30 m of coastal waters, where they apparently
settle or migrate from the 0-10 m stratum to the 10-20 m
stratum at night (Moser and Watson 1990). Conversely,
larvae over shallow water (13 m bottom depth) tend to
move downward during the day (Barnett et al. 1984).
Juveniles settle in shallow water on the open coast, but
are more abundant in bays (Allen 1988, Moser and
Watson 1990). Juveniles remain in bays for about two
years until they emigrate to the coast where they settle
at water depths less than 100 m, with greatest abundance
at depths less than 30 m (Miller and Lea 1972, Allen
Population Status and Influencing Factors
Catch records indicate that the abundance of California
halibut within its historic range was high in the late
1960s, declined in the 1970s, and increased in the 1980s.
The intense El Niño in 1982-83 coincided with higher
abundance and landings of halibut (Jow 1990). Overall, however, California halibut populations seem to be
undergoing a long-term decline. This decline may be related to large-scale changes in the marine environment,
overfishing, alterations and destruction of estuarine habitat, or a shift in location of population centers (Plummer
et al. 1983). Pollution has been shown to reduce hatching success, reduce size of larvae at hatching, produce
morphological and anatomical abnormalities, and reduce
feeding and growth rates (MBC Applied Environmental Sciences 1987). By comparison, thermal effluents
from California coastal power plants do not seemingly
inhibit growth and may be advantageous to California
halibut (Innis 1990).
Early records indicate that California halibut were
uncommon in San Francisco Bay. Alpin (1967) sampled
the Central Bay with bottom trawls during 1963-1966
and reported catching only three California halibut (two
in the spring and one in June). Ganssle (1966) reported
catching only two adult California halibut (May 1963,
1964) while fishing bottom trawls in San Pablo Bay.
Recently, consistent high salinities probably have contributed to increased abundance of California halibut in
the bay. Moreover, recent data suggest that successful
year classes in 1983, 1987, and 1990 have contributed
to increased abundance in the bay (CDWR 1991). These
were years with warm water ocean events, and it is hypothesized that California halibut abundance in the San
Francisco Bay increased because of increased local
Chapter 2 —
Estuarine Fish and Associated Invertebrates
145
Fish
California halibut feed initially on small invertebrates,
but later switch almost exclusively to feeding on fish
(Haaker 1975). Although the diet of larvae has not been
examined, they probably feed on tiny planktonic organisms (Allen 1990). Small juveniles in three southern California embayments fed mostly on harpacticoid copepods and gammaridean amphipods, with some polychaetes, mysids, small fish, and crab megalopae also being
taken (Haaker 1975, Allen 1988). In Anaheim Bay, California, large juveniles and small adults ate bay shrimp,
topsmelt, California killifish, and gobies, whereas subadults and adults more than 23.0 cm SL consumed
mostly northern anchovy, croaker, and other larger fishes
(Haaker 1975). Other forage taxa in the diets included
ostracods and acteonid snails. In Tomales Bay, adult
California halibut (65.4-83.3 cm SL) fed on Pacific
saury, Pacific herring, sanddabs, white sea perch, and
California market squid (Bane and Bane 1971). The
California halibut is an ambush predator (Haaker 1975).
During foraging it lies partially buried on the sandy
bottom and waits until its prey is close enough to seize.
1982). Larger juveniles (greater than 20 mm in length)
may move from open coastal areas to resettle in bays
(Kramer 1990).
Adults move inshore during spring and summer,
and offshore during winter (Ginsburg 1952, Haaker
1975). Although the inshore movements are associated
with spawning, they may also be influenced by seasonal
patterns in forage fish abundance. For example, during
spring and summer, California grunion (Leuresthes
tenuis) are abundant near the surf zone (Feder et al.
1974), whereas northern anchovy (Engraulis mordax) are
abundant in bays and estuaries (Tupen 1990).
California halibut are occasionally found in Central and South San Francisco Bay (Alpin 1967, Pearson
1989) and San Pablo Bay (Ganssle 1966). Recently, eggs
of a description similar to those of California halibut
were collected in San Francisco Bay; however, their identity was not verified (Wang 1986). Both larval and juvenile California halibut have been captured in San Francisco and San Pablo bays (Wang 1986).
Fish
spawning, higher survival of larvae, or migration of juveniles from more southern coastal areas with warmer
ocean waters (Hieb and Baxter 1994).
Abundance indices (determined from trawl samples)
for California halibut in San Francisco Bay increased
from 1989 to 1992 (Hieb and Baxter 1994). The 1992
index was the highest since the study began in 1980.
Also, most halibut collected in San Francisco Bay are age
two and older, whereas other flatfishes are caught primarily as young-of-the-year. Nevertheless, California
halibut abundance indices are still very low relative to
other common species of flatfish in the Bay (Hieb and
Baxter 1994).
In an attempt to increase California halibut numbers, natural production has been augmented by hatchery production (Crooke and Taucher 1988). Although
this effort could increase future recruitment, negative
effects of the hatchery program include a possible reduction in genetic variability within natural populations and
the high cost producing fish (Hobbs et al.1990).
shallow water where temperatures approximate 13-15°C,
although successful spawning may also occur at temperatures approaching 22° C (Gadomski and Caddell 1991).
Favorable characteristics for bays and estuaries that serve
as nursery areas include productive habitats with abundant food supplies and shallow areas that allow juveniles
to avoid predators, including adult halibut (Plummer et
al. 1983). Juveniles and adults prefer sandy bottoms and
water temperatures between 10-25° C, with a preference
for 20.8° C (Ehrlich et al. 1979). Juveniles are relatively
tolerant of reduced dissolved oxygen and increased water temperatures (Waggoner and Feldmeth 1971).
Higher water temperatures induces faster growth rates
and decreases the time to settlement for most young-ofthe-year halibut (Gadomski et al.1990). Eggs, larvae, and
adults are found in euhaline waters, but juveniles often
occur in oligohaline to euhaline conditions (Haaker
1975).
Trophic Levels
Ahlstrom, E.H. and H.G. Moser. 1975. Distributional
atlas of fish larvae in the California current region:
flatfishes, 1955 through 1960. Ca. Coop. Ocean.
Fish Invest., Atlas No. 23, 207 pp.
Ahlstrom, E.H., K. Amaoka, D.A. Hensley, H.G. Moser
and B.Y. Sumida. 1984. Pleuronectiformes; development. In: H.G. Moser (chief ed). Ontogeny and
systematics of fishes, p. 640-670. Allen Press, Inc.,
Lawrence, KS.
Allen, L.G. 1988. Recruitment, distribution, and feeding habits of young-of-the-year California halibut
(Paralichthys californicus) in the vicinity of Alamitos
Bay-Long Beach Harbor, California, 1983-1985.
Bull. Southern Ca. Acad. Sci. 87:19-30.
______. 1990. Open coast settlement and distribution
of young-of-the-year California halibut, Paralichthys
californicus, along the southern California coast
between Point Conception and San Mateo Point,
June-October, 1988. In: C.W. Haugen (ed). The
California halibut, Paralichthys californicus, resource
and fisheries. Ca. Dept. Fish and Game, Fish Bull.
174: 145-152.
Allen, M.J. 1982. Functional structure of soft-bottom
fish communities of the Southern California shelf.
Ph.D. Diss., Univ. Ca., San Diego, CA. 577 pp.
______. 1990. The biological environment of the California halibut, Paralichthys californicus. In: C.W.
Haugen (ed). The California halibut, Paralichthys
californicus, resource and fisheries. Ca. Dept. Fish
and Game, Fish Bull. 174.
Alpin, J.A. 1967. Biological survey of San Francisco Bay,
1963-1966. Ca. Dept. Fish and Game. MRO Ref.
67-4. 131 pp.
Bane, G.W. and A.W. Bane. 1971. Bay fishes of northern
California. Mariscos Publ., Hampton Bays, NY. 143 pp.
Proximal Species
Predators: Thornback (important predator on settling
juveniles), California sea lions (predator on large juveniles and adults), northern sea lions, Pacific angel shark,
Pacific electric, bottlenose dolphin.
Prey:
Plankton—major prey item for larvae.
Harpacticoid copepods, gammaridean amphipods—major prey item for young juveniles.
Polychaetes, mysids, and crab—minor prey item for
young juveniles.
Mysids—major prey item for juveniles.
Gobies—prey item for juveniles and adults.
Bay shrimp, ghost shrimp—prey item for older juveniles.
Topsmelt, California killifish—prey item for older juveniles and adults.
Northern anchovy—major prey item for adults.
White croakers, hornyhead turbot—prey item for large
adults.
Octopus, squid, California grunion—prey item for
adults.
Parasites: Trematodes, cestodes, and nematodes (endoparasites); copepods and isopods (ectoparasite).
Competitors: Speckled sanddab (potentially important).
Good Habitat
Good spawning habitat for California halibut is limited
to inshore waters or bays and estuaries in moderately
146
Baylands Ecosystem Species and Community Profiles
Plants
Amphibians &
Reptiles
Larvae, juveniles, and adults are carnivorous (secondary
and higher order consumers).
References
nia halibut, Paralichthys californicus. Fishery Bull.
89:567-576.
Gadomski, D.M., S.M. Caddell, L.R. Abbott and T.C.
Caro. 1990. Growth and development of larval and
juvenile California halibut, Paralichthys californicus, reared in the laboratory. In: C.W. Haugen (ed).
The California halibut, Paralichthys californicus,
resource and fisheries. Ca. Dept. Fish and Game,
Fish Bull. 174:85-98.
Gadomski, D.M. and J.H. Petersen. 1988. Effects of
food deprivation on the larvae of two flatfishes.
Mar. Ecol. Prog. Ser. 44:103-111.
Ganssle, D. 1966. Fishes and decapods of San Pablo
and Suisun bays. In: D.W. Kelley (ed). Ecological
studies of the Sacramento-San Joaquin Estuary, Part
I. Ca. Dept. Fish and Game, Fish Bull. 133:64-94.
Ginsburg, I. 1952. Flounders of the genus Paralichthys
and related genera in American waters. Fish Bull.,
U.S. 71:1-351.
Haaker, P.L. 1975. The biology of the California halibut, Paralichthys californicus (Ayres), in Anaheim
Bay, California. In: E.D. Lane and C.W. Hill (eds).
The marine resources of Anaheim Bay. Ca. Fish
and Game, Fish Bull. 165:137-151.
Hieb, K. and R. Baxter. 1994. Interagency ecological
studies program 1992 annual report for the Sacramento San Joaquin estuary (P.L. Herrgesell,
comp.). Ca. Dept Fish and Game, Ca. Dept. of
Water Res., U.S. Bureau Reclam., U.S. Fish and
Wildl. Serv., p. 95-106.
Hobbs, R.C., L.W. Botsford and R.G. Kope. 1990.
Bioeconomics evaluation of the culture/stocking
concept for California halibut. In: C.W. Haugen,
editor. The California halibut, Paralichthys californicus, resource and fisheries. Ca. Dept. Fish and
Game, Fish Bull. 174.
Innis, D.B. 1990. Juvenile California halibut,
Paralichthys californicus, growth in relation to thermal effluent. In: C.W. Haugen (ed). The California halibut, Paralichthys californicus, resource and
fisheries. Ca. Dept. Fish and Game, Fish Bull.
174:153-165.
Jow, T. 1990. The California halibut trawl fishery. In:
C.W. Haugen (ed). The California halibut,
Paralichthys californicus, resource and fisheries. Ca.
Dept. Fish and Game, Fish Bull. 174.
Kramer, S.H. 1990. Distribution and abundance of juvenile California halibut, Paralichthys californicus,
in shallow water of San Diego county. In: C.W.
Haugen (ed). The California halibut, Paralichthys
californicus, resource and fisheries. Ca. Dept. Fish
and Game, Fish Bull. 174.
Lavenberg, R.J., G.E. McGowen, A.E. Jahn, J.H. Petersen
and T.C. Sciarrotta. 1986. Abundance of southern
California nearshore ichthyoplankton: 1978-1984.
Ca. Coop. Oceanic Fish. Invest. Rep. 27:53-64.
Chapter 2 —
Estuarine Fish and Associated Invertebrates
147
Fish
Barnett, A.M., A.E. Jahn, P.D. Sertic and W. Watson.
1984. Distribution of ichthyoplankton off San
Onofre, California, and methods for sampling very
shallow coastal waters. Fish. Bull., U.S. 82:97-111.
Barsky, K.C. 1990. History of the commercial California halibut fishery. In: C.W. Haugen (ed). The
California halibut, Paralichthys californicus, resource
and fisheries. Ca. Dept. Fish and Game Fish Bull.,
No. 174: 217-227.
Caddell, S.M., D.M. Gadomski and L.R. Abbott. 1990.
Induced spawning of the California halibut,
Paralichthys californicus, (Pisces: Paralichthyidae)
under artificial and natural conditions. In: C.W.
Haugen (ed). The California halibut, Paralichthys
californicus, resource and fisheries. Ca. Dept. Fish
and Game, Fish Bull. 174.
California Department of Fish and Game (CDFG).
1988. Review of some California fisheries for 1987.
Ca. Dept. Fish and Game. Ca. Coop. Ocean. Fish.
Invest. Rep. 29:11-20.
California Department of Water Resources (CDWR).
1991. Interagency ecological studies program 1991
annual report for the Sacramento San Joaquin estuary (P.L. Herrgesell, comp.). Ca. Dept Fish and
Game, Ca. Dept. of Water Res., U.S. Bureau
Reclam., U.S. Fish & Wildl. Serv. p.104-105.
Crooke, G.A. and C. Taucher. 1988. Ocean hatcheries—wave of the future? Outdoor Ca.. 49(3):10-13.
Ehrlich, K.F., J.H. Hood, S. Muszynski and G.E.
McGowen. 1979. Thermal behavior responses of
selected California littoral fishes. Fish Bull., U.S.
76(4):837-849.
Emmett, R.L., S.L. Stone, S.A. Hinton, and M.E. Monaco. 1991. Distribution and abundance of fishes
and invertebrates in west coast estuaries, Vol. II:
species life history summaries. ELMR Rep. No. 8.
NOAA/NOS Strategic Environmental Assessments
Div., Rockville, MD, p. 250-255.
Eschmeyer, W.N., E.S. Herald and H. Hammann. 1983.
A field guide to Pacific coast fishes of North America.
Houghton Mifflin Co., Boston, MA, 336 pp.
Feder, H.M., C.H. Turner and C. Limbaugh. 1974.
Observations on fishes associated with kelp beds
in southern California. Ca. Fish and Game, Fish
Bull. 160:1-144.
Fitch, J.E. 1965. Offshore fishes of California. 3rd revision. Ca. Dept. Fish and Game, Sacramento, CA.
80 pp.
Fitch, J.E. and R.J. Lavenberg. 1971. Marine food and
game fishes of California. Univ. Ca. Press, Berkeley, CA. 179 p.
Frey, H.W. 1971. California’s living marine resources
and their utilization. Ca. Dept. Fish and Game,
Sacramento, CA, 148 pp.
Gadomski, D.M. and S.M. Caddell. 1991. Effects of
temperature on early life-history-stages of Califor-
148
Baylands Ecosystem Species and Community Profiles
Fish
Amphibians &
Reptiles
Personal Communications
Randal Baxter, California Department of Fish and
Game, Stockton.
Starry Flounder
Platichthys stellatus
Kurt F. Kline
General Information
The starry flounder is in the family Pleuronectidae, or
right-eyed flounders. Pleuronectids are generally found
in temperate marine environments, with only a few species found in the tropics or sub-tropics. There are 22
species found along the coast of California. The starry
flounder is one of the few pleuronectids commonly
found in brackish and freshwater (Orcutt 1950, Haertel
and Osterberg 1967). While placed in the
Pleuronectidae, the starry flounder is commonly right or
left-eyed. However, it is quite distinguishable from other
flatfishes due to the alternating dark gray and
orange-yellow bands on the dorsal, anal, and caudal fins.
Many of the pleuronectids support commercial and
sport fisheries. The starry flounder is a minor sport species in San Francisco Bay and most fish are taken from
boats when fishing for California halibut, sturgeon, or
striped bass. It common in the commercial fishery, but
as a by-catch to targeted species such as petrale sole and
California halibut. In recent years, nearshore gear restrictions have resulted in a decrease in starry flounder landings, as this species is most common within a few miles
of shore (Haugen 1992).
Reproduction
Spawning occurs in winter in shallow coastal areas near
the mouths of rivers and sloughs (Orcutt 1950, Wang
1986, Baxter 1999). Some researchers have suggested
that spawning may occur within San Francisco Bay
Plants
MBC Applied Environmental Sciences. 1987. Ecology
of important fisheries species offshore California.
Rep. to Min. Manag. Serv., U.S. Dept. Int., Washington, D.C., 251 pp. (Contract No. MMS 1412-0001-30294).
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Fish and Game,
Fish Bull. No. 157, 249 p.
Moser, H.G. and W. Watson. 1990. Distribution and
abundance of early life history stages of the California halibut, Paralichthys californicus, and comparison with the fantail sole, Xystreurys liolepis. In:
C.W. Haugen (ed). The California halibut,
Paralichthys californicus, resource and fisheries. Ca.
Dept. Fish and Game, Fish Bull. 174.
Moyle, P.B. and J.J. Cech, Jr. 1988. Fishes, an introduction to ichthyology, 2nd ed., p. 312. Prentice-Hall,
Inc., Englewood Cliffs, NJ.
Pearson, D.E. 1989. Survey of fishes and water properties of South San Francisco Bay, California, 197382. NOAA Tech. Report NMFS 78. 21 pp.
Petersen, J.H., A.E. Jahn, R.J. Lavenberg, G.E.
McGowen and R.S. Grove. 1986. Physical-chemical characteristics and zooplankton biomass on the
continental shelf off southern California. Ca. Coop.
Oceanic Fish. Invest. Rep. 27:36-52.
Plummer, K.M., E.E. DeMartini, and D.A. Roberts.
1983. The feeding habits and distribution of juvenile-small adult California halibut (Paralichthys californicus) in coastal waters off northern San Diego
county. Calif Coop. Ocean Fish. Invest. Rep.
24:194-201.
Schultze, D.L. 1986. Digest of California commercial
fish laws, January 1, 1986. Ca. Dept. Fish and
Game, Sacramento, CA, 40 pp.
Tupen, J.W. 1990. Movement and growth of tagged
California halibut, Paralichthys californicus, off the
central coast of California. In: C.W. Haugen (ed).
The California halibut, Paralichthys californicus,
resource and fisheries. Ca. Dept. Fish and Game,
Fish Bull. 174.
U.S. Dept. of Commerce (USDC). 1986. Marine recreational fishery statistics survey, Pacific coast. U.S.
Dept. Comm., Nat. Ocean. Atm. Adm., Current
Fish. Stat. No. 8328, 109 p.
Waggoner, J.P., III, and C.R. Feldmeth. 1971. Sequential mortality of the fish fauna impounded in construction of a marina at Dana Point, CA. Ca. Dept
Fish and Game 57(3):167-176.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and adjacent waters, California: a
guide to the early life histories. Tech. Rep. No. 9,
prepared for the Interagency Ecological Study Program for the Sac.-San Joaquin Estuary. Ca. Dept.
Water Res., Ca. Dept. Fish and Game, U.S. Bureau Reclam. U.S. Fish Wildl. Serv.
(Radtke 1966, Moyle 1976); however, neither ripe female starry flounder nor mature flounder eggs or
pre-flexion larvae were collected from San Francisco Bay
in the early 1980s (B. Spies, pers. comm., Wang 1986).
Growth and Development
Food and Feeding
Population Status and Influencing Factors
There is evidence of a long-term decline in the San Francisco Bay starry flounder population from the Commercial Passenger Fishing Vessel log book data. Both catch/
hour (CPUE) and total catch of starry flounder declined
in the mid-1970s from a peak in the late 1960s and early
1970s (CDFG 1992). This decline in CPUE and catch continued at least through the early 1990s. Additionally, juvenile starry flounder abundance indices from San Francisco
Bay steadily declined from the early to the late 1980s (Figure 2.10). Abundance remained very low through 1994 and
increased somewhat from 1995-99. Outflow related mechanisms have been proposed to control recruitment of age-0
starry flounder to the Bay (CDFG 1992, Hieb and Baxter
1993). The increase in the abundance of age-0 fish from
1995 to 1999 supports this hypothesis.
Hydrologic factors and other environmental conditions in San Pablo and Suisun bays are important in
determining the distribution of juvenile starry flounder.
The San Francisco Estuary is close to the southern limit
of the distribution for starry flounder and long-term
In Monterey Bay and Elkhorn Slough, the smallest starry
flounder (10-99 mm SL) fed primarily on copepods and
amphipods. Larger juveniles (100-199 mm SL) fed on
larger amphipods, polychaetes, and bivalves (especially
siphon tips). Fish >199 SL mm fed on whole crabs and
bivalves, sand dollars, brittle stars, and occasionally fish
(Orcutt 1950). In San Francisco Bay, a large portion of
the diet of starry flounders > 199 mm was bivalves (primarily Mya, Ischadium, Tapes, Solen, Mytilus, and
Gemma), polychaetes, and crustaceans (especially
Upogebia, Cancer magister, C. gracilis, and Hemigraphsus
oregonensis) (CDFG, unpubl. data).
Distribution
Starry flounder range from Santa Barbara, California
northward to arctic Alaska, then southwesterly to the Sea
of Japan (Miller and Lea 1972). Adult starry flounder
inhabit shallow coastal marine water, whereas juveniles
rear in bays and estuaries (Orcutt 1950, Moyle 1976,
Wang 1986). Emmett et al. (1991) state that juvenile
starry flounder are found almost exclusively in estuaries.
Figure 2.10 Annual Abundance Indices of Starry
Flounder: A. Age-0, May-October; B. Age-1,
February-October (CDFG Otter Trawl data)
Chapter 2 —
Estuarine Fish and Associated Invertebrates
149
Fish
Eggs and larvae are pelagic and found mostly in the upper
water column (Orcutt 1950, Wang 1986). Starry flounder larvae are approximately 2 mm long at hatching and
settle to the bottom about two months after hatching,
at approximately 7 mm standard length (SL) (Policansky
and Sieswerda 1979, Policansky 1982). Larvae depend
upon favorable ocean currents to keep them near their
estuarine nursery areas before settlement. Transforming
larvae and juveniles migrate from the coast to brackish
or freshwater nursery areas, where they rear for 1 or more
years (Haertel and Osterberg 1967, Wang 1986, Hieb
and Baxter 1993). As they grow, juvenile starry flounder move to higher salinity, but appear to remain in estuaries through at least their second year of life (Haertel
and Osterberg 1967, Hieb and Baxter 1993).
Most males mature by the end of their second year
of life (220-276 mm SL), while females mature at 3 or
4 (239-405 mm SL) (Orcutt 1950). During the late fall
and winter, mature starry flounder probably migrate to
shallow coastal waters to spawn (Orcutt 1950). After
spawning, some adult starry flounder return to the Bay
for feeding, and are most common in the Bay from late
spring through early fall (Ganssle 1966). They reach a
maximum length of 915 mm (Miller and Lea 1972)
In San Francisco Bay, there is a shift in distribution with growth. Age-0 fish are found more commonly
in fresh to brackish water (Suisun Bay, Suisun Marsh,
and the delta), while age-1 and older juveniles are more
commonly associated with brackish to marine waters
(Suisun and San Pablo bays). Throughout their time in
the San Francisco Bay, juvenile starry flounder are commonly found in shallow water, including shoals, intertidal areas, and tidal marshes (Woods 1981, Moyle et al
1986, Baxter 1999, CDFG, unpubl. data).
Fish
changes in the oceanic environment (particularly temperature) may also affect recruitment. Ocean temperatures have been above average for the region for much
of the 1980s and 1990s and it is possible that adult populations moved northward into cooler waters. Temperature can also influence spawning and early development,
as increased temperatures may result in decreased hatching success and larval survival.
Trophic Levels
Primary to secondary carnivore. Feeds primarily on large
benthic invertebrates and rarely on fish.
Proximal Species
Prey: Benthic invertebrates including bivalves, polychaetes, and crustaceans.
Good Habitat
Acknowledgments
Some of the materials in this report are summarized from
the Plueronectiformes chapter of IEP Technical Report
63 which is referenced below (Baxter 1999).
References
Baxter, R.. 1999. Starry flounder. In: J. Orsi (ed). Report on the 1980-1995 fish, shrimp, and crab sampling in the San Francisco Estuary, California. Interagency Ecological Program for the SacramentoSan Joaquin Estuary. Tech. Rept. No. 63.
California Department of Fish and Game (CDFG).
1992. Estuary dependent species, Exhibit #6. In:
State Wat. Qual. Contr. Bd. 1992. Wat. Qual./
Wat. Rights Proceedings on San Fran. Bay and the
Sac./San Joaquin Delta. 97 pp.
Emmett, R.L., S.A Hinton, S.L. Stone and M.E. Monaco. 1991. Distribution and abundance of fish
and invertebrate in the west coast estuaries. Vol. II:
Species life history summaries. NOAA/NOS Strategic Envir. Assessment Div., Rockville, MD
ELMR Rep. No. 8, 329 pp.
Ganssle, D. 1966. Fishes and decapod of San Pablo and
Suisun Bays. In: D.W. Kelly (ed). Ecological studies of the Sacramento San Joaquin Estuary, Part 1.
Ca. Dept. Fish and Game Fish Bull. 133:64-94.
Garrison, K.J. and B.S. Miller. 1982. Review of the early life
history of Puget Sound fishes. Univ. of Washington. Fish.
Res. Inst. Seattle, Washington. UW 8216: 729p.
150
Baylands Ecosystem Species and Community Profiles
Personal Communications
Bob Spies, Applied Marine Sciences, Livermore, California.
Plants
Amphibians &
Reptiles
Suitable habitat includes shallow to deep subtidal mud
and sand flats. Juvenile rearing occurs in the shallow areas
of Suisun and San Pablo bays. Open deeper waters with
higher salinity are generally more acceptable for adults.
Haertel, L. and C. Osterberg. 1967. Ecology of zooplankton, benthos, and fishes in the Columbia River
Estuary. Ecology 48(3): 459-472.
Haugen, C.W. 1992. Starry flounder. In: W.S. Leet,
C.W. Dewees, and C.W. Haugen (eds). California’s
living marine resources and their utilization. California Sea Grant.
Hieb, K. and R. Baxter. 1993. Delta Outflow/San Francisco Bay Study. In: P.L. Herrgesell (compiler).
1991 Annual Report, Interagency Ecological Studies Program for the Sacramento-San Joaquin Estuary, pp. 101-116
Interagency Ecological Studies Program for the Sacramento-San Joaquin Estuary (IESP) 1993. 1991 Annual Report.
P.L. Herrgesell, compiler. Sacramento, Ca. 150 pp.
Miller, D.J. and R.N. Lea. 1972. Guide to the coastal
marine fishes of California. Ca. Fish and Game
Fish Bull. 157, 249 pp.
Moyle, P.B. 1976. Inland Fishes of California. Univ. of
California Press, Berkeley, Ca., 405 pp.
Moyle, P.B., R.A. Daniels, B. Herbold, and D.M. Baltz.
1986. Patterns in distribution and abundances of
a noncoevolved assemblage of estuarine fishes in
California. Fishery Bulletin 84(1):105-117.
Orcutt, H.G. 1950. The life history of the starry flounder, Platichthys stellatus (Pallus). Ca. Fish Game
Fish Bull. 78, 64 pp.
Policansky, D. 1982. Influence of age, size, and temperature on metamorphosis in starry flounder, Platichthys
stellatus. Can. J. Aquatic Sci. 39(3):514- 517.
Policansky, D. and P. Sieswerda. 1979. Early life history
of the starry flounder, Platichthys stellatus, reared
through metamorphosis in the laboratory. Trans.
Amer. Fish. Soc. 108(3):326 327.
Radtke, L.D. 1966. Distribution of smelt, juvenile sturgeon, and starry flounder in the Sacramento-San
Joaquin Delta with observations on food of sturgeon.
In: J.L. Turner and D.W. Kelley (Comp). Ecological studies of the Sacramento-San Joaquin Delta, Part II. Ca.
Dept. Fish and Game, Fish Bull. 136: 115-129.
Wang, J.C.S. 1986. Fishes of the Sacramento-San
Joaquin Estuary and adjacent waters, California:
A guide to the early life histories. Interagency Ecological Study Program for the Sac.-San Joaquin
Estuary, Tech. Rep. 9, 345 pp.
Woods, E. 1981. Fish Utilization. In: The Hayward
Regional Shoreline marsh restorations: Biological
succession during the first year following dike removal. T. Niesen and M. Josselyn, eds. Tiburon
Center for Environmental Studies, Tech. Rept.
1:35-46.
3
Invertebrates
Franciscan Brine Shrimp
Artemia franciscana Kellogg
their effects on this species are currently being investigated (Maiss and Harding-Smith 1992).
General Information
The Franciscan brine shrimp, Artemia franciscana (formerly salina) (Bowen et al. 1985, Bowen and Sterling
1978, Barigozzi 1974), is a small crustacean found in
highly saline ponds, lakes or sloughs that belong to the
order Anostraca (Eng et al. 1990, Pennak 1989). They
are characterized by stalked compound eyes, an elongate
body, and no carapace. They have 11 pairs of swimming
legs and the second antennae are uniramous, greatly enlarged and used as a clasping organ in males. The average length is 10 mm (Pennak 1989). Brine shrimp commonly swim with their ventral side upward. A. franciscana
lives in hypersaline water (70 to 200 ppt) (Maiss and
Harding-Smith 1992).
In the Bay area, the optimum temperature for A.
franciscana is 21-31°C. In the winter, when temperatures
fall below this range, brine shrimp populations decline
and their growth becomes stunted (Maiss and HardingSmith 1992). Other environmental factors such as wind,
salinity, and the quantity and quality of phytoplankton
may also affect Bay area populations of A. franciscana and
Artemia franciscana has two types of reproduction, ovoviviparous and oviparous. In ovoviviparous reproduction,
the fertilized eggs in a female can develop into free-swimming nauplii, which are set free by the mother. In oviparous reproduction, however, the eggs, when reaching the
gastrula stage, become surrounded by a thick shell and
are deposited as cysts, which are in diapause (Sorgeloos
1980). In the Bay area, cysts production is generally
highest during the fall and winter, when conditions for
Artemia development are less favorable. The cysts may
persist for decades in a suspended state. Under natural
conditions, the lifespan of Artemia is from 50 to 70 days.
In the lab, females produced an average of 10 broods,
but the average under natural conditions may be closer
to 3-4 broods, although this has not been confirmed.
Each brood contains from 30 to 100 offspring which
mature in 10-25 days (Maiss and Harding-Smith 1992).
The larva grows and differentiates through approximately 15 molts (Sorgeloos 1980).
Food and Feeding
Artemia franciscana feed on phytoplankton and bluegreen algae that occur in Bay area salt ponds (Maiss and
Harding-Smith 1992).
Distribution
Artemia franciscana occurs in highly saline waters
throughout western North America, Mexico, and in the
Caribbean (Bowen et al. 1985). In California, A. franciscana occurs from sea level to 1,495m and in many
parts of the state, but its distribution is spotty because
of this species salinity requirements (Eng et al. 1990).
Historically in the Bay area they were found in salt
pannes and sloughs were hypersaline conditions occurred. Currently they occur in salt ponds in the northern and southern portion of San Francisco Bay that are
Chapter 3 — Invertebrates
151
Invertebrates
Reproduction, Growth, and Development
Brita C. Larsson
used for the commercial production of salt. Salt ponds
cover approximately 111 km and in the North bay 36
Km off the Bay’s shoreline (Lonzarich 1989). The distribution of Artemia in these salt ponds is limited by the
salinity of the ponds. The optimum salinity range for
Artemia is 70 ppt to 175 ppt (Carpelan 1957). They do
not occur where the salinity is above 200 ppt.
Invertebrates
Population Status and Influencing Factors
Commercial salt production in San Francisco Bay is currently an active industry, so habitat for A. franciscana is
not limited and populations are large due to ample
amounts of habitat. Donaldson et al. (1992) sampled a
496 acre salt pond in the San Francisco Bay National
Wildlife Refuge and estimated the highest winter adult
population at 40 billion and the lowest winter population at 4.5 billion. Brine shrimp populations are lowest
in the winter and peak in the summer months when
their optimal temperatures occur so these numbers are
conservative for a maximum population value for the
pond. Current populations of the brine shrimp probably
far exceed historic populations because the salt ponds in
which they occur are manmade. Salt ponds occurred
naturally and there is even some evidence that the
Ohlone Indians manipulated a portion of the Bay shoreline for salt production but never was there as much salt
pond habitat for brine shrimp as currently occurs in the
Bay area.
Trophic Level
Artemia franciscana is a primary consumer.
Proximal Species
Anderson (1970) lists sightings of 55 bird species using
salt ponds in San Francisco Bay. Mallards, California
gulls, whimbrels, Wilson’s phalarope, eared grebes and
American avocets are several species which feed on A.
franciscana. Western and least sand pipers, willets, greater
yellow legs and Bonaparte’s gulls are commonly seen
roosting and feeding in the salt pond environment and
most likely feed on Artemia in these ponds (Maiss and
Harding-Smith 1992).
Good Habitat
Brine shrimp occur in salt ponds adjacent to San Francisco Bay that have salinities ranging from 70 to 200 ppt
but are most common when the range is between 90 and
150 ppt (Maiss and Harding-Smith 1992). Harvey et al.
(1988) reported that up to 46% of the 23,465 acres
of active salt ponds in South Bay are within the 70200 ppt salinity range in the summer and contain
brine shrimp.
152
Baylands Ecosystem Species and Community Profiles
References
Anderson, W. 1970. A preliminary study of the relationship of saltponds and wildlife-South San Francisco Bay. Calif. Fish and Game 56(4):240-252.
Barigozzi, C. 1974. Artemia: a survey of its significance
in genetic problems. In: T. Tobshansky, M. K.
Hecht and W.C. Steere (eds). Evolutionary Biology
(7). Plenum Press, New York, NY. Pp. 221-252.
Bowen, S.T. and G. Sterling. 1978. Esterase and malate
dehydrogenase isozyme polymorphism in S.
artemia populations. Comp. Biochem. and Physio.
61B:593-595.
Bowen, S.T., E.A. Fogarino, K.N. Hitchner, G.L. Dana,
V.H.S. Chow, M.R. Buoncristiani and J.R. Carl.
1985. Ecological isolation in Artemia: population
differences in tolerance of anion concentrations.
J. Crustacean Biology 5:106- 129.
Carpelan, L.H. 1957. Hydrobiology of the Alviso salt
ponds. Ecology 38:375-390.
Donaldson, M.E., D.E. Conklin, and T.D. Foin. 1992.
Population dynamics of Artemia Franciscana in the
San Francisco Bay National Wildlife Refuge: Phase
II. Interim Report #2.
Eng, L.L. D. Belk, and C.H. Eriksen. 1990. California
Anostraca: distribution, habitat, and status. J. Crustacean Biology 10:247-277.
Harvey, T.E., P.R. Kelly, R.W. Towe, and D. Fearn. 1988.
The value of saltponds for Waterbirds in San Francisco Bay and considerations for future management. Presentation at the Wetlands ’88: Urban
Wetlands and Riparian Habitat Conference, June
26-29, Oakland, CA.
Lonzarich, D. 1989. Life History and Patterns of Distribution in Salt pond Fishes: A Community Level
Study. M.S. thesis San Jose University, Ca.
Maiss, F.G. and E.K. Harding-Smith. 1992. San Francisco Bay National Wildlife Refuge final environmental assessment of commercial brine shrimp harvest. U.S. Fish and Wildlife Service, San Francisco
Bay National Wildlife Refuge, Newark, Ca. 27p.
Pennak, R.W. 1989. Freshwater invertebrates of the
United States. 3rd edition. Protozoa to Mollusca.
John Wiley and Sons, Inc. New York, NY. 628 p.
Sorgeloos, P. 1980. Life history of the brine shrimp
Aretmia. In: G. Persoone, P. Sorgeloos, O. Roels
and E. Jaspers (eds). The Brine Shrimp Artemia.
Universa Press Wetteren, Belgium. pp.xxi-xxiii.
California Vernal Pool Tadpole
Shrimp
Lepidurus packardi Simon
Brita C. Larsson
General Information
Reproduction, Growth, and Development
Much of what is known about the reproduction, growth,
and development of L. packardi comes from studies by
Ahl (1991) and Longhurst (1955). Their life history is
dependent on ephemeral freshwater pools. In California,
vernal pools are generally hydrated during the rainy season, which extends from winter to early spring. Populations of tadpole shrimp are reestablished from diapaused
eggs when winter rains rehydrate vernal pools. Once a
pool rehydrates, the eggs hatch over a three week period,
some hatching within the first four days. It takes another
three to four weeks for the tadpole shrimp to become
sexually reproductive. Populations consist of both males
and females, though late in the season, pools are often
dominated by males. After copulation, fertilized eggs
descend into the foot capsule of the female (Desportes
and Andrieux 1944). The eggs are sticky and when they
are deposited they adhere to plant matter and sediment
particles (Federal Register 1994). A female can have up
Food and Feeding
Tadpole shrimp feed on organic detritus and living organisms such as fairy shrimp and other invertebrates
(Pennak 1989, Fryer 1987).
Distribution
L. packardi is endemic to vernal pools in the Central Valley, coast ranges and a limited number of sites in the
Transverse Range and Santa Rosa Plateau (Federal Register 1994). The distribution of this species is not well
known for the Bay area. Recently, L. packardi was collected at the Warm Springs Seasonal Wetland which is
a part of the Don Edwards San Francisco Bay National
Wildlife Refuge (Caires et al 1993). Other populations
have been found north of the eastern half of Potrero Hills
in the North Bay (S. Forman, Pers. obs. ). Seasonal wetlands occur sporadically in both the North and South
Bay and may provide additional habitat for this species.
Surveys in seasonal wetlands surrounding San Francisco
Bay may contribute and increase information on the distribution of this species.
Population Status and Influencing Factors
Current status of the population of tadpole shrimp in
the Bay area is not known. Loss of seasonal wetland habitat in the Bay area may be significantly affecting the
population of this species especially since distribution information for the Bay area is so limited.
Trophic Level
Lepidurus packardi is most likely a secondary consumer.
Proximal Species
Waterfowl, western spadefoot toad, and tadpoles.
Dr. J.L. King
Good Habitat
Lepidurus packardi inhabits vernal pools. They have been
found in pools ranging in size from 5 square meters to
36 hectares. The water in the pools can be clear to turbid. The pools often have low conductivity, TDS, and
alkalinity (Federal Register 1994, Eng et al. 1990). The
pools dry up in the late spring and are dry in the summer and fall then fill with rain water in the winter and
early spring. Vernal pool formations occur in grass bot-
Chapter 3 — Invertebrates
153
Invertebrates
The California vernal pool tadpole shrimp is a small
crustacean found in ephemeral freshwater pools that
belong to the order Notostraca. They are characterized by sessile compound eyes, a shield-like carapace
covering the head and much of the trunk, and a telson that is a flat and paddle-shaped protuberance.
They can reach a length of 50 mm and have approximately 35 pairs of legs and two long cercopods
(Pennak 1989). Tadpole shrimp are primarily benthic
organisms that swim with their legs down. They can
also climb or scramble over objects and plow through
bottom sediments (Federal Register 1994). Information about the biology of this species is limited and
incomplete (Ahl 1991).
to six clutches of eggs, totaling about 861 eggs during
her lifetime (Ahl 1991). Depending on the depth and
persistence of water in a pool, some eggs hatch immediately. The remainder inter diapause and lie dormant in the
sediment during the dry portion of the year (Ahl 1991).
Reticulate Water Boatman
Trichocorixa reticulata Guerin
Wesley A. Maffei
Description and Systematic Position
Trichocorixa reticulata is a small hemipteran, approximately 3-5mm in length, that belongs to the family
Corixidae. This insect, also known as the salt marsh water boatman, can be recognized by the fine network of
lines on its hemelytra (outer wing covers), the 10-11 dark
transverse bands on the pronotum, and the pala of front
legs not exceeding two-thirds the width of an eye along
the ventral margin (Figure 3.1).
Distribution
References
Ahl, J.S. 1991. Factors affecting contributions of the
tadpole shrimp, Lepidurus packardi, to its over summering egg reserves. Hydrobiologia 212:137-143.
Caries, T.D. Dawn, D. DiNunzio, A. Harris, N. Kogut,
M. Kutiled, S.H. Ladd, J. Stanziano, M. Stickler
and A. Webber. 1993. Sur la biologie de Lepidurus
apus. Bull. Soc. Zool. Fr. 69:61-68.
Desportes, C. and L.H. Andrieux. 1944. Sur la biologie
de Lepidurus apus. Bull. Soc. Zool. Fr. 69:61-68.
Eng, L.L., D. Belk and C.H. Eriksen. 1910. California
Anostraca: distribution, habitat, and status. J. crustacean Biology 10:247-277.
Federal Register. 1994. Endangered and threatened
plants: determination of endangered status for the
conservancy fairy shrimp. Longhorn fairy shrimp,
and vernal pool tadpole shrimp; and threatened
status for the vernal pool fairy shrimp. Fed. Reg.
59:48136-48153.
Fryer, G. 1987. A new classification of the branchiopod
Crustacea. Zool. J. Linn. Soc. 91:357-383.
Gallagher, S.P. 1996. Seasonal occurrence and habitat
characteristics of some vernal pool Branchiopods
in Northern California, U.S.A. J. Crustacean Biology 16(2):323-329.
Longhurst, A.R. 1955. A review of the Notostraca. Bull.
Br. Mus. Nat. Hist. Zool. 3:1-57.
Pennak, R.W. 1989. Freshwater invertebrates of the
United States. 3rd edition. Protozoa to Mollusca.
John Wiley and Sons, Inc. New York, NY. pp628.
Sailer (1948) states that this insect is found along the
Pacific Coast from northern San Francisco Bay south to
Peru. Populations from Kansas, New Mexico, Texas,
Florida, and the Hawaiian Islands have also been recorded. One isolated record was reported in China but
this has been unconfirmed. Within the San Francisco
Bay environs this water boatman can be found in mid
to upper marsh tidal pools and man-made salt ponds.
Figure 3.2 shows the locations around the Bay Area
where T. reticulata have been collected, and Table 3.1
shows the collection dates.
Suitable Habitat
T. reticulata prefers saline environments. Cox (1969)
found this insect in southern California coastal salt ponds
with salinities ranging from brackish up to 160 ‰ and
Jang (1977) states that this water boatman can occur in
ponds with salinities up to 170 ‰. Carpelan (1957)
found the Alviso population in Cargill salt ponds that
ranged from 23 ‰ up to 153 ‰. In all instances it was
found that the greatest numbers of individuals and the
most reproduction occurred in saline environments with
a salinity range of 35-80 ‰.
Actual Size
3-5 mm
Personal Communication
Steve Forman, LSA Associates, Incorporated
Figure 3.1 Reticulate Water Boatman –
Trichocorixa reticulata
154
Baylands Ecosystem Species and Community Profiles
Wes Maffei
Invertebrates
tomed swales of grasslands in old alluvial soils, underlain by hardpan or in mud bottomed pools (Federal Register 1994). Pools with cobblely hardpan bottoms also
serve as habitat (Gallagher 1996). Gallagher (1996)
found that the depth, volume, and duration of inundation of a pool was important for the presence of L.
packardi in vernal pools when compared to the needs of
other branchiopods. He found L. packardi did not reappear in ponds that dried and rehydrated during the study
period, while other Branchiopod species did. L. packardi
needs deeper and longer-lasting pools if they are to persist
over a rainy season in which both wet and dry periods occur. Temperature variation in pools where L. packardi have
been found to vary from 3 to 23°C (Gallagher 1996). Salinity, conductivity, dissolved solids, and pH of the water
in vernal pools are also important in determining the distribution of tadpole shrimp (Federal Register 1994).
Table 3.1 Known Collection Sites For
Trichocorixa reticulata 1
Location
Date Specimen(s) Collected
Sausalito
29 Oct 1921
Redwood City
15 Jun 1922, 24 Apr 1923,
8 May 1923
Berkeley
18 Apr 1962
Oakland
14 Apr 1930
Baumberg Tract, Hayward
8 Oct 1989
Coyote Hills Park
25 Oct 1988, 11 May 1989
Mowry Slough
25 Sep 1997
Alviso (Coyote Creek)
12 Aug 1980
General Sample
Location
Figure 3.2 Known Trichocorixa reticulata Localities Within San Francisco Bay Tidal and Diked
Marshes
Biology
Sailer (1942) believes that all species of Trichocorixa over
winter as adults. Scudder (1976) states that Tones
(unpub.) has found that in Saskatchewan, Trichocorixa
verticalis interiores over winters in the egg stage.
Eggs are laid singly on submerged vegetation or
objects on the bottom substrate. Developmental time for
eggs and immatures can very considerably with temperature.
Adult water boatman are both herbivorous and
predatory feeding on algal cells and various microorganisms. Although Corixids are aquatic in all life stages, the
adults are capable of leaving the water and dispersing by
flight. Maffei (unpub.) has noted that south San Francisco Bay populations are attracted to dark colored objects, with adults landing in large numbers on the hoods
of green or burgundy colored vehicles while adjacent
white vehicles had few if any specimens.
Reproduction
Cox (1969) and Carpelan (1957) have noted that peak
reproduction occurs in saline environments with salinities ranging between 35 ‰ and almost 80 ‰. Egg laying is continuous during spring, summer and fall with
the greatest number of nymphs occurring during April
and May. Cox (1969) has also found that crowding of
adults led to increased egg production in females.
Balling and Resh (1984) have reported, that the
number of generations per year for the Petaluma Marsh
population was at least in part dependent on the longev-
ity of the tidal ponds. They found that ponds which
dried during late summer contained over wintering, nonreproducing adults while water filled ponds would produce another generation. Reproduction does occur yearround but Cox (1969) states that salinity and adult
densities influence the number of eggs laid and the
maturation rates of the immature stages. Balling and
Resh (1984) noted that the time between generations of
the Petaluma Marsh population was also affected by
variable egg development times, variable instar development rates, and inter-pond differences in recruitment of
adults. In general, it has been determined that environmental conditions can cause water boatmen to either
accelerate or delay their development and production of
subsequent generations.
Significance to Other Wetlands Taxa
This insect is considered an important prey item for
shore birds. Howard (1983) studied the esophageal contents of 35 Ruddy Ducks, Oxyura jamaicensis, at the
Alviso salt ponds and found that this water boatman
comprised 12.6% of the total food volume. Howard also
examined the gizzard contents of 53 Ruddy Ducks and
found that 25.5% of the total food volume was water
boatmen. Anderson (1970) analyzed the stomach contents of 10 Ruddy Ducks and found that water boatman,
snails and Widgeon grass seeds were the primary components of their diet. He also found Least Sandpipers,
Wilson’s Phalarope and Northern Phalarope’s utilized
this insect as part of their diets.
Conservation Needs and Limiting Factors
Salinity and the length of time tidal marsh ponds contain water seem to be the primary driving forces affect-
Chapter 3 — Invertebrates
155
Invertebrates
1
Information assembled from specimens contained within the California Academy of Sciences Insect Collection, University of California
Berkeley Essig Museum, University of California Bohart Museum, San
Jose State University Edwards Museum, San Mateo County Mosquito
Abatement District Insect Collection, and private collections of Dr. J.
Gordon Edwards and Wesley A. Maffei.
ing both developmental rates and reproduction.
Anderson, W. 1970. A preliminary study of the relationship of salt ponds and wildlife—South San
Francisco Bay. Calif. Dept. Fish and Game
56(4):240-252.
Balling, S.S. and V. Resh. 1984. Life history and variability in the water boatman Trichocorixa reticulata
(Hemiptera: Corixidae) in San Francisco Bay salt
marsh ponds. Ann. Ent. Soc. Amer. 77(1):14-19.
Carpelan, L.H. 1957. Hydrobiology of the Alviso Salt
Ponds. Ecology 38:375-390.
Cox, M.C. 1969. The biology of the euryhaline water
boatman Trichocorixa reticulata (Guerin-Meneville)
M.S. Thesis, San Diego State Univ. 84pp.
Howard, J.A. 1983. Feeding ecology of the ruddy duck
on the San Francisco Bay National Wildlife Refuge. M.A. Thesis, San Jose State Univ. 53 pp.
Jang, E.B. 1977. Hydromineral regulation in the saline
water corixid Trichocorixa reticulata. M.S. Thesis,
Hayward State Univ. 79 pp.
Maffei, W. 1989-1996. Unpublished field notes.
Sailer, R.I. 1942. The genus Trichocorixa (Hemiptera:
Corixidae). Phd Thesis, Univ. of Kansas, Lawrence,
Kansas. 118pp.
Scudder, G.E. 1976. Water-boatmen of saline waters
(Hemiptera: Corixidae). pp. 263-289. In: L. Cheng
(ed). Marine Insects. North-Holland Publ.,
Amsterdam. 581pp.
Additional Readings
Hungerford, H.B. 1948. The Corixidae of the western
hemisphere (Hemiptera). Univ. Kans. Sci. Bull. 32:
827pp.
Usinger, R.L. (ed). 1956. Aquatic insects of California.
Univ. Calif. Press, Berkeley. 508pp.
Tiger Beetles
Cicindela senilis senilis, C. oregona,
and C. haemorrhagica
Wesley A. Maffei
Description and Systematic Position
Cicindela senilis senilis, C. oregona, and C. haemorrhagica
are moderate sized beetles, approximately 10-15mm in
length, that belong to the family Cicindelidae (Figure
3.3). These beetles, also known as tiger beetles, can be
easily identified by their large, bulging eyes and long,
sickle-shaped mandibles that bear small teeth. Adults of
C. senilis and C. oregona are usually shining metallic blue
to green on the ventral surface with the dorsum dull coppery brown and bearing small yellowish-white irregular
markings. Cicindela haemorrhagica is similar in appearance to both C. senilis and C. oregona except that the
ventral surface of the abdomen is usually bright red. The
larvae are S-shaped, yellowish-white, have the head and
the first thoracic segment flattened, an enlarged hump
on the fifth abdominal segment with hooks, and large
mandibles that are similar to the adults.
Distribution
Historically the San Francisco Estuary, including the
beaches just outside of the Golden Gate Bridge, was
home to four species of tiger beetles. These were:
Cicindela haemorrhagica, C. hirticollis, C. oregona oregona
and C. senilis senilis. Only two species, C. haemorrhagica
and C. senilis senilis are present today with C. haemorrhagica in decline within or near the tidal areas of the
San Francisco Bay Estuary. Cicindela oregona oregona
may still be present within the estuary but the last known
population was destroyed in 1996.
The dominant tiger beetle, C. senilis senilis, is currently found throughout the south and central portions
of the estuary with one population having been identified from Grizzly Island in 1991. Museum records in-
Actual Size
10 - 15 mm
Wes Maffei
Invertebrates
References
Figure 3.3 Tiger Beetle – Cicindela senilis senilis
156
Baylands Ecosystem Species and Community Profiles
Figure 3.4 Known Tiger
Beetle Localities Within
San Francisco Bay Tidal
and Diked Marshes
Invertebrates
dicate that this beetle was also found in San Rafael,
Martinez and Port Costa but these sites have not been
sampled in over 40 years. C. haemorrhagica, has become
increasingly scarce as its habitat continues to be altered
for human needs. This beetle is currently found at Trojan Marsh (San Leandro), Hayward Landing (Hayward),
Salt Ponds west of Newark and the Richmond Field Station (Richmond). Historically this beetle had a broader
distribution with sites as far north as Martinez and south
throughout most of the south San Francisco Bay. The
populations at Alameda, Bayfarm Island and Oakland no
longer exist and other sites identified from museum
records have apparently not been sampled in at least three
or more decades (Maffei, unpub.). C. oregona is probably no longer present within the tidal and diked marshes
of the San Francisco Bay. The last known population was
at Bayfarm Island and was extirpated in 1996 when the
site was graded in preparation for development. Figure
3.4 shows the locations around the Bay Area where C.
senilis senilis, C. oregona, and C. haemorrhagica have been
collected, and Table 3.2 shows the collection dates.
Suitable Habitat
San Francisco Bay tiger beetles are commonly found
along open, muddy margins of creeks and streams and
also along the muddy margins of salt pannes that are
occasionally inundated by high tides. High, dry banks
of channels and open areas of levees associated with salt
ponds and muted tidal marshes tend to be favored sites
for C. senilis senilis. Habitat utilized by both adults and
larvae can be characterized as having extensive areas of
fine silt or sandy clay-like soil, exposed to full sun, with
minimal to moderate vegetation, and being located near
water. C. haemorrhagica and C. oregona oregona have
shown a preference for wet, sandy beach-like areas that
may or may not be influenced by fresh water from creeks
and canals.
Biology
The specific biology of San Francisco Bay tiger beetles is
not well known. The information that follows is a general-
Chapter 3 — Invertebrates
157
Table 3.2 Known Collection Sites for Tiger Beetle Populations1
Location
Date Specimen(s) Collected
Cicindela haemorrhagica
Date Specimen(s) Collected
Cicindela senilis senilis
Martinez
28 Aug 1959, 21 Sep 1959
24 Apr 1993
San Rafael (*)
1951
23 May 1941, 20 May
Richmond Field Station
Alameda
1930,
23 Aug 1930, 24 Aug
Milbrae
1 Sep 1912, 2 Jun 1912,
3 Oct 1914
4 Jul 1932
San Mateo
24 Oct 1952
San Francisco
31 Jan 1944 (***)
Redwood City (Salt Marsh)
Burlingame
7 Oct 1969
16 Sep 1951, 15 Jul 1951,
15 Jun 1952
Lake Merritt
Jul 1906
Redwood City (Harbor)
15 Jun 1952
Oakland
15 Aug 1902
Redwood City
15 Apr 1952, 26 Sep 1952
Redwood City (nr Yt. Harbor)
15 Jun 1952
Bair Island
9 Mar 1997
31 Jul 1951
East Palo Alto (Marsh)
13 Jul 1951
23 May 1921
Redwood City (Saltmarsh)
29 Jun 1969
Palo Alto (Salt Marsh)
East Palo Alto
31 Jul 1951
Grizzly Island (wildlife area)
10 Oct 1991
Milpitas
15 Jul 1966, 26 Jul 1966
Port Costa
21 Sep 1947
Newark (2 mi west of)
25 Jun 1975, 24 Jul 1980
Martinez
28 Aug 1955, 28 Sep 1955
Bayfarm Island
21 Jun 1990
Emeryville
20 Aug 1936
Russell Salt Marsh, Hayward
30 Jul 1996, 27 May 1997
Lake Merritt
Trojan Marsh
2 Aug 1997
4 Oct 1904, 9 Oct 1904,
12 Sep 1907, 12 Apr 1909
Alameda
Jun 1901, 16 Aug 1902,
9 May 1907
Palo Alto Yacht Harbor
Invertebrates
Location
Cicindela oregona
Bayfarm Island
May 1939
San Francisco Beach
14 Apr 1957
Oliver Salt Ponds, Hayward
5 Aug 1989, 2 Jul 1990
Burlingame
22 May 1952
Bayfarm Island
11 Apr 1972, Jul 1989,
21 Jun 1990, Jul 1993,
Aug 1993, 12 Apr 1993,
1 Sep 1995
Whale’s Tail Marsh (Hayward) 12 Apr 1993, 8 Apr 1993,
11 Mar 1993
Oakland (*)
Jun 1906, 16 Aug 1902
Concord (*)
27 Apr 1935
Cicindela hirticollis (data from Graves 1988)
Oakland
no date
San Francisco
1907?
Baumberg Salt Ponds
Mar 1989, 1 Apr 1990,
13 Jun 1989, 11 Mar 1997
Patterson Hill Marsh, Fremont 11 Apr 1989
Newark (2 mi west of)
25 Jun 1975, 24 Jul 1980
Dumbarton Bridge (Newark) 9 May 1952
Newark Salt Flats
17 Jun 1966
W. End Mowry Slough
19 Sep 1997
Brinker Marsh
Mar 1989, 10 Mar 1997
E. End Albrae Slough, Fremont
12 Mar 1997
Dixon Rd, Milpitas
23 Jun 1956
Milpitas (wet sand)
1 May 1966, 12 Oct 1966
Alviso
21 Mar 1947, 22 Mar 1947,
27 Mar 1947, 12 Apr 1947,
Apr 1954, 14 Apr 1955,
15 Apr 1955, 12 May 1959,
19 May 1959, 8 Jun 1980
* May or may not be within the confines of the Ecosystem Goals Project.
*** Probably a dubious record, suspect mislabeled specimen.
1
Information assembled from specimens contained within the California Academy of Sciences Insect Collection, University of California Berkeley Essig
Museum, University of California Bohart Museum, San Jose State University Edwards Museum, San Mateo County Mosquito Abatement District Insect
Collection, and private collections of Dr. J. Gordon Edwards and Wesley A. Maffei
ized biology for these insects drawn from the studies of
other species and a summary article by Pearson (1988).
Adults of these beetles are active on hot, sunny days
and are exceedingly quick both in flight and on the
ground. When approached these insects will run away
or fly for a short distance, land, and then face their pur-
158
Baylands Ecosystem Species and Community Profiles
suer. Larvae and adults are predators, feeding on other
insects. Prey items for San Francisco Bay tiger beetles
include but are not limited to the Brine Flies Ephydra
cinerea, Ephydra millbrae, Lipochaeta slossonae, and
Mosillus tibialis, and various beetles belonging to the
families Carabidae and Tenebrionidae. Pearson and
C. haemorrhagica do not emerge until mid to late June
and are usually present through September.
Actual Size
2.5 mm
Wes Maffei
Reproduction
Figure 3.5 Cicindela senilis senilis Larva in Burrow
Significance to Other Wetlands Taxa
These beetles may be a potential prey item for shore
birds. Cramp and Simmons (1983) cite a stomach content analysis study of the European race of Snowy Plover, Charadrius alexandrinus alexandrinus, in Hungary
which revealed the presence of 28 tiger beetles. Swarth
(1983) noted that these beetles were occassionally eaten
by Snowy Plovers found at Mono Lake. Marti (1974)
found tiger beetle parts in burrowing owl pellets that
were studied in the northeastern part of Larimer County,
Colorado.
Conservation Needs and Limiting Factors
Nagano (1982) has stated that some tiger beetles are considered to be good indicators of coastal wetlands distur-
Chapter 3 — Invertebrates
159
Invertebrates
Mury (1979) found that adults of some species of tiger
beetles also fed on dead organisms. Faasch (1968) and
Swiecimski (1956) found the adults located live prey
visually while dead prey were found tactilely. Adult
beetles tend to frequent the muddy margins of their
habitat where prey items are readily encountered while
the immature stages tend to be found in the drier areas.
The eggs, larvae and pupae are subterranean, with
the larvae living in vertical burrows and waiting near the
top to seize any prey that passes by (Figure 3.5). Prey
items are captured with the mandibles and pulled down
to the bottom of the burrow where it is ingested. Faasch
(1968) found that a dark object against a light background released the prey-catching behavior. Burrows are
enlarged by loosening the soil with the mandibles and
using the head and pronotum to push the soil to surface. At the surface, the soil is flicked off by flipping the
head and pronotum backward (Shelford 1908, Willis
1967). The depth of larval tunnels has been found to
range between 15 and 200 cm depending on the age of
the larva, the species of tiger beetle, the season and soil
type and conditions (Criddle 1910, Willis 1967, Zikan
1929).
Larvae undergo three molts with the time for development lasting one to four years and averaging about
two years (Willis 1967). Pearson and Knisley (1985)
found that the availability of food effected rate of development and was therefore a limiting resource in the life
cycle of tiger beetles. They found that ample prey shortened the developmental time from egg to adult with 60
days total developmental time having been observed for
some laboratory reared beetles. Prior to pupation, the last
instar larva plugs the tunnel entrance and excavates a
chamber or pupal cell. The period for pupation is usually short, lasting no more than 30 days.
Larvae can be found throughout the year while
adults are present from March through October. Peak
adult activity for the south San Francisco Bay Cicindela
senilis senilis populations is from late April through June
(Maffei, unpub.). Blaisdell (1912) noted that adults of
C. senilis , which emerged in the fall, would hibernate.
Males initiate copulation by approaching a female in
short sprints which is similar to the intermittent sprinting used when foraging. Once close enough to a potential mate, the male leaps onto their back, grasping the
thorax with his mandibles and the elytra with his front
and middle legs. The male’s hind legs remain on the
ground and the coupling sulci of the female receives his
mandibles. Males frequently mount both males and females of any tiger beetle species present. Females try to
dislodge intruding males by rolling on their backs, lurching and then running out into bright sunshine. It is
believed that the fit of the male mandibles into the female sulci may be species specific and that this feature
allows other males, and females of other species, to rid
themselves of unwanted mates (Freitag 1974).
Oviposition usually occurs when the female
touches the ground with her antennae and bites the soil
with her mandibles. The ovipositor is then extended and
with a thrusting motion of the abdomen a hole up to 1
cm is excavated. One egg is deposited in the hole and it
is then covered over so that no evidence of disturbance
exists. The choice of soil type for oviposition has been
found to be extremely critical for many species (Knisley
1987, Leffler 1979, Shelford 1912, Willis 1967).
Availability of prey has been found to directly affect female mortality and the number of eggs produced.
Adult beetles in prey poor habitats were only found to
approach maximum fecundity during years of high rainfall and high prey populations (Pearson and Knisley
1985). Prey availability for larvae was found to affect the
size of later instars, which ultimately affected the size of
the adults produced and individual fecundity (Hori
1982a, Pearson and Knisley 1985).
Invertebrates
bance, with the least disturbed habitats having the greatest species diversity. San Francisco Bay tidally influenced
wetlands appear to have two species of tiger beetle, with
those sites that have had minimal disturbance or that
have not seen much human activity for long periods of
time having the highest populations (Maffei, unpub.).
Unfortunately, few sites exist that have not been subjected to human activity. This has resulted in a loss of
species diversity, with potential tiger beetle habitat usually having only a single species present and having small
disjunct populations. Historically, there were sites that
had more than one species present within a given habitat (ie. Lake Merritt, Bayfarm Island and Burlingame).
San Francisco Bay populations of Cicindela senilis
senilis and C. haemorrhagica prefer to be near permanent
or semi-permanent bodies of water utilizing tidal pannes
with sizable unvegetated flats and/or nearby minimally
vegetated levees. Cicindela haemorrhagica has shown a
preference for sandy beach-like sites but can utilize dry,
fine silty sites as is evidenced by the population at Russell
Salt Marsh, Hayward. Both species of beetles need to
have fine silty clay-like or sandy clay soils, that are unvegetated or sparsely vegetated, within in which to breed.
Bright sunshine and minimal flooding are also important factors.
The immature stages of other species of tiger
beetles have been found to inhabit a smaller range of the
habitat than the adults and are not capable of tolerating
as much variation in physical factors such as soil moisture, soil composition and temperature (Hori 1982b,
Knisley 1987, Knisley 1984, Knisley and Pearson 1981,
Shelford 1912, Shelford 1908). The length and duration
of flooding can also be important, although what the
specifics of these parameters are for San Francisco Estuary tiger beetles is not clear.
Larochelle (1977) found that many species of adults
are readily attracted to lights. What impact this might
have on San Francisco Bay Tiger Beetles with respect to
dispersal and survival is unknown.
References
Blaisdell, F.S. 1912. Hibernation of Cicindela senilis (Coleoptera). Ent. News. 23:156-159.
Cramp, S. and K. Simmons (eds). 1983. The birds of
the western palearctic. Vol. III. Oxford Univ. Press,
Oxford. pp. 153-165.
Criddle, N. 1910. Habits of some mannitoba tiger
beetles (Cicindelidae). II. Can. Ent. 42:9-15.
Faasch, H. 1968. Beobachtungen zur biologie und zum
verhalten von Cicindela hybrida L. und Cicindela
campestris L. und experimentelle analyse ihres
beutefangverhaltens. Zool. Jarhb. Abt. Syst. Oekol.
Geogr. Tiere 95:477-522.
Freitag, R. 1974. Selection for a non-genitalic mating
structure in female tiger beetles of the genus
160
Baylands Ecosystem Species and Community Profiles
Cicindela (Coleoptera: Cicindelidae). Can. Ent.
106:561-568.
Hori, M. 1982a. The biology and population dynamics
of the tiger beetle, Cicindela japonica (Thunberg).
Physiol. Ecol. Jpn. 19:77-212.
______. 1982b. The vertical distribution of two species
of tiger beetles at Sugadaira (Mt. Neko-Dake),
Nagano Prefecture, with special reference to their
habitat preferences. Cicindela 14:19-33.
Knisley, C.B. 1987. Habitats, food resources and natural enemies of a community of larval Cicindela in
Southeastern Arizona (Coleoptera: Cicindelidae).
Can. J. Zool. 65:191-200.
______. 1984. Ecological distribution of tiger beetles
(Coleoptera: Cicindelidae) in Colfax County, New
Mexico. Southwest. Nat. 29:93-104.
Knisley, C.B. and D.L. Pearson. 1981. The Function of
turret building behaviour in the larval tiger beetle,
Cicindela willistoni (Coleoptera: Cicindelidae).
Ecol. Ent. 6:401-410.
Larochelle, A. 1977. Cicindelidae caught at Lights.
Cicindela 9:50-60.
Leffler, S.R. 1979. Tiger beetles of the pacific northwest (Coleoptera: Cicindelidae). Unpub. PhD Thesis. Univ. Wash., Seattle. 731 pp.
Maffei, W. 1989-1996. Unpublished field notes.
Marti, C.D. 1974. Feeding ecology of four sympatric
owls. The Condor. 76(1):45-61.
Nagano, C.D. 1982. The population status of seven
species of insects inhabiting Tijuana Estuary National Wildlife Refuge, San Diego County, California. Report to the Office of Endangered Species.
Pearson, D.L. 1988. Biology of tiger beetles. Ann. Rev.
Ent. 33:123-147.
Pearson, D.L. and C.B. Knisley. 1985. Evidence for food
as a limiting resource in the life cycle of tiger beetles
(Coleoptera: Cicindelidae). Oikos 45:161-168.
Pearson, D.L. and E.J. Mury. 1979. Character divergence and convergence among tiger beetles (Coleoptera: Cicindelidae). Ecology 60:557-566.
Shelford, V.E. 1908. Life histories and larval habits of
the tiger beetles (Cicindelidae). Zool. J. Linn. Soc.
30:157-184.
______. 1912. Ecological succession. Biol. Bull. Woods
Hole Mass. 23:331-370.
Swarth, C.W. 1983. Foraging ecology of snowy plovers
and the distribution of their arthropod prey at
Mono Lake, California. Master’s Thesis, Calif State
Univ., Hayward.
Swiecimski, J. 1956. The role of sight and memory in
food capture by predatory beetles of the species
Cicindela hybrida L. (Coleoptera: Cicindelidae).
Pol. Pismo Ent. 26:205-232.
Willis, H.L. 1967. Bionomics and zoogeography of tiger beetles of saline habitats in the Central United
States (Coleoptera: Cicindelidae). Univ. Kans. Sci.
Bull. 47:145-313.
Zikan, J.J. 1929. Zur Biologie der Cicindeliden
brasiliens. Zool. Anz. 82:269-414.
Additional Readings
Western Tanarthrus Beetle
Tanarthrus occidentalis Chandler
Distribution
Tanarthrus occidentalis was first collected in 1976 and
subsequently described as a new taxon by Chandler in
1979. Specimens were collected from the Cargill salt
pans, now part of the San Francisco Bay National Wildlife Refuge, adjacent to Dum-barton Bridge, Alameda
County, California. Additional populations have been
identified from the salt pans of the Baumberg tract, Hayward, California, and from Bayfarm Island, Alameda,
California. In 1996 the Bayfarm Island population was
extirpated due to modification of their habitat in preparation for anticipated development. Surveys of the south
and central San Francisco Bay area have revealed no
other populations at this time (Maffei, unpub.). Figure
3.7 shows the locations around the Bay Area where T.
occidentalis specimens have been collected, and Table 3.3
shows the collection dates.
Wesley A. Maffei
Description and Systematic Position
Tanarthrus occidentalis is a small beetle, approximately
3-5mm in length, that belongs to the family Anthicidae
(Figure 3.6). The head, pronotum, legs and abdomen
are reddish-orange and the elytra are usually brown or
black with the apical and basal third sometimes reddish
or yellowish in color. This beetle can be separated from
similar bay area Anthicid beetles by noting the distinct
medial constriction of the eleventh antennal segment. It
can further be separated from Formicilla spp., a similar
Wes Maffei
Actual Size
3 - 5 mm
Figure 3.6 Western Tanarthrus Beetle –
Tanarthrus occidentalis
General Sample
Location
Figure 3.7 Known Tanarthrus occidentalis Localities
Within San Francisco Bay Tidal and Diked Marshes
Chapter 3 — Invertebrates
161
Invertebrates
Arnett, R.H. 1968. The beetles of the United States (A
manual for identification). Ann Arbor, Mich.: The
American Entomological Institute, xii + 1112pp.
Dunn, G.W. 1892. Coleoptera and mollusca of the
Ocean Beach at San Francisco. ZOE 2(4):310-312.
Dunn, G.W. 1891. Tiger beetles of California. ZOE
2(2):152-154.
Graves, R.C. 1988. Geographic distribution of the North
American tiger beetle Cicindela hirticollis Say.
Cicindela 20(1):1-21.
Willis. 1968. Artificial key to the species of cicindela of
North America north of Mexico. J. Kans. Ent. Soc.
41(3):303-317.
appearing bay area Anthicid of marshes and grasslands,
by examining the posterior margin of the mesepisternum
which lacks a posterior fringe of long hairs.
Chandler (1979) has indicated that this beetle is
very similar to T. iselini, which is found only in central
New Mexico, but can readily be separated by antennal
morphology.
Table 3.3 Known Collection Sites For Tanarthrus
occidentalis 1
Location
2 mi W. Newark, off
Dumbarton Bridge (salt Pans)
1978
Oliver South #2 Salt Pond,
Hayward
Baumberg Salt Pond #11,
Hayward
Date Specimen(s) Collected
27 May 1976, 15 May
5 Aug 1989
2 Jun 1989, 13 Jun 1989,
5 Aug 1989, 8 Aug 1989,
10 Jul 1997
Reproduction
Unknown.
Significance to Other Wetlands Taxa
This beetle has been identified as part of the immature
Snowy Plover Diet (Page et al. 1995, Feeney and Maffei
1991). Its relationship to other taxa, other than Dictynid
spiders, that utilize abandoned salt crystallizers is unknown at this time.
1
Invertebrates
Information assembled from specimens contained within the California Academy of Sciences Insect Collection, the University of California
Berkeley Essig Museum, the University of California Bohart Museum, the
San Jose State University Edwards Museum, the San Mateo County
Mosquito Abatement District Insect Collection, and the private collections of Dr. J. Gordon Edwards and Wesley A. Maffei.
Suitable Habitat
Tanarthrus occidentalis has been found in no other locality except for abandoned crystallizer ponds and salt
pannes of southern San Francisco Bay. In all instances
these sites remain dry for most of the year except during late winter when temporary pools of rainwater form.
Habitat can be characterized as having extensive areas
of salt crystals interspersed with open areas of fine silt
and very little or no vegetative cover.
Biology
The biology of this beetle is not fully understood. Maffei
(unpub.) has observed the Baumberg tract population
and found that the adults commonly occur out on inactive, salt encrusted crystallizer ponds. These beetles were
observed feeding on the carcasses of the brine flies
Ephydra cinerea and Lipochaeta slossonae (family
Ephydridae) which were still in the webs of unidentified Dictynid spiders. They appeared to function as
“house cleaners” being able to move freely about the web
site unmolested by the resident spider. Peak adult activity
is May through September.
The immature stages of this beetle have not been
located at this time. Larvae of other members of the
beetle family Anthicidae feed on detritus and one species has been recorded as a predator.
162
Baylands Ecosystem Species and Community Profiles
Conservation Needs and Limiting Factors
The conservation needs and limiting factors associated
with this beetle are not very clear. Its association only
with salt encrusted areas, other than the margins of salt
ponds, that remain dry for most of the year appears to
be the primary limiting factor.
References
Chandler, D.S. 1979. A new species of tanarthrus from
California (Coleoptera: Anthicidae). Pan-Pacific
Ent. 55(2):147-148.
Feeney, L.R. and W.A. Maffei. 1991. Snowy plovers and
their habitat at the Baumberg Area and Oliver Salt
Ponds, Hayward, California. Prepared for the City
of Hayward. 162pp.
Maffei, W.A. 1989-1995. Unpublished field notes.
Page, G.W., J.S. Warriner, J.C. Warriner and P.W.C.
Patton. 1995. Snowy plover (Charadrius alexandrinus). In: A. Poole and F. Gill (eds). The birds of
North America, # 154 The Academy of Natural
Sciences, Philadelphia, PA, and The American
Ornithologists’ Union, Washington, D.C.
Additional Readings
Arnett, R.H. 1968. The beetles of the united states (a
manual for identification). Ann Arbor, Mich.: The
American Entomological Institute, xii + 1112pp.
Chandler, D.S. 1975. A revision of Tanarthrus LeConte
with a presentation of its mid-cenozoic speciation
(Coleoptera: Anthicidae). Trans. Amer. Ent. Soc.
101:319-354.
Inchworm Moth
Perizoma custodiata
Wesley A. Maffei
Description and Systematic Position
Distribution
Coastal areas from central northern California south
along the coast of Baja California and including the Gulf
of California. Found throughout San Francisco Bay tidal
and diked salt marshes. Figure 3.9 shows the locations
around the Bay Area where Perizoma custodiata have
been collected, and Table 3.4 shows the collection dates.
Reproduction
The number of generations per season and the number
of eggs per female is apparently unknown for San Francisco Bay populations.
Significance to Other Wetlands Taxa
Snowy plovers have been observed consuming adult
moths at the Baumberg Tract in Hayward, California
(Feeney and Maffei 1991). This insect may also be a part
of other shore bird and passerine bird diets.
The digger wasp, Ammophila aberti, has been observed provisioning its nests with the larvae of this moth
(Maffei, unpub.).
Adult moths are pollinators of Frankenia salina and
are probably pollinators for many of the other flowering
plants within diked and tidal marshes.
Suitable Habitat
Upper middle to high marsh that has berms or levees with
adequate populations of Alkali Heath (Frankenia salina).
Biology
Adults are on the wing from March through November,
with peak adult populations occurring during late spring
and early summer.
Actual Size
3 - 5 mm
Wes Maffei
22 - 29 mm
Figure 3.8 Inchworm Moth – Perizoma custodiata
General Sample
Location
Figure 3.9 Known Perizoma custodiata Localities
Within San Francisco Bay Tidal and Diked
Marshes
Chapter 3 — Invertebrates
163
Invertebrates
Perizoma custodiata is a small moth, with a wingspan of
approximately 22-29mm, that belongs to the family
Geometridae. This moth, commonly known as a measuring worm or inch worm moth, has an alternating pattern of vertical light and dark bands on the fore wings
with plain, pale tan hind wings (Figure 3.8). The variation in width and intensity of the fore wing banding has
caused different entomologists to describe this moth as
a new taxon on four different occasions (Guenee 1857,
Hulst 1896, Packard 1876). Wright (1923) noted the
difficulty in separating examples of the “different species” of the Pacific Coast recognized at that time, stating that they intergrade so much that he found it difficult to tell one from another.
Larvae are a uniform light green or tan in color and
attain a maximum size of approximately 30mm.
Larvae have been observed feeding on Frankenia
salina (Maffei, unpub.) and Packard (1876) has noted
that the larvae of other members of the genus Perizoma
live on low growing plants with the pupa being subterranean. Caterpillars have been observed on Alkali Heath
that was inundated by high tides of 6.3 or greater at the
Whale’s Tail Marsh, Hayward, California. The eggs and
larvae have not been found during the winter months,
and it is presumed that these moths over winter as pupae.
Table 3.4 Known Collection Sites For Perizoma custodiata 1
Invertebrates
Location
Date Specimen(s) Collected
West Pittsburg
15 Feb 1957, 21 Mar 1957,
19 Sep 1957
Martinez
30 Aug 1962
Richmond
18 Jun 1956, 12 Apr 1959
Berkeley
11 Mar 1923, 3 Nov 1923
Berkeley (Bayshore)
27 Jul 1916, 16 May 1955
Alameda
1920
12 May 1918, 13 May
Dumbarton Marsh
22 Jul 1968, 20 Sep 1968,
22 Nov 1968
Shoreline Int. Ctr. (Hwyd)
2 Jul 1990
Baumberg Tract (Hayward)
24 Feb 1990, 1 Apr 1990
Napa
5 May (no year)
Petaluma
13 May 1936, 15 May 1938
Mill Valley (Slough)
Mill Valley
Location
Date Specimen(s) Collected
South Marin Co. Shore
12 Apr 1950
San Francisco
1 Sep 1909, 25 Sep 1909,
9 Oct 1909, 15 Jun 1919,
5 Oct 1919, 21 Oct 1919,
9 Nov 1919, 30 Sep 1920,
4 Oct 1920, 22 Oct 1920,
24 Oct 1920, 11 Dec 1920,
30 Dec 1920, 4 Jan 1921,
6 Sep 1921, 17 Oct 1921,
26 Oct 1922, 14 Jul 1925,
15 Sep 1925
San Francisco (Dunes)
7 Apr 1961
Millbrae
10 Sep 1914
San Mateo
3 Oct 1920
Palo Alto
12 Jun 1933, 27 Jun 1933,
22 Jul 1933, 11 Aug 1933,
26 Apr 1954
17 Jun 1950
E. Palo Alto
May 1978
23 Mar 1920, 5 Sep 1923,
26 Nov 1924, 3 Oct 1926
Bair Island
1 Mar 1987, 9 Mar 1997
17 Apr 1939, 12 May 1940
1
Information assembled from specimens contained within the California Academy of Sciences Insect Collection, University of California Berkeley Essig
Museum, University of California Bohart Museum, San Jose State University Edwards Museum, San Mateo County Mosquito Abatement District Insect
Collection, and private collections of Dr. J. Gordon Edwards and Wesley A. Maffei.
Conservation Needs and Limiting Factors
Frankenia salina has been identified as the larval host
plant for this moth (Maffei, unpub.). Upper middle to
high marsh areas with small dense patches of this plant
support fairly high numbers of this organism. Its wide
distribution along the Pacific Coast would seem to preclude this organism from any immediate danger of extirpation.
Hulst, G.D. 1896. Classification of the Geometrina of
North America. Trans. Amer. Ent. Soc.23:245-386.
Maffei, W.A. 1989-1996. Unpublished field notes.
Packard, A.S. 1876. A Monograph of the geometrid
moths or phalaenidae of the United States. Report
of the U.S. Geological Survey of the Territories.
10:1-607.
Wright, W.S. 1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921.
Proc. Calif. Acad. Sci. 4th Series. 12(9):113-115.
References
Feeney, L.R. and W.A. Maffei. 1991. Snowy plovers and
their habitat at the Baumberg Area and Oliver Salt
Ponds, Hayward, California. Prepared for the City
of Hayward. 162pp.
Guenee, M.A. 1857. Histoire naturelle des insectes. species general des lepidopteres par M.M. Boisduval
et Guenee. Tome 9. Uranides et Phalenites. Vol.
2, 584pp.
164
Baylands Ecosystem Species and Community Profiles
Additional Readings
Hodges, R.W. (ed). 1983. Check list of the Lepidoptera
of America North of Mexico. E.W. Classey Ltd,
Oxfordshire, England. 284pp.
Holland, W.J. 1903. The moth book. Doubleday, Page
and Co., New York. 479pp.
Actual Size
Pygmy Blue Butterfly
egg = 1 mm
Brephidium exilis Boisduval
Wesley A. Maffei
larvae - 13 mm
Description and Systematic Position
Figure 3.11 Brephidium exilis Egg and larva (from
Comstock 1927)
Suitable Habitat
Prefers lowland areas such as alkali flats, salt marshes,
vacant lots, roadsides and desert prairie with various
Chenopodiaceae and Aizoaceae.
Biology
The adult flight period for San Francisco Bay populations is late February through October, with peak abundance occurring in September (Comstock 1927, Garth
and Tilden 1986, Tilden 1965).
Larvae feed on most parts of the host plant. Recorded larval hosts are: Atriplex canescens, A. coulteri, A.
serenana, A. leucophylla, A. patula hastata, A. semibaccata,
A. rosea, A. cordulata, A. hymenelytra, A. coronata, A.
lentiformis breweri, Suaeda fruticosa, S. californica, S.
torreyana, Salicornia virginica, Chenopodium album, C.
Distribution
Brephidium exilis is found from southwestern Louisiana
and Arkansas westward to California and south to Venezuela (Howe 1975, Scott 1986). Strays have been noted
as far north as Kansas and Idaho. This butterfly is widely
distributed throughout the San Francisco Bay, being
particularly abundant in salt marshes (Tilden 1965). Figure 3.12 shows the locations around the Bay Area where
B. exilis have been collected, and Table 3.5 shows the
collection dates.
Actual Size
8 mm
Figure 3.10 Adult Pygmy Blue Butterfly –
Brephidium exilis.
Wes Maffei
13 - 20 mm
General Sample
Location
Figure 3.12 Known Brephidium exilis Localities
Within San Francisco Bay Tidal and Diked
Marshes
Chapter 3 — Invertebrates
165
Invertebrates
Brephidium exilis, also known as the Pygmy Blue, is a
small butterfly, with a wingspan measuring approximately 13-20mm (Figure 3.10). Adult butterflies have
the dorsal surface of the wings brown with the basal third
to half light blue. The ventral surface of the wings are
grayish white with pale brown bands and a row of iridescent black and silver spots along the outer edge of the
hind wing. The eggs are flattened, light bluish-green in
color, and have a fine raised white mesh on the surface.
Larvae are pale green or cream colored and have a finely
punctate surface with white tipped tubercles, a yellowish
white dorsal line, and a bright yellow substigmatal line (Figure 3.11). Some specimens may lack the lateral substigmatal line but all mature larvae have a frosted appearance which resembles the ventral surface of salt bush
leaves or the flower heads of pigweed. The pupae can
be quite variable in color but are usually light brownish
yellow, have a dark brown dorsal line, and have the wing
pads pale yellowish green in color sprinkled with brownish dots.
Three subspecies of this butterfly have been recognized with Brephidium exilis noted as the western subspecies (Scott 1986).
Table 3.5 Known Collection Sites For Brephidium exilis 1
Location
Location
Date Specimen(s) Collected
West Pittsburg
15 Apr 1957
Oakland
8 Apr 1938
Avon
27 Aug 1972
San Leandro
14 Aug 1935
Richmond Point
3 Oct 1964
Milpitas
29 Nov 1974
Richmond
10 Aug 1953
Alviso
1 Nov 1985, 11 Jun 1986
Berkeley (Shoreline)
8 Jun 1915, 22 Jun 1989,
18 Oct 1995
Palo Alto
West Berkeley
20 Jun 1987, 31 Oct 1987,
23 Nov 1987, 25 Jun 1988,
23 Jun 1990
4 Oct 1908, 8 Jun 1909,
1 Oct 1935, Aug 1937,
10 Jul 1967
East Palo Alto
14 Jun 1952
Menlo Park
20 Sep 1958, 9 Oct 1958
12 May 1918, 17 May
Redwood City
28 Jul 1963
Alameda
1918
Larkspur
Invertebrates
Date Specimen(s) Collected
San Mateo
4 Oct 1955, 10 Oct 1955
San Carlos Airport
11 Aug 1977
20 Sep 1958
1
Information assembled from specimens contained within the California Academy of Sciences Insect Collection, University of California Berkeley Essig
Museum, the University of California Bohart Museum, San Jose State University Edwards Museum, San Mateo County Mosquito Abatement District Insect
Collection, and private collections of Dr. J. Gordon Edwards and Wesley A. Maffei.
leptophyllum, Salsola iberica, S. kali tenuifolia, Halogeton
glomeratus, Trianthema portulacastrum, and Sesuvium
verrucosum (Comstock 1927, Garth and Tilden 1986,
Howe 1975, Scott 1986, Tilden 1965).
Nagano and coworkers (1981) found this butterfly to be an indicator of saline soils.
Reproduction
This butterfly has many generations within one season, with one generation often overlapping the next
(Howe 1975). Scott (1986) states that males patrol
all day over the host plants in search of females.
Eggs are laid singly and can be found anywhere on
the host plant, but are usually on the upper surfaces
of leaves. The number of eggs produced per female is
unknown.
Significance to Other Wetlands Taxa
Most likely a prey item for birds utilizing the marshes
of the estuary. Larvae may also be a food item for insectivorous vertebrates. South bay populations of this butterfly are parasitized by the small black tachinid fly
Aplomya theclarum (Maffei, unpub.).
Conservation Needs and Limiting Factors
None.
166
Baylands Ecosystem Species and Community Profiles
References
Comstock, J.A. 1927. Butterflies of California. Los Angeles. Privately Published. 334pp.
Garth, J.S. and J.W. Tilden. 1986. California butterflies.
University of California Press, Berkeley. 246pp.
Howe, W.H. 1975. The butterflies of North America.
Doubleday and Co., New York. 633pp.
Maffei, W.A. 1995. Unpublished field notes.
Nagano, C.D., C.L. Hogue, R.R. Snelling and J.P.
Donahue. 1981. The insects and related terrestrial
arthropods of the Ballona Creek Region. In: The
Biota of the Ballona Region, Los Angeles County,
California. Report to the Los Angeles County Dept.
of Regional Planning. pp. E1-E89.
Scott, J.A. 1986. The butterflies of North America.
Stanford University Press, Stanford. 583pp.
Tilden, J.W. 1965. Butterflies of the San Francisco Bay
Region. University of California Press, Berkeley. 88pp.
Additional Readings
Hodges, R.W. (ed). 1983. Checklist of the Lepidoptera
of America North of Mexico. E.W. Classey
Ltd.Oxfordshire, England. 284 pp.
Summer Salt Marsh Mosquito
Aedes dorsalis (Meigen)
Wesley A. Maffei
Description and Systematic Position
Distribution
This mosquito can be found throughout most of the
United States, southern Canada, Europe and Asia (Carpenter and LaCasse 1955, Darsie and Ward 1981).
Within California, this mosquito can be found in coastal
salt marshes and the brackish waters of the Sacramento
and San Joaquin Delta (Bohart 1956, Bohart and
Washino 1978).
Suitable Habitat
Larvae are found in a variety of brackish and freshwater
habitats throughout their world range (Carpenter and
LaCasse 1955). Within San Francisco Bay A. dorsalis are
usually encountered in temporarily flooded tidal marsh
pannes, heavily vegetated ditches and brackish seasonal
wetlands. Adults prefer open habitats such as grasslands,
open salt marsh and the edges of woodlands.
Biology
Adults are aggressive day biting mosquitoes that have
been found capable of traveling distances of more than
Reproduction
Male mating swarms have been observed occurring over
low growing bushes, prominent objects and open fields
(Dyar 1917, Garcia et al. 1992). Both observations noted
that swarming activity began at sunset and that the
swarms were not more than two to three meters above
the ground. Swarming and mating usually occurs on the
marsh within a few days of adult emergence and is followed by random dispersal of host seeking adults.
Chapter 3 — Invertebrates
167
Invertebrates
The summer salt marsh mosquito, Aedes dorsalis, is a medium sized mosquito measuring approximately 5-6 mm
in length. Freshly emerged adults are one of the most
brightly colored marsh mosquitoes found within the San
Francisco Estuary. These insects are brilliant gold in
color, have a dorsal white band running the length of
the abdomen and have broad white bands on the tarsal
segments of the legs. Older specimens may be yellow or
yellowish-brown in color and the markings on the abdomen may be incomplete if the scales have been rubbed
off. The immature stages can be identified by insertion
of the siphon tuft at or beyond the middle of the siphon
tube, a broadly incomplete anal saddle, presence of a
weak saddle hair and moderate to short anal papillae. The
presence of single upper and lower head hairs has been
used as an additional diagnostic feature but this can be
inconsistent, especially in later instar larvae.
The similarity of this mosquito to Aedes melanimon
Dyar has resulted in some confusion with early efforts to
identify both adults and larvae. Detailed studies of different populations of both of these mosquitoes have helped
to clarify and verify the systematic position of both of these
insects (Bohart 1956, Chapman and Grodhaus 1963).
30 miles (Rees and Nielsen 1947). Flights of adults in
Alameda County have been known to disperse distances
of more than five miles from their larval source (Maffei,
unpub.). Garcia and Voigt (1994) studied the flight potential of this mosquito in the lab and found that the
adults exhibited strong flight characteristics which they
believed helped them to adapt to the strong winds encountered in their preferred open habitats. Females are
readily attracted to green, grassy fields and will rest there
waiting for available hosts (Maffei, unpub.).
Host studies have shown that large mammals are
preferred, especially cattle and horses (Edman and
Downes 1964, Gunstream et al. 1971, Shemanchuk et
al. 1963, Tempelis et al. 1967). The effects of adult feeding activity on livestock can be severe resulting in reduced feeding and in some instances injury to animals
attempting to evade severe attacks. Recent adult activity within the San Francisco Estuary has impacted outdoor school activities, businesses and residents, resulting in at least two instances where medical attention was
required for people reacting to multiple bites (Maffei,
unpub.).
Eggs are deposited individually on the mud along
the edges of tidal pools or the receding water line of
brackish seasonal wetlands. Winter is passed in the egg
stage and hatching occurs with the first warm weather
of spring. Additional hatches occur with subsequent
refloodings of the larval habitat. Eggs can remain viable
for many years with only part of any given brood hatching during any single flooding event.
The larval stage can last from four to fourteen days
with duration being primarily dependent on temperature. Other factors that can regulate rate of larval development include competition for space and quality and
availability of nutrients. Rees and Nielsen (1947) found
larvae that completed their development in saline pools
of the Great Salt Lake with salt concentrations as high
as 120 ‰. Washino and Jensen (1990) reared larvae,
from Contra Costa County salt marshes, in solutions
simulating 0, 10, 50 and 100% concentrations of seawater and found that survivorship improved as salt content approached that of seawater.
Total developmental time, from egg to adult, has
been observed to occur in less than one week (Maffei,
unpub.).
The number of gonotrophic cycles and eggs produced per female remains unclear for San Francisco Bay
populations. Early work by Telford (1958) found that
12 broods and approximately eight generations occurred
during one breeding season at Bolinas in Marin County.
The number of generations per year does vary with respect to weather and tidal conditions.
Invertebrates
Significance to Other Wetlands Taxa
This species of mosquito is commonly found in association with the tidal pool brine fly Ephydra millbrae and
the water boatman Trichocorixa reticulata. Both the brine
fly and the water boatman have been identified as food
sources for shorebirds and waterfowl (Anderson 1970;
Feeney and Maffei 1991; Howard 1983; Maffei, unpub.;
Martin and Uhler 1939). The larvae of this mosquito
may also be a food source for these birds and adults may
be a food source for swallows.
Conservation Needs and Limiting Factors
This mosquito, like other species of mosquitoes, is extremely opportunistic. Care must be taken when altering or restoring seasonal or tidal wetlands. Sites that
drain poorly will create habitat that can readily produce
very large numbers of aggressive biting adults. Plans for
long term maintenance of seasonal and tidal wetlands
should include resources for mosquito control as the
need arises. The dynamic nature of these types of habitats
coupled with human activities can easily convert a nonbreeding site into a major mosquito producing source.
References
Anderson, W. 1970. A preliminary study of the relationship of salt ponds and wildlife—South San
Francisco Bay. Calif. Dept. Fish and Game
56(4):240-252.
Bohart, R.M. 1948. Differentiation of larvae and pupae
of Aedes dorsalis and Aedes squamiger. Proc. Ent.
Soc. Wash. 50:216-218.
______. 1956. Identification and distribution of Aedes
melanimon and Aedes dorsalis in California. Proc.
C.M.C.A. 24:81-83.
Bohart, R.M. and R.K. Washino. 1978. Mosquitoes of
California. Third Ed. Univ. Calif. Div. Agr. Sci.,
Berkeley, Publ. 4084. 153 pp.
Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of
North America. Univ. Calif. Press, Berkeley. 360 pp.
168
Baylands Ecosystem Species and Community Profiles
Chapman, H.C. and G. Grodhaus. 1963. The separation of adult females of Aedes dorsalis (Meigen) and
Ae. melanimon Dyar in California. Calif. Vector
Views 10(8):53-56.
Darsie, R.F. and R.A. Ward. 1981. Identification and
geographical distribution of the mosquitoes of
North America, North of Mexico. Mosq. Syst.
Suppl. 1:1-313.
Dyar, H.G. 1917. Notes on the Aedes of Montana
(Diptera: Culicidae). Ins. Ins. Mens. 5:104-121.
Edman, J.D. and A.E.R. Downe. 1964. Host blood
sources and multiple-feeding habits of mosquitoes
in Kansas. Mosq. News 24:154-160.
Feeney, L.R. and W.A. Maffei. 1991. Snowy plovers and
their habitat at the Baumberg Area and Oliver Salt
Ponds, Hayward, California. Prepared for the City
of Hayward. 162pp.
Garcia, R. and W.G. Voigt. 1994. Flight potential of three
salt marsh mosquitoes from San Francisco Bay. Ann.
Rep. Mosq. Cont. Res., Univ. Calif. pp. 46-48.
Gunstream, S.E., R.M. Chew, D.W. Hagstrum and
C.H. Tempelis. 1971. Feeding patterns of six species of mosquitoes in arid Southeastern California. Mosq. News 31:99-101.
Howard, J.A. 1983. Feeding ecology of the ruddy duck
on the San Francisco Bay National Wildlife Refuge. M.A. Thesis, San Jose State University.53 pp.
Maffei, W.A. 1990-1997. Unpublished field notes.
Martin, A.C. and F.M. Uhler. 1939. Food of game ducks
in the United States and Canada. USDA. Tech.
Bull. #634. 156pp.
Rees, D.M. and L.T. Nielsen. 1947. On the biology
and control of Aedes dorsalis (Meigen) in Utah.
Proc. N.J. Mosq. Exterm. Assoc. 34:160-165.
Shemanchuk, J.A., A.E.R. Downe and L. Burgess. 1963.
Hosts of mosquitoes (Diptera: Culicidae) from the
irrigated pastures of Alberta. Mosq. News.
23(4):336-341.
Telford, A.D. 1958. The pasture Aedes of Central and
Northern California. Seasonal History. Ann. Ent.
Soc. Amer. 51:360-365.
Tempelis, C.H., D.B. Francy, R.O. Hayes and M.F. Lofy.
1967. Variations in feeding patterns of seven culicine mosquitoes on vertebrate hosts in Weld and
Larimer Counties, Colorado. Amer. J. Trop. Med.
Hyg. 7:561-573.
Washino, R.K. and T. Jensen. 1990. Biology, ecology
and systematics of Aedes dorsalis, Ae. melanimon,
and Ae. campestris in Western North America. Ann.
Rep. Mosq. Cont. Res., Univ. Calif. pp 40-41.
lar, with additional sites having been identified along the
shoreline of the East Bay.
Winter Salt Marsh Mosquito
Aedes squamiger (Coquillett)
Suitable Habitat
Wesley A. Maffei
Description and Systematic Position
Distribution
This mosquito is found along the Pacific Coast region
from Marin and Sonoma counties, California, south to
Baja California, Mexico (Bohart and Washino 1978,
Carpenter and LaCasse 1955, Darsie and Ward 1981,
Freeborn and Bohart 1951). Figure 3.15 shows the distribution of Aedes squamiger in 1950. The current distribution within the San Francisco Bay area is very simi-
Biology
Eggs hatch as early as late September and can continue
to hatch with the accumulation of rainfall from each
successive storm event. Maffei (unpub.) found larvae that
hatched from the incidental flooding of a marsh by a
duck club as early as late September. Bohart, et. al.
(1953) states that three to six major hatches of eggs occur during the fall months. It is believed that only part
of the eggs laid during the prior spring season hatch with
a decreasing percentage of the remaining eggs hatching
during successive years. Garcia, et al. (1991) found that
as many as four floodings were necessary to hatch all of
the eggs from field collected samples. Bohart and
Washino (1978) state that the eggs are usually dormant
from April through September and that this obligatory
diapause is terminated by the decreasing fall temperatures that fall below 7°C. Garcia et al. (1991) found that
hatching does not occur until the eggs have been exposed
to temperatures that are less than 10°C. Voigt (pers
comm.) believes that once the eggs have been thermally
Actual Size
12 mm
Figure 3.13 Adult Winter Salt Marsh Mosquito –
Aedes squamiger
Wes Maffei
Wes Maffei
8 mm
Figure 3.14 Terminal Abdominal Segment of a
Fourth Instar Larva
Chapter 3 — Invertebrates
169
Invertebrates
Aedes squamiger is a medium-sized to large mosquito,
measuring approximately 6-9mm in length, that belongs
to the fly family Culicidae (Figure 3.13). Adults have a
distinctive black and white speckled appearance and
large, flat scales along the wing veins which separates this
fly from other San Francisco Bay mosquitoes. Larvae can
be identified by the presence of an incomplete anal
saddle, a siphon tuft distal to the pecten row, an anal
saddle hair as long or longer than the anal saddle, and
upper and lower head hairs that are usually branched
(Figure 3.14).
This mosquito was described as a new taxon by
Coquillett in 1902 from specimens collected from the
cities of Palo Alto and San Lorenzo, California. Bohart
(1948) differentiated the larvae and pupae of Aedes dorsalis and Aedes squamiger thereby providing a means of
separating the immature stages of these two species
which are very similar in appearance. In 1954, Bohart
described and provided keys to the first stage larvae of
California Aedes and further clarified the differences
between these two mosquitoes.
Preferred habitat consists primarily of coastal pickle weed
tidal and diked marshes, especially salt marsh pools that
are diluted by winter and early spring rains. Cracked
ground of diked wetlands and old dredge disposal sites
are also a favorite habitat for deposition of eggs and development of larvae. This mosquito prefers brackish or
saline habitats and has not been found in truly fresh
water marshes. Bohart, et. al. (1953) found larvae of
various stages in pools with salinities ranging from 1.2
‰ to 35 ‰. Studies by Garcia and coworkers (1992,
1991) indicated that optimal larval development occurred at salinities between 5 ‰ and 15 ‰.
Figure 3.15 Aedes
Squamiger Distribution
in the San Francisco Bay
Area, 1950
Invertebrates
From Aarons, 1954
conditioned that hatching can then occur anytime in the
future following submersion. This may possibly help to
explain summer hatches following flooding of sites by
inadvertant human activity (Maffei, unpub.).
Larvae are principally found in salt marsh pools that
are diluted by fall and winter rainfall. Bohart and coworkers (1953) found that a minimum of 48 days were required for the development of the aquatic stages before
adult emergence, with the first pupae having been found
during the first week of February. Under “normal” conditions pupae are usually found from the last two weeks
of February through the beginning of March. Estimates
of the number of larvae per acre vary from 1.65 million
to 1.45 billion depending on environmental factors
(Aarons 1954, Aarons et al 1951, Lowe 1932). Larvae
are capable of remaining submerged for extended periods of time where they browse on vegetation and mud.
Garcia, et al (1990) calculated the minimum developmental threshold for development of larvae to the adult
170
Baylands Ecosystem Species and Community Profiles
stage to be 4.4°C. Additional studies by Garcia and coworkers (1991) found that first and second instar larvae
had developmental thresholds that were 2-4°C lower
than the later instars. From these data , they concluded
that the lower developmental thresholds of the earlier
instars allowed larvae from later hatching installments
to emerge as adults in closer synchrony with those larvae that hatched earlier in the season. They also noted
that larvae and pupae could survive in the mud at sites
that underwent periodic draw-down of the water. Garcia,
et al. (1990) also studied the diapause habit of the last
instar larvae and concluded that this interesting trait
probably contributed in some degree to the partial synchronous emergence of the adults.
Adults usually emerge during the last week of February through the end of March. Emergence of adults
in April has occurred from unusually heavy late winter
and early spring rains that have caused late egg hatches
with rapid larval development. Adults usually fly to ar-
Reproduction
Observations on mating swarms have shown that Aedes
squamiger tends to swarm approximately one hour before to one-half hour after sunset (Garcia et. al. 1992).
Swarms can consist of a few to several thousand indi-
viduals that hover over prominent objects such as trees
or large bushes and can occur at heights ranging from
six to approximately 50 feet (Bohart and Washino 1978,
Garcia et. al. 1992). Garcia et al. (1992, 1983) found
that adults traveled back and forth to the marshes quite
readily producing a new batch of eggs with each trip. He
also found that the highest parous condition observed
was seven, with average parity rates ranging between 3
and 5.4. Garcia, et al. (1992) found a direct correlation
between wing length and the number of eggs produced
with larger females producing more eggs. The maximum
number of eggs produced per female was less than 250.
Garcia, et al. (1990) also found that temperature played
an important role in longevity, ovarian development and
oviposition. Females held at 15°C were still alive 50 days
after their last blood meal and average longevity was
about 35 days when kept at 20°C. The minimum temperature threshold for ovarian development or oviposition was found to be about 15°C.
Significance to Other Wetlands Taxa
Aedes squamiger larvae are frequently found in association with larvae of the Summer Salt Marsh Mosquito,
Aedes dorsalis, and the Winter Marsh Mosquito, Culiseta
inornata. The adults of these mosquitoes may be a possible food source for swallows and the larvae may be a
food source for waterfowl.
Conservation Needs and Limiting Factors
This mosquito, like other species of mosquitoes, is extremely opportunistic. Care must be taken when altering or restoring seasonal or tidal wetlands. Sites that
drain poorly will create habitat that can readily produce
very large numbers of aggressive biting adults. Plans for
long term maintenance of seasonal and tidal wetlands
should include resources for mosquito control as the
need arises. The dynamic nature of these types of habitats coupled with human activities can easily convert a
non-breeding site into a major mosquito producing source.
References
Aarons, T. 1954. Salt marsh mosquito survey in the San
Francisco Bay Area 1950-1953. Proc. C.M.C.A.
22:75-78.
Aarons, T., J.R. Walker, H.F. Gray and E.G. Mezger.
1951. Studies of the flight range of Aedes
Squamiger. Proc. C.M.C.A. 19:65-69.
Bohart, R.M. 1948. Differentiation of larvae and pupae
of Aedes dorsalis and Aedes squamiger. Proc. Ent.
Soc. Wash. 50:216-218.
______. 1954. Identification of first stage larvae of California Aedes (Diptera: Culicidae). Ann. Ent. Soc.
Amer. 47:356-366.
Chapter 3 — Invertebrates
171
Invertebrates
eas away from their breeding sites, using ravines and
natural or man made waterways from the marshes to the
local hills as passageways. From these passageways the
adults spread laterally into the wind protected areas of
the surrounding community (Freeborn 1926). It is believed that at these protected sites adults mate and and
seek blood meals (Telford 1958). Gray (1936) noted that
this mosquito flew the longest distance of any California mosquito from its larval source. Aarons (1954) noted
that adults were found in Saratoga, some 10 miles from
the nearest known larval source. Other workers have
found that adults of this mosquito are capable of traversing distances of more than 15 miles from any possible
larval site (Aarons, et. al. 1951, Krimgold and Herms
1934, Lowe 1932, Stover 1931, Stover 1926). Biting
activity begins in April and usually ends by early June.
Rabbit baited traps in the east bay have collected adults
from 16 March to 28 June (Garcia et al. 1983). Adults
are known to be aggressive day and early dusk biting
mosquitoes. This species along with the Summer Salt
Marsh Mosquito, Aedes dorsalis, were the first mosquitoes to become the primary focus of organized mosquito
control efforts in California. The first mosquito control
campaigns were undertaken at San Rafael in 1903 and
also at Burlingame in 1904. The earliest written record
of what is believed to be the attacks of Aedes squamiger
and Aedes dorsalis on humans was in a diary entry of
Father Juan Crespi in April of 1772 (Bolton 1927). In
his diary he describes the vicious attacks of mosquitoes
that sorely afflicted his party while traveling along the
eastern side of San Francisco Bay. Aarons, et al. (1951)
states that there is reason to believe that the salt marsh
mosquitoes made certain times of the year almost unbearable for the early Indians.
Females oviposit in those parts of the marshes that
are not under water. Eggs are laid on plants and along
the muddy margins of ponds close to the water line. Most
of the eggs are located in these higher areas of the
marshes and will therefore not hatch without a combination of tides and rainfall. For diked marshes, at least
a few inches of rainfall must occur to inundate the eggs
and stimulate hatching. Maffei (unpub.) has found that
the runoff of as little as one inch of rainfall from city
streets into marshes used as flood control basins can flood
a marsh sufficiently to hatch eggs and produce larvae.
Females that oviposit in late spring will deposit eggs in
the lower portions of the marshes and it is these eggs that
hatch first with tidal activity only or ponding of early rain
water runoff.
Invertebrates
Bohart, R.M. and R.K. Washino. 1978. Mosquitoes of
California. Third Edition. Univ. Calif. Div. Agr.
Sci., Berkeley, Publ. 4084. 153pp.
Bohart, R.M., E.C. Mezger and A.D. Telford. 1953.
Observations on the seasonal history of Aedes
squamiger. Proc. C.M.C.A. 21:7-9.
Bolton, H.E. 1927. Journal of Father Juan Crespi. Univ.
Calif. Press, Berkeley.
Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of
North America. Univ. Calif. Press, Berkeley. 360pp.
Coquillett, D.W. 1902. New Diptera from North
America. Proc. U.S.N.M. 25:83-126.
Darsie, R.F. and R.A. Ward. 1981. Identification and
geographical distribution of the mosquitoes of
North America, North of Mexico. Mosq. Syst.
Suppl. 1:1-313.
Freeborn, S.B. 1926. The mosquitoes of California.
Univ. Calif. Publ. Ent. 3:333-460.
Freeborn, S.B. and R.M. Bohart. 1951. The mosquitoes of California. Bull. Calif. Insect Survey. 1:2578.
Garcia, R., B. Des Rochers and W. Tozer. 1983. Biology
and ecology of Aedes squamiger in the San Francisco Bay Area. Ann. Rep. Mosq. Cont. Res., Univ.
Calif. pp. 29-31.
Garcia, R., W.G. Voigt and A.K. Nomura. 1992. Ecology of Aedes Squamiger in the Northern San Francisco Bay Area. Ann. Rep. Mosq. Contr. Res., Univ.
Calif. pp. 53-57.
Garcia, R., W.G. Voigt, A.K. Nomura and A. Hayes.
1991. Biology of Aedes squamiger. Ann. Rep. Mosq.
Cont. Res., Univ. Calif. pp 51-52.
______. 1992. Biology of Aedes squamiger. Unpub.
Progress Report for Alameda County Mosquito
Abatement District. 7 pp.
Gray, H.F. 1936. Control of pest mosquitoes for comfort. Civil Eng. 6(10):685-688.
Krimgold, D.B. and H.P. Herms. 1934. Report on salt
marsh breeding and migration of the San Francisco Bay Area in March 1934. Unpublished Report.
Lowe, H.J. 1932. Studies of Aedes Squamiger in the San
Francisco Bay Region. Proc. C.M.C.A., Paper No.
1.
Maffei, W.A. 1990-1996. Unpublished field notes.
Stover, S.E. 1926. Eradication of salt marsh mosquitoes. Weekly Bull., Calif. State Board of Health.
Vol. V(44).
______. 1931. Conference notes and discussion on flight
habits of salt marsh mosquitoes. Proceedings of
the California Mosquito Control Association. 2:11
& 14.
Telford, A.D. 1958. The pasture Aedes of Central and
Northern California. Seasonal History. Ann. Ent.
Soc. Amer. 51:360-365.
172
Baylands Ecosystem Species and Community Profiles
Additional Readings
Meyer, R.P. and S.L. Durso. 1993. Identification of the
mosquitoes of California. Calif. Mosquito and
Vector Control Assoc., Sacramento. 80pp.
Quayle, H.J. 1906. Mosquito control work in California. Univ. Calif. Agric. Exp. Sta. Bull. #178, pp 155.
Washino’s Mosquito
Aedes washinoi Lansaro and Eldridge
Wesley A. Maffei
Description and Systematic Position
Aedes washinoi was described as a new taxon by Lanzaro
and Eldridge in 1992 and was determined to be a sibling species of Aedes clivis and Aedes increpitus. Prior to
1992, all three species of mosquitoes were known as
Aedes increpitus. Adults of this mosquito are almost impossible to separate from its sibling species, when using
morphological features, and can also sometimes be confused with Aedes squamiger. The easiest way to distinguish Ae. squamiger and Ae. washinoi is to examine the
wing scales. Aedes squamiger has very broad, flat, platelike scales on the wings whereas Ae. washinoi will have
the usual thin, pointed wing scales. The wings of Ae.
washinoi will also tend to be uniformly dark with a concentration of pale scales on the anterior wing veins. In
all other respects, both Ae. squamiger and Ae. washinoi
share a similar black and white speckled appearance. The
larvae of this mosquito can be difficult to separate but
Darsie (1995) has provided additions to Darsie and
Wards 1981 keys to facilitate identification.
Distribution
This mosquito is found from Portland, Oregon south
to Santa Barbara, California and eastward into the lower
Sierra Nevada mountains. Populations of this mosquito
have also been found along the eastern Sierra Nevada
Range at Honey Lake.
Suitable Habitat
Within the San Francisco Estuary the preferred habitat
is shallow ground pools and upland fresh to slightly
brackish water sites that are next to salt marshes or in
riparian corridors. These habitats also tend to be dominated by willow or cotton wood trees and/or black berry
vines.
Reproduction
Adults have been observed swarming under or near the
tree canopy of their larval habitat (Garcia, et al. 1992).
Significance to Other Wetlands Taxa
Unknown.
Conservation Needs and Limiting Factors
This mosquito, like other species of mosquitoes, is extremely opportunistic. Care must be taken when altering or restoring seasonal wetlands or riparian corridors.
Sites that have shallow ground pools and willow or cotton wood trees or blackberry vines will create habitat that
can readily produce very large numbers of aggressive biting adults. The restoration of historical willow groves
should not occur if homes are within two miles of the
project site.
References
Darsie, R.F. 1995. Identification of Aedes tahoensis, Aedes
clivis and Aedes washinoi using the Darsie/Ward
keys (Diptera: Culicidae). Mosq. Syst. 27(1):4042.
Darsie, R.F. and R.A. Ward. 1981. Identification and
geographical distribution of the mosquitoes of
North America, north of Mexico. Mosq. Syst.
Suppl. 1:1-313.
Garcia, R., W.G. Voigt and A.K. Nomura. 1992. Ecology of Aedes squamiger in the northern San Francisco Bay Area. Ann. Rep. Mosq. Contr. Res., Univ.
Calif. pp. 53-57.
Additional Readings
Bohart, R.M. and R.K. Washino. 1978. Mosquitoes of
California. Third Edition. Univ. Calif. Div. Agr.
Sci., Berkeley, Publ. 4084. 153 pp.
Freeborn, S.B. and R.M. Bohart. 1951. The mosquitoes
of California. Bull. Calif. Insect Survey. 1:25-78.
Meyer, R.P. and S.L. Durso. 1993. Identification of the
mosquitoes of California. Calif. Mosquito and Vector Control Assoc., Sacramento. 80 pp.
Invertebrates
Larvae usually hatch during early winter after a series of
successive storm events has filled ground depressions
with water. Additional hatches of larvae can occur if late
winter and early spring rains refill drying larval sites.
Larvae of this mosquito also exhibit a late fourth instar
diapause and partial synchronous adult emergence similar to that observed in Aedes squamiger. Adults emerge
during late winter and early spring and can persist
through early June, depending on weather conditions.
Females are aggressive day biting mosquitoes that
tend not to travel far from their larval sources. Maffei
(unpub.) found that adult mosquitoes traveled a maximum distance of one and one-half miles from their larval habitat and that local, man made canals were used
as a passageway into the surrounding community.
Eggs are deposited in the muddy margins adjacent
to the receding water line of the larval habitat and hatch
the following winter when reflooded.
Lanzarro, G.C. and B.F. Eldridge. 1992. A classical and
population genetic description of two new sibling
species of Aedes (Ochlerotatus) increpitus Dyar.
Mosq. Syst. 24(2):85-101.
Maffei, W.A. 1990-1995. Unpublished field notes.
Western Encephalitis Mosquito
Culex tarsalis Coquillett
Wesley A. Maffei
Description and Systematic Position
The western encephalitis mosquito is a medium sized
mosquito measuring approximately 5-6 mm in length.
This fly was described in 1896 as a new taxon by
Coquillett from specimens gathered in the Argus Mountains of Inyo County, California (Belkin et al. 1966).
Adults can be identified by using the following
morphological features: legs with bands of pale scales
overlapping the tarsal joints; femur and tibia of the hind
legs with a pale stripe or row of pale spots on the outer
surface; proboscis with a complete median pale band;
ventral abdominal segments with v-shaped patches of
darkened scales; and the inner surface of the basal antennal segment with patches of pale scales. The larvae
can be recognized by the four to five pairs of ventrally
located siphon tufts that are nearly in line with each
other (Figure 3.16) and the 3-branched lateral abdominal hairs found on segments III to VI.
Wes Maffei
Biology
Figure 3.16 Terminal Abdominal Segment of
C. tarsalis larva
Chapter 3 — Invertebrates
173
The immature stages are found in all types of fresh water habitats except treeholes. Poorly drained pastures, rice
fields, seepages, marshes and duck club ponds are especially favored as breeding habitat for this mosquito.
Telford (1958) found larvae in salt marsh pools with salinities up to 10 ‰. Urban sources include poorly maintained swimming pools, ornamental ponds, storm drains,
flood control canals, ditches, waste water ponds and most
man made containers (Beadle and Harmston 1958,
Bohart and Washino 1978, Harmston et al. 1956, Meyer
and Durso 1993, Sjogren 1968).
Adults rest by day in shaded or darkened areas such
as mammal burrows, tree holes, hollow logs, under
bridges, in caves, in eves and entry ways of residences,
brush piles and in dense vegetation (Mortenson 1953,
Loomis and Green 1955, Harwood and Halfill 1960,
Price et al. 1960, Rykman and Arakawa 1952).
periods of low temperatures or unseasonably warm winters can vary the time spent in diapause.
Flight range studies indicate that this mosquito will
readily disperse from its larval source. Reeves et al. (1948)
found that adults generally dispersed two miles or less,
although prevailing winds helped to distribute marked
females up to three miles. Bailey et al. (1965) studied
the dispersal patterns of Yolo County, California populations and found that prevailing winds were important
to adult dispersal with significant numbers of adults
having traveled seven miles within two nights. The
maximum distance traveled was recorded at 15.75 miles.
From their studies they concluded that the likely dispersal distance of Sacramento Valley populations was
probably about 20-25 miles. It was further concluded
that most locally controlled mosquito sources are repeatedly reinfested during the summer because these mosquitoes travel so readily with the wind.
The larval stages feed on a wide variety of microorganisms, unicellular algae and microscopic particulate
matter. The amount of time required to complete development from egg to adult varies depending on water
temperature, availability of food and crowding. Bailey
and Gieke (1968) found that water temperatures of 69°F
to 86°F were optimal for larval development. Beyond
86°F, the larval stage lasted about eight days but mortality was very high. Mead and Conner (1987) found the
average developmental rates from egg to adult to be 18.7
days at 67°F and 7.4 days at 88°F.
Biology
Reproduction
Adult females of this species usually feed at night. Precipitin tests indicate a wide variety of hosts consisting
of various birds and mammals with an occasional reptile or amphibian (Anderson et al. 1967, Edman and
Downe 1964, Gunstream et al. 1971, Hayes et al.
1973, Reeves and Hammon 1944, Rush and Tempelis
1967, Shemanchuk et al. 1963, Tempelis 1975, Tempelis et al. 1967, Tempelis et al. 1965, Tempelis and
Washino 1967). Reeves (1971) states that host availability and season are probably the most important considerations in the adult host feeding pattern. The availability
of nesting birds during spring and early summer may
account for the preponderance of identified, early season, avian blood meals. With the progression of the summer season, availability and behaviour of bird hosts varies and a switch to mammal hosts occurs (Hammon et
al. 1945, Hayes et al. 1973, Reeves and Hammon
1944, Reeves et al. 1963, Tempelis et al. 1967, Tempelis and Washino 1967). Adults pass the winter months
in facultative diapause which is triggered by short day
length and low ambient temperatures. In the warmer
parts of southern California adults are active year round
while in San Francisco Bay populations inactivity usually occurs from December through February. Additional
Male mating swarms occur shortly before to just after
sunset. Harwood (1964) found that initiation of the
mating swarm was related to changes in the light intensity and that light levels of approximately 7 foot candles
would initiate crepuscular flight activity. He further
found that lab colonized males could be induced to
swarm when abrupt changes in light intensity occurred.
Lewis and Christenson (1970) studied female ovipositional behaviour and found that the initial search for
oviposition sites by females occurs close to the lowest
available surface. Groups of eggs, also known as egg rafts,
are deposited directly onto the water with the average
number of eggs per raft varying between 143 to 438
(Bock and Milby 1981, Buth et al. 1990, Reisen et al.
1984). Environmental factors such as water temperature,
crowding and availability of food have been found to
affect development of the immature stages, which in
turn, affects the size of the female mosquito and ultimately the number of eggs and egg rafts produced. Logan and Harwood (1965) studied the effects of photoperiod on ovipositional behaviour of a Washington strain
of Culex tarsalis and found that peak oviposition occurred
within the first hour of darkness and light.
Distribution
This mosquito has been found in central, western and
southwestern United States, southwestern Canada
and northwestern Mexico (Carpenter and LaCasse
1955, Darsie and Ward 1981). Within California, this
fly has been found in every county from elevations below sea level to almost 10,000 feet (Bohart and
Washino 1978, Meyer and Durso 1993).
Invertebrates
Suitable Habitat
174
Baylands Ecosystem Species and Community Profiles
Autogeny, or the development of eggs without a
blood meal, does occur with this mosquito. Moore
(1963) found that autogenous Culex tarsalis, from Sacramento Valley, California, produced an average of 116
eggs per female with an observed maximum of 220. He
also found that the level of autogeny decreased from
spring to summer. Spadoni et al. (1974) also studied
autogeny in Culex tarsalis populations from the same
region finding similar results and detecting autogeny as
early as April. They further found that no autogenous
egg development was observed in overwintering females from November through February and that the
mean number of eggs produced per autogenous female
was 144.
Significance to Other Wetlands Taxa
Conservation Needs and Limiting Factors
Sound water management practices should include consultations with local public health and mosquito or vector control agencies to prevent or at least minimize the
production of this mosquito from managed, restored or
newly created wetlands. Adequate resources need to be
provided in all short and long term management plans
to help protect humans and horses from the encephalitis viruses that can be vectored by this mosquito.
References
Andersen, D.M., G.C. Collett and R.N. Winget. 1967.
Preliminary host preference studies of Culex tarsalis
Coquillett and Culiseta inornata (Williston) in
Utah. Mosq. News 27:12-15.
Bailey, S.F. and P.A. Gieke. 1968. A study of the effect
of water temperatures on rice field mosquito development. Proc. Calif. Mosq. Cont. Assoc. 36:5361.
Bailey, S.F., D.A. Eliason and B.L. Hoffman. 1965.
Flight and dispersal of the mosquito Culex tarsalis
Coquillett in the Sacramento Valley of California.
Hilgardia 37(3):73-113.
Chapter 3 — Invertebrates
175
Invertebrates
This mosquito is the primary vector of Western Equine
Encephalitis (WEE) and Saint Louis Encephalitis (SLE)
viruses for most of the western United States (Brown
and Work 1973, Longshore et al. 1960, Reeves and
Hammon 1962, Work et al. 1974). Rosen and Reeves
(1954) have also determined that this fly is an important vector of avian malaria.
Larvae of the Winter Marsh Mosquito, Culiseta
inornata, are frequently found with the immature
stages of this mosquito during fall and spring. The
larvae of this insect may be a possible food source for
waterfowl.
Beadle, L.D. and F.C. Harmston. 1958. Mosquitoes in
sewage stabilization ponds in the Dakotas. Mosq.
News 18(4):293-296.
Belkin, J.N., R.X. Schick and S.J. Heinemann. 1966.
Mosquito studies (Diptera: Culicidae) VI. Mosquitoes originally described from North America.
1(6):1-39.
Bock, M.E. and M.M. Milby. 1981. Seasonal variation
of wing length and egg raft size in Culex tarsalis.
Proc. C.M.V.C.A. 49:64-66.
Bohart, R.M. and R.K. Washino. 1978. Mosquitoes of
California. Third Edition. Univ. Calif. Div. Agr.
Sci., Berkeley, Publ. 4084. 153 pp.
Brown, D. and T.H. Work. 1973. Mosquito transmission of arboviruses at the Mexican Border in Imperial Valley, California 1972. Mosq. News 33:381385.
Buth, J.L., R.A. Brust and R.A. Ellis. 1990. Development time, oviposition activity and onset of diapause in Culex tarsalis, Culex restuans and Culiseta
inornata in Southern Manitoba. J. Amer. Mosq.
Cont. Assoc. 6(1):55-63.
Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of
North America. Univ. Calif. Press, Berkeley. 360pp.
Darsie, R.F. and R.A. Ward. 1981. Identification and
geographical distribution of the mosquitoes of
North America, North of Mexico. Mosq. Syst.
Suppl. 1:1-313.
Gunstream, S.E., R.M. Chew, D.W. Hagstrum and
C.H. Tempelis. 1971. Feeding patterns of six species of mosquitoes in arid Southeastern California. Mosq. News 31:99-101.
Hammon, W. McD., W.C. Reeves and P. Galindo. 1945.
Epidemiologic studies of encephalitis in the San
Joaquin Valley of California 1943, with the isolation of viruses from mosquitoes. Amer. J. Hyg.
42:299-306.
Harmston, F.C., G.B. Schultz, R.B. Eads and G.C.
Menzies. 1956. Mosquitoes and encephalitis in the
irrigated high plains of Texas. Publ. Hlth. Repts.
71(8):759-766.
Harwood, R.F. 1964. Physiological factors associated
with male swarming of the mosquito Culex tarsalis
Coq. Mosq. News 24:320-325.
Harwood, R.F. and J.E. Halfill. 1960. Mammalian burrows and vegetation as summer resting sites of the
mosquitoes Culex tarsalis and Anopheles freeborni.
Mosq. News 20:174-178.
Hayes, R.O., C.H. Tempelis, A.D. Hess and W.C.
Reeves. 1973. Mosquito host preference studies
in Hale County, Texas. Amer. J. Trop. Med. Hyg.
22:270-277.
Lewis, L.F. and D.M. Christenson. 1970. Ovipositional
site selection in cages by Culex tarsalis as influenced by container position, water quality, and female age. Proc. Utah Mosq. Abate. Assoc. 23:27-31.
Invertebrates
Logen, D. and R.F. Harwood. 1965. Oviposition of the
mosquito Culex tarsalis in response to light cues.
Mosq. News 25(4):462-465.
Longshore, W.A., E.H. Lennette, R.F. Peters, E.C.
Loomis and E.G. Meyers. 1960. California encephalitis surveillance program. Relationship of
human morbidity and virus isolation from mosquitoes. Amer. J. Hyg. 71:389-400.
Loomis, E.C. and D.H. Green. 1955. Resting habits of
adult Culex tarsalis Coquillett in San Joaquin
County, California, November, 1953 Through
November, 1954. A preliminary report. Proc. Calif. Mosq. Cont. Assoc. 23:125-127.
Mead, S.S. and G.Conner. 1987. Temperature related
growth and mortality rates of four mosquito species. Proc. C.M.V.C.A. 55:133-137.
Meyer, R.P. and S.L. Durso. 1993. Identification of the
mosquitoes of California. Calif. Mosq. Cont.
Assoc., Sacramento, Calif. 80 pp.
Moore, C.G. 1963. Seasonal variation in autogeny in
Culex tarsalis Coq. in Northern California. Mosq.
News. 23(3):238-241.
Mortenson, E.W. 1953. Observations on the overwintering habits of Culex tarsalis Coquillett in nature.
Proc. Calif. Mosq. Cont. Assoc. 21:59-60.
Price, R.D., T.A. Olson, M.E. Rueger and L.L.
Schlottman. 1960. A survey of potential overwintering sites of Culex tarsalis Coquillet in Minnesota. Mosq. News 20:306-311.
Reeves, W.C. 1971. Mosquito vector and vertebrate host
interaction: the key to maintenance of certain arboviruses. In: A.M. Falls (ed). Ecology and physiology of parasites. Univ. of Toronto Press, Toronto,
pp 223-230.
Reeves, W.C. and W. McD. Hammon. 1944. Feeding
habits of the proven and possible mosquito vectors of western equine and Saint Louis encephalitis in the Yakima Valley, Washington. Amer. J. Trop.
Med. 24:131-134.
_____. 1962. Epidemiology of the arthropod-borne viral encephalitides in Kern County. Univ. Calif.
Publ. Hlth. 4:1-257.
Reeves, W.C., B. Brookman and W. McD. Hammon. 1948.
Studies of the flight range of certain Culex mosquitoes, using a fluorescent dye marker, with notes on
culiseta and anopheles. Mosq. News 8:61-69.
Reeves, W.C., C.H. Tempelis, R.E. Bellamy and
M.F.Lofy. 1963. Observations on the feeding habits of Culex tarsalis in Kern County, California using precipitating antisera produced in birds. Amer.
J. Trop. Med. Hyg. 12:929-935.
Reisen, W.K., M.M. Milby and M.E. Bock. 1984. The
effects of immature stress on selected events in the life
history of Culex tarsalis. Mosq. News 44(3):385-395.
Rosen, L. and W.C. Reeves. 1954. Studies on avian
malaria in vectors and hosts of encephalitis in Kern
176
Baylands Ecosystem Species and Community Profiles
County, California III. The comparative vector
ability of some of the local culicine mosquitoes.
Amer. J. Trop. Med. Hyg. 3:704-708.
Rush, W.A. and C.H. Tempelis. 1967. Biology of Culex
tarsalis during the spring season in Oregon in relation to western encephalitis virus. Mosq. News
27:307-315.
Rykman, R.E. and K.Y. Arakawa. 1952. Additional collections of mosquitoes from wood rat’s nests. PanPac. Ent. 28:105-106.
Shemanchuk, J.A., A.E.R. Downe and L. Burgess. 1963.
Hosts of mosquitoes (Diptera: Culicidae) from the
irrigated pastures of Alberta. Mosq. News.
23(4):336-341.
Sjogren, R.D. 1968. Notes on Culex tarsalis Coquillett
Breeding in Sewage. Calif. Vector Views 15(4):4243.
Telford, A.D. 1958. The pasture Aedes of Central and
Northern California. Seasonal history. Ann. Ent.
Soc. Amer. 51:360-365.
Tempelis, C.H. 1975. Host feeding patterns of mosquitoes, with a review of advances in analysis of blood
meals by serology. J. Med. Ent. 11(6):635-653.
Tempelis, C.H. and R.K. Washino. 1967. Host feeding
patterns of Culex tarsalis in the Sacramento Valley,
California, with notes on other species. J. Med.
Ent. 4:315-318.
Tempelis, C.H., W.C. Reeves, R.F. Bellamy and M.F.
Lofy. 1965. A three-year study of the feeding habits of Culex tarsalis in Kern County, California.
Amer. J. Trop. Med. Hyg. 14:170-177.
Tempelis, C.H., D.B. Francy, R.O. Hayes and M.F. Lofy.
1967. Variations in feeding patterns of seven culicine mosquitoes on vertebrate hosts in Weld and
Larimer counties, Colorado. Amer. J. Trop. Med.
Hyg. 7:561-573.
Work, T.H., M. Jozan and C.G. Clark. 1974. Differential patterns of western equine and Saint Louis
encephalitis virus isolation from Culex tarsalis
mosquitoes collected at two sites in Imperial Valley. Proc. C.M.C.A. 42:31-35.
Additional Readings
Anderson, A.W. and R.F. Harwood. 1966. Cold tolerance in adult female Culex tarsalis (Coquillett).
Mosq. News 26(1):1-7.
Bellamy, R.E. and W.C. Reeves. 1963. The winter biology of Culex tarsalis in Kern County, California.
Ann. Ent. Soc. Amer. 56:314-323.
Edman, J.D. and A.E.R. Downr. 1964. Host-blood
sources and multiple-feeding habits of mosquitoes
in Kansas. Mosq. News 24(2):154-160.
Harwood, R.F. and J.E. Halfill. 1964. The effect of photoperiod on fatbody and ovarian development of
Culex tarsalis. Ann. Ent. Soc. Amer. 57:596-600.
Wes Maffei
Harwood, R.F. and N. Takata. 1965. Effect of photoperiod and temperature on fatty acid composition of the
mosquito Culex tarsalis. J. Ins. Physiol. 11:711-716.
Winter Marsh Mosquito
Culiseta inornata (Williston)
Wesley A. Maffei
Figure 3.18 Terminal Abdominal Segment of a
Fourth Instar Larva
Description and Systematic Position
Distribution
This mosquito can be found throughout the United
States, southern Canada and northern Mexico over a
wide range of elevations and habitats (Carpenter and
LaCasse 1955). Populations of the winter marsh mosquito have been found throughout California except in
Mariposa County (Meyer and Durso 1993).
Suitable Habitat
Wes Maffei
The immature stages can be found in a wide variety of
habitats ranging from duck club ponds, ditches, seepages, rainwater pools, salt marshes and manmade con-
Figure 3.17 Wing of an Adult Cs. inornata
tainers. Telford (1958) found larvae in Marin County
marshes with salinities ranging from 8 ‰ to 26 ‰.
Adults are usually found resting near their larval
habitats during their breeding season while summer aestivating adults are presumed to utilize animal burrows
in upper marshes and adjacent uplands (Barnard and
Mulla 1977, Shemanchuk 1965).
Biology
Adults are present fall, winter and spring and enter facultative diapuase in the summer as a means of surviving
the hot, dry California summers. Aestivating females are
thought to emerge from mammalian burrows and shelters in the fall following decreased temperatures and the
first fall rains. Meyer, et al. (1982a, 1982b) found that
optimal flight activity occurred between temperatures of
48°F and 64°F, with a sharp decrease below 43°F and
above 64°F. Washino, et al. (1962) studied populations
of this mosquito in Kern County, California and found
that small numbers of adult females persisted throughout
the summer period.
Adult female mosquitoes feed primarily on large
domestic mammals although populations associated with
brackish marshes have been significantly pestiferous to
humans within the San Francisco Estuary (Bohart and
Washino 1978; Maffei, unpub.). Precipitin tests have
shown that the primary hosts are cattle, sheep, horses
and pigs (Bohart and Washino 1978, Edman and Downe
1964, Edman et al. 1972, Gunstream et al. 1971, Reeves
and Hammon 1944, Shemanchuk et al. 1963, Tempelis 1975, Tempelis et al. 1967, Tempelis and Washino
1967, and Washino et al. 1962).
Flight range studies have found that the maximum
distance traveled was 14 miles (Clarke 1943). Adults of
San Francisco Bay populations tend to stay close to their
larval source, usually traveling less than two miles for a
blood meal. Wind and proximity of available hosts are
probably important factors affecting adult dispersal and
may help account for the variability observed between
different populations of this mosquito.
Chapter 3 — Invertebrates
177
Invertebrates
The winter marsh mosquito was described from specimens collected in the Argus Mountains, Inyo County,
California, in 1893 (Belkin, et al 1966). This insect is
one of California’s largest mosquitoes, measuring approximately 8-10 mm in length. Adults are generally
light brown to reddish-brown in color and lack any unusual or distinctive markings. Diagnostic features of the
imagines include: tip of the abdomen bluntly rounded;
wings with the radial and medial cross veins nearly in
line with each other; anterior wing veins with intermixed
light and dark scales; and wings without distinct patches
of dark scales (Figure 3.17). Larvae can be identified by
the presence of only one tuft of hairs inserted near the
base of the pecten row on the siphon and by having the
lateral hairs of the anal saddle distinctly longer than the
anal saddle (Figure 3.18).
Invertebrates
Adults can be attracted to lights. Bay area mosquito
abatement Districts monitor adult populations of this
mosquito by using New Jersey light traps. Barnard and
Mulla (1977) found that the trapping efficiency of New
Jersey light traps could be improved by incresing the intensity of the incandescent light bulbs used from 25W
to 100W.
Studies of lab colonized females by Owen (1942)
found that the average life expectancy for adults was
about 97 days with a maximum of 145 days. Weather
conditions, specifically temperature and humidity, and
availability of nutrients will affect adult longevity.
Total developmental time from egg to adult has
been studied by Shelton (1973) and Mead and Conner
(1987) and both found that water temperatures above
78°F were lethal to larval development. Average developmental times ranged from 48 days at 51°F to 13 days
at 74°F. Shelton (1973) also noted that as water temperature increased beyond 68°F, average body weight
and adult survivorship decreased markedly.
Reproduction
Rees and Onishi (1951) found that adults usually do not
swarm and that freshly emerged females are mated by
waiting males. Copulation usually occurs end to end vertically, with the female above the male, and is completed
in about 3.5 to 6.5 hours.
Groups of eggs, also known as egg rafts, are deposited directly on the water. Buxton and Breland (1952)
studied the effects of temporary dessication and found
that eggs were still viable after three to four days exposure in damp leaves at various temperatures. They also
found that the eggs tolerated exposure to temperatures
as low as 17.6°F and had a hatch rate as high as 98%.
The survival of larvae hatched from eggs exposed at
17.6°F was low varying from 50% to 100% mortality
following 24 and 48 hours exposure respectively.
Significance to Other Wetlands Taxa
Winter Marsh Mosquito larvae are frequently found in
association with larvae of Aedes squamiger and the Encephalitis Mosquito, Culex tarsalis. The larvae of this
mosquito may be a possible food source for waterfowl.
Conservation Needs and Limiting Factors
This mosquito, like other species of mosquitoes, is extremely opportunistic. Care must be exercised when
managing, altering or restoring seasonal wetlands. Sites
that pond water will produce very large numbers of
adults. Care must be exercised when manipulating water levels in diked marshes. The fall flooding of these
types of wetlands for waterfowl management can produce
enormous numbers of adults. The proximity of human
178
Baylands Ecosystem Species and Community Profiles
habitation or recreational facilities can be seriously affected by the biting activity of these mosquitoes.
References
Barnard, D.R. and M.S. Mulla. 1977. Postaestival adult
activity in Culiseta inornata (Williston) in relation
to larval breeding and changes in soil temperature.
Proc. Calif. Mosq. Cont. Assoc. 45:183-185.
Belkin, J.N., R.X. Schick and S.J. Heinemann. 1966.
Mosquito studies (Diptera: Culicidae) VI. Mosquitoes originally described from North America.
1(6):1-39.
Bohart, R.M. and R.K. Washino. 1978. Mosquitoes of
California. Third Edition. Univ. Calif. Div. Agr.
Sci., Berkeley, Publ. 4084. 153 pp.
Buxton, J.A. and O.P. Breland. 1952. Some species of
mosquitoes reared from dry materials. Mosq. News.
12(3):209-214.
Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of
North America. Univ. Calif. Press, Berkeley. 360pp.
Clarke, J.L. 1943. Studies of the flight range of mosquitoes. J. Econ. Ent. 36:121-122.
Edman, J.D. and A.E.R. Downe. 1964. Host blood
sources and multiple-feeding habits of mosquitoes
in Kansas. Mosq. News 24:154-160.
Edman, J.D., L.A. Weber and H.W. Kale II. 1972. Hostfeeding patterns of florida mosquitoes II. Culiseta.
J. Med. Ent. 9:429-434.
Gunstream, S.E., R.M. Chew, D.W. Hagstrum and
C.H. Tempelis. 1971. Feeding patterns of six species of mosquitoes in arid southeastern California.
Mosq. News 31:99-101.
Kliewer, J.W., T. Miura, R.C. Husbands and C.H. Hurst.
1966. Sex pheromones and mating behaviour of
Culiseta inornata (Diptera: Culicidae). Ann. Ent.
Soc. Amer. 59(3): 530-533.
Maffei, W. 1990-1995. Unpublished field notes.
Mead, S. and G. Conner. 1987. Temperature-related
growth and mortality rates of four mosquito species. Proc. C.M.V.C.A. 55:133-137.
Meyer, R.P. and S.L. Durso. 1993. Identification of the
mosquitoes of California. Calif. Mosq. Cont.
Assoc., Sacramento, Calif. 80 pp.
Meyer, R.P., R.K. Washino and T.L. McKenzie. 1982a.
Studies on the biology of Culiseta inornata
(Diptera: Culicidae) in three regions of Central
California, USA. J. Med. Ent. 19(5):558-568.
______. 1982b. Comparisons of factors affecting
preimaginal production of Culiseta inornata
(Williston) (Diptera: Culicidae) in two different
habitats of Central California. Env. Ent.
11(6):1233-1241.
Owen, W.B. 1942. The biology of Theobaldia inornata
Williston in captive colony. J. Econ. Ent. 35:903907.
Additional Readings
Brine Flies
Diptera: Ephydridae
Wesley A. Maffei
Description and Systematic Position
There are numerous species of brine flies (Diptera:
Ephydridae) that can be found within the confines of
the San Francisco Bay region. Three are exceptionally
numerous within the bay’s tidal and diked seasonal wetlands. These are: Ephydra cinerea, Ephydra millbrae (Figure 3.19), and Lipochaeta slossonae (Figure 3.20). Adults
can readily be recognized by the following features:
head—lacking oral vibrissae, having a swollen protruding face, and having small diverging postvertical
setae; wings -with the costa broken near the subcosta
and humeral crossvein, and lacking an anal cell.
Adult flies are small in size (E. cinerea 2-3 mm
in length, E. millbrae 4-5 mm in length, and L.
slossonae 2-3 mm in length) and have unpatterned
wings. The coloration for each is as follows: E. cinerea—opaque bluish-grey with a greenish tinge and
legs with knees and most tarsal segments yellow; E.
millbrae- brownish grey with brown legs; and L.
slossonae—whitish grey with a black-brown thoracic
dorsum and legs having yellow tarsal segments.
The immature stages are small yellowish-white
larvae bearing eight pairs of ventral prolegs with two
or three rows of hooks. The last pair of prolegs are
enlarged and have opposable hooks and the last abdominal segment bears elongate respiratory tubes with
terminal spiracles. The puparium is similar in shape
to the last larval stage and is generally dark yellow to
brown in color (Figure 3.21).
Distribution
Ephydra millbrae is found throughout the San
Francisco Bay Area in mid to upper marsh tidal pools
that are infrequently affected by the tides. E. cinerea
Darsie, R.F. and R.A. Ward. 1981. Identification and
geographical distribution of the mosquitoes of
North America, north of Mexico. Mosq. Syst.
Suppl. 1:1-313.
Wes Maffei
Actual Size
5 mm
Figure 3.19 Adult Ephydra millbrae (Adapted
from Jones (1906) and Usinger (1956))
Chapter 3 — Invertebrates
179
Invertebrates
Rees, D.M. and K. Onishi. 1951. Morphology of the
terminalia and internal reproductive organs and
copulation in the mosquito Culiseta inornata
(Williston). Proc. Ent. Soc. Wash. 53:233-246.
Reeves, W.C. and W. McD. Hammon. 1944. Feeding
habits of the proven and possible mosquito vectors of western equine and Saint Louis encephalitis in the Yakima Valley, Washington. Amer. J. Trop.
Med. 24:131-134.
Telford, A.D. 1958. The pasture Aedes of Central and
Northern California. Seasonal History. Ann. Ent.
Soc. Amer. 56:409-418.
Tempelis, C.H. 1975. Host-feeding patterns of mosquitoes, with a review of advances in analysis of blood
meals by serology. J. Med. Ent. 11(6):635-653.
Tempelis, C.H. and R.K. Washino. 1967. Host feeding
patterns of Culex tarsalis in the Sacramento Valley,
California, with notes on other species. J. Med.
Ent. 4:315-318.
Tempelis, C.H., D.B. Francy, R.O. Hayes and M.F. Lofy.
1967. Variations in feeding patterns of seven culicine mosquitoes on vertebrate hosts in Weld and
Larimer counties, Colorado. Amer. J. Trop. Med.
Hyg. 7:561-573.
Shelton, R.M. 1973. The effect of temperature on the
development of eight mosquito species. Mosq.
News 33(1):1-12.
Shemanchuk, J.A. 1965. On the hibernation of Culex
tarsalis Coquillett, Culiseta inornata Williston, and
Anopheles earlei Vargas, (Diptera: Culicidae) in
Alberta. Mosq. News 25(4):456-462.
Shemanchuk, J.A., A.E.R. Downe and L. Burgess. 1963.
Hosts of mosquitoes (Diptera: Culicidae) from the
irrigated areas of Alberta. Mosq. News. 23(4):336341.
Washino, R.K., R.L. Nelson, W.C. Reeves, R.P. Scrivani
and C.H. Tempelis. 1962. Studies on Culiseta
inornata as a possible vector of encephalitis virus
in California. Mosq. News. 22(3):268-274.
Actual Size
Actual Size 5 mm
8 mm
Figure 3.20 Adult Lipochaeta slossonae
(Adapted from Jones (1906) and Usinger (1956))
Wes Maffei
Wes Maffei
10 mm
Invertebrates
Figure 3.21 Ephydra millbrae Larva and Pupa
(Adapted from Jones (1906) and Usinger (1956)
is closely associated with hypersaline environments,
especially salt ponds of the south and north bay.
Lipochaeta slossonae is commonly found in or near
crystallizer ponds of the south bay and possibly also
in salt ponds with salt concentrations somewhat above
that of sea water in other parts of the San Francisco
Bay region. Figure 3.22 shows the locations around
the Bay Area where brine flies have been collected, and
Table 3.6 shows the collection dates.
Figure 3.22 Known Brine
Fly Localities Within San
Francisco Bay Tidal and
Diked Marshes
180
Baylands Ecosystem Species and Community Profiles
Suitable Habitat
Saline and hypersaline environments.
Biology
Simpson (1976) has summarized marine Ephydrid fly
biology and a modified portion of that is presented here.
Eggs are deposited in the water and hatch after one to
Lipochaeta slossonae adults have the peculiar habit of resting with the wings and legs held very close to the body,
giving the appearance of a tube or torpedo. Should the
wind cause them to lose their footing, they simply roll
freely across the substrate until stopped by some object
such as large salt crystals of crystallizer ponds or a spiders web. Dictynid spiders frequently build webs on crystallizer ponds collecting large numbers of these flies
(Maffei, unpub.).
Precise food habits have been determined for only
a few species of Ephydrids with adults of E. cinerea
known to feed on masses of blue-green algae and the alga
Enteromorpha sp. while L. slossonae utilizes various diatoms and dinoflagellates. Cheng and Lewin (1974) observed that L. slossonae would fluidize the silt or sandy
substrate by vigorously shaking their bodies, thereby freeing some of the microorganisms upon which they feed.
Table 3.6 Known Collection Sites For Brine Flies 1
Location
Date Specimen(s) Collected
Ephydra cinerea
Location
Date Specimen(s) Collected
Ephydra millbrae
Oakland (Tide Flat)
20 Jul 1937
Sears Pt. (Solano Co.)
29 Jun 1951
San Leandro
19 Nov 1947
Mill Valley (Slough)
17 Apr 1950
Baumberg Tract (Hayward)
25 May 1989, 2 Jun 1989,
8 Jun 1989
Tiburon
5 Jul 1927
San Francisco
22 May 1915
Colma (Colma Creek)
5 May 1974
4 Jul 1968, 19 Jul 1968,
3 Aug 1968, 17 Aug 1968,
20 Aug 1968, 15 Sep 1968,
20 Sep 1968, 3 Oct 1968,
4 Nov 1968, 9 Nov 1968
Millbrae
20 Mar 1908, 1 Sep 1912
Newark
13 Aug 1930
Alvarado
2 Aug 1931
Alviso (Artesian Slough)
Alviso
Milpitas
29 Nov 1974
San Mateo
3 Oct 1920, 4 Aug 1925,
10 May 1931
Fremont (Mouth of
Coyote Hills Slough)
Dumbarton Marsh
15 Jul 1976
San Mateo
3 Oct 1920
Foster City
20 Mar 1973
Redwood City
Apr 1923, 10 Apr 1923
Menlo Park
31 Jul 1955
1 Jun 1980
Dumbarton Dr. (San
Mateo Co.)
30 Dec 1947
2 Oct 1969, 18 Nov 1971
Palo Alto
28 Jul 1894, 6 Aug 1894,
30 Jun 1915
Palo Alto (Salt Marshes)
2 Apr 1906
Mountain View
12 May 1915, 18 May 1915,
12 Jul 1924
Lipochaeta slossonae
Pittsburg
25 Nov 1923
Oliver Salt Ponds (Hayward)
5 Aug 1989
Martinez
31 Aug 1962
4 Jun 1989
Berkeley
29 Mar 1929, 26 Sep 1947
Oakland
20 Jun 1949
San Leandro
19 Nov 1947
Baumberg Tract (Hayward)
29 May 1989, 24 Feb 1990
Alviso
29 Mar 1942, 10 Apr 1969
Baumberg Tract (Hayward)
Alviso Yacht Harbor
26 Feb 1971
Milpitas
29 Nov 1974
San Jose
21 Oct 1977
1
Information assembled from specimens contained within the California Academy of Sciences Insect Collection, the University of California Berkeley
Essig Museum, the University of California Bohart Museum, the San Jose State University Edwards Museum, the San Mateo County Mosquito Abatement
District Insect Collection, and the private collections of Dr. J. Gordon Edwards and Wesley A. Maffei.
Chapter 3 — Invertebrates
181
Invertebrates
five days. The larva immediately begins feeding and will
pass through three instars. First and second instar larvae shed their cuticles in order to pass on to the next
larval stage. The cuticle of the last larval instar is not shed
but instead forms the protective pupal covering, also
known as the puparium. Adults emerge three to ten days
after the onset of pupation by inflating a balloon-like
ptilinum inside their heads. The ptilinum forces a circular cap off of the front of the puparium allowing the
adults to emerge. Deflation of the ptilinum and attainment of normal adult coloration occurs within 0.5 to
1.5 hours. Total developmental time from deposition
of eggs to emergence of adults ranges from two to five
weeks.
Adults are generally reluctant to fly and when disturbed will usually fly very close to the ground for very
short distances (Simpson 1976 and Wirth 1971).
Invertebrates
Larvae apparently feed on the same organisms as the
adults (Brock, et al. 1969).
The known salinity tolerances for the different
brine flies varies. Jones (1906) observed that E. millbrae
will occur in salt water pools with salinities up to 42 ‰.
Ephydra cinerea and L. slossonae seem to prefer saline environments much higher than 42 ‰ but are not entirely
restricted to these hypersaline habitats (Maffei, unpub.).
Nemenz (1960) studied the ability of immature
E. cinerea to maintain proper water balance in high
saline environments and found that the larvae had a
normal osmotic pressure of 20.4 atmospheres in their
haemolymph. He concluded that the adaptation to
highly concentrated salt solutions was partly due to a
relatively impermeable cuticle and probably also to
active osmotic regulation.
Reproduction
Females begin laying eggs one to two weeks after they
emerge. Ephydra cinerea has been observed to walk down
stems of aquatic vegetation or emergent objects to oviposit
underwater. The other Ephydrid flies oviposit on the water surface, where the eggs quickly sink to the bottom. Jones
(1906) states that the eggs of E. millbrae are deposited on
the floating mats of its puparia. Females deposit between
10 and 60 eggs and may require up to 20 days to complete
deposition of their eggs.
Significance to Other Wetlands Taxa
These insects are an important prey item of shore birds
and game ducks (Martin and Uhler 1939). Feeney and
Maffei (1991) observed Snowy Plovers and Maffei
(unpub.) observed California Gulls, Black Necked Stilts
and American Avocets charging through large assemblages of brine flies catching disturbed adults as they attempted to fly away. Murie and Bruce (1935) have observed populations of the Western Sandpiper, Calidris
mauri, feeding on Brine Flies near the Dumbarton
bridge. Anderson (1970) found Lesser Scaups, Dunlins,
Avocets, Western Sandpipers and Northern Phalaropes
feeding on Ephydra cinerea in the salt ponds of southern Alameda County.
These flies are a common prey item of spiders,
especially the Dictynidae and Salticidae. The tiger
beetle, Cicindela senilis senilis, will catch these flies,
and the adults of the Anthicid beetle, Tanarthrus
occidentalis, utilizes the carcasses of these flies as a
food source.
Conservation Needs and Limiting Factors
Ephydra cinerea seems to prefer the hypersaline environs
of salt ponds and has shown poor ability to adapt to the
tidal pools of mid elevation tidal marshes. The larvae of
182
Baylands Ecosystem Species and Community Profiles
this fly are also easily out competed by E. millbrae in salt
marsh tidal pools.
The frequency of flooding and duration of flooding or
drying periods limits the reproductive success of E. millbrae.
References
Anderson, W. 1970. A preliminary study of the relationship of saltponds and wildlife—South San
Francisco Bay. Calif. Fish and Game 56(4):240252.
Brock, M.L., R.G. Weigert and T.D. Brock. 1969. Feeding by Paracoenia and Ephydra on the microorganisms of hot springs. Ecology 50:192-200.
Cheng, L. and R.A. Lewin. 1974. Fluidization as a feeding mechanism in beach flies. Nature
250(5462):167-168.
Feeney, L.R. and W.A. Maffei. 1991. Snowy plovers and
their habitat at the Baumberg Area and Oliver Salt
Ponds, Hayward, California. Prepared for the City
of Hayward. 162pp.
Jones, B.J. 1906. Catalogue of the Ephydridae, with bibliography and description of new species. Univ.
Calif. Publ. Ent. 1(2):153-198.
Maffei, W. 1989-1996. Unpublished field notes.
Martin, A.C. and F.M. Uhler. 1939. Food of game ducks
in the United States and Canada. U.S.D.A. Tech.
Bull. #634. 156pp.
Murie, A. and H.D. Bruce. 1935. Some feeding habits
of the western sandpiper. The Condor. 37:258259.
Nemenz, H. 1960. On the osmotic regulation of the
larvae of Ephydra cinerea. J. Insect Physiol. 4:3844.
Simpson, K.W. 1976. Shore flies and brine flies (Diptera:
Ephydridae). Pages 465-495 In: L. Cheng (ed).
Marine insects. North-Holland Publ. Co.,
Amsterdam.
Usinger, R.L. (ed). 1956. Aquatic insects of California.
Univ. Calif. Press, Berkeley. 508pp.
Wirth, W.W. 1971. The brine flies of the genus Ephydra
in North America (Diptera: Ephydridae). Ann.
Ent. Soc. Amer. 64(2):357-377.
Additional Readings
Wirth, W.W., W. Mathis and J.R. Vockeroth. 1987.
Ephydridae. Pages 1027-1047 In: J.F. McAlpine
(ed). Manual of nearctic Diptera. Research Branch
Agriculture Canada, Ontario.
Jamieson’s Compsocryptus Wasp
Compsocryptus jamiesoni Nolfo
Wesley A. Maffei
Description and Systematic Position
Invertebrates
Compsocryptus jamiesoni is a moderate sized wasp, approximately 15-25mm in length, that belongs to the family Ichneumonidae, tribe Mesostenini. Overall body
ground color is rusty red-brown with the middle of the
face, vertex and occiput of the head, apical third of the
antennae, and the thoracic sutural markings black. The
wings are light brownish-yellow with three dark brown
transverse bands, the apical pair of bands merging near
the posterior margin of the wing (Figure 3.23). Females
have an ovipositor measuring approximately 6mm in
length and the base of the third abdominal tergite black.
Nolfo (1982) has indicated that this wasp is very similar to both Compsocryptus calipterus brevicornis and Compsocryptus aridus, which have been found within the confines of the San Francisco Bay Region exclusive of its salt
marshes. Males of this wasp are very similar to Compsocryptus calipterus brevicornis but can readily be separated
by the absence of any dark markings on the apex of the
hind femur. Females are similar to Compsocryptus aridus
but differ in having the body color rusty red-brown
rather than brownish-yellow and the dark markings of
the wings broader.
General Sample
Location
Figure 3.24 Known Compsocryptus jamiesoni
Localities Within San Francisco Bay Tidal and
Diked Marshes
where Compsocryptus jamiesoni have been collected, and
Table 3.7 shows the collection dates.
Distribution
This wasp was first collected in 1981 and subsequently
described as a new taxon by Nolfo in 1982 from specimens collected at the salt marshes in Alviso, Santa Clara
County, California. Additional populations have been
identified from the salt marshes of the eastern San Francisco Bay as far north as San Leandro, California (Maffei,
unpub.). Surveys for this wasp from other parts of the
San Francisco Estuary have not been done at this time.
Figure 3.24 shows the locations around the Bay Area
Wes Maffei
Actual Size
20 mm
Figure 3.23 Jamieson’s Compsocryptus Wasp –
Compsocryptus jamiesoni
Suitable Habitat
Compsocryptus jamiesoni have only been found on short
grass or herbage in or near tidal and muted tidal marshes.
Table 3.7 Known Collection Sites For
Compsocryptus jamiesoni 1
Location
Date Specimen(s) Collected
Trojan Marsh (San Leandro)
11 Sep 1997
Oliver Salt Ponds (Hayward)
23 Sep 1989
Baumberg Tract (Hayward)
4 Jun 1989
Shoreline Int. Ctr. (Hwyd)
1 Jul 1990, 2 Jul 1990
Ecology Marsh
24 Aug 1994
Hetch-Hetchy Marsh
16 Jul 1997
Alviso (Triangle Marsh)
3 Jun 1980
Santa Clara (Topotype)
2 Sep 1928
San Jose (Topotype)
16 Aug 1982
1 Information assembled from specimens contained within the California Academy of Sciences Insect Collection, University of California Berkeley Essig Museum, University of California Bohart Museum, San Jose State
University Edwards Museum, San Mateo County Mosquito Abatement
District Insect Collection, and private collections of Dr. J. Gordon
Edwards and Wesley A. Maffei.
Chapter 3 — Invertebrates
183
Biology
Little is known concerning the biology of this wasp.
Other members of the tribe Mesostenini are known to
be parasitic in cocoons of lepidoptera and other ichneumonids, puparia of diptera and other wasps, and the egg
sacs of spiders (Townes 1962). Adults regularly utilize
dew or rainwater from foliage and nectar from flowers
when available and can be found from April through
October. The peak flight period for C. jamiesoni is June
through August (Maffei, unpub.).
Reproduction
Invertebrates
Unknown.
Significance to Other Wetlands Taxa
Unknown.
Conservation Needs and Limiting Factors
Unknown.
References
Maffei, W. 1989-1996. Unpublished field notes.
Nolfo, S. 1982. Compsocr yptus jamiesoni, new
ichneumonid from California (Hymenoptera:
Ichneumonidae). Ent. News. 93(2):42
Townes, H. and M. Townes. 1962. Ichneumon-flies
of America north of Mexico: Part III. subfamily
Gelinae, tribe Mesostenini. Bull. U.S.N.M.
216(3):1-602.
184
Baylands Ecosystem Species and Community Profiles
A Note on Invertebrate Populations
of the San Francisco Estuary
Wesley A. Maffei
The study of San Francisco Bay invertebrate populations
and their interrelationships has usually been given low
priority or altogether neglected during the planning and
implementation of enhancement or restoration projects.
Environmental assessments of habitat quality and health
have frequently forgotten about the terrestrial or semiaquatic invertebrates that are usually very sensitive to environmental changes. Arthropods, especially insects, are
sensitive indicators of environmental disturbance or
change (Lenhard and Witter 1977, Hellawell 1978,
Hawkes 1979).
A survey of the literature shows that few studies
have been done on the biology and ecology of the terrestrial and semi-aquatic invertebrates within the San
Francisco Estuary. What is known about these organisms generally comes from studies of invertebrate populations well outside of this geographic area. For many
of the common species, this is probably adequate. Unfortunately, little information exists about what species
are found within the different wetland habitats, and less
still is known about the impacts of wetlands projects on
the existing invertebrate populations. Those species that
are pests (i.e., mosquitoes) are fairly well known, while
taxa such as Jamieson’s compsocryptus wasp or the western tanarthrus beetle, which were described as new to
science within the last twenty years, have poorly known
or completely unknown biologies. This lack of basic information, specifically what species exist where, coupled
with an understanding of their basic biologies, warrants
careful consideration and research. The fact that unknown populations of organisms, or unique, sensitive,
or threatened and endangered taxa do exist within or
near the tidal reaches of the Bay suggests that more care
should be taken when planning enhancement or restoration projects. The relationship of some invertebrate
species to the success of other organisms (i.e. plants or
invertebrates) needs to be clarified.
Some invertebrates are known to play a significant
role in the life cycles of other organisms. Functioning
as pollinators, herbivores, scavengers, predators, and
prey, terrestrial and semi-aquatic invertebrates are a significant component of any habitat or community. It
became apparent through the course of the Goals Project
that the experts on many of the key species of fish and
wildlife were not always clear about the roles played
by invertebrates with respect to the survival of their
target species or communities. This prompted the
construction of some graphic displays, in this case
food webs, by which to illustrate what little is known
about the roles performed by the largest and most
cluded in these webs are those routinely found in association with the plant or plants that are indicated by the
boxes with the thickened black borders. Figures 3.25,
3.26, and 3.27 are examples of partial webs developed
to illustrate the relationship of some of the organisms
associated with the plant species alkali heath (Frankenia
salina), common pickleweed (Salicornia virginica), and
willow (Salix lasiolepis), respectively. Figures 3.28 and
3.29 are examples of partial webs that illustrate the relationships of organisms within mid-marsh pans and
crystallizer pond habitats. The web for the organisms associated with old crystallizer ponds was included to illustrate that even in this inhospitable habitat, webs of
life can and do exist. When known vertebrate relationships for most of the webs have been included. Table
3.8 is a brief summary of the descriptions and biologies
of some of the invertebrates from the alkali heath web.
Table 3.9 is a listing of the scientific names associated
with a major common name category. It is hoped that
these tables might help the reader better visualize the nature of the relationships shown for the different organisms included in the webs.
Figure 3.25 A Partial Web of the Organisms Associated With Alkali Heath (Frankenia salina) in San
Francisco Tidal Marshes
Chapter 3 — Invertebrates
185
Invertebrates
easily overlooked group of organisms in our estuary,
the invertebrates.
Food webs are frequently used to illustrate the complex relationships between organisms within a given area
or habitat. Unfortunately, they cannot hope to tell the
entire story. Factors such as the seasonality of the organisms, length of time and time of year the studies were
performed, the limited number of organisms that can be
included in the web, and the complexity of the habitat
or ecosystem being studied tend to result in webs that
over generalize what actually exists or has been observed.
The following sample invertebrate webs are undoubtedly incomplete. They have been assembled from
many hours of field observation in the southern portion
of the San Francisco Estuary, and from an exhaustive
search of the literature. The most notable feature of all
of these webs is the delicate relationships that exist between all of the organisms involved. The potential reduction or loss of one member of the web clearly illustrates how its associates could be impacted. It should
be noted that not all of the organisms that have been
found or studied are represented. The organisms in-
Table 3.8 Partial Summary of Organisms Associated with Alkali Heath.
Bees and Wasps
Butterflies and Moths
Bombus vosnesenskii – A moderate to large sized bumblebee that is mostly black with a small amount of
yellow on the thorax and posterior portion of the
abdomen. Adults tend to nest in abandoned rodent
burrows along levees and adjacent upland habitat.
Perizoma custodiata – A moderate sized moth belonging to the family of moths known as measuring worms,
or Geometridae. Adults are tan gray or brown in color
and have dark geometric bands across the forewings.
Larvae are about one inch long, light green in color
and feed on the leaves of Frankenia. Adults are
present throughout the year, with peak populations
occurring from spring through fall.
Invertebrates
Anthophora spp. – A moderate sized native bee,
belonging to the family Anthophoridae, that is light
brown to grayish brown in appearance and has long
antennae. Adults collect pollen from flowers, are
solitary, and dig fairly deep burrows in the ground.
Burrows are usually lined with a waxy substance.
Frequently visited plants are Brassica spp., Frankenia
sp. and hemlock.
Melissodes spp. – A small to moderate sized native
bee, belonging to the family Anthophoridae, which is
grayish in color. Pollen collecting habits are similar to
Anthophora spp.
Osmia spp. – A bluish–black bee with smoky colored
wings, that belongs to the family of bees known as leaf
cutting bees, or Megachilidae. This genus of bees is
commonly known as mason bees because of their
habit of building small earthen cells on or under stones,
in abandoned burrows, in holes in boards, twigs and
logs, and in plant galls.
Ammophila spp. – A long, slender solitary digger wasp
belonging to the family Sphecidae. These wasps build
simple, vertical burrows, that are provisioned with moth
caterpillars. Nests usually occur in fine, silty or sandy soil
with minimal vegetation.
Aporinellus completus and Aporinellus taeniatus –
Small black spider wasps (family Pompilidae) that
provision their nests with jumping spiders (genus
Phidippus).
Chrysura pacifica – A small iridescent bluish–purple to
bluish–green wasp, measuring up to 10mm in length.
This wasp parasitizes the leaf cutting bee Osmia.
Parnopes edwardsi – A moderate sized brilliant light
green wasp, measuring about 10–13mm, that parasitizes the sand wasp Bembix americana.
Bembix americana – A large sand wasp that is bluish
gray in color with pale white markings on the abdomen. The eyes are usually bright yellow to yellowish–
green in color. Adult wasps provision their ground nests
with adult flies.
Dasymutilla aureola – A golden yellow to bright orange
insect known as a velvet ant. These insects are not
closely related to ants but do have the appearance of
looking like an ant. Velvet ants provision their burrows
with ground nesting bees and wasps.
Beetles
Formicilla sp. – A very small, brown to tan colored
beetle, known as an Ant–like flower beetle. These
beetles are known to feed on decaying vegetation
and can sometimes be very common at the bases of
Frankenia sp.
Stink Bugs
Chlorochroa sayi – A moderately sized (one–half inch)
stink bug that is pale to deep green in color. This insect
is known for releasing a foul smelling odor when
disturbed or threatened.
186
Baylands Ecosystem Species and Community Profiles
Brephidium exilis – A very small brown and blue
butterfly that is a frequent visitor of Frankenia.
Synanthedon bibionipennis – A small moth, belonging
to the family of moths known as clear wing moths, or
Sesiidae. Adults emerge in late May to early June and
can be found through late September. These insects
are frequently associated with Frankenia sp. It is
believed that the larvae may feed on the roots and
the bases of Frankenia sp. plants. Currently, this is the
only clear wing moth known to inhabit the levees of
mid to upper tidal marshes within the San Francisco
Estuary.
Flies
Gymnosoma fuliginosum – A small, bright orange and
black fly that is parasitic on the green stink bug,
Chlorochroa sayi.
Physocephala texana – A bright red and black fly,
about one–half an inch long, that parasitizes the
bumblebee Bombus vosnesenskii.
Aplomya theclarum – A very tiny black fly, with a bright
silver face, that parasitizes the larvae of the pygmy
blue butterfly.
Acrosticta dichroa – A small, bright green and red fly
with one brown spot at the tips of the wings. This fly is
frequently seen walking up and down the stems of
Frankenia holding it’s wings outstretched and rotating
them in opposite directions. Biology unknown.
Exoprosopa spp. and Villa spp. – Small to moderate
sized, fuzzy looking flies that are commonly known as
bee flies. Villa spp. is light brown in color with clear
wings and Exoprosopa spp. is brown and white
banded with brightly patterned brown and clear
wings. Both species of flies are parasites of immature
sand wasps of the genus Bembix.
Eristalinus aeneus –A moderate sized, shiny olive
green fly that is commonly known as a hover fly or
flower fly. The larvae of this fly are known as rat–
tailed maggots and are found in somewhat saline or
brackish pools of tidal marshes. Adult flies are an
important food source for Bembix sand wasps and
spiders.
Lejops curvipes – A moderate sized flower fly, measuring about 10–15mm, that is bright reddish–orange, with
a central black stripe on the abdomen and mostly
black legs.
Spiders
Phidippus spp. – Two species are common within our
marshes. One is solid black with the top of the
abdomen bright red and can reach a size up to one–
third of an inch. The other is dark gray with variegated white lines and reaches a size of about a
quarter of an inch.
Invertebrates
Figure 3.26 A Partial Web of the Organisms Associated With Common Pickleweed (Salicornia virginica) in San Francisco Bay Lower High Tidal Marshes
The need for terrestrial invertebrate surveys has become more apparent with the increase in wetland enhancement and restoration projects. The conversion of
one habitat type to another “more valuable” or “more
improved” habitat type can and usually does have significant impacts on the often-unnoticed invertebrate
populations that exist within them. In some cases these
impacts can be positive, while in other instances the opposite is true. Table 3.10 lists by site and date(s) those
known terrestrial and semi-aquatic invertebrate surveys
or species studies.
It is hoped that these preliminary illustrations and
discussions will shed a small amount of light on the complexity of the commonly overlooked micro fauna that exists within the tidal and diked habitats of our estuary. It
is further hoped that this glimpse might stimulate others to investigate further the biology and ecology of the
terrestrial micro fauna within these habitats. We must
improve our understanding of the importance of invertebrates to the survival of the other bayland organisms
if we are to make better-informed decisions about the
future of habitats and organisms of the San Francisco
Bay.
References
Anderson, W. 1970. A preliminary study of the relationship of salt ponds and wildlife - South San
Francisco Bay. Calif. Dept. Fish and Game. 56:240252.
Anderson, J., A. Bryant, L. Heinemann, V. Jennings,
M. Kilkenny, R. Owens, L. Rogers, B. Summers,
M. Savinsky, J. Trapani, R. Williamson and S.
Zimmer. 1980. An ecological study of a South San
Francisco Bay salt marsh. Unpubl. Report, San Jose
State Univ. 199 pp.
Balling, S.S. 1974. The influence of mosquito control
recirculation ditches on aspects of San Francisco
Bay salt marsh arthropod communities. Unpubl.
PhD Dissertation, Univ. Calif., Berkeley. 292 pp.
Balling, S.S. and V.H. Resh. 1982. Arthropod community response to mosquito control recirculation
ditches in San Francisco Bay salt marshes. Environ.
Ent. 11(4):801-808.
______. 1983. Mosquito control and salt marsh management: factors influencing the presence of Aedes
larvae. Mosq. News 43(2):212-218.
Chapter 3 — Invertebrates
187
Invertebrates
Figure 3.27 A Partial Web of the Organisms Associated With Willow (Salix lasiolepis)
Figure 3.28 Partial Web of Organisms Associated With Mid-Tidal Marsh Pans
188
Baylands Ecosystem Species and Community Profiles
______. 1984. Life history and variability in the water
boatman Trichocorixa reticulata (Hemiptera:
Corixidae) in San Francisco Bay salt marsh ponds.
Ann. Ent. Soc. Amer. 77(1):14-19.
______. 1991. Seasonal patterns in a San Francisco Bay,
California, Salt Marsh Arthropod Community.
Pan. Pac. Ent. 67(2):138-144.
Barnby, M.A. and V.H. Resh. 1980. Distribution of arthropod populations in relation to mosquito control recirculation ditches and natural channels in
the Petaluma salt marsh of San Francisco Bay. Proc.
C.M.V.C.A. 48:100-102.
Barnby, M.A., J.N. Collins and V.H. Resh. 1985.
Aquatic and macroinvertebrate communities of
natural and ditched potholes in a San Francisco
Bay salt marsh. Estuarine, Coastal and Shelf Science. 20:331-347.
Bergey, E.A., S.F. Balling, J.N. Collins, G.A. Lamberti
and V.H. Resh. 1992. Bionomics of invertebrates
within an extensive Potamogeton pectinatus bed of
a California marsh. Hydrobiologia 234:15-24.
Caires, T., D. Dawn, D. DiNunzio, A. Harris, N. Kogut,
M. Kutilek, S.H. Ladd, J. Stanziano, M. Stickler
and A. Webber. 1993. Preliminary survey of
biodiversity in the Warm Springs seasonal wetland,
Alameda County, California. Report for the U.S.
Fish and Wildl. Serv. 46 pp.
Cameron, G.N. 1969. Environmental determination of
insect species diversity in two salt marsh communities. Unpub. PhD Dissertation, Univ. Calif.,
Davis. 136 pp.
______. 1972. Analysis of insect trophic diversity in
two salt marsh communities. Ecology. 53:58-73.
Carpelan, L.H. 1957. Hydrobiology of the Alviso salt
ponds. Ecology 38:375-390.
Chandler, D.S. 1979. A new species of Tanarthrus from
California (Coleoptera: Anthicidae) Pan Pac. Ent.
55(2):147-148.
Collins, J.N., S.S. Ballindo and V.H. Resh. 1983. The
Coyote Hills Marsh model: calibration of interactions among floating vegetation, waterfowl, invertebrate predators, alternative prey, and anopheles
mosquitoes. Proc. C.M.V.C.A. 51:69-73.
Coquillett, D.W. 1902. New Diptera from North
America. Proc. U.S.N.M. 25:83-126.
Doane, R.W. 1912. New western tipula. Ann. Ent. Soc.
Amer. 5:41-61.
Donaldson, .E., D.E. Conklin and T.D. Foin. 1992.
Population dynamics of Artemia franciscana in the
San Francisco Bay National Wildlife Refuge: Phase
II. Interim Report #2.
Feeney, L.R. and W.A. Maffei. 1991. Snowy plovers and
their habitat at the Baumberg Area and Oliver Salt
Ponds, Hayward, California, March 1989 through
May 1990. Report Prepared for the City of Hayward, Calif. 162 pp.
Feeney, L.R., J.A. Alvarez and W.A. Maffei. 1996. Oakland deep draft navigation improvements protection plan for burrowing owls. Pt III. results of the
implementation of the proposed mitigation plan.
Report prepared for the Port of Oakland Environmental Dept. 92 pp. Plus Indices.
Chapter 3 — Invertebrates
189
Invertebrates
Figure 3.29 A Partial Web of the Organisms in the Baumberg and Oliver Brothers Salt Crystallizer Ponds,
Hayward, California
Table 3.9 Food Web Taxa by Major Common Name Category
Butterflies and Moths
Bugs
Brephidium exilis
Agrilus sp.
Chlorochroa sayi
Nymphalis antiopa
Amara spp.
Trichocorixa reticulata
Papilio rutulus
Bembidion spp.
Perizoma custodiata
Cicindela senilis senilis
Synanthedon bibionipennis
Cryptocephalus castaneus
Hoppers and Psyllids
Aphalara sp.
Aphids
Cixius praecox
Oliarius dondonius
Psyllids
Scale Insects
Invertebrates
Beetles
Enochlerus eximius
Lacewings
Chrysoperla plorabunda
Sympherobius bifasciatus
Spiders
Enochrus diffusus
Erynephala morosa
Dictynid Spider
Formicilla spp.
Lycosa spp.
Ochthebius rectus
Pardosa sp.
Pachybrachus melanostictus
Phidippus spp.
Powder post Beetles
Birds
Psyllobora vigintimaculata
Flies
Great Blue Heron
Synaphaeta guexi
Acrocera steyskali
Great Egret
Tanarthrus occidentails
Acrocera fasciata
Salt Marsh Yellow Throat
Tecnophilus croceicollis
Acrosticta dichroa
Snowy Egret
Xylotrechus insignis
Snowy Plover
Aedes dorsalis
Aedes squamiger
Aedes washinoi
Aplomya theclarum
Argyra californica
Ephydra millbrae
Eristalinus aeneus
Exoprosopa spp.
Gymnocarcelia ricinorum
Gymnosoma fuliginosum
Helophilus latifrons
Lejops curvipes
Lipochaeta slossonae
Lispe approximata
Pegomya spp.
Peleteria sp.
Physocephala texana
Ravinia sp.
Ants, Wasps and Bees
Song Sparrows
Western Gull
Ammophila spp.
Anthophora spp.
Mammals
Aporinellus completus
Feral Cat
Aporinellus taeniatus
Red Fox
Bembix americana comata
Reithrodontomys raviventris
Bombus vosnesenskii
Bombus occidentalis
Plants
Cerceris californicus
Cuscuta salina
Chrysura pacifica
Dunaliella sp.
Coelioxys spp.
Enteromorpha sp.
Dasymutilla aureola
Frankenia salina (= grandifolia)
Megachile spp.
Salicornia virginica
Melissodes spp.
Salix lasiolepis
Osmia spp.
Fungi
Parnopes edwardsi
Mildew type fungus
Pontania californica
Villa spp.
Fowler, B.H. 1977. Biology and life history of the salt
marsh snail, Assiminea californica, (Tryon, 1865)
(Mesogastropoda: Rissoacea). Unpubl. M.A. thesis, San Jose State Univ., San Jose, Calif. 143 pp.
Garcia, R., W.G. Voigt and A.K. Nomura. 1992. Ecology of Aedes squamiger in the northern San Francisco Bay Area. Ann. Rep. Mosq. Cont. Res., Univ.
Calif. pp. 53-57.
Garcia, R., W.G. Voigt, A.K. Nomura and A. Hayes.
1992. Biology of Aedes squamiger. Unpub. progress
190
Baylands Ecosystem Species and Community Profiles
report for Alameda County Mosquito Abatement
District. 7 pp.
Gustafson, J.F. and R.S. Lane. 1968. An annotated bibliography of literature on salt marsh insects and
related arthropods in California. Pan. Pac. Ent.
44(4):327-331.
Gustafson, J.F., R.L. Peterson and V.F. Lee. 1973. Additional references to previous lists of salt marsh
arthropods. Unpublished paper, California Academy of Sciences, Entomology Dept.
Table 3.10 Known Terrestrial or Semi-aquatic Invertebrate Surveys or Studies of Selected Invertebrate Taxa1
Locale
Reference(s)
Alviso Salt Ponds, Charlston to Alviso Slough
1951-1952
L.H. Carpelan (1957)
Outer Coyote Creek Tributary
1980
J. Anderson, et. al. (1980)
Warm Springs Seasonal Wetlands, Fremont
1993
T. Caires, et. al. (1993)
1995
W. Maffei, (unpub. field notes)
Dumbarton Point Marsh, Fremont
1968
R.S. Lane (1969)
Coyote Hills Marsh, Fremont
1983-1984
E.A. Bergey, et. al. (1992)
Ecology Marsh, Fremont
1994
C. Daehler and D. Strong (1995)
1996-1996
W. Maffei (unpub. field notes)
1989-1990
L. Feeney and W. Maffei (1991)
1997-1997
W. Maffei (unpub. field notes)
Baumberg and Oliver Salt Ponds
L. Feeney, et al.. (1996)
Richmond Field Station
1992-1994
J.A. Powell (unpub. Paper, 1994)
Petaluma Marsh
1977-1978
S. Balling and V. Resh (1991, 1982)
M. Barnby and V. Resh (1980)
V. Resh and S. Balling (1983)
1980-1980
S. Balling and V. Resh (1984)
1979
S. Balling (1982)
1981-1981
M. Barnby (1985)
1995(?)
M. Paquin (unpub.)
Suisun Marsh, Between Cutoff and Suisun Sloughs 1978-1979
S. Balling (1982)
S. Balling and V. Resh (1982)
V. Resh and S. Balling (1983)
Suisun Marsh
A.M. Shapiro (1975a, b, 1974)
1972-1973
The studies shown pertain primarily to insects and arachnids and do not include the numerous biological studies on mosquitoes.
Hellawell, J.M. 1978. Biological surveillance of rivers.
National Environment Research Council and Water Research Centre, Stevenage, England. 332 p.
______. 1986. Biological indicators of freshwater pollution and environmental management. Elsevier,
London, England. 546 p.
Jones, B.J. 1906. Catalogue of the Ephydridae, with bibliography and description of new species. Univ.
Calif. Publ. Ent. 1(2):153-198.
Josselyn, M.A. 1983. The ecology of San Francisco Bay
tidal marshes: a community profile. U.S.Fish and
Wildl. Serv., Biol. Services, Washington D.C.
FWS/OBS-83-23. 102 pp.
Josselyn, M. and J. Buchholz. 1984. Marsh restoration
in San Francisco Bay: A guide to design and planning. Paul F. Romberg Tiburon Center for Environmental Studies, Tech. Rep. #3, San Francisco
State Univ. 103 pp.
Lamberti, G.A. and V.H. Resh. 1984. Seasonal patterns
of invertebrate predators and prey in Coyote Hills
Marsh. Proc. C.M.V.C.A. 52:126-128.
Lane, R.S. 1969. The insect fauna of a coastal salt marsh.
M.A. thesis. San Francisco State Univ. San Francisco, Calif. 78pp.
Lanzarro, G.C. and B.F. Eldridge. 1992. A classical and
population genetic description of two new sibling
species of Aedes (Ochlerotatus) increpitus Dyar.
Mosq. Syst. 24(2):85-101.
Lonzarich, D.G. 1989. Temporal and spatial variations
in salt pond environments and implications for fish
and invertebrates. M.A. thesis, San Jose State Univ.
81 pp.
Madrone Associates. 1977. The natural resources of
Napa Marsh. Coastal Wetland Ser. #19. Calif.
Dept. Fish and Game, Sacramento. 96 pp.
Maffei, W.A. 1989-1997. Unpublished field notes.
Mason, H.L. 1969. A Flora of the marshes of California. Univ. of Calif. Press, Berkeley. 878 pp.
Nichols, F.H. 1973. A review of benthic faunal surveys in
San Francisco Bay. U.S. Geol. Surv. Cir. #677. 20 pp.
Chapter 3 — Invertebrates
191
Invertebrates
Oakland Airport, Burrowing Owl Mitigation Area 1995
Cullinan Ranch, USGS Survey
1
Date of Study
Invertebrates
Nolfo, S. 1982. Compsocryptus jamiesoni, new
Ichneumonid from California (Hymenoptera:
Ichneumonidae). Ent. News. 93(2):42.
Powell, J.A. 1994. Richmond Field Station Lepidoptera.
Unpublished report, Essig Museum, Univ. Calif.
Berkeley. 9 pp.
Quayle, H.J. 1906. Mosquito control work in California. Univ. Calif. Agric. Exp. Sta. Bull. #178, pp 155.
Race, M. 1981. Field ecology and natural history of
Cerithidea californica (Gastropoda: Prosobranchia)
in San Francisco Bay. Veliger 24(1):18-27.
Resh, V.H. and S.S. Balling. 1983a. Ecological impact
of mosquito control recirculation ditches on San
Francisco Bay marshlands: study conclusions and
192
Baylands Ecosystem Species and Community Profiles
management recommendations. Proc. C.M.V.C.A.
51:49-53.
______. 1983b. Tidal circulation alteration for salt marsh
mosquito control. Environ. Manag. 7(1):79-84.
Shapiro, A.M. 1974a. A salt marsh population of Lycaena
helloides (Lepidoptera: Lycaenidae) feeding on Potentilla (Rosaceae). Ent. News. 85:40-44.
______. 1974b. Butterflies of the Suisun Marsh, California. J. Res. on the Lepidoptera. 13(3):191-206.
______. 1975. Supplementary records of butterflies in
the Sacramento Valley and Suisun Marsh, lowland
central California. J. res. on the Lepidoptera.
14(2):100-102.
Tilden, J.W. 1965. Butterflies of the San Francisco Bay
Region. University of California
4
Amphibians and Reptiles
California Tiger Salamander
Ambystoma californiense
Mark R. Jennings
General Information
Reproduction
Brad Shaffer
Most adults probably reach sexual maturity in two years,
but some individuals may take longer during periods of
unfavorable conditions such as annual droughts (Shaffer
et al. 1993). Adults migrate during the night from subterranean refuge sites (small mammal burrows) to breed-
Growth and Development
Eggs hatch after 2-4 weeks (Storer 1925, Twitty 1941),
and gilled aquatic larvae take a minimum of at least 10
weeks to successfully reach metamorphosis (Anderson
1968, Feaver 1971). Larvae lack legs upon hatching at
10.5 mm total length, but quickly grow four legs within
1-2 weeks. Larvae generally are about 75 mm in total
length and weigh about 10 grams at metamorphosis into
juveniles, although they may remain in water (for up to
six months) and grow to much larger sizes with a better
chance of survival after metamorphosis (Jennings and
Hayes 1994). Overwintering of larvae, which is common
with several species of Ambystoma (see Stebbins 1985),
is unusual with A. californiense because of the temporary nature of its natural breeding habitat (Shaffer et al.
1993). All records of overwintering or aseasonal metamorphosis are based on salamanders in artificially-created habitats (Jennings, unpubl. data).
Upon metamorphosis (usually early May-through
July), juveniles disperse in mass at night away from desiccating breeding ponds into terrestrial habitats (Zeiner
et al. 1988, Loredo and Van Vuren 1996, Loredo et al.
1996). Juveniles have also been known to disperse during periods of unfavorable conditions (e.g., August) re-
Chapter 4 — Amphibians and Reptiles
193
Amphibians &
Reptiles
The California tiger salamander (Family: Ambystomatidae) is a large (75-125 mm SVL) terrestrial salamander
with several white or yellow spots or bars on a jet-black
field (Stebbins 1985). Although often referred to as a
subspecies of the more widespread tiger salamander (A.
tigrinum; e.g., see Frost 1985, Stebbins 1985, Zeiner et
al. 1988), the California tiger salamander is currently
recognized as a full species (Jones 1989, Shaffer et al.
1993, Barry and Shaffer 1994). In 1992, the California
tiger salamander was petitioned for listing as an endangered species (Long 1992) based on concerns about
population declines due to the extensive loss of habitat,
introductions of non-native aquatic predators, and interbreeding with introduced salamanders originally
brought in as live fish bait (Long 1992; see also Jennings
and Hayes 1994). The U.S. Fish and Wildlife Service
ruled that the petition was warranted but precluded by
pending listing actions on higher priority species (Sorensen 1994).
ing ponds after the onset of relatively warm winter rains
(late November-early March) where courtship and egg
deposition occurs (Twitty 1941, Barry and Shaffer 1994,
Loredo and Van Vuren 1996). Males may precede females to breeding ponds (Shaffer et al. 1993, Loredo and
Van Vuren 1996) and distances travelled by adults from
refuge sites to breeding sites may be up to 1.6 km (Austin and Shaffer 1992). Females lay single or small groups
of 2-4 eggs (8.5-12 mm in diameter) on detritus, submerged vegetation, or on the benthos of relatively shallow rain pools (Storer 1925). The number of eggs laid
per female is unknown. During periods of low rainfall,
California tiger salamanders may not reproduce (Jennings and Hayes 1994). After reproducing, adults return
to subterranean refuge sites, some to the same small
mammal burrows they emerged from earlier in the year
(Shaffer et al. 1993).
sulting in mass mortality (Holland et al. 1990). Both
adults and juveniles seek refuge in small mammal burrows (especially those of California ground squirrels
(Spermophilus beecheyi) and Botta’s pocket gophers
(Thomomys bottae) [Barry and Shaffer 1994, Loredo et
al. 1996]) and spend most of the year underground until the onset of cooler and wetter surface conditions (Jennings and Hayes 1994). Juveniles probably feed on invertebrates in subterranean mammal burrows and grow
throughout the year. During the winter months however, both juveniles and adults emerge from burrows and
forage at night on the surface for extended periods of
time, although adults appear to do all their foraging after completing their reproductive activities (Shaffer et
al. 1993).
California tiger salamanders are relatively long-lived
animals, reaching ages of 20 years or more in captivity
(Jennings, unpubl. data). The average life span of adults
in the wild is unknown.
Amphibians &
Reptiles
Food and Feeding
Larval California tiger salamanders subsist on aquatic
invertebrates (Oligochaetes, Cladocera, Conchostraca,
Ostracoda, Anostraca, Notostraca, Chironomids, etc.),
as well as the larvae of western spadefoots (Scaphiopus
hammondii), California toads (Bufo boreas halophilus),
and Pacific treefrogs (Hyla regilla), if the latter are present
in breeding ponds (Anderson 1968; Feaver 1971; Jennings, unpubl. data). Larval salamanders are also highly
cannibalistic (Jennings, unpubl. data). Good numbers
of food organisms in breeding ponds appear to be important for the survival and rapid growth of salamander
larvae to metamorphosis (Jennings, unpubl. data).
Juvenile and adult salamanders subsist on terrestrial invertebrates (Oligochaetes, Isopoda, Orthoptera,
Coleoptera, Diptera, Araneida, Gastropoda, etc.; Stebbins 1972; Morey and Guinn 1992; Jennings, unpubl.
data). There is no evidence of adult salamanders feeding in aquatic environments (Jennings, unpubl. data).
In the Bay Area, California tiger salamanders have
disappeared from almost all of the lower elevation areas
(<50 m), save one small site on the San Francisco Wildlife Refuge near Fremont, Alameda County (Jennings,
unpubl. data). There are scattered populations currently
inhabiting vernal pool and stockpond habitats in hills
surrounding the South Bay (Jennings, unpubl. data), to
the nort of Coyote Hills in Suisun, and in northern
Contra Costa. A group of relict populations is also
present in the North Bay region in vernal pool habitats
near Petaluma (Shaffer et al. 1993) (Figure 4.1).
Current Status and Factors Influencing
Population Numbers
Based on the data presented in Shaffer et al. (1993) and
Jennings and Hayes (1994), California tiger salamanders
appear to have disappeared from approximately 58% and
55% (respectively), of their historic range in the state
(Sorensen 1994). This salamander is most affected by
land use patterns and other anthropogenic events which
fragment habitat and create barriers between breeding
and refuge sites (Jennings and Hayes 1994). Some of the
more important factors negatively influencing salamander populations include: conversion and isolation of
vernal pool habitats (and surrounding oak woodland and
grasslands) to agriculture and urbanization (Barry and
Shaffer 1994); lowering of the groundwater table by
Distribution
The historical distribution of the California tiger salamander ranged from the vicinity of Petaluma, Sonoma
County and Dunnigan, Colusa-Yolo County line (Storer
1925) with an isolated outpost north of the Sutter Buttes
at Gray Lodge, Butte County (Hayes and Cliff 1982) in
Central Valley, south to vernal pools in northwest Tulare
County, and in the South Coast Range south to ponds
and vernal pools between Bulleton and Lompoc in the
Santa Ynez drainage, Santa Barbara County (Jennings
and Hayes 1994). The known elevational range extends
from 3 m-1054 m (Shaffer et al. 1993). The species has
disappeared from about 55% of its historic range (Jennings and Hayes 1994).
194
Baylands Ecosystem Species and Community Profiles
Figure 4.1 California Tiger Salamander – Some
Current Locations
overdraft (Jennings and Hayes 1994); mortality of juvenile and adult salamanders by vehicles on roads (Twitty
1941); the introduction of non-native predators such as
mosquitofish (Gambusia affinis), bullfrogs (Rana catesbeiana) and crayfish (specifically Procambarus clarkii)
into breeding habitats (Shaffer et al. 1993); the widespread poisoning of California ground squirrels and other
burrowing rodents (Loredo et al. 1996); and interbreeding with introduced salamanders originally brought in
as live fish bait (Shaffer et al. 1993). Juvenile and adult
salamanders have also been found in a number of human-created habitats such as septic tank lines, pipes,
wells, wet basements, and permanent irrigation ponds
(Jennings and Hayes 1994). Such habitats may not be
suitable for the long-term survival or successful reproduction of local salamander populations.
Trophic Levels
Larval and post-metamorphic life stages are secondary
consumers.
Proximal Species
Good Habitat
The best habitats for California tiger salamanders are
vernal pool complexes with colonies of California ground
squirrels or Botta’s pocket gophers within the complex
or nearby (Shaffer et al. 1993). Such habitats are normally associated with grasslands or oak woodlands
(Barry and Shaffer 1994). Additionally, there needs
to be abundant invertebrate resources and other native amphibian larvae in the vernal pools used by
breeding salamanders.
References
Anderson, P.R. 1968. The reproductive and developmental history of the California tiger salamander.
Unpubl. M.A. Thesis, Fresno State College, Fresno,
Calif. vii+82 p.
Austin, C.C. and H.B. Shaffer. 1992. Short-, medium, and long-term repeatability of locomotor performance in the tiger salamander Ambystoma californiense. Functional Ecology, 6(2):145-153.
Chapter 4 —
Amphibians and Reptiles
195
Amphibians &
Reptiles
Predators: Common [=San Francisco] garter snake,
Coast garter snake, Central Coast garter snake, California red-legged frog, bullfrog, shrews, striped skunk, opossum, herons, and egrets. Ducks and predacious aquatic
insects prey on larvae only.
Prey: Oligochaetes, snails, and terrestrial insects. Zooplankton and aquatic insects are prey for larvae.
Habitat: California ground squirrel and valley pocket
gopher (maintain tiger salamander’s terrestrial habitats)
Barry, S.J. and H.B. Shaffer. 1994. The status of the
California tiger salamander (Ambystoma californiense) at Lagunita: A 50-year update. J. of Herpetology, 28(2):159-164.
Feaver, P.E. 1971. Breeding pool selection and larval
mortality of three California amphibians: Ambystoma tigrinum californiense Gray, Hyla regilla
Baird and Girard, and Scaphiopus hammondii
Girard. Unpubl. M.A. Thesis, Fresno State College, Fresno, Calif. v+58 p.
Frost, D.R. (ed). 1985. Amphibian species of the world:
A taxonomic and geographical reference. Allen
Press, and the Association of Systematics Collections, Lawrence, Kansas. v+732 p.
Hayes, M.P. and F.S. Cliff. 1982. A checklist of the
herpetofauna of Butte County, the Butte Sink, and
Sutter Buttes, Calif. Herpetological Review,
13(3):85-87.
Holland, D.C., M.P. Hayes and E. McMillan. 1990.
Late summer movement and mass mortality in the
California tiger salamander (Ambystoma californiense).
The Southwestern Naturalist, 35(2):217-220.
Jennings, M.R. and M.P. Hayes. 1994. Amphibian and
reptile species of special concern in California. Final report to the Ca. Dept. Fish and Game, Inland
Fisheries Div., Rancho Cordova, Ca., under Contract (8023). iii+255 p.
Jones, T.R. 1989. The evolution of macrogeographic and
microgeographic variation in the tiger salamander
Ambystoma tigrinum (Green). Unpubl. Ph.D. Dissertation, Arizona State Univ., Tempe, Az. xiii+173 p.
Long, M.M. 1992. Endangered and threatened wildlife
and plants; 90-day finding and comment of status
review for a petition to list the California tiger salamander. Fed. Reg., 57(224):54545-54546. [Thursday, November 19, 1992].
Loredo, I., D. Van Vuren, and M.L. Morrison. 1996.
Habitat use and migration behavior of the California tiger salamander. J. of Herpetology
30(2):282-285.
Loredo, I. and D. Van Vuren. 1996. Reproductive ecology of a population of the California tiger salamander. Copeia, 1996(4):895-901.
Morey, S.R. and D.A. Guinn. 1992. Activity patterns,
food habits, and changing abundance in a community of vernal pool amphibians. Pages 149-158.
In: D. F. Williams, S. Byrne and T. A. Rado (eds).
Endangered and sensitive species of the San Joaquin
Valley, California: Their biology, management, and
conservation. The Calif. Energy Commission, Sacramento, Calif., and the Western Section of The
Wildl. Society. xv+388 p.
Shaffer, H.B., R.N. Fisher and S.E. Stanley. 1993. Status report: The California tiger salamander (Ambystoma californiense). Final report to the Ca. Dept.
Fish and Game, Inland Fisheries Div., Rancho
Cordova, Calif., under Contracts (FG 9422 and
FG 1383). 92 p.
Sorensen, P.C. 1994. Endangered and threatened wildlife and plants; 12-month petition finding for the
California tiger salamander. Fed. Reg.,
59(74):18353-18354. [Monday, April 18, 1994].
Stebbins, R.C. 1972. Amphibians and reptiles of California. California Natural History Guide (31).
Univ. of Ca. Press, Berkeley, Los Angeles, and London. 152 p.
______. 1985. A field guide to western amphibians and
reptiles. Second edition, revised. Houghton Mifflin
Company, Boston, Massachusetts. xiv+336 p.
Storer, T.I. 1925. A synopsis of the Amphibia of California. Univ. of Calif. Publications in Zoology,
27:1-342.
Twitty, V.C. 1941. Data on the life history of Ambystoma
tigrinum californiense Gray. Copeia, 1941(1):1-4.
Zeiner, D.C., W.F. Laudenslayer, Jr. and K.E. Mayer (compiling editors). 1988. California’s wildlife. Volume I.
Amphibians and reptiles. California Statewide Wildlife Habitat Relationships System, Ca. Dept. of Fish
and Game, Sacramento, Ca. ix+272 p.
Reproduction
California toads breed between January and July with
higher altitude populations delaying breeding until June
or July (Storer 1925). At lower elevations, toads are active all year, but at higher elevations adults emerge from
hibernation sites immediately before reproducing (Stebbins 1951). Males and females congregate at night
around aquatic breeding sites such as stockponds, temporary roadside pools, cement water reservoirs, and the
margins of flowing streams where males call, amplexus
occurs, and females lay up to 16,500 eggs in two long
strings wrapped around vegetation at water depths <150
mm (Storer 1925, Livezey and Wright 1947). Eggs
strings are about 5 mm in diameter and the inclusive eggs
1.7 mm in diameter (Storer 1925). After reproducing,
adults generally disperse back into the surrounding terrestrial habitats such as meadows and woodlands where
they use almost any sort of cover (e.g., trees, low vegetation, beds of leaves, small mammal burrows, rocks,
pieces of concrete, downed logs, etc.) that provides a
slight amount of moisture and protection from the drying effects of the sun and wind (Storer 1925).
Amphibians &
Reptiles
Growth and Development
California Toad
Bufo boreas halophilus
Mark R. Jennings
General Information
The California toad (Family: Bufonidae) is a moderatesized (62-125 mm SUL) toad with prominent oval
parotoid glands and a light middorsal stripe (Stebbins
1985). Dorsal coloration is normally dusky, gray, or
greenish, with warts set in black patches (Storer 1925).
Natural intergrades with boreal toads (B. b. boreas) in
northern California and hybrids with Yosemite toads (B.
canorus) in the Sierra Nevada have been recorded (Storer
1925, Karlstrom 1962).
Eggs hatch within four days to a few weeks (depending
on the prevailing water temperature; Storer 1914) and
the resulting larvae normally comprise schools composed
of one or more clutches (Jennings, unpubl. data). Larvae grow rapidly and usually metamorphose in 2-3
months (from April-August) at 19-52 mm (Storer 1925,
Wright and Wright 1949). Recent metamorphs are 1215 mm in total length (Wright and Wright 1949) and
are often observed around the immediate margin of the
breeding pond under any cover that protects them from
the wind and sun (Storer 1925). The number of newly
metamorphosed toads at such breeding sites can number in the thousands (Storer 1925). Young toads grow
rapidly and probably reach sexual maturity in two years
at lower elevations and somewhat longer at mid-higher
elevations (Stebbins 1951). Both juveniles and adults are
largely crepuscular, although an occasional individual
will be observed during the day in wet or overcast conditions (Storer 1925).
Adults may live 10 years or more in captivity
(Bowler 1977) but the longevity of toads in the wild is
unknown.
Food and Feeding
Rick Fridell
Larvae are thought to be algal grazers (Stebbins 1951),
but the foraging ecology of larval California toads is unknown. Juveniles and adults feed on a wide variety of
terrestrial and flying invertebrates including: Oligochaetes, Isopoda, Diplopoda, Orthoptera, Plecoptera,
196
Baylands Ecosystem Species and Community Profiles
Dermaptera, Hemiptera, Homoptera, Coleoptera,
Trichoptera, Lepidoptera, Diptera, Arachnids, and Gastropoda (Storer 1914; Eckert 1934; Stebbins 1951,
1972; Morey and Guinn 1992; Jennings, unpubl. data).
Cannibalism can also occur (Stebbins 1972).
Distribution
California toads are found over most of California (except for the northernmost counties where they are replaced by boreal toads, and almost all of the Mojave and
Colorado deserts where they are replaced by other toad
species) from sea level to over 3,050 meters in the Sierra Nevada (Stebbins 1972). This toad is replaced at
higher elevations in the central and southern Sierra
Nevada by the Yosemite toad (Bufo canorus) [Karlstrom
1962]. California toads are widespread in the Bay Area
(Stebbins 1959) (Figure 4.2) and are still relatively common in stockponds and other aquatic habitats in the surrounding foothills (Jennings, unpubl. data).
Current Status and Factors Influencing
Population Numbers
Trophic Levels
Larval stages are primary consumers and post-metamorphic life stages are secondary consumers.
Proximal Species
Predators: Common [=San Francisco] garter snake,
coast garter snake,central coast garter snake, bullfrog, introduced predatory fishes, herons, egrets, raccoon,
striped skunk, and opossum. Predacious aquatic insects
prey on larvae.
Prey: Aquatic insects, Oligochaetes, Gastropoda, Isopoda,
and terrestrial insects.
Habitat: Willows, cattails, tules, sedges, blackberries,
riparian vegetation.
Good Habitat
California toads inhabit grasslands, woodlands, meadows, gardens, golf courses, and parks—in fact, anywhere
where a permanent source of moisture is present and
breeding ponds of at least two months duration are available (Storer 1925). The largest populations of toads seem
to be found around stockponds or reservoirs that have
an abundance of invertebrate prey, many small mammal
burrows and objects (or vegetation) that are available for
cover, and a lack of introduced predators (fishes and
bullfrogs [Rana catesbeiana]) in aquatic habitats.
References
Figure 4.2 California Toad – Presumed Bay Area
Distribution
Bowler, J.K. 1977. Longevity of reptiles and amphibians in North American collections. Society for
the Study of Amphibians and Reptiles, Miscellaneous Publications, Herpetological Circular
(6):1-32.
Eckert, J.E. 1934. The California toad in relation to
the hive bee. Copeia. 1934(2):92-93.
Chapter 4 —
Amphibians and Reptiles
197
Amphibians &
Reptiles
California toads are still present throughout most of their
native range in California, although they are now rare
in many urban areas where they were formerly common
(such as in the Los Angeles Basin; Jennings unpubl.
data). The possible reasons for the localized declines are
insecticides used in eradicating introduced Mediterranean fruit flies (Ceratitis capitata), changing land use
patterns by agriculture and urban communities which
now leave less sites containing permanent water and areas of dense vegetation (such as tule-lined canals, low
ground cover, etc.), and habitat fragmentation by roads
and dense regions of urbanization (Jennings, unpubl.
data). In the Bay Area, California toads are still relatively
abundant in natural and moderately-altered habitats
(Stebbins 1959; Jennings, unpubl. data). The factors
most associated with toad survival include local breeding ponds that last for at least two months, and sufficient cover (vegetative and small mammal burrows) that
provide places for toads to feed and grow, as well as escape predators and desiccating conditions.
Amphibians &
Reptiles
Karlstrom, E.L. 1962. The toad genus Bufo in the Sierra Nevada of California: Ecological and systematic relationships. Univ. of Calif. Publications in
Zoology. 62(1):1-104.
Livezey, R.L. and A.H. Wright. 1947. A synoptic key to
the salientian eggs of the United States. The American Midland Naturalist. 37(1):179-222.
Morey, S.R. and D.A. Guinn. 1992. Activity patterns,
food habits, and changing abundance in a community of vernal pool amphibians. In: D.F. Williams, S. Byrne and T.A. Rado (eds). Endangered
and sensitive species of the San Joaquin Valley,
California: Their biology, management, and conservation. The Calif. Energy Commission, Sacramento, Calif., and the Western Section of The
Wildl. Society. xv+388 p.
Stebbins, R.C. 1951. Amphibians of western North
America. Univ. of Calif. Press, Berkeley and Los
Angeles. ix+539 p.
______. 1959. Reptiles and amphibians of the San Francisco
Bay region. California Natural History Guide (3). Univ.
of Calif. Press, Berkeley and Los Angeles. 72 p.
______. 1972. Amphibians and reptiles of California.
California Natural History Guide (31). Univ. of Calif. Press, Berkeley, Los Angeles, and London. 152 p.
______. 1985. A field guide to western amphibians and
reptiles. Second ed., revised. Houghton Mifflin
Co., Boston, Massachusetts. xiv+336 p.
Storer, T.I. 1914. The California toad, an economic asset. U.C. J. of Agriculture, 2(3):89-91.
______. 1925. A synopsis of the Amphibia of California.
Univ. of Calif. Publications in Zoology, 27:1-342.
Wright, A.H. and A.A. Wright. 1949. Handbook of frogs
and toads of the United States and Canada. Third
ed. Comstock Publ. Co., Inc., Ithaca, NY. xii+640 p.
198
Baylands Ecosystem Species and Community Profiles
Pacific Treefrog
Hyla regilla
Mark R. Jennings
General Information
The Pacific treefrog (Family: Hylidae) is a small (19-50
mm SUL) frog with toe pads and a black eye stripe (Stebbins 1985). The dorsal coloration is highly variable—
green, tan, reddish, gray, brown, or black—sometimes
with dark dorsal spots (Wright and Wright 1949,
Resnick and Jameson 1963, Stebbins 1972); however
green or shades of brown are the usual colors observed
(Nussbaum et al. 1983). For a time, these treefrogs were
lumped with chorus frogs of the genus Pseudacris (see
Hedges 1986). However, recent work has shown that
Pacific treefrogs are not chorus frogs, hence the reversion to the old genus Hyla (Crocroft 1994).
This frog has the most notable voice of the frog
world as its call has been used as a natural background
sound in innumerable movies produced by Hollywood
(Myers 1951).
Reproduction
Pacific treefrogs can become sexually mature in one year,
but most become sexually mature in two years (Jameson
1956). At lower elevations, treefrogs are active all year,
but at higher elevations adults emerge from hibernation
sites immediately before reproducing (Stebbins 1951).
From late November to July (beginning with the first
warm rainfall), males congregate at night around any
suitable shallow pond of water (or at the shallow edges
of deep water ponds or reservoirs) and chorus to attract
receptive females (Storer 1925, Brattstrom and Warren
1955, Schaub and Larsen 1978, Nussbaum et al. 1983),
and also call to space themselves from one another
(Snyder and Jameson 1965, Allan 1973). Groups of two
or three males tend to call in sequence during these choruses and the sequence is consistently started by one frog
known as the bout leader (Whitney and Krebs 1975).
The choruses may continue into daylight hours (Jameson
1957) and can be deafening if hundreds or thousands
of calling males are involved (Stebbins 1959; Jennings,
unpubl. data). Females attracted to these calls usually
select the bout leader to mate with (Whitney and Krebs
1975). Upon amplexus, females lay approximately 2025 packets containing 9-70 (usually 22-25) eggs on submerged aquatic vegetation or on the bottom of shallow
pools (Smith 1940, Livezey and Wright 1947), generally at depths >100 mm (Storer 1925). Egg masses are
normally laid close together, one against another, or separated by <25 mm (Stebbins 1951). The eggs are about
1.3 mm in diameter and females may lay from 500 to
1,250 total eggs (Storer 1925, Smith 1940). After egg
deposition, males and females remain the vicinity of the
breeding pond for up to one and three months respectively, and then return to surrounding terrestrial habitats (Jameson 1957, Nussbaum et al. 1983). Females
may also breed up to three times during the year (Perrill
and Daniel 1983).
Growth and Development
Distribution
Pacific treefrogs are found over most of California (except drier parts of the Mojave and Colorado deserts) from
sea level to around 3,670 m in the Sierra Nevada (Stebbins 1972, 1985). In the Bay Area they are very abundant (Stebbins 1959; Jennings, unpubl. data), and found
throughout the region (Figure 4.3).
Current Status and Factors Influencing
Population Numbers
Pacific treefrogs have always been abundant throughout
most of their native range (e.g., see Storer 1925, Stebbins 1951, Nussbaum et al. 1983), and they still remain
so even in the Sierra Nevada (Jennings 1996, contra
Drost and Fellers 1996). Treefrogs are especially common in the Bay Area (Stebbins 1959) because of their
ability to utilize habitats created by humans—especially
in urban areas. Although populations are negatively influenced by the premature drying of breeding ponds and
continued loss of many individuals through predation,
Food and Feeding
Larvae are thought to be algal grazers (Storer 1925), but
the foraging ecology of larval Pacific treefrogs is unknown. Juveniles and adults feed on a wide variety of
terrestrial and flying invertebrates including: Oligocha-
Figure 4.3 Pacific Treefrog – Presumed Bay Area
Distribution
Chapter 4 —
Amphibians and Reptiles
199
Amphibians &
Reptiles
Eggs hatch in four days to two weeks, depending on the
prevailing water temperatures and the resulting larvae
(6.0-7.5 mm total length) grow rapidly (Storer 1925).
Larvae are also known to aggregate into large groups of
several hundred individuals (Brattstrom and Warren
1955). Metamorphosis is generally within two months
at anytime between February-late August (Storer 1925;
Jennings, unpubl. data), at total lengths between 45 and
55 mm (Storer 1925, Wright and Wright 1949). Postmetamorphs are about 12-15 mm and grow rapidly
within the first two months often doubling their size
(Jameson 1956). For the first few months, postmetamorphs remain in the immediate vicinity of the
breeding pond utilizing almost any cover present (rocks,
vegetation, leaves, etc.) for protection from the drying
effects of the sun and wind (Jameson 1956). After several months, juveniles disperse out into surrounding terrestrial habitats and seek places that contain moisture and
are protected by the elements. Such places include small
mammal burrows, rock fissures, tree cavities, dense vegetation, piles of debris, buildings, artificial drains, etc.,
that may be 0.8 km or more from the nearest standing
water (Storer 1925, Jameson 1957).
Adults may live four years or more in captivity (Jennings, unpubl. data). Longevity in the wild is apparently
somewhat over three years (Jameson 1956).
etes, Oniscidea, Orthoptera, Hemiptera, Homoptera,
Coleoptera, Lepidoptera, Diptera, Hymenoptera, Arachnids, and Gastropoda (Needham 1924; Stebbins 1951;
Brattstrom and Warren 1955; Nussbaum et al. 1983;
Morey and Guinn 1992; Jennings, unpubl. data).
treefrogs are able to successfully reproduce in numbers
to overcome these set backs (Jameson 1956, 1957).
Trophic Levels
Larval stages are primary consumers and post-metamorphic life stages are secondary consumers.
Proximal Species
Habitat: Willows, cattails, tules, and sedges.
Predators: Common [=San Francisco] garter snake,
Coast garter snake, Central Coast garter snake, California red-legged frog, bullfrog, introduced predatory fishes,
herons and egrets, raccoon, striped skunk, and opossum.
California tiger salamander and various predacious
aquatic insects prey on larvae only.
Prey: Aquatic and terrestrial insects.
Amphibians &
Reptiles
Good Habitat
Pacific treefrogs can essentially inhabit almost any place
that contains sufficient moisture and protection from the
wind and sun, and has suitable nearby breeding sites.
They can reproduce in temporary aquatic environments
as small as a jar of water as long as the water remains
present for two months or more (Jennings, unpubl. data)
and the water temperature is below 35-38° C
(Schechtman and Olson 1941). The largest populations
seem to be present in complexes of shallow ponds (lacking fishes and other aquatic predators) surrounded by
growths of tules (Scirpus sp.) and other aquatic vegetation (Jameson 1956, 1957), although Pacific treefrogs
also seem to do well in golf courses, city parks, and other
places that have permanent aquatic habitats and places
with riparian vegetation (Jennings, unpubl. data).
References
Allan, D.M. 1973. Some relationships of vocalization
to behavior in the Pacific treefrog, Hyla regilla.
Herpetologica, 29(4):366-371.
Brattstrom, B.H. and J.W. Warren. 1955. Observations
on the ecology and behavior of the Pacific treefrog,
Hyla regilla. Copeia, 1955(3):181-191.
Crocroft, R.B. 1994. A cladistic analysis of chorus frog
phylogeny (Hylidae: Pseudacris). Herpetologica,
50(4):420-437.
Drost, C.A. and G.M. Fellers. 1996. Collapse of a regional frog fauna in the Yosemite area of the California Sierra Nevada, USA. Conservation Biology,
10(2):414-425.
Hedges, S.B. 1986. An electrophoretic analysis of Holarctic
hylid frog evolution. Systematic Zoology, 35(1):1-21.
Jameson, D.L. 1956. Growth, dispersal and survival of
the Pacific tree frog. Copeia, 1956(1):25-29.
200
Baylands Ecosystem Species and Community Profiles
______. 1957. Population structure and homing responses
in the Pacific tree frog. Copeia, 1957(3):221-228.
Jennings, M.R. 1996. Status of amphibians. Pages 311—31-24. In: Sierra Nevada Ecosystem Project:
Final Report to Congress, Volume II, Assessments
and Scientific Basis for Management Options. Centers for Water and Wetland Resources, Univ. of
Calif., Davis, Calif.
Livezey, R.L. and A.H. Wright. 1947. A synoptic key to
the salientian eggs of the United States. The American Midland Naturalist, 37(1):179-222.
Morey, S.R. and D.A. Guinn. 1992. Activity patterns,
food habits, and changing abundance in a community of vernal pool amphibians. Pages 149-158.
In: D.F. Williams, S.Byrne, and T. A. Rado (eds).
Endangered and sensitive species of the San Joaquin
Valley, California: Their biology, management, and
conservation. The Calif. Energy Commission, Sacramento, Calif., and the Western Section of The
Wildl. Society. xv+388 p.
Myers, G.S. 1951. The most widely heard amphibian
voice. Copeia, 1951(2):179.
Needham, J.G. 1924. Observations of the life of the
ponds at the head of Laguna Canyon. J. of Entomology and Zoology, 16(1):1-12.
Nussbaum, R.A., E.D. Brodie, Jr. and R.M. Storm.
1983. Amphibians and reptiles of the Pacific Northwest. Univ. of Idaho Press, Moscow, Idaho. 332 p.
Perrill, S.A. and R.E. Daniel. 1983. Multiple egg clutches
in Hyla regilla, H. cinerea and H. gratiosa. Copeia,
1983(2):513-516.
Resnick, L.E. and D.L. Jameson. 1963. Color polymorphism in Pacific tree frogs. Science, 142(3595):10811083.
Schaub, D.L. and J.H. Larsen. 1978. The reproductive
ecology of the Pacific treefrog (Hyla regilla).
Herpetologica, 34(4):409-416.
Schechtman, A.M. and J.B. Olson. 1941. Unusual temperature tolerance of an amphibian egg (Hyla
regilla). Ecology, 22(4):409-410.
Smith, R.E. 1940. Mating and oviposition in the Pacific Coast tree toad. Science, 92(2391):379-380.
Snyder, W.F. and D.L. Jameson. 1965. Multivariate geographic variation of mating call in populations of
the Pacific tree frog (Hyla regilla). Copeia,
1965(2):129-142.
Stebbins, R. C. 1951. Amphibians of western North
America. Univ. of Calif. Press, Berkeley and Los
Angeles. ix+539 p.
______. 1959. Reptiles and amphibians of the San Francisco Bay region. California Natural History Guide (3).
Univ. of Ca. Press, Berkeley and Los Angeles. 72 p.
______. 1972. Amphibians and reptiles of California.
California Natural History Guide (31). Univ. of
Calif. Press, Berkeley, Los Angeles, and London.
152 p.
______. 1985. A field guide to western amphibians and
reptiles. Second edition, revised. Houghton Mifflin
Company, Boston, Massachusetts. xiv+336 p.
Storer, T.I. 1925. A synopsis of the Amphibia of California. Univ. of Calif. Publications in Zoology,
27:1-342.
Whitney, C. L. and J.R. Krebs. 1975. Spacing and calling in Pacific tree frogs, Hyla regilla. Canadian J.
of Zoology, 53(11):1519-1527.
Wright, A.H. and A.A. Wright. 1949. Handbook of frogs
and toads of the United States and Canada. Third
edition. Comstock Publishing Company, Inc.,
Ithaca, New York. xii+640 p.
California Red-Legged Frog
Rana aurora draytonii
Mark R. Jennings
General Information
Dan Holland
Adults generally reach sexual maturity in their second
year for males and third year for females (Jennings and
Hayes 1985), although sexual maturity may be reached
earlier during years of abundant food resources (Jennings, unpubl. data). During extended periods of
drought, frogs may take 3-4 years to reach maturity (Jennings and Hayes 1994a). Reproduction generally occurs
at night in permanent ponds or the slack water pools of
streams during the winter and early spring (late November-through April) after the onset of warm rains (Storer
1925, Hayes and Jennings 1988, Jennings and Hayes
1994a). California red-legged frogs can only successfully
reproduce in aquatic environments with water temperatures <26° C and salinities <4.5% as developing embryos
cannot tolerate conditions higher than this (Jennings,
unpubl. data). Larvae can tolerate somewhat higher
water temperatures and salinities (Jennings, unpubl.
data). Males generally appear at breeding sites from 2-4
weeks before females (Storer 1925). At breeding sites,
males typically call in small, mobile groups of 3-7 individuals that attract females (Jennings and Hayes 1994a).
Females amplex with males and attach egg masses containing approximately 2,000-6,000 moderate-sized (2.02.8 mm diameter) eggs to an emergent vegetation brace
at depths usually from 75-100 mm (Storer 1925). Egg
masses are normally laid at the surface of the water
(Livezey and Wright 1947). California red-legged frogs
are explosive breeders, usually depositing their egg
masses within 3-4 week period after large rainfall events
(Hayes and Miyamoto 1984). After reproduction, males
usually remain at the breeding sites for several weeks
before removing to foraging habitats, while females immediately remove to these foraging habitats (Jennings,
unpubl. data). There is no evidence of double clutching with this species (Jennings, unpubl. data).
Growth and Development
Eggs hatch after 6-14 days (depending on the prevailing water temperature), and the resulting larvae (8.810.3 mm total length) require 3.5-7 months to attain
metamorphosis at 65-85 mm total length (Storer 1925;
Jennings, unpubl. data). Larvae, which are solitary and
almost never overwinter, typically metamorphose between July and September (Storer 1925, Jennings and
Hayes 1994a).
Juvenile frogs are 25-30 mm total length at metamorphosis and commonly sun themselves during the day
at the edge of the riparian zone next to the breeding site.
As they grow, they gradually shift from diurnal and nocturnal periods of activity, to largely nocturnal activity
(Hayes and Tennant 1986). During periods of rainfall,
both juveniles and a few adults may disperse away from
breeding sites and may be found some distance (up to
Chapter 4 —
Amphibians and Reptiles
201
Amphibians &
Reptiles
The California red-legged frog (Family: Ranidae) is a
large (85-138 mm SUL) brown to reddish brown frog
with prominent dorsolateral folds and diffuse moderatesized dark brown to black spots that sometimes have light
centers (Storer 1925, Jennings and Hayes 1994a). The
species is the largest native frog in the state and there
are data to support elevation as a separate species from
the northern red-legged frog (R. a.aurora) [see Hayes and
Miyamoto 1984, Green 1985]; however, there is also
large zone of intergradation along the Pacific slope of the
North Coast Range (Hayes and Kremples 1986). In
1993, the California red-legged frog was petitioned for
listing as an endangered species by the U.S. Fish and
Wildlife Service (Sorensen 1993) based on a significant range reduction and continued threats to surviving populations (Miller 1994). The frog was subsequently listed as Threatened by the U.S. Fish and
Wildlife Service (Miller et al. 1996).
Reproduction
0.8 km) away from the nearest water (Jennings, unpubl.
data). Along the lower reaches of streams on the Central Coast of California which tend to almost completely
dry up during the late summer, subadult and adult frogs
have been found to occupy small mammal burrows under leaf litter or dense vegetation in the riparian zone
(Rathbun et al. 1993). These frogs make overland trips
every few days or so to isolated stream pools to rehydrate
themselves although one frog remained in riparian habitat for 77 days (Rathbun in litt. 1994 as cited in Miller
et al. 1996). Based on these observations, frogs found
in coastal drainages appear to be rarely inactive, whereas
those found in interior sites probably hibernate (Storer
1925).
Based on limited field data, California red-legged
frogs appear to live about 8-10 years in the wild (Jennings, unpubl. data).
Amphibians &
Reptiles
Food and Feeding
Larvae are thought to be algal grazers (Storer 1925), but
the feeding ecology of larval California red-legged frogs
is unknown. Juvenile and adult frogs have a highly variable animal food diet that includes: Amphipods, Isopods,
Orthoptera, Isoptera, Hemiptera, Homoptera, Neuroptera, Coleoptera, Lepidoptera, Diptera, Hymenoptera,
Arachnids, Gastropoda, small fishes, amphibians, and
small mammals (Stebbins 1972, Hayes and Tennant
1985, Baldwin and Stanford 1987). Most prey that can
be swallowed that are not distasteful are eaten, with
larger frogs capable of taking larger prey (Jennings and
Hayes 1994a). Small red-legged frogs, Pacific treefrogs
(Hyla regilla), and California mice (Peromyscus californicus) may contribute significantly to the diet of subadults
and adults (Arnold and Halliday 1986, Hayes and
Tennant 1985).
1950s it was still considered to be present in much of
the San Francisco Bay region (Stebbins 1959). However,
earlier overexploitation, subsequent habitat loss from agriculture and urbanization, and the introduction of exotic aquatic predators have presently reduced red-legged
frog distribution to scattered locations in the foothills
and mountains of the San Francisco Bay region (Jennings, unpubl. data) (Figure 4.4).
Current Status and Factors Influencing
Population Numbers
Based on the data presented in Jennings and Hayes
(1994a, 1994b), California red-legged frogs appear to
have disappeared from approximately 70-75%, of their
historic range in the state (Miller et al. 1996). This frog
is most affected by land use patterns and other anthropogenic events which fragment high quality habitat and
create environments unsuitable for the continued survival of the species (Jennings and Hayes 1994a, 1994b).
Some of the more important factors negatively influencing frog populations include: conversion and isolation
of perennial pool habitats (and surrounding riparian
zones) to agriculture; reservoir construction projects; urbanization; lowering of the groundwater table by overdraft; overgrazing by domestic livestock; extended
drought; mortality of juvenile and adult frogs by vehicles
on roads; and the introduction of non-native predators
such as mosquitofish (Gambusia affinis), bullfrogs (Rana
Distribution
Historically, California red-legged frogs were found
throughout the Pacific slope drainages from the vicinity of Redding, Shasta County (Storer 1925), inland and
at least to Point Reyes, Marin County (Hayes and
Kremples 1986), California (coastally) southward to the
Santo Domingo River drainage in Baja California,
Mexico (Linsdale 1932). They also historically occurred
in a few desert slope drainages in southern California
(Jennings and Hayes 1994b). California red-legged frogs
generally occurred below 1370 m in the Sierra Nevada
foothills (Jennings 1996) and 1520 m in southern California, although some of the populations toward the
upper limit of the range of this frog may represent translocations (Jennings and Hayes 1994a). In the Bay Area,
this frog was historically abundant enough to support
an important commercial fishery just before the turn of
the century (Jennings and Hayes 1985) and up to the
202
Baylands Ecosystem Species and Community Profiles
Figure 4.4 California Red-Legged Frog – Some
Current Locations
catesbeiana) and crayfish (specifically Procambarus clarkii)
into breeding habitats (Miller et al. 1996). Juvenile and
adult frogs have also been found in a number of humancreated habitats such as irrigation canals, golf course
ponds, sewage treatment ponds, gravel pits, and intermittent irrigation ponds (Jennings and Hayes 1994a;
Jennings, unpubl. data). Such habitats may not be suitable for the long-term survival or successful reproduction of local frog populations, especially near urban areas where predators such as bullfrogs and raccoons (Procyon
lotor) are able to build up large populations as a result
of human activities (Jennings, unpubl. data).
Trophic Levels
Larval stages are primary consumers and post-metamorphic life stages are secondary/tertiary consumers.
Proximal Species
Good Habitat
Although California red-legged frogs can occur in
ephemeral or artificially-created ponds devoid of vegetation, the habitats that have been observed to have the
largest frog populations are perennial, deep (>0.7 m)
water pools bordered by dense, shrubby riparian vegetation (Jennings 1988, Hayes and Jennings 1986). This
dense riparian vegetation is characterized by arroyo willows (Salix lasiolepis) intermixed with an understory of
cattails (Typha sp.), tules (Scirpus sp.), or bulrushes (Scirpus sp.) [Jennings 1988].
References
Arnold, S.J. and T.Halliday. 1986. Life history notes:
Hyla regilla, predation. Herpetological Review,
17(2):44.
Baldwin, K.S. and R.A. Stanford. 1987. Life history
notes: Ambystoma tigrinum californiense, predation.
Herpetological Review, 18(2):33.
Green, D.M. 1985. Differentiation in heterochromatin
amount between subspecies of the red-legged frog,
Rana aurora. Copeia, 1985(4):1071-1074.
Hayes, M.P. and M.R. Jennings. 1988. Habitat correlates of distribution of the California red-legged
frog (Rana aurora draytonii) and the foothill yellow-legged frog (Rana boylii): implications for man-
Chapter 4 —
Amphibians and Reptiles
203
Amphibians &
Reptiles
Predators: Common [=San Francisco] garter snake,
Coast garter snake, Central Coast garter snake, bullfrog,
heron, egret, raccoon, and introduced predatory fishes.
Predacious aquatic insects prey on larvae only.
Prey: California tiger salamander, Pacific treefrog, California mouse, bullfrog, and aquatic and terrestrial insects.
Habitat: Willows, cattails, tules, sedges, and blackberries.
agement. Pages 144-158. In: R.C. Szaro, K.E.
Severson, and D.R. Patton (technical coordinators).
Management of Amphibians, Reptiles, and Small
Mammals in North America. Proceedings of the
Symposium, July 19-21, 1988, Flagstaff, Arizona.
U.S. Forest Serv., Rocky Mountain Forest and
Range Experiment Station, Fort Collins, Colorado.
General Tech. Report (RM-166):1-458.
Hayes, M.P. and D.M. Kremples. 1986. Vocal sac variation among frogs of the genus Rana from western
North America. Copeia, 1986(4):927-936.
Hayes, M.P. and M.M. Miyamoto. 1984. Biochemical,
behavioral and body size differences between the
red-legged frogs, Rana aurora aurora and Rana
aurora draytonii. Copeia, 1984(4):1018-1022.
Hayes, M.P. and M.R. Tennant. 1986. Diet and feeding behavior of the California red-legged frog, Rana
aurora draytonii (Ranidae). The Southwestern
Naturalist, 30(4): 601-605.
Jennings, M.R. 1988. Natural history and decline of
native ranids in California. Pages 61-72. In: H.F.
De Lisle, P.R. Brown, B.Kaufman, and B. McGurty
(eds). Proceedings of the Conference On California Herpetology. Southwestern Herpetologists Society, Special Publ. (4):1-143.
Jennings, M.R. 1996. Status of amphibians. Pages 311—31-24. In: Sierra Nevada Ecosystem Project:
Final Report to Congress, Volume II, Assessments
and Scientific Basis for Management Options. Centers for Water and Wetland Resources, Univ. of
Calif., Davis, Calif.
Jennings, M.R. and M.P. Hayes. 1985. Pre-1900 overharvest of the California red-legged frog (Rana aurora draytonii): The inducement for bullfrog (R. catesbeiana) introduction. Herpetologica, 41(1):94-103.
______. 1994a. Amphibian and reptile species of special concern in California. Final report to the Ca.
Dept. Fish and Game, Inland Fisheries Div.,
Rancho Cordova, Calif., under Contract (8023).
iii+255 p.
______. 1994b. The decline of native ranid frogs in the
desert southwest. Pages 183-211. In: P.R. Brown
and J.W. Wright (eds). Herpetology of the North
American Deserts: Proceedings of a Symposium.
Southwestern Herpetologists Society, Special Publ.
(5):iv+300 p.
Linsdale, J.M. 1932. Amphibians and reptiles from
Lower California. Univ. of Calif. Publications in
Zoology, 38(6):345-386.
Livezey, R.L. and A.H. Wright. 1947. A synoptic key to
the salientian eggs of the United States. The American Midland Naturalist, 37(1):179-222.
Miller, K.J. 1994. Endangered and threatened wildlife
and plants; proposed endangered status for the
California red-legged frog. Fed. Reg., 59(22):48884895. [Wednesday, February 2, 1994].
Western Pond Turtle
Clemmys marmorata
Mark R. Jennings
General Information
The western pond turtle (Family: Emydidae) is a moderate-sized (120-210 mm CL), drab brown or khaki-colored turtle often lacking prominent markings on its carapace (Bury and Holland, in press). Carapace coloration
is usually dark brown or dull yellow-olive, with or without darker streaks or vermiculations radiating from the
centers of the scutes (Ernst et al. 1994, Jennings and
Hayes 1994).
There are two poorly differentiated subspecies of
the western pond turtle (C. m. marmorata and C. m.
pallida) with a wide zone of intergradation in central
California (Bury 1970). Based on a morphological evaluation, Holland (1992) found three distinct evolutionary
groups within this taxon. However, Gray (1995) found
through DNA fingerprinting of C. marmorata samples
from the extreme southern and northern edges of its
range support the original designation of two distinct
subspecies. Bury and Holland (in press) indicate that
more comprehensive genetic studies are currently underway to determine the taxonomic status of this taxon.
In 1992, the western pond turtle was petitioned for
listing as an endangered species (Sorensen and Propp
1992) based on concerns about widespread population
declines due to the extensive loss of habitat, overexploitation, introductions of non-native aquatic predators
(Sorensen and Propp 1992; see also Jennings and Hayes
1994). The U.S. Fish and Wildlife Service subsequently
ruled that the petition was not warranted (USFWS 1993)
and this turtle remains a candidate 2 species (Drewry
1994).
Reproduction
In California, sexual maturity in western pond turtles
occurs at between seven and 11 years of age at approximately 110-120 mm CL with males maturing at slightly
Joe DiDonato
Amphibians &
Reptiles
Miller, K.J., A. Willy, S. Larsen, and S. Morey. 1996.
Endangered and threatened wildlife and plants;
determination of threatened status for the California red-legged frog. Fed. Reg, 61(101):2581325833. [Thursday, May 23, 1996].
Rathbun, G.B., M.R. Jennings, T.G. Murphey, and N.R.
Siepel. 1993. Status and ecology of sensitive aquatic
vertebrates in lower San Simeon and Pico Creeks,
San Luis Obispo County, California. Final report
prepared for the Calif. Dept. of Parks and Recreation, San Simeon Region, through Cooperative
Agreement (14-16-0009-01-1909). U.S. Fish and
Wildl. Serv., Natl. Ecology Research Center,
Piedras Blancas Research Station, San Simeon,
Calif. ix+103 p.
Sorensen, P.C. 1993. Endangered and threatened wildlife and plants; finding on petition to list the California red-legged frog. Fed. Reg. 58(136):38553.
[Monday, July 19, 1993].
Stebbins, R.C. 1959. Reptiles and amphibians of the
San Francisco Bay region. California Natural History Guide (3). Univ. of Calif. Press, Berkeley and
Los Angeles. 72 p.
______. 1972. Amphibians and reptiles of California.
California Natural History Guide (31). Univ. of
Calif. Press, Berkeley, Los Angeles, and London.
152 p.
Storer, T.I. 1925. A synopsis of the Amphibia of California. Univ. of Calif. Publications in Zoology,
27:1-342.
204
Baylands Ecosystem Species and Community Profiles
smaller sizes and ages than females (Jennings and Hayes
1994). Sexual maturity is delayed in turtles that experience drought conditions and in more northerly populations (Jennings and Hayes 1994). Adult turtles typically
mate in late April or early May, although mating can
occur year-around (Holland 1985a, 1992). The nesting
season is from late April to early August (Storer 1930,
Buskirk 1992, Rathbun et al. 1992, Jennings and Hayes
1994, Goodman 1997a). Females emigrate from the
aquatic habitats to an unshaded, upland location that
may be a considerable distance (400 m or more) from
riparian zones to nest (Storer 1930; Rathbun et al. 1992;
Bury and Holland, in press). However, most nest locations are close to riparian zones if nesting substrates and
exposures are suitable (Jennings, unpubl. data). Once a
suitable site is located, females deposit from 1-13 eggs
that have a thin, but hard (calcified) outer shell in a shallow (ca. 10-12 cm deep) nest (Rathbun et al. 1992,
1993)—usually in well-drained clay or silt soils (Jennings
and Hayes 1994). Eggs laid in excessively moist substrates have a high probability of failing (Feldman 1982).
Females can lay more than one clutch of eggs a year and
may dig several “ false” nests lacking eggs to deter potential predators (Rathbun et al. 1993, Goodman 1997b).
Young turtles hatch at lengths of 25-29 mm CL (Ernst
et al. 1994) after an incubation period of 3-4.5 months
(Buskirk 1992; Bury and Holland, in press) and are
thought to overwinter in the nest because there are only
a few records of hatchling turtle emergence in the early
fall in southern and central California (Buskirk 1992,
Jennings and Hayes 1994). Most hatchling turtles are
thought to emerge from the nest and move to aquatic
sites in the spring (Buskirk 1992) where they typically
double their length the first year and grow relatively rapidly over the next 4-5 years (Storer 1930, Holland
1985a). Young turtles spend most of their time feeding
in shallow water dominated by relatively dense vegetation of submergents, short emergents, or algal mats
(Buskirk 1992, Jennings and Hayes 1994). Juveniles and
adults prefer slack- or slow-water aquatic habitats with
basking sites such as rocks and logs (Bury 1972, Reese
1996). Water temperatures >15° C markedly increase
turtle activity so many western pond turtles are probably
active year around in coastal locations (Reese and Welsh
1998) and only active from March or April-October or
November in interior locations (Bury and Holland, in
press). Juveniles and adults seem to remain in pond environments except when ponds dry up or at higher elevations when turtles may disperse into terrestrial environments to hibernate (Jennings and Hayes 1994;
Goodman 1997a, Bury and Holland, in press). In stream
environments, juveniles and adults show considerable
variation with regards to movements and the timing of
Food and Feeding
Juvenile and adult western pond turtles feed largely on
the same items although juveniles feed more on smaller
aquatic invertebrates (Bury 1986). Food items found in
turtle stomachs include: algae, aquatic plants, Nematomorpha, Cladocera, Decapoda, Isopoda, Ephemeroptera
(nymphs only), Odonata (nymphs only), Orthoptera,
Hemiptera (nymphs and adults), Neuroptera (larvae
only), Coleoptera (larvae and adults), Trichoptera (larvae only), Diptera (larvae and adults), Araneae, Gastropoda, fishes, and amphibians (Carr 1952, Holland
1985b, Bury 1986, Ernst et al. 1994, Goodman 1997a).
These turtles are dietary generalists and highly opportunistic (Ernst et al. 1994). They will consume almost
anything that they are able to catch and overpower. The
relatively slow pursuit of these turtles results in their diet
being dominated by relatively slow-moving aquatic invertebrates and carrion, although aquatic vegetation may
also be eaten (Evenden 1948, Bury 1986, Baldwin and
Stanford 1987), especially by females having recently laid
eggs (Jennings and Hayes 1994).
Distribution
The western pond turtle historically occurred in most
Pacific slope drainages from Klickitat County, Washington along the Columbia River (Slater 1962) south to
Arroyo Santa Domingo, northern Baja California,
Chapter 4 —
Amphibians and Reptiles
205
Amphibians &
Reptiles
Growth and Development
movements into terrestrial environments (Rathbun et al.
1993, Reese and Welsh 1998). Some turtles will leave
the stream during the summer when water conditions
are low and water temperatures are elevated (>35° C),
while others will not. However, almost turtles seem to
leave streams during the winter months when large flood
events are common (Rathbun et al. 1993). Additionally,
some turtles will move considerable distances (e.g., 350
m) to overwinter in terrestrial habitats such as leaf litter
or under the root masses of trees (Rathbun et al. 1992,
1993). Some individual turtles have displayed site fidelity for hibernation sites from year to year (Bury and
Holland, in press).
Western pond turtles often move about from pool
to pool in stream situations, sometimes on a daily basis
during seasons of activity (Bury 1972, Reese and Welsh
1998). Distances moved along streams can be up to 5
km (Bury and Holland, in press). These turtles also have
the ability to move several kilometers if their aquatic
habitat dries up (Reese 1996) and they can tolerate at
least seven days without water (Jennings and Hayes
1994; Bury and Holland, in press).
Western pond turtles are known to live over 42
years in the wild (Jennings and Hayes 1994) although
most individuals have a much shorter life span of around
20-25 years (Bury 1972).
Mexico (Jennings and Hayes 1994). Isolated populations
are also known from Carson, Humboldt, and Truckee
drainages in western Nevada (LaRivers 1962, Banta
1963). In California the species is known from most
Pacific slope drainages between the Oregon and Mexican borders below 1430 m (Jennings and Hayes 1994).
Turtle observations from above this elevation are thought
to be introductions (Jennings and Hayes 1994). Western pond turtles are present throughout the Bay Area
(Stebbins 1959) (Figure 4.5), although at much lower
numbers and at fewer localities than previously—especially in urban areas (Jennings, unpubl. data).
Amphibians &
Reptiles
Current Status and Factors Influencing
Population Numbers
The western pond turtle is declining in population size
and numbers throughout its range, particularly in southern California and the San Joaquin Valley (Bury and
Holland, in press). Many turtle populations in these areas
of decline are now composed almost entirely of old adults
without any successful recruitment (Jennings and Hayes
1994). The reasons for these declines are largely due to
urbanization, agricultural development, flood control
projects, exotic diseases, exploitation for the food and pet
trade, extended drought, and the introduction of exotic
predatory species such as largemouth bass (Micropterus
salmoides) and bullfrogs (Rana catesbeiana) which also
compete for the availability of prey items—especially
with young turtles (Brattstrom 1988; Buskirk 1992; Jennings and Hayes 1994; Reese 1996; Goodman 1997a;
Bury and Holland, in press). Turtle nests and gravid
females moving overland are especially vulnerable to
predation by raccoons (Procyon lotor), whose populations
have greatly increased in many rural areas due to the
increase in human populations in these areas (Bury and
Holland, in press; Jennings, unpubl. data). Additionally,
some of the largest turtle populations in the Central
Valley and southern California are found in sewage treatment ponds. Unfortunately, such habitats are probably
unsuitable for the long term survival of the species because of the lack of suitable habitat for nest sites and
increased vulnerability of adult turtles to predation by humans, raccoons, and other animals (Jennings, unpubl. data).
Trophic Levels
Young turtles are essentially secondary consumers; adults
are primary and secondary consumers.
Proximal Species
Predators: Raccoon, bullfrog, black bear, humans, and
introduced predatory fishes. Striped skunk and opossum
prey on eggs and hatchlings, and herons and egrets prey
on young turtles.
206
Baylands Ecosystem Species and Community Profiles
Figure 4.5 Western Pond Turtle – Presumed Bay
Area Distribution
Prey: Aquatic Insects, aquatic vegetation, and California tiger salamander larvae.
Habitat: Aquatic vegetation.
Good Habitat
The largest western pond turtle populations have been
observed in warm water (15-35° C), slack- or slow-water habitats, which have abundant basking sites and underwater refugia. The presence of dense stands of
submergent or emergent vegetation, and abundant
aquatic invertebrate resources, as well suitable nearby
nesting sites and the lack of native and exotic predators,
are also important components (Bury 1972; Jennings and
Hayes 1994; Bury and Holland, in press).
References
Baldwin, K.S. and R.A. Stanford. 1987. Life history
notes: Ambystoma tigrinum californiense, predation.
Herpetological Review, 18(2):33.
Banta, B.H. 1963. On the occurrence of Clemmys
marmorata (Reptilia, Testudinata) in western Nevada. The Wasmann J. of Biology, 21(1):75-77.
Brattstrom, B.H. 1988. Habitat destruction in California with special reference to Clemmys marmorata:
A perpesctive [sic]. Pages 13-24. In: H.F. De Lisle,
P.R. Brown, B.Kaufman, and B. McGurty (eds).
Proceedings of the Conference On California Her-
______. 1992. Level and pattern in morphological variation: A phylogeographic study of the western pond
turtle (Clemmys marmorata). Unpubl. Ph.D. Dissertation, The Univ. of Southwestern Louisiana,
Lafayette, Louisiana. vii+124 p.
Jennings, M.R. and M.P. Hayes. 1994. Amphibian and
reptile species of special concern in California. Final report to the Ca. Dept. Fish and Game, Inland
Fisheries Div., Rancho Cordova, Calif., under
Contract (8023). iii+255 p.
LaRivers, I. 1962. Fishes and fisheries of Nevada. Nevada State Fish and Game Commission, Carson
City, Nevada. 782 p.
Rathbun, G.B., N. Siepel, and D.C. Holland. 1992.
Nesting behavior and movements of western pond
turtles (Clemmys marmorata). The Southwestern
Naturalist, 37(3): 319-324.
Rathbun, G.B., M.R. Jennings, T.G. Murphey, and N.R.
Siepel. 1993. Status and ecology of sensitive aquatic
vertebrates in lower San Simeon and Pico Creeks,
San Luis Obispo County, California. Final report
prepared for the Calif. Dept. of Parks and Recreation, San Simeon Region, through Cooperative
Agreement (14-16-0009-01-1909). U.S. Fish and
Wildl. Serv., Natl. Ecology Research Center,
Piedras Blancas Research Station, San Simeon,
Calif. ix+103 p.
Reese, D.A. 1996. Comparative demography and habitat use of western pond turtles in northern California: The effects of damming and related alterations. Unpubl. Ph.D. Dissertation, Univ. of Ca.,
Berkeley. xvi+253 p.
Reese, D.A. and H.H. Welsh, Jr. 1998. Habitat use by
western pond turtles in the Trinity River, Calif. J.
of Wildl. Management, 62(3):242-253.
Slater, J.R. 1962. Variations and new range of Clemmys
marmorata. Occasional Papers of the Dept. of Biology, Univ. of Puget Sound (20):204-205.
Sorensen, P. and L.J. Propp. 1992. Endangered and
threatened wildlife and plants; 90-day finding and
commencement of status reviews for a petition to
list the western pond turtle and California redlegged frog. Fed. Reg., 57(193):4561-45762.
[Monday, October 5, 1992].
Stebbins, R.C. 1959. Reptiles and amphibians of the
San Francisco Bay region. California Natural History Guide (3). Univ. of Ca. Press, Berkeley and
Los Angeles. 72 p.
Storer, T.I. 1930. Notes on the range and life-history of the
Pacific fresh-water turtle, Clemmys marmorata. Univ. of
Ca. Publications in Zoology, 35(5):429-441.
U.S. Fish and Wildlife Service (USFWS). 1993. Endangered and threatened wildlife and plants; notice of 1-year petition finding on the western pond
turtle. Fed. Reg., 58(153):42717-42718. [Wednesday, August 11, 1993].
Chapter 4 —
Amphibians and Reptiles
207
Amphibians &
Reptiles
petology. Southwestern Herpetologists Society,
Special Publ. (4):1-143.
Bury, R.B. 1970. Clemmys marmorata. Catalogue of
American Amphibians and Reptiles:100.1-100.3.
______. 1972. Habits and home range of the Pacific
pond turtle, Clemmys marmorata. Unpubl. Ph.D.
Dissertation, Univ. of Ca., Berkeley, Calif. 205 p.
______. 1986. Feeding ecology of the turtle, Clemmys
marmorata. J. of Herpetology, 20(4):515-521.
Bury, R.B. and D.C. Holland (in press). Clemmys
marmorata (Baird and Girard 1852). Conservation
Biology of Freshwater Turtles, 2.
Buskirk, J.R. 1992. An overview of the western pond
turtle, Clemmys marmorata. Pages 16-23. In: K. R.
Beaman, F. Caporaso, S. McKeown, and M. Graff
(eds). Proceedings of the First International Symposium on Turtles and Tortoises: Conservation and
Captive Husbandry. California Turtle and Tortoise
Club, Van Nuys, Calif. 172 p.
Carr, A.F. 1952. Handbook of turtles: The turtles of the
United States, Canada, and Baja California. Cornell
Univ. Press, Ithaca, New York. xv+542 p.
Drewry, G. (ed). 1994. Endangered and threatened wildlife and plants; animal candidate review for listing
as endangered or threatened species. Dept. of the
Interior, Fish and Wildl. Serv. Fed. Reg.
59(219):58982-59028. [Tuesday, November 15,
1994].
Ernst, C.H., J.E. Lovich, and R.W. Barbour. 1994.
Turtles of the United States and Canada.
Smithsonian Institution Press, Washington and
London. xxxviii+578 p.
Evenden, F.G. 1948. Distribution of the turtles of western Oregon. Herpetologica, 4(6): 201-204.
Feldman, M.1982. Notes on reproduction in Clemmys
marmorata. Herpetological Review, 13(1):10-11.
Goodman, R.H., Jr. 1997a. The biology of the southwestern pond turtle (Clemmys marmorata pallida)
in the Chino Hills State Park and the West Ford of
the San Gabriel River, M.S. Thesis, Calif. State
Polytechnic Univ., Pomona, Calif. viii+81 p.
______. 1997b. Occurrence of devoie clutching in the
southwestern pond turtle, Clemmys marmorata
pallida, in the Los Angeles Basin. Chelonian Conservation and Biology, 2(3):419-421.
Gray, E.M. 1995. DNA fingerprinting reveals a lack of
genetic variation in northern populations of the
western pond turtle (Clemmys marmorata). Conservation Biology, 9(5):1244-1255.
Holland, D.C. 1985a. An ecological and quantitative
study of the western pond turtle (Clemmys
marmorata) in San Luis Obispo County, California. Unpubl. M.A. Thesis, Fresno State Univ.,
Fresno, Calif. viii+181 p.
______. 1985b. Life history notes: Clemmys marmorata,
feeding. Herpetological Review, 16(4):112-113.
Growth and Development
California Alligator Lizard
Elgaria multicarinata multicarinata
Kevin MacKay
Mark R. Jennings
General Information
Reproduction
California alligator lizards are egg layers (Smith 1946)
that probably reach sexual maturity in two years at about
73 mm SVL for males and about 92 mm SVL for females
(Goldberg 1972). Mating apparently occurs over a relatively long period (up to 26 hours or more), but most
copulation events are considerably shorter than this
(Fitch 1935). Based on data from closely related E. m.
webbii in southern California, adults emerge from hibernation and mate from late February-late May to August-mid September, and eggs are probably laid in small
mammal burrows (Stebbins 1954) or under rocks from
June-mid July (Goldberg 1972) or later into August
(Stebbins 1954) or early September (Burrage 1965).
Clutch sizes are 5-41 (average 13) and females can lay
more than one clutch a year (Burrage 1965).
208
Food and Feeding
California alligator lizards probably consume the same
food items taken by E. m. webbii. Food items recorded
in the latter include: Isopoda, Orthoptera, Isoptera,
Hemiptera, Homoptera, Coleoptera (larvae and adults),
Lepidoptera (larvae and adults), Diptera (larvae and
adults), Scorpionida, Araneida [including the egg cases
and adults of the black widow spider (Latradectus
mactans)], Gastropoda, lizards (Sceloporus occidentalis, S.
graciosus, and E. multicarinata), small mammals, and the
eggs and young of small birds (Fitch 1935, Cowles 1937,
Stebbins 1954, Cunningham 1956).
Distribution
California alligator lizards are found in the Sacramento
Valley and surrounding foothills, from Shasta County
south through the North Coast Range (MendocinoMarin counties), the San Francisco Bay region and the
South Coast Range to Ventura County (Fitch 1938).
The elevational range is from sea level to around 1830
m in the Sierra Nevada (Basey and Sinclear 1980). This
lizard is apparently absent from most of the San Joaquin
Valley proper, but it is found on the northern Channel
Islands (Fitch 1938). It intergrades with the E. m.
scincicauda in Mendocino and Trinity counties in the north
and E. m. webbii in Ventura County in the south and El
Dorado County in the east (Stebbins 1985). They are found
throughout the Bay Area (Stebbins 1959) (Figure 4.6).
Jens V. Vindum, Academy of Sciences
Amphibians &
Reptiles
The California alligator lizard (Family: Anguidae) is a
large (100-125 mm SVL) alligator lizard with a broad
head, keeled scales, and a reddish-blotched dorsum
marked with nine or more dusky crossbands between the
head and hindlimbs (Stebbins 1959, 1985). The top of
the head is often mottled (Fitch 1938). There is a longitudinal stripe or row of dashes down the middle of each
scale row on the belly (Stebbins 1985).
All alligator lizards in the western United States
were formerly placed in the genus Gerrhonotus (e.g.,
Smith 1946; Stebbins 1958, 1959, 1972, 1985; Lais
1976). However, recently revised alligator lizard systematics places these species in the genus Elgaria (Waddick
and Smith 1974; Gauthier 1982; Good 1987a, 1987b,
1988).
Based on data from closely related E. m. webbii in southern California, incubation of the eggs probably takes 4257 days (Atsatt 1952, Burrage 1965) and hatchlings appear from mid August-early October at 30-36 mm SVL
and 0.5-0.6 g (Burrage 1965, Goldberg 1972). Juveniles
grow rapidly the next season, reaching sexual maturity
after about 18 months (Goldberg 1972). The longevity
of California alligator lizards in the wild is unknown, but
marked lizards have been recaptured after four years (Jennings, unpubl. data).
Both juveniles and adults are active in the daytime,
at dusk, and at night, and have a relatively low preferred
temperature range (Brattstrom 1965, Cunningham
1956, Kingsbury 1994). Because of this, they do not
bask. Instead, they prefer very dense cover and often
position themselves under warmed objects such as rocks
or pieces of wood during certain times of the day
(Kingsbury 1994). Alligator lizards frequent riparian
zones where their prehensile tails are used in climbing
trees and other vegetation in pursuit of prey
(Cunningham 1955; Stebbins 1959, 1972). They are
also found under debris such as woodpiles, brush heaps,
old logs, etc. (Stebbins 1954).
Baylands Ecosystem Species and Community Profiles
Current Status and Factors Influencing
Population Numbers
California alligator lizards are still present in good numbers over almost all of their historic range because of their
ability to survive (and even thrive) in urban environments. The most important predator of these lizards in
such modified habitats is the domestic cat (Felis cattus).
In more natural habitats, alligator lizards are eaten by a
number of reptile, avian, and mammal predators (Fitch
1935). They are still very abundant in the foothills of
the Bay Area (Jennings, unpubl. data).
Trophic Levels
California alligator lizards are secondary/tertiary consumers.
Proximal Species
Figure 4.6 California Alligator Lizard – Presumed
Bay Area Distribution
California alligator lizards occupy many habitats from
pickleweed flats to open grasslands, to oak woodlands,
to mixed coniferous forest, to urban environments (Fitch
1935; Lais 1976; Stebbins 1954, 1985). However, the
largest observed populations are in the riparian zones of
oak woodlands and in coastal sage scrub near beaches
(Jennings, unpubl. data).
References
Atsatt, S.R. 1952. Observations on the life history of
the lizards Sceloporus graciosus vandenburghianus
and Gerrhonotus multicarinatus webbi. Copeia,
1952(4):276.
Basey, H.E. and D.A. Sinclear. 1980. Amphibians and
reptiles. Pages 13-74. In: J. Verner and A.S. Boss
(technical coordinators). California Wildlife and
Their Habitats: Western Sierra Nevada. U.S. Forest Serv., Pacific Southwest Forest and Range Experiment Station, Berkeley, Calif. General Tech.
Report (PSW-37):iii+439 p.
Brattstrom, B.H. 1965. Body temperatures of reptiles.
The American Midland Naturalist, 73(2):376-422.
Burrage, B.R. 1965. Notes on the eggs and young of
the lizards Gerrhonotus multicarinatus webbi and
G. m. nanus. Copeia, 1965(4):512.
Cowles, R.B. 1937. The San Diegan alligator lizard and
the black widow spider. Science, 85(2195):99-100.
Cunningham, J.D. 1955. Arboreal habits of certain reptiles and amphibians in southern California.
Herpetologica, 11(3):217-220.
______. 1956. Food habits of the San Diego alligator
lizard. Herpetologica, 12(3):225-230.
Fitch, H.S. 1935. Natural history of the alligator lizards. Transactions of the Academy of Science of
Saint Louis, 29(1):3-38.
______. 1938. A systematic account of the alligator lizards (Gerrhonotus) in the western United States and
Lower Calif. The American Midland Naturalist,
20(2): 381-424.
Gauthier, J. A. 1982. Fossil xenosaurid and anguid lizards from the early Eocene Wasatch Formation,
southeast Wyoming, and a revision of the
Anguioidea. Contributions to Geology, Univ. of
Wyoming, 21(1):7-54.
Goldberg, S.R. 1972. Reproduction in the southern alligator lizard Gerrhonotus multicarinatus.
Herpetologica, 28(3):267-273.
Good, D.A. 1987a. An allozyme analysis of anguid
subfamilial relationships (Lacertilia: Anguidae).
Copeia, 1987(3):696-701.
______. 1987b. A phylogenetic analysis of cranial osteology in the Gerrhonotine lizards. J. of Herpetology, 21(4):285-297.
Chapter 4 —
Amphibians and Reptiles
209
Amphibians &
Reptiles
Habitat: Pickleweed, riparian vegetation, blackberries,
willows.
Predators: Domestic cat, striped skunk, opossum, raccoon, heron, egret, hawks, coyote, red fox, bullfrog,
common garter snake, and Coast garter snake.
Prey: Terrestrial insects, oligochaetes, and arachnids.
Good Habitat
Central Coast Garter Snake
Thamnophis atratus atratus
Mark R. Jennings
General Information
The central coast garter snake (Family: Colubridae) is a
medium-sized (60-102 cm TL) garter snake with eight
upper labial scales and a highly variable dorsal color
throughout its range (Bellemin and Stewart 1977).
Snakes usually have a dark olive to black dorsum and
single yellow to orange dorsal stripe and sometimes lateral stripes of pale yellow (Stebbins 1985). The throat
is a bright yellow. Both T. a. atratus and T. e. terrestris
have similar dorsal and ventral colorations in habitats
occupied along the central coast of California (Bellemin
and Stewart 1977, Stebbins 1985). Boundy (1990) considers what is currently T. a. atratus, to be be actually
composed of two different subspecies. However, his
proposed subspecies from the mountains of the East Bay
region and the South Coast Range (south of Santa Cruz
County) has not been formally published.
Garter snake taxonomy has undergone a considerable number of revisions during this century, especially
during the past 40 years. The snake T. a. atratus is often referred to as T. elegans atratus, or T. couchii atratus
in the literature (e.g., see Fitch 1940, 1984; Fox 1948a,
1948b, 1951; Stebbins 1954, 1972; Fox and Dessauer
1965; Lawson and Dessauer 1979). Rossman and
Stewart (1987) were the first to convincingly elevate T.
atratus as a separate species and this arrangement has
been followed by others (e.g., see Lawson 1987).
Reproduction
Central coast garter snakes are live-bearers. Females give
birth from 4-14 (or more) young (ave. 8.6) in the fall
(August-September) [Fox 1948a, 1948b]. Adults probably mate annually during the spring (March-April) [Fox
1948b, 1952a], but females have the ability to store
sperm for up to 53 months (Stewart 1972).
Denise Loving
Amphibians &
Reptiles
______. 1988. Allozyme variation and phylogenetic relationships among the species of Elgaria (Squamata:
Anguidae). Herpetologica, 44(2):154-162.
Kingsbury, B.A. 1994. Thermal constraints and
eurythermy in the lizard Elgaria multicarinata.
Herpetologica, 50(3):266-273.
Lais, P.M. 1976. Gerrhonotus multicarinatus. Catalogue
of American Amphibians and Reptiles:187.1187.4.
Smith, H.M. 1946. Handbook of lizards; lizards of the
United States and of Canada. Comstock Publishing Co., Inc., Ithaca, New York. xxi+557 p.
Stebbins, R.C. 1954. Amphibians and reptiles of western North America. McGraw-Hill Book Company,
Inc., New York, Toronto, and London. xxii+536
p.
______. 1958. A new alligator lizard from the Panamint
Mountains, Inyo County, California. American
Museum Novitates (1883):1-27.
______. 1959. Reptiles and amphibians of the San Francisco Bay region. California Natural History Guide
(3). Univ. of Ca. Press, Berkeley and Los Angeles.
72 p.
______. 1972. Amphibians and reptiles of California.
California Natural History Guide (31). Univ. of
Ca. Press, Berkeley, Los Angeles, and London. 152
p.
______. 1985. A field guide to western amphibians and
reptiles. Second edition, revised. Houghton Mifflin
Company, Boston, Massachusetts. xiv+336 p.
Waddick, J.W. and H.M. Smith. 1974. The significance
of scale characters in evaluation of the lizard genera Gerrhonotus, Elgaria, and Barisia. The Great
Basin Naturalist, 34(4):257-266.
210
Baylands Ecosystem Species and Community Profiles
Growth and Development
Proximal Species
Unknown. If similar to other garter snakes on the central coast of California, neonates are present from late
August through November (Rathbun et al. 1993) and
juveniles grow rapidly during the first year of their lives (Fox
1948a). Sexual maturity is reached in about 2-3 years (Fox
1948a). Longevity in the wild is unknown, but adults probably live for at least 4-5 years (Jennings, unpubl. data).
Predators: Racoon, herons, egrets, hawks, and bullfrogs.
Prey: threespine stickleback, sculpins, Pacific treefrog,
California toad, foothill yellow-legged frog, California
red-legged frog, bullfrog, coast range newt.
Food and Feeding
Juvenile and adult snakes feed almost entirely on fishes
(e.g., Gasterosteus aculeatus, Hesperoleucus symmetricus,
and Cottus spp.), newts (larvae and adults of Taricha
torosa), toads (Bufo boreas halophilus), and frogs (e.g.,
larvae, juveniles, and adults of Hyla regilla, Rana aurora
draytonii, R. boylii, and R. catesbeiana) [Fitch 1941; Fox
1951, 1952b; Bellemin and Stewart 1977; Boundy 1990;
Barry 1994; Jennings, unpubl. data].
Distribution
Coast garter snakes are most abundant in riparian habitats with shallow ponds containing abundant numbers
of native fishes and amphibians, and dense thickets of
vegetation nearby (Jennings, unpubl. data). Such habitats are most common in natural sag ponds and artificial stock ponds (Barry 1994).
References
Barry, S.J. 1994. The distribution, habitat, and evolution of the San Francisco garter snake, Thamnophis sirtalis tetrataenia. Unpubl. M.A. Thesis, Univ.
of Ca., Davis, Calif. iii+140 p.
Bellemin, J.M. and G. R.Stewart. 1977. Diagnostic characters and color convergence of the garter snakes
Thamnophis elegans terrestris and Thamnophis
couchii atratus along the central California coast.
Bull. of the Southern Calif. Academy of Sciences,
76(2):73-84.
Current Status and Factors Influencing
Population Numbers
Central coast garter snakes are negatively affected by
habitat alteration, especially by agriculture and urbanization which often results in intermittent aquatic habitats unsuitable for this species. These snakes are also
probably negatively affected by the introduction of exotic predators such as bullfrogs (Rana catesbeiana) and
largemouth bass (Micropterus salmoides), which are
known to eat garter snakes (Schwalbe and Rosen 1988).
However, these central coast garter snakes are still relatively abundant in aquatic habitats located in the foothills surrounding the Bay Area where urban development
is less intrusive (Jennings, unpubl. data).
Trophic Levels
Central coast garter snakes are tertiary consumers.
Figure 4.7 Central Coast Garter Snake – Presumed Bay Area Distribution
Chapter 4 —
Amphibians and Reptiles
211
Amphibians &
Reptiles
Central coast garter snakes inhabit small streams, ponds,
and other aquatic habitats in the San Francisco Peninsula and the East Bay Hills, Contra Costa County (south
of the Sacramento River), southward through the South
Coast Range to Point Conception, Santa Barbara
County, and east to the western edge of the San Joaquin
Valley (Fox 1951, Bellemin and Stewart 1977). Snakes
north of San Francisco Bay are T. a. aquaticus (Fox 1951,
Stebbins 1985). Their elevational distribution is from
near sea level to 1290 m on Mount Hamilton (Fox
1951). They are relatively common East Bay and South
Bay regions of the San Francisco Estuary (Stebbins 1959;
Jennings, unpubl. data) (Figure 4.7).
Good Habitat
212
Baylands Ecosystem Species and Community Profiles
Flagstaff, Arizona. U.S. Forest Serv., Fort Collins,
Colorado. General Tech. Report (RM-166):1-458.
Stebbins, R.C. 1954. Amphibians and reptiles of western North America. McGraw-Hill Book Company,
Inc., New York, Toronto, and London. xxii+536 p.
______. 1959. Reptiles and amphibians of the San Francisco Bay region. California Natural History Guide (3).
Univ. of Ca. Press, Berkeley and Los Angeles. 72 p.
______. 1972. Amphibians and reptiles of California.
California Natural History Guide (31). Univ. of Ca.
Press, Berkeley, Los Angeles, and London. 152 p.
______. 1985. A field guide to western amphibians and
reptiles. Second edition, revised. Houghton Mifflin
Company, Boston, Massachusetts. xiv+336 p.
Stewart, G.R. 1972. An unusual record of sperm storage in a female garter snake (genus Thamnophis).
Herpetology, 28(4):346-347.
Coast Garter Snake
Thamnophis elegans terrestris
Kevin MacKay
Mark R. Jennings
General Information
The coast garter snake (Family: Colubridae) is a medium-sized (45-107 cm TL) garter snake with eight
upper labial scales and a highly variable dorsal color
throughout its range (Bellemin and Stewart 1977, Stebbins 1985). Snakes usually have a reddish to solid black
dorsum (sometimes with a checkerboard of dark spots
or bars), and single pale to bright yellow dorsal stripe,
and two lateral stripes of yellow to salmon (Fitch
1983, Stebbins 1985). The throat and belly are usually tinged with orange flecks (Fox 1951). Both T. a.
atratus and T. e. terrestris have similar dorsal and ven-
Dr. Alan Francis
Amphibians &
Reptiles
Boundy, J. 1990. Biogeography and variation in southern populations of the garter snake Thamnophis
atratus, with a synopsis of the T. couchii complex.
Unpubl. M.A. Thesis, San Jose State Univ., San
Jose, Calif. 105 p.
Fitch, H.S. 1940. A biogeographical study of the
ordinoides artenkries of garter snakes (genus Thamnophis). Univ. of Ca. Publications in Zoology,
44(1):1-150.
______. 1941. The feeding habits of California garter
snakes. Ca. Dept. Fish and Game, 27(2):1-32.
______. 1984. Thamnophis couchii. Catalogue of American Amphibians and Reptiles:351.1-351.3.
Fox, W. 1948a. The relationships of the garter snake
Thamnophis ordinoides. Copeia, 1948(2):113-120.
______. 1948b. Effect of temperature on development
of scutellation in the garter snake, Thamnophis
elegans atratus. Copeia, 1948(4):252-262.
______. 1951. Relationships among the garter snakes
of the Thamnophis elegans rassenkreis. Univ. of Ca.
Publications in Zoology, 50(5):485-530.
______. 1952a. Seasonal variation in the male reproductive system of Pacific Coast garter snakes. J. of
Morphology, 90(3):481-554.
______. 1952b. Notes on feeding habits of Pacific Coast
garter snakes. Herpetologica, 8(1):4-8.
Fox, W. and H.C. Dessauer. 1965. Collection of garter
snakes for blood studies. American Philosophical
Society Yearbook for 1964:263-266.
Lawson, R. 1987. Molecular studies of thamnophine
snakes: 1. The phylogeny of the genus Nerodia. J.
of Herpetology, 21(2):140-157.
Lawson, R. and H.C. Dessauer. 1979. Biochemical genetics and systematics of garter snakes of the Thamnophis elegans-couchii-ordinoides complex. Occasional Papers of the Museum of Zoology, Louisiana State Univ. (56):1-24.
Rathbun, G.B., M.R. Jennings, T.G. Murphey, and N.R.
Siepel. 1993. Status and ecology of sensitive aquatic
vertebrates in lower San Simeon and Pico Creeks,
San Luis Obispo County, California. Final report
prepared for the Calif. Dept. of Parks and Recreation, San Simeon Region. U.S. Fish and Wildl.
Serv., Natl. Ecology Research Center, San Simeon,
Calif. ix+103 p.
Rossman, D.A. and G.R. Stewart. 1987. Taxonomic
reevaluation of Thamnophis couchii (Serpentes,
Colubridae). Occasional Papers of the Museum of
Zoology, Louisiana State Univ. (63):1-25.
Schwalbe, C.R. and P.C. Rosen. 1988. Preliminary report on effect of bullfrogs on wetland herpetofaunas in southeastern Arizona. Pages 166-173. In:
R.C. Szaro, K.E. Severson, and D.R. Patton (technical coordinators). Management of Amphibians,
Reptiles, and Small Mammals in North America.
Proceedings of the Symposium, July 19-21, 1988,
tral colorations in habitats occupied along the central
coast of California (Bellemin and Stewart 1977, Stebbins 1985).
Reproduction
Coast garter snakes are live-bearers. Females give birth
to from 4-14 young (average 8.6) in the fall (August-September) [Fox 1948, Stebbins 1954]. Adults probably
mate annually during the spring (March-July) [Fox 1948,
1952a, 1956], but females have the ability to store sperm
for up to 53 months (Stewart 1972).
Growth and Development
Unknown. If similar to other garter snakes on the central coast of California, neonates are present from late
August through November (Rathbun et al. 1993) and
juveniles grow rapidly during the first year of their lives
(Fox 1948). Sexual maturity is reached in about 2-3 years
(Fox 1948). Longevity in the wild is unknown, but adults
probably live for at least 4-5 years (Jennings, unpubl.
data).
Figure 4.8 Coast Garter Snake – Presumed Bay
Area Distribution
Coast garter snakes subsist largely on slugs (Arion sp.,
Ariolimax columbianus, and others), California slender
salamanders (Batrachoseps attenuatus), ensatinas (Ensatina eschscholtzii), arboreal salamanders (Aneides
lugubris), Pacific treefrogs (Hyla regilla), western fence
lizards (Sceloporus occidentalis), California voles (Microtus californicus), deer mice (Peromyscus maniculatus),
young brush rabbits (Sylvilagus bachmani), harvest mice
(Rheithrodontomys spp.), nestling white-crowned sparrows (Zonotrichia leucophrys nuttalli), and nestling song
sparrows (Melospiza melodia) [Fitch 1941; Fox 1951,
1952b; James et al. 1983; Barry 1994]. Fox (1951) also
records at least one instance of cannibalism in the wild.
There is a heavy preference for slugs, rodents, and nestling birds in some areas inhabited by this snake (Fox
1951, 1952b; James et al. 1983). Coast garter snakes will
also eat a wide variety of fishes and amphibians if the
occasion arises (see Fox 1952b).
Current Status and Factors Influencing
Population Numbers
Distribution
Predators: Raccoon, herons, egrets, hawks, California
kingsnake, and bullfrog.
Prey: Pacific treefrog, California red-legged frog, bullfrog, Coast Range newt, oligochaetes, California mouse,
California vole, white-crowned sparrow, brush rabbit
(young only), shrews, slugs.
Coast garter snakes inhabit the North and South Coast
Ranges from just north of the Oregon border, south to
Point Conception, Santa Barbara County (Fox 1951,
Bellemin and Stewart 1977, Stebbins 1985). They intergrade with T. e. elegans at mid-elevations of the North
Coast Range (Stebbins 1985). The elevational range is
from near sea level to around 350 m (Fox 1951). Coast
garter snakes are widely distributed in the Bay Area
(Stebbins 1959) (Figure 4.8).
Coast garter snakes are negatively affected by habitat alteration, especially by agriculture and urbanization
which often results in disturbed or open habitats unsuitable for this species. Because these snakes do not require
permanent aquatic habitats for long term survival like
other garter snake taxa in the Bay Area, they are less affected overall by human activities. Coast garter snakes are
still relatively abundant in terrestrial habitats located in the
foothills surrounding the Bay Area (Jennings, unpubl. data).
Trophic Levels
Coast garter snakes are tertiary consumers.
Proximal Species
Good Habitat
Coast garter snakes inhabit meadows (such as grasslands)
and clearings with second growth in the fog belt and also
Chapter 4 —
Amphibians and Reptiles
213
Amphibians &
Reptiles
Food and Feeding
Amphibians &
Reptiles
References
Barry, S.J. 1994. The distribution, habitat, and evolution of the San Francisco garter snake, Thamnophis sirtalis tetrataenia. Unpubl. M.A. Thesis, Univ.
of Ca., Davis, Calif. iii+140 p.
Bellemin, J.M. and G.R. Stewart. 1977. Diagnostic characters and color convergence of the garter snakes
Thamnophis elegans terrestris and Thamnophis
couchii atratus along the central California coast.
Bull. of the Southern Calif. Academy of Sciences,
76(2):73-84.
Fitch, H.S. 1941. The feeding habits of California garter snakes. Calif. Fish and Game, 27(2):1-32.
______. 1983. Thamnophis elegans. Catalogue of American Amphibians and Reptiles:320.1-320.4.
Fox, W. 1948. The relationships of the garter snake
Thamnophis ordinoides. Copeia, 1948(2):113-120.
______. 1951. Relationships among the garter snakes
of the Thamnophis elegans rassenkreis. Univ. of Ca.
Publications in Zoology, 50(5):485-530.
______. 1952a. Seasonal variation in the male reproductive system of Pacific Coast garter snakes. J. of
Morphology, 90(3):481-554.
______. 1952b. Notes on feeding habits of Pacific Coast
garter snakes. Herpetologica, 8(1):4-8.
______. 1956. Seminal receptacles of snakes. The Anatomical Record, 124(3):519-540.
James, D.K., L. Petrinovich, T.L. Patterson and A.H.
James. 1983. Predation on white-crowned sparrow
nestlings by the western terrestrial garter snake in
San Francisco, California. Copeia, 1983(2):511-513.
Rathbun, G.B., M. R. Jennings, T.G. Murphey, and
N.R. Siepel. 1993. Status and ecology of sensitive
aquatic vertebrates in lower San Simeon and Pico
Creeks, San Luis Obispo County, California. Final report prepared for the Calif. Dept. of Parks
and Recreation, San Simeon Region, through Cooperative Agreement (14-16-0009-01-1909). U.S.
Fish and Wildl. Serv., Natl. Ecology Research Center, Piedras Blancas Research Station, San Simeon,
Calif. ix+103 p.
Stebbins, R.C. 1954. Amphibians and reptiles of western
North America. McGraw-Hill Book Company, Inc.,
New York, Toronto, and London. xxii+536 p.
______. 1959. Reptiles and amphibians of the San Francisco Bay region. California Natural History Guide (3).
Univ. of Ca. Press, Berkeley and Los Angeles. 72 p.
214
Baylands Ecosystem Species and Community Profiles
______. 1972. Amphibians and reptiles of California. California Natural History Guide (31). Univ. of Ca. Press,
Berkeley, Los Angeles, and London. 152 p.
______. 1985. A field guide to western amphibians and
reptiles. Second edition, revised. Houghton Mifflin
Company, Boston, Massachusetts. xiv+336 p.
Stewart, G.R. 1972. An unusual record of sperm storage in a female garter snake (genus Thamnophis).
Herpetology, 28(4):346-347.
San Francisco Garter Snake
Thamnophis sirtalis tetrataenia
Mark R. Jennings
General Information
The San Francisco garter snake (Thamnophis sirtalis
tetrataenia; Family Colubridae) is a medium sized (46122 cm TL), snake with seven upper labial scales and a
wide dorsal stripe of greenish yellow edged with black,
bordered on each side by a broad red stripe followed by
a black one (Barry 1978, Stebbins 1985). The belly is a
bright greenish blue (often turquoise) and the top of the
head is red (Stebbins 1985, Barry 1993). This snake was
one of the first reptiles to be listed as Endangered by the
U.S. Fish and Wildlife Service in 1967 (U.S. Fish and
Wildlife Service 1985).
Although the name of this snake has been stable
since Fox (1951) solved the mystery regarding the original collection of T. s. tetrataenia in 1855, Boundy and
Rossman (1995) recently proposed that the nomenclature of T. s. tetrataenia be revised because the holotype
of T. s. infernalis was found to actually be a specimen of
T. s. tetrataenia. This proposal of substituting T. s.
infernalis for T. s. tetrataenia and T. s. concinnus for T.
s. infernalis (sensu lato Fox 1951), has been followed by
Rossman et al. (1996). However, a petition has been
Ted Brown, Academy of Sciences
chaparral (Stebbins 1972). They are often abundant in
canyons with coast live oaks (Quercus agrifolia), California bay (Umbellularia californica) and numerous shrubs,
as well as riparian zones or other areas of dense vegetation (such as blackberries (Rubus discolor and R. ursinus),
thimbleberries (R. parviforus) and Baccharis (Baccharis
spp.)) next to more open areas (Fox 1951).
received and published by the International Commission
on Zoological Nomenclature to conserve the usage of T.
s. infernalis and T. s. tetrataenia and designate a neotype
for T. s. infernalis (Barry and Jennings 1998). Thus, the
existing usage of the Fox (1951) nomenclature should
be followed until a ruling is made on the case.
Reproduction
San Francisco garter snakes are live bearers which mate
during the spring (March-April) and also during the fall
(September-November), the latter often in breeding aggregations of several males and one female (Fox 1952a,
1954, 1955). Neonates (18-20 cm total length) are normally born in litters of 1-35 (average 16) during late July
to early August (Fox et al. 1961; Cover and Boyer 1988;
Barry 1993, 1994), although a few litters are born as late
as early September (Larsen 1994). Females have the ability to store sperm for up to 53 months (Stewart 1972).
eat California voles (Microtus californicus), even when
they are abundantly available (Barry 1993).
Distribution
San Francisco garter snakes are a Bay Area endemic that
are essentially restricted to San Mateo County, California (Stebbins 1959, Barry 1978) (Figure 4.9). Historically, they occurred in aquatic habitats and adjacent
uplands along the San Andreas Rift Zone from near
Pacifica, southeast to the Pulgas Water Temple, and
along an arc from the San Gregorio-Pescadero highlands,
west to the coast, and south to Point Año Nuevo (Barry
1978, 1994; McGinnis 1984). At least two recent records
just south of Point Año Nuevo—from the mouth of
Waddell Creek, Santa Cruz County—are questionable
(Barry 1993, 1994). Intergrades with T. s. infernalis have
been recorded in eastern San Mateo County (southeast
of the Pulgas Water Temple) and extreme western Santa
Clara County (Fox 1951, Barry 1994).
Growth and Development
Current Status and Factors Influencing
Population Numbers
San Francisco garter snakes have disappeared from significant portions of their native range due to habitat loss
from agriculture and urbanization—especially from housing developments and freeway construction (Medders
Food and Feeding
Subadult and adult San Francisco garter snakes feed
largely on the larvae and post-metamorphic life stages of
Pacific treefrogs (Hyla regilla) and California red-legged
frogs (Rana aurora draytonii). California toads (Bufo
boreas halophilus), introduced bullfrogs (R. catesbeiana),
introduced mosquitofish (Gambusia affinis), and threespine sticklebacks (Gasterosteus aculeatus) are also taken
(Fox 1951, Wharton 1989, McGinnis 1984, Barry
1994). Juvenile snakes feed largely on newts (Taricha
spp.), earthworms, and Pacific treefrogs (Barry 1993) and
will refuse other most non-amphibian items offered to
them (Fox 1952b, Larsen et al. 1991). Adult snakes rarely
Figure 4.9 San Francisco Garter Snake – Current
Known Location Restricted to San Mateo County
Chapter 4 —
Amphibians and Reptiles
215
Amphibians &
Reptiles
Snakes are most active from March to September although they can be observed during any month of the
year (Wharton 1989, Barry 1994, Larsen 1994). Juveniles grow rapidly during their first year, spending much
of their time feeding in riparian zones or aquatic habitats (Barry 1994). Males and females probably reach
sexual maturity in two years (at about 46 cm and 55 cm
total length respectively), although some slower growing snakes reach sexual maturity in three years (Barry
1994). During the summer months, subadult and adult
snakes may disperse away riparian areas into adjacent
habitats to feed on amphibians in rodent burrows (Barry
1993). During the winter months, juvenile and adult
snakes hibernate in small mammal burrows in adjacent
upland habitats (Larsen 1994). Some snakes can move
large distances (>2 km) over short periods of time
(Wharton 1989), but limited radio tracking data indicate that most movements are considerably shorter than
this distance (Larsen 1994).
Amphibians &
Reptiles
1976; USFWS 1985; Barry 1978, 1993). Historically,
the largest known population of snakes was at a series
of sag ponds (locally referred to as the “ Skyline Ponds” )
along Hwy 35 in the vicinity of Pacifica, Daly City, San
Bruno, and South San Francisco (Barry 1978, 1993,
1994; USFWS 1985). Today, this complex of ponds has
been completely covered by urbanization. The large Bay
Area population of snakes studied by Wharton (1989)
has extensively declined due to the loss of several prey
species from saltwater intrusion into the marsh (see
Larsen 1994) and this population may now be close to
extinction (Jennings, unpublished data). Besides the
above, declines also resulted from large numbers of
snakes being collected for the pet trade (especially overseas) and T. s. tetrataenia continues to be illegally collected for pets despite stiff penalties for doing so (e.g.,
see Bender 1981). Today, about 70% of the current remaining San Francisco garter snake habitat is composed
of artificially constructed aquatic sites such as farm
ponds, channelized sloughs, and reservoir impoundments (Barry 1993). Such habitats are often managed in
ways that are detrimental to the snake and its preferred
prey, the California red-legged frog (Barry 1993, 1994;
Larsen 1994).
Current estimates put the number of San Francisco
garter snakes at about 65 “ permanent” reproductive
populations of around 1500 total snakes >1 year of age
(Barry 1993). About half the known populations are
protected to some extent by refuges such as water preserves or state parks (Barry 1993). The key to preserving the species is to set aside adequate amounts of habitat and manage these areas for T. s. tetrataenia and its
prey, especially California red-legged frogs (Barry 1993,
Larsen 1994).
Trophic Levels
San Francisco garter snakes are tertiary consumers.
Proximal Species
Prey: Coast Range newt, California red-legged frog,
threespine stickleback, Pacific treefrog, bullfrog.
Predators: Hawks, herons, egrets, bullfrog, striped
skunk, opossum, and raccoon.
Good Habitat
San Francisco garter snakes are most abundant in natural sag ponds or artificial waterways that have been allowed to develop a dense cover of vegetative (Barry
1993). This is due to the presence of large amphibian
populations (=prey base) and many basking sites for juvenile and adult snakes which are relatively secure from
potential predators (Barry 1994). The presence of adjacent upland areas with abundant numbers of small mam-
216
Baylands Ecosystem Species and Community Profiles
mal burrows are also important as hibernation sites for
snakes during the winter (Larsen 1994).
References
Barry, S.J. 1978. Status of the San Francisco garter snake.
Ca. Dept. Fish and Game, Inland Fisheries Endangered Species Program, Special Publ. (78-2):121.
______. 1993. The San Francisco garter snake: protection is the key to recovery. Tideline, 13(4):1-3; 15.
______. 1994. The distribution, habitat, and evolution
of the San Francisco garter snake, Thamnophis
sirtalis tetrataenia. Unpubl. M.A. Thesis, Univ. of
Ca., Davis, Calif. iii+140 p.
Barry, S.J. and M.R. Jennings 1998. Coluber infernalis
Blainville, 1835 and Eutaenia sirtalis tetrataenia
Cope in Yarrow, 1875 (currently Thamnophis s.
tetrataenia and T. s. infernalis; Reptilia, Serpentes):
proposed conservation of the subspecific names by
the designation of a neotype for T. s. infernalis. The
Bull. of Zoological Nomenclature, 55(4):in press.
Bender, M. 1981. “ Sting” operation reveals massive illegal trade. Endangered Species Tech. Bull., 6(8):1; 4.
Boundy, J. and D.A. Rossman. 1995. Allocation and
status of the garter snake names Coluber infernalis
Blainville, Eutaenia sirtalis tetrataenia Cope and
Eutaenia imperialis Coues and Yarrow. Copeia,
1995(1):236-240.
Cover, J.F., Jr. and D.M. Boyer. 1988. Captive reproduction of the San Francisco garter snake Thamnophis sirtalis tetrataenia. Herpetological Review,
19(2):29-33.
Fox, W. 1951. The status of the gartersnake, Thamnophis sirtalis tetrataenia. Copeia, 1951(4):257-267.
______. 1952a. Seasonal variation in the male reproductive system of Pacific Coast garter snakes. J. of
Morphology, 90(3):481-554.
______. 1952b. Notes on feeding habits of Pacific Coast
garter snakes. Herpetologica, 8(1):4-8.
______. 1954. Genetic and environmental variation in
the timing of the reproductive cycles of male garter snakes. J. of Morphology, 95(3):415-450.
______. 1955. Mating aggregations of garter snakes.
Herpetologica, 11(3):176.
Fox, W., C. Gordon, and M.H. Fox. 1961. Morphological effects of low temperatures during the embryonic development of the garter snake, Thamnophis elegans. Zoologica, 46(2):57-71.
Larsen, S.S. 1994. Life history aspects of the San Francisco garter snake at the Millbrae habitat site.
Unpubl. M.S. Thesis, Calif. State Univ., Hayward,
Calif. ix+105 p.
Larsen, S.S., K.E. Swaim, and S.M. McGinnis. 1991.
Innate response of the San Francisco garter snake
and the Alameda whipsnake to specific prey items.
Transactions of the Western Section of the Wildl.
Society, 27:37-41.
McGinnis, S.M. 1984. The current distribution and
habitat requirements of the San Francisco garter
snake, Thamnophis sirtalis tetrataenia, in coastal San
Mateo County. Final report of work conducted
under Interagency Agreement C-673 and prepared
for the Ca. Dept. Fish and Game, Inland Fisheries
Div., Rancho Cordova, Calif. 38 p.
Medders, S. 1976. Serpent or supermarket? Natl. Parks
and Conservation Magazine, 50(4):18-19.
Rossman, D.A., N.B. Ford, and R.A. Seigel. 1996. The
garter snakes; evolution and ecology. Univ. of Oklahoma Press, Norman, Oklahoma. xx+332 p.
Stebbins, R.C. 1959. Reptiles and amphibians of the
San Francisco Bay region. California Natural His-
tory Guide (3). Univ. of Ca. Press, Berkeley and
Los Angeles. 72 p.
______. 1985. A field guide to western amphibians and
reptiles. Second edition, revised. Houghton Mifflin
Company, Boston, Massachusetts. xiv+336 p.
Stewart, G.R. 1972. An unusual record of sperm storage in a female garter snake (genus Thamnophis).
Herpetology, 28(4):346-347.
Wharton, J.C. 1989. Ecological and life history aspects
of the San Francisco garter snake (Thamnophis
sirtalis tetrataenia). Unpubl. MA Thesis, San Francisco State Univ., San Francisco, Calif. x+91 p.
U.S. Fish and Wildlife Service (USFWS). 1985. Recovery plan for the San Francisco garter snake (Thamnophis sirtalis tetrataenia). U.S. Fish and Wildl.
Serv., Portland, Oregon. 77 p.
Amphibians &
Reptiles
Chapter 4 —
Amphibians and Reptiles
217
Amphibians &
Reptiles
218
Baylands Ecosystem Species and Community Profiles
5
Mammals
Salt Marsh Harvest Mouse
Reithrodontomys raviventris
Howard S. Shellhammer
Life History
Historical and Modern Distribution
USFWS
SMHM are composed of two subspecies. The northern
subspecies, R. r. haliocoetes, is found on the upper portions of the Marin Peninsula; in the Petaluma, Napa and
Suisun marshes; as well as a disjunct series of populations on the northern Contra Costa County coast. The
southern subspecies, R. r. raviventris, is found in the
Chapter 5 — Mammals
219
Mammals
Salt marsh harvest mice (SMHM) are small, native rodents which are endemic to the salt marshes and adjacent diked wetlands of San Francisco Bay and are listed
as an endangered species by the U.S. Fish and Wildlife
Service and the State of California (Shellhammer 1982).
They range in total length from 118 to 175 millimeters
and in weight from 8 to 14 grams. They are vegetarians
that can drink water ranging from moderately saline to
sea water. They swim calmly and well. They do not
burrow, but will build ball-like nests of dry grasses and
other vegetation on the ground or up in the pickleweed
(Fisler 1965). Their behavior is placid, so much so that
their behavior is used as a secondary criterion in identifying them to the species level.
more highly developed portions of the Bay from the
Richmond area, down around the South San Francisco
Bay (primarily south of a line between Redwood City and
Hayward), and a disjunct series of small populations on
the Marin Peninsula. Some modern distributions are
indicated in Figure 5.1 and a listing of available trapping data are included in Appendix 5.1.
Their chromosome number and morphology have
been studied by Shellhammer, and the two subspecies
show some differences in chromosome shape indicating that genetic isolating mechanisms are beginning
to form between them. No recent and modern genetic
studies have been completed at the present time,
hence nothing is known about the genetic variability
of this species and whether or not it faces problems
of inbreeding and random genetic drift as its average
population size decreases.
The major threats to their habitat include filling, diking, subsidence, and changes in water salinity (Shellhammer 1982, 1989). Various estimates have
been made that at least 75% of all the tidal marshes
around the Bay have been filled in or otherwise destroyed over the last 150 years. Most of the remaining marshes have been back-filled or diked-off, and
hence most of the remaining tidal marshes are narrow strips along the bay side of the levees. Those strip
marshes and most of the remaining larger marshes
have lost their upper and part of their middle zones,
such that there is little escape cover from high tides
available. In the southern end of the South San Francisco Bay, the combination of subsidence caused by
water drawdown and the freshening of that part of the
Bay by massive amounts of non-saline, treated sewage effluent has changed the saline vegetation of that
area to brackish and freshwater species such as bulrushes (Scirpus sp.), cattails (Typha sp.), and peppergrass (Lepidium latifolium), species not used by
SMHM (Duke et al. 1990; Shellhammer 1982, 1989).
Because of these influences, SMHM has disappeared from many marshes and is present in very low
numbers in most others. The highest consistent populations are found in relatively large marshes along the
eastern edge of San Pablo Bay and in old dredge spoil
Figure 5.1 Salt Marsh
Harvest Mouse – Some
Current Locations and
Suitable Habitat
Mammals
Note: Mice are likely
present in areas identified
as “suitable” habitat based
on current information
regarding habitat types.
Mice may also be present
in other areas.
disposal ponds on former Mare Island Shipyard property;
most of these marshes are in or will be included in the
San Pablo Bay unit of the San Francisco Bay National
Wildlife Refuge (Bias and Morrison 1993, Duke et al.
1995). Other areas supporting large populations include
some parts of the Contra Costa County coastline (Duke
et al. 1990, 1991), some parts of the Petaluma Marshes,
and the Calaveras Point Marsh in the South San Francisco Bay (Duke et al. 1990), although the latter area is
deteriorating because of the declining salinity and correlated changes in vegetation.
Diked wetlands adjacent to the Bay have grown in
importance as the tidal marshes bayward of their outboard dikes have decreased in size and quality (Shellhammer 1989). Most of such diked marshes in the South San
Francisco Bay are being threatened by urban and industrial development along their borders. In addition, most
220
Baylands Ecosystem Species and Community Profiles
of these diked marshes are not managed to provide adequate vegetative cover of halophytic species or to maintain their salinity over time (Duke et al. 1990, Shellhammer 1989).
Suitable Habitat
SMHM are dependent on the thick, perennial cover of
salt marshes and move in the adjacent grasslands only
in the spring and summer when the grasslands provide
maximum cover (Fisler 1965). Their preferred habitats
are the middle and upper portions of those marshes, i.e.,
the pickleweed (Salicornia virginica) and peripheral
halophyte zones, and similar vegetation in diked wetlands adjacent to the Bay (Shellhammer et al. 1982,
1988). Some areas of known suitable habitat are
shown in Figure 5.1.
References
There are many questions that need to be addressed in
order to properly manage the SMHM. They include the
following: (1) Little is known about the degree of genetic
heterozygosity and polymorphism of this species. Is it
variable, and hence is the SMHM resistant to increasing isolation, genetic drift, and potential increased inbreeding, or is it a species that has survived a series of
genetic bottlenecks and become monomorphic and lacks
resilience? Without information on its population genetics, the only prudent course of action is to argue for the
largest possible population sizes of SMHM. Much more
needs to be known about the population genetics of this
species if it is to be properly managed over the long run.
(2) It is not known how much upland edge constitutes
enough of a buffer to protect SMHM from alien predators (especially cats) and human disturbance. The U.S.
Fish and Wildlife Service Endangered Species biologists
recommend 100 feet, but 100 feet of grassland, for example, may not be enough of a barrier to keep out dogs,
cats, red foxes, or humans. (3) The impact of introduced red foxes is not known, but they have had a
great impact on the California clapper rail, which is
found in the same marshes with SMHM.
Control of red foxes is being carried out in those
marshes in which there are rails and mice, but not in all
marshes potentially containing SMHM alone. Actually,
very little is known about the effects of predators on the
SMHM, including the effects of the rail. (4) Little is
known of the interactions between various species of
rodents in diked marshes. Geissel et al. (1988) demonstrated seasonal displacement of SMHM from optimal
habitat by California voles. Elaine Harding of U.C. Santa
Cruz is studying (as of 1997-98) rodent interactions and
has concerns that certain management practices in diked
wetlands might work against SMHM. (5) Little is known
about the impact of peppergrass on SMHM numbers.
SMHM remain in mixed pickleweed-peppergrass
communities (Duke et al. 1990, 1991), but no studies have been carried out in areas of 100% peppergrass, a condition that is becoming increasingly common in the southern end of the South San Francisco
Bay. (6) Lastly, there is the strong possibility that
youthful pickleweed marshes are more productive of
SMHM than older ones. That is certainly the case
reported by Bias (1994) and Bias and Morrison (1993)
at Mare Island Naval Shipyards and in the marshes
bordering the adjacent San Pablo Bay, a marsh that
has been growing actively by accretion for decades.
The effect of the relative youth of marshes (or possible the lack of their senescence), needs to be looked
at along with the potential effects of toxics, the depth
of buffer zones when marshes are bordered by either
urban and industrial development, and other concerns
spelled out previously in this document.
Bias, M.A. 1994. Ecology of the salt marsh harvest mouse
in San Pablo Bay. Ph.D. dissert., Univ. of Ca.,
Berkeley, Calif. 243 pp.
Bias, M. A. and M. L. Morrison. 1993. Final report: salt
marsh harvest mouse on Mare Island Naval Shipyard, 1989-1992. Unpubl. rpt. to Natural Resources
Mgmt. Branch, Western Div., Naval Facilities Engineering Command, San Bruno, CA., 223 pp.
Duke, R.R., H.S. Shellhammer, R.A. Hopkins and E.
Steinberg. 1990. San Jose Permit Assistance Program salt marsh harvest mouse trapping surveys,
Spring and Summer, 1990. Prepared for CH2M
Hill, Emeryville, CA by H.T. Harvey and Assoc..
Alviso, CA. Project No. 477-11.
Duke, R. R., S. B. Terrill, R. A. Hopkins, E. Steinberg and
E. K. Harding. 1991. Concord Weapons Station small
mammal characterization. Prepared for PRC Environmental Mgmt. Co. by H.T. Harvey and Assoc.,
Alviso, CA. Project 505-03.
Fisler, G.F. 1965. Adaptations and speciation in harvest
mice of the marshes of San Francisco Bay. Univ.
Ca. Publ. Zool. 77: 1-108.
Geissel, W.H., H.S. Shellhammer and H.T. Harvey.
1988. The ecology of the salt marsh harvest mouse
(Reithrodontomys raviventris) in a diked salt marsh.
J. Mammalogy. 69: 696-703.
Shellhammer, H.S. 1982. Reithrodontomys raviventris.
Mammalian Species, No. 169: 1-3. The American
Society of Mammalogists.
______. 1989. Salt marsh harvest mice, urban development, and rising sea levels. Conservation Biology
3: 59-65.
Shellhammer, H.S., R. Jackson, W. Davilla, A.M. Gilroy,
H.T. Harvey and L. Simons. 1982. Habitat preferences of salt marsh harvest mice (Reithrodontomys raviventris). Wasmann J. Bio. 40: 102-114.
Shellhammer, H.S., R.R. Duke, H.T. Harvey, V. Jennings,
V. Johnson and M. Newcomer. 1988. Salt marsh
harvest mice in the diked salt marshes of Southern
San Francisco Bay. Wasmann J. Bio. 46: 89-103.
Additional Readings
Duke, R.R., H.S. Shellhammer, R.A. Hopkins, E.
Steinberg, G. Rankin. 1995. Mare Island Naval
Shipyard salt marsh harvest mouse 1994 trapping
surveys. Prepared for Dames and Moore. Tucson.
AZ by H.T. Harvey and Assoc., Alviso, CA. Project
921-01.
U.S. Fish and Wildlife Service (USFWS). 1984. Salt
marsh harvest mouse and California clapper rail
recovery plan. Portland, Oregon. 141 pp.
Chapter 5 —
Mammals
221
Mammals
Conservation and Management
Appendix 5.1 Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled by Elaine
Harding from a USFWS database.
Mammals
Location
Alameda County
Albrae Slough
Albrae Slough
Albrae Slough
Audubon Marsh
Audubon Marsh
Baumberg Tract
Cabot Boulevard
Calaveras Point
Calaveras Point
Calaveras Point
Calaveras Point
Coyote Creek
Coyote Creek
Coyote Creek
Coyote Hills
Coyote Hills
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Coyote Hills Slough
Drawbridge
Drawbridge
Dumbarton
Dumbarton
Dumbarton
Dumbarton
Dumbarton
Durham Road Marsh
EBRPD SMHM Preserve
EBRPD SMHM Preserve
EBRPD SMHM Preserve
Emeryville Crescent
Emeryville Crescent
Fremont Redevelopment
Hayward
Hayward Marsh
Hayward Marsh
Ideal Marsh
Ideal Marsh
Irvington STP
Johnson Landing
Johnson Landing
Leslie-Lincoln
Leslie Quarry Site
222
Subarea
Trap
Nites
100
2600
200
2700
300
Coyote Cr
north of
Newby Isl
east
area A
area D
area CH
area A
area PA
area CH
area 4
railroad
Caltrans
1046
100
100
400
1000
300
892
500
710
100
100
400
200
200
200
200
200
200
400
400
200
100
0
25
100
400
200
500
200
540
600
725
540
1500
900
300
900
1075
100
200
200
900
1950
200
300
Baylands Ecosystem Species and Community Profiles
Mice
2
15
4
4
7
9
6
0
0
22
104
0
2
0
17
4
0
0
2
0
0
6
0
1
7
6
3
0
4
3
0
13
0
6
0
4
10
7
6
0
13
1
21
40
1
0
1
21
15
0
2
Mice Per
Trap Nite
0.02
0.006
0.02
0.001
0.023
0.006
0
0
0.055
0.104
0
0.002
0
0.024
0.04
0
0
0.01
0
0
0.03
0
0.005
0.018
0.015
0.015
0
0.12
0
0.033
0
0.012
0
0.007
0.017
0.01
0.011
0
0.014
0.003
0.023
0.037
0.01
0
0.005
0.023
0.008
0
0.007
Year
1974, 1975
1983
1984
1984
1985
1985
1989
1974, 1975
1974, 1975
1990
1990
1985
1990
1990
1975
1980
1974, 1975
1983
1983
1983
1964
1984
1984
1984
1984
1985
1985
1974, 1975
1978
1971
1974, 1975
1978
1980
1990, 1991
1980
1983
1984
1985
1982
1986
1985
1985
1982
1990
1974, 1975
1980
1985
1982
1983
1985
1985
Author
Cummings, E.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Cummings, E.
Cummings, E.
Duke, R.
Duke, R.
Duke, R.
Duke, R.
Zetterquist
Gilroy, A.
Cummings, E.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Cummings, E.
Schuat, D.B.
Cummings, E.
Leitner
Gilroy, A.
Gilroy, A.
Kobetich
Kobetich
Kobetich
Olsen, D.
Kobetich
Jennings, V.R.
Shellhammer, H.
Cummings, E.
Gilroy, A.
Kobetich
Kobetich
Kobetich
Jennings, V.R.
Shellhammer, H.
Appendix 5.1 (continued) Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled
by Elaine Harding from a USFWS database.
Location
north of
north of
northeast
northwest
north
e. of Dra
w. of Dra
central
east
headquart
SFC
Whale’s T
Caltrans
area B
E. Palo A
King and Ly
King and Ly
Trap
Nites
1200
7410
3120
200
75
100
200
400
300
400
871
200
300
1350
600
200
100
365
950
300
5850
600
800
1350
1350
500
100
200
400
300
900
2400
200
817
817
4350
1240
300
200
200
100
1020
600
600
1000
160
900
450
900
1350
200
Mice
41
23
36
2
2
0
1
0
2
5
9
4
0
34
9
0
0
1
36
4
20
0
6
0
0
0
0
0
3
2
5
27
2
2
2
126
28
0
1
1
0
17
2
2
5
0
1
7
13
0
2
Mice Per
Trap Nite
0.034
0.003
0.012
0.01
0.027
0
0.005
0
0.007
0.013
0.01
0.02
0
0.025
0.015
0
0
0.003
0.038
0.013
0.003
0
0.008
0
0
0
0
0
0.008
0.007
0.006
0.011
0.01
0.002
0.002
0.029
0.023
0
0.005
0.005
0
0.017
0.003
0.003
0.005
0
0.001
0.016
0.014
0
0.01
Year
1985
1988
1988-89
19851971
1974, 1975
1980
1980
1985
1985
1985
1985
1986
1985
1978
1980
1980
1980
1982
1983
1983
1984
1985
1985
1985
1990
1974, 1975
1980
1980
1985
1985
1986
1985
1983
1983
1987
1990
1985
1985
1985
1974, 1975
1986
1983
1984
1985
1985
1985
1989
1985
1988
1985
Chapter 5 —
Author
Shellhammer, H.
Johnson, V.
Shellhammer, H.
Schaub, D.B.
Cummings, E.
Gilroy, A.
Gilroy, A.
Shellhammer, H.
Jennings, V.R.
Anderson, J.
Shellhammer, H.
Gilroy, A.
Gilroy, A.
Gilroy, A.
Newcomer, M.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Kobetich
Shellhammer, H.
Xucera, T.E.
Cummings, E.
Gilroy, A.
Gilroy, A.
Kobetich
Shellhammer, H.
Shellhammer, H.
Kobetich
Shellhammer, H.
Shellhammer, H.
Jennings, V.R.
Cummings, E.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Jennings, V.R.
Shellhammer, H.
Foerster, K.
Shellhammer, H.
Klinger, R.C.
Shellhammer, H.
Mammals
223
Mammals
Alameda County (continued)
Mayhew’s Landing
Mayhew’s Landing
Mayhew’s Landing
Meadow Gun Club
Mowry Slough
Mowry Slough
Mowry Slough
Mowry Slough
Mowry Slough
Mt. Eden Creek
Mt. Eden Creek
Mud Slough
Mud Slough
Munster Site
Newark Slough
Newark Slough
Newark Slough
Newark Slough
Newark Slough
Newark Slough
Newark Slough
Newark Slough
Newark Slough
Oakland Airport
Oakland Airport
Oakland Airport
Old Alameda Creek
Old Alameda Creek
Old Alameda Creek
Old Alameda Creek
Old Fremont Airport
Old Fremont Airport
?? Gun Club
Roberts Landing
Roberts Landing
Roberts Landing
Roberts Landing
Sulphur Creek
Sulphur Creek
Thornton Ave.
Turk Island
Union City 511 Areas
Union City Marsh
Union City Marsh
Union City Marsh
University Ave
Warm Springs Mouse Pas
Warm Springs Mouse Pas
Warm Springs Seasonal
Warm Springs Seasonal
Whistling Wings Duck Club
Subarea
Appendix 5.1 (continued) Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled
by Elaine Harding from a USFWS database.
Mammals
Location
Contra Costa County
Antioch Point
Castro Creek Marsh
Concord Naval Weapons
Concord Naval Weapons
Concord Naval Weapons
Concord Naval Weapons
Concord Naval Weapons
Hastings Slough
Hoffman Marsh
Martinez East
Payten Shough
Pittsburg
Pittsburg East
Pittsburg West
Point Edith
Point Edith
Point Edith
Richmond Dump
San Pablo Creek
San Pablo Creek
San Pablo Creek
Shell marsh
Shell marsh
Stockton Ship Channel
Stockton Ship Channel
Stockton Ship Channel
Stockton Ship Channel
Marin County
Bahia
Bahia
Bahia
Black John Slough
China Camp State Park
Corte Madera
Corte Madera
Corte Madera
Corte Madera
Corte Madera
Corte Madera
Corte Madera
Gallinas Creek
Gallinas Creek
Gallinas Creek
Hamilton Air Force Base
John F. McInnis Park
Larkspur ferry Marsh
Muzzi Marsh
Novato Creek
Petaluma Creek
Petaluma Sewage Treatm
Pickleweed Park: San Rafael
224
Subarea
Trap
Nites
672
150
123
447
1800
2890
1200
80
200
900
2800
100
100
CWWS #22
CWWS #20
CWWS #21
CWWS #24
south
south, no
north
Mahoney S
north ban
south ban
south
800
800
200
125
100
2480
2270
800
400
200
200
400
930
3000
300
200
100
100
100
200
672
1412
750
100
100
672
300
1050
480
430
100
200
100
1094
Baylands Ecosystem Species and Community Profiles
Mice
21
51
12
4
19
22
200
37
0
0
22
64
0
0
25
5
5
1
13
0
81
6
1
0
0
6
0
31
68
3
16
2
3
6
0
2
19
0
0
1
1
34
1
4
0
0
1
0
0
37
Mice Per
Trap Nite
0.076
0.08
0.033
0.043
0.012
0.069
0.031
0
0
0.024
0.023
0
0
0.006
0.006
0.005
0.104
0
0.033
0.003
0.001
0
0
0.03
0
0.033
0.023
0.01
0.01
0.03
0.06
0
0.01
0.028
0
0
0.01
0.01
0.051
0.003
0.004
0
0
0.01
0
0
0.034
Year
1985
1981
1971
1979
1979
1985
1991
1988
1976
1980
1988
1978
1980
1980
1987
1988
1988
1980
1971
1974, 1975
1986
1988
1990
1980
1980
1980
1980
1984
1987
1989
1987
1980
1971
1974, 1975
1976
1980
1981
1983
1990
1974, 1975
1974, 1975
1981
1982
1986
1988
1986
1974, 1975
1980
1974, 1975
1990
Author
Mishaga, R.
Schaub, D.B.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Simons, L.
Shellhammer, H.
Simons, L.
Simons, L.
Botti, F.
Shellhammer, H.
Shellhammer, H.
Simons, L.
Schaub, D.B.
Cummings, E.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Duke, R.
Bott, F.
Simons, L.
Schaub, D.B.
Cummings, E.
Simons, L.
Mishaga, R.
Shellhammer, H.
Freas, K.E.
Cummings, E.
Cummings, E.
Mishaga, R.
Newcomer, M.
Shellhammer, H.
Cummings, E.
Simons, L.
Cummings, E.
Bias, M.A.
Appendix 5.1 (continued) Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled
by Elaine Harding from a USFWS database.
Location
south end
northeast
w of brid
east
Corkscrew
southwest
east
Corkscrew
southwest
marina si
south
north
south of
west
east
south of
ITT marsh
Trap
Nites
Mice
Mice Per
Trap Nite
Year
Author
1200
1200
11
0
0.009
0
1990
1991
100
0
0
1974, 1975
Cummings, E.
200
100
100
100
2
14
0
1
0.01
0.14
0
0.01
1980
1980
1980
1980
Simons, L.
Simons, L.
Simons, L.
Simons, L.
500
100
100
100
300
300
220
200
100
100
150
116
900
900
100
100
150
150
250
900
800
978
100
500
200
100
100
100
1344
3
1
0
1
7
3
19
0
0
1
3
0
0
0
2
0
0
3
5
0
1
42
0
8
1
0
0
0
37
0.006
0.01
0
0.01
0.023
0.01
0.086
0
0
0.01
0.02
0
0
0
0.02
0
0
0.02
0.02
0
0.001
0.043
0
0.016
0.005
0
0
0
0.028
1971
1974, 1975
1974, 1975
1974, 1975
1985
1985
1988
1980
1974, 1975
1974, 1975
1989
1978
1985
1985
1971
1974, 1975
1980
1980
1976
1984
1985
1989-90
1974, 1975
1990, 1991
1980
1974, 1975
1974, 1975
1974, 1975
1990
Schaub, D.B.
Cummings, E.
Cummings, E.
Cummings, E.
100
200
100
100
100
200
100
900
200
300
4200
0
0
0
0
4
0
0
11
6
0
54
0
0
0
0
0.04
0
0
0.012
0.03
0
0.013
1975
1975
1974, 1975
1974, 1975
1974, 1975
1986
1975
1985
1990
1985
1988
Cummings, E
Cummings, E
Cummings, E
Anderson, J.
Malenson, M.A.
Shellhammer, H.
Duke, R.
Chapter 5 —
Mammals
Botti, F.
Gilroy, A.
Cummings, E.
Cummings, E.
McGinnis, S.M.
Johnston, D.S.
Shellhammer, H.
Duke, R.
Schaub, D.B.
Cummings, E.
Gilroy, A.
Gilroy, A.
Shellhammer, H.
Shellhammer, H.
Cummings, E.
Gilroy, A.
Cummings, E.
Cummings, E.
Cummings, E.
Flannery, A.W.
Malenson, M.A.
Johnson, V.
225
Mammals
Marin County (continued)
Spinnaker Lagoon
Spinnaker Lagoon
Marin/Sonoma County
Petaluma River Mouth
Napa County
Coon Island
Fagan Marsh
Deman Slough
Napa Slough
San Mateo County
Bair Island
Bair Island
Bair Island
Bair Island
Bair Island
Bair Island
Bair Island
Bay Slough
Belmont Slough
Bird Island
East Third Street
Foster City
Foster City
Foster City Marina Sit
Greco Island
Greco Island
Greco Island
Greco Island
Ideal Cement Marsh
Ideal Cement Marsh
Ideal Cement Marsh
Ideal Cement Marsh
Laumeister Marsh
Laumeister Marsh
Palo Alto Yacht Harbor
Phelps Slough
Ravenswood Slough
Redwood Shores
San Rafael Canal
Santa Clara County
Alviso
Alviso Dump
Alviso Marina
Alviso Slough
Alviso Slough
Artesian Slough
Calabazas Creek
Coyote Creek
Coyote Creek
Crittenden Marsh
Emily Renzel Marsh
Subarea
Appendix 5.1 (continued) Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled
by Elaine Harding from a USFWS database.
Mammals
Location
Santa Clara County (continued)
New Chicago Marsh
New Chicago Marsh
New Chicago Marsh
New Chicago Marsh
New Chicago Marsh
New Chicago Marsh
New Chicago Marsh
New Chicago Marsh
Owens Corning Landfill
Palo Alto Baylands
Palo Alto Baylands
Palo Alto Baylands
Palo Alto Baylands
Palo Alto Baylands
Palo Alto Flood Basin
Palo Alto Flood Basin
Palo Alto Flood Basin
Ravensweed Area
Sunnyvale
Sunnyvale Baylands Park
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Triangle Marsh
Solano County
ACME Landfill Site
Benicia
Benicia State Park
Chabot Creek Outfall M
Collinsville
Collinsville
Collinsville
Collinsville
Collinsville, Rail Cor
Cordelia Dike
Cullinan Ranch
Denverton Highway
Ehaann Duck Club
Figueras Tract
Figueras Tract
Gentry/Pierce Property
Gold Hills Road Overcr
Grizzly Bay 1
226
Subarea
Sammis si
Grey Goos
Marine Te
S. Dutchm
Trap
Nites
Mice
Mice Per
Trap Nite
Year
100
1152
300
400
392
2820
1400
705
800
40
2058
100
300
1500
100
220
100
800
1200
540
20
4376
100
200
922
300
182
384
300
600
500
1500
2
14
0
0
11
65
4
8
6
1
196
0
1
32
0
1
0
1
3
0
0
71
23
0
12
2
0
2
2
5
10
35
0.02
0.012
0
0
0.028
0.023
0.003
0.011
0.008
0.025
0.095
0
0.003
0.021
0
0.005
0
0.001
0.003
0
0
0.016
0.23
0
0.013
0.007
0
0.005
0.007
0.008
0.02
0.023
1974, 1975
1975
1978
1980
1985
1986
1987
1988
1990
1971
1972
1974, 1975
1985
1990
1974, 1975
1975
1975
1990
1990
1987
1971
1974
1974, 1975
1976
1977-1978
1983
1984
1985
1986
1986
1990
1990
1200
160
200
483
1296
1296
1536
2350
640
150
2385
150
800
100
100
1800
500
300
9
2
0
4
32
32
8
2
0
0
5
0
3
2
2
10
3
5
0.008
0.013
0
0.008
0.025
0.025
0.005
0.001
0
0
0.002
0
0.004
0.02
0.02
0.006
0.006
0.017
1989
1979
1980
1989
1978
1978
1979
1980
1979
1980
1983
1980
1988
1974, 1975
1974
1986
1990
1980
Baylands Ecosystem Species and Community Profiles
Author
Cummings, E
Zetterquist,
Gilroy, A., a
Shellhammer, H.
Shellhammer, H.
Duke, R.
Shellhammer, H.
Duke, R.
Schaub, D.B.
Wondolleck, E
Cummings, E
Cummings, E
Zetterquist,
Malenson, M.A.
Duke, R.
Duke, R.
Schaub, D.B.
Rice, V.C.
Cummings, E
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Anderson, J.
Anderson, J.
Duke, R.
Duke, R.
Foster, J.
Michaels, J.L.
Simons, L., a
Ford, K.
Envirodyne En
Envirodyne En
Shellhammer, H
Shellhammer, H
Shellhammer, H
Shellhammer, H
Cummings, E.
Lindeman, E.
Duke, R.
Shellhammer, H
Appendix 5.1 (continued) Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled
by Elaine Harding from a USFWS database.
Location
Windmill
Dump
wildlife
powerline
footbridge
west bank
3 areas
Brown’s Is
Ryer Is
Bryan Par
Trap
Nites
300
74
150
150
200
300
98
150
50
300
300
300
100
150
420
300
100
1384
2114
1764
14672
9383
20502
300
300
1296
1800
1200
300
150
400
880
980
30
800
90
90
800
50
600
1200
1109
375
700
400
400
195
300
600
300
Mice
2
19
0
0
11
1
20
0
9
1
0
2
12
20
0
3
2
296
140
240
1005
336
1427
2
2
32
17
21
0
3
0
7
0
0
1
6
8
3
21
7
4
1
30
2
18
0
0
6
4
0
0
Mice Per
Trap Nite
0.007
0.257
0
0
0.055
0.003
0.204
0
0.18
0.003
0
0.007
0.12
0.133
0
0.01
0.02
0.214
0.066
0.136
0.068
0.036
0.07
0.007
0.007
0.025
0.009
0.018
0
0
0.018
0
0
0.033
0.008
0.089
0.033
0.026
0.14
0.007
0.001
0.027
0.005
0.026
0
0
0.031
0.013
0
0
Year
1980
1971
1980
1980
1981
1985
1971
1980
1971
1980
1980
1985
1974, 1975
1976
1990
1986
1974, 1975
1985
1986
1987
1989
1990
1991
1980
1980
1978
1978
1991
1980
1972
1980
1986
1987
1987
1985
1988
1985
1985
1982
1971
1980
1985
1987
1986
1990
1980
1980
1980
1985
1990
1980
Chapter 5 —
Author
Shellhammer, H.
Schuab, D.B.
Shellhammer, H.
Shellhammer, H.
Schuab, D.B.
Shellhammer, H.
Schuab, D.B.
Shellhammer, H.
Shellhammer, H.
Cummings, E.
Cummings, E.
Kovach, S.D.
Kovach, S.D.
Kovach, S.D.
Bias et al.
Bias et al.
Bias et al.
Shellhammer, H.
Shellhammer, H.
Duke, R.
Shellhammer, H.
Rollins, G.
Shellhammer, H.
Kovach, S.D.
Shellhammer, H.
Kovach, S.D.
Kovach, S.D.
Newcomer, M.
Schuab, D.B.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Mammals
227
Mammals
Solano County (continued)
Grizzly Bay 2
Grizzly Island
Hill Slough
Hill Slough
Hill Slough
Hill Slough
Island #1
Jackspine Wetland
Joice Island
Joice Island
Joice Island
Joice Island
Leslie Intake
Leslie Intake
Opes Road Marsh
Uco Slough
Mare Island Naval Ship
Mare Island Naval Ship
Mare Island Naval Ship
Mare Island Naval Ship
Mare Island Naval Ship
Mare Island Naval Ship
Mare Island Naval Ship
Meins Landing
Meins Landing Mound
Montezuma Site
Montezuma Site
Montezuma Site
Morrow Island
Napa River
Nurse Slough
Nurse Slough
Park Place
Park Place Shopping Ce
Rayer Island
Roe Island
Roe Island (east)
Roe Island (west)
Sears Point 1
Simmons Island
Simmons Island
Simmons Island
Southern Solano Annexa
Southhampton Bay (outb
Southhampton Marsh
Stockton Ship Channel
Stockton Ship Channel
Suisun Marsh Club No. 2
Suisun Slough
Sulphur Springs Creek
Teal Boathouse
Subarea
Appendix 5.1 (continued) Important Data Sets for Salt Marsh Harvest Mouse (1971 - 1991). Compiled
by Elaine Harding from a USFWS database.
Location
Building
Stockgate
Decoy
Bayside
1 m. upst
east
mouth
Trap
Nites
228
Mice
Mice Per
Trap Nite
Year
Author
300
150
300
300
150
300
800
2
0
3
0
0
0
7
0.007
0
0.01
0
0
0
0.009
1980
1980
1980
1980
1980
1985
1988
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
Shellhammer, H.
100
100
256
100
672
100
100
100
100
100
200
100
750
205
9
4
5
0
80
7
0
1
2
2
12
4
33
8
0.09
0.04
0.02
0
0.119
0.07
0
0.01
0.02
0.02
0.06
0.04
0.044
0.039
1971
1974, 1975
1979
1980
1988
1971
1974, 1975
1980
1974, 1975
1980
1982
1980
1982
1980
Schaub, D.B.
Cummings, E.
Moss, J.G.
Simons, L.
Stroud, M.C.
Schaub, D.B.
Cummings, E.
Simons, L.
Cummings, E.
Simons, L.
Newcomer, M.
Simons, L.
Newcomer, M.
Simons, L.
Mammals
Solano County (continued)
Teal Slough
Vennink
Vennink
Vennink
Vennink
West Grizzly Island
Wildwings Duck Club (M
Sonoma County
Lower Tubbs Island
Lower Tubbs Island
Lower Tubbs Island
Lower Tubbs Island
Mare Island Naval Ship
Petaluma Creek
Petaluma River
Petaluma River Mouth
Sonoma Creek
Sonoma Creek
Sonoma Creek
Tolay Creek Mouth
Tubbs Island 1
Tubbs island Accessory
Subarea
Baylands Ecosystem Species and Community Profiles
Shellhammer, H.
California Vole
Microtus californicus
William Z. Lidicker, Jr.
Life History
Historical and Modern Distributions
The taxonomic status of San Francisco Bay voles is complex. Marsh inhabiting voles from Point Isabel (Contra
Costa County) south on the east side of the Bay and
around to the west side as far north as Redwood City
have been described as the subspecies paludicola. Thaeler
(1961) examined these populations in detail and concluded that at least the East Bay populations could not
be distinguished from the upland subspecies californicus. Voles from the Marin County side of the Bay are
placed in M.c. eximius, and those from Grizzly Island
(Solano County) and eastward into the Delta represent
the large, dark subspecies aestuarinus. Of special interest and concern, Thaeler (1961) described the vole population inhabiting the marshes around the mouth of San
Pablo Creek (Contra Costa County) as M.c.
sanpabloensis. This subspecies is viewed as a species of
special concern by the State of California (Williams
1986). It is darker and yellower than the adjacent populations of M.c. californicus. Further, its palatines are
deeply excavated along their posterior borders, the rostrum is narrow, and the auditory bullae relatively inflated.
J.K. Clark; Courtesy UC IPM Project
Suitable Habitat
Habitat use extends from adjacent upland grasslands into
both salt and freshwater marshes, at least into those
where flooding does not occur regularly. Voles are good
swimmers, however, and can survive occasional inundation. Some known current locations and potential suitable habitats are shown in Figure 5.2.
Conservation and Management
Efforts to conserve wetlands should be aware of the endemic form M.c. sanpabloensis and attempt to achieve
Chapter 5 —
Mammals
229
Mammals
California voles are common inhabitants of the San Francisco Bay wetlands. They are vegetarians, feeding extensively on Salicornia and other marsh vegetation. They
make runways through the vegetation, burrow extensively in non-flooded areas, and often utilize driftwood
for cover. They are critically important prey species for
a wide variety of mammalian and avian predators.
The population dynamics of voles has been studied intensively in adjacent upland grasslands (Cockburn
and Lidicker 1983; Krebs 1966; Lidicker 1973; Pearson
1966, 1971; Salvioni and Lidicker 1995), but little is
known about marsh populations. It is not known, for
example, if most marsh populations are merely extensions of upland ones or independent demographic units.
An exception is the San Pablo Creek vole; see below.
Grassland populations around the Bay exhibit annual or
multi-annual cycles in numbers, but the demographic
behavior of salt marsh populations is unknown. Similarly, we know that grassland voles breed mainly in the
wet season, and especially intensely from February
through May. Voles in marshes may well be different,
perhaps breeding mostly in the summer and very little
during the flood-prone winters.
California voles are keystone species in grassland
communities by virtue of their importance as a major
prey species (Pearson 1985) and their potentially great
effect on vegetation (Lidicker 1989). Thus, if similar
roles are played in San Francisco Bay wetlands, these
rodents may be vital to the health of the wetland communities. Because they are known to exhibit strong fluctuations in numbers (four orders of magnitude), suitable
habitat patches must be large enough for the species to
survive low-density bottlenecks. These voles are also
known to exhibit strong non-trophic interactions with
other species of mammals. The introduced house mouse
(Mus musculus) is strongly affected negatively by the
presence of voles (DeLong 1966, Lidicker 1966). Interactions with Western harvest mice (Reithrodontomys
megalotis) are more complex (Heske et al. 1984). At
moderate Microtus densities harvest mice are positively
influenced, presumably because the harvest mice make
effective use of vole runways. However, at high vole
densities, the Reithrodontomys are strongly negatively
impacted. It is possible that salt marsh harvest mice
(Reithrodontomys raviventris) may interact in a similarly complex way with voles. Geissel et al. (1988)
demonstrated seasonal displacement of salt marsh
harvest mice by voles. More subtle indirect effects may
also be important. For example, if voles sustain populations of red fox (Vulpes vulpes), an indirect negative
effect on clapper rails (Rallus longirostris obsoletus) may
be manifest.
Figure 5.2 California Vole –
Some Current Locations
and Suitable Habitat
Mammals
Note: Voles are likely
present in areas identified
as “ suitable” habitat based
on current information
regarding habitat types.
Voles may also be present
in other areas.
representation of the other three currently recognized
subspecies in the Bay Area as well. Because of their role
as a major prey species, California voles are likely keystone species in the health of Bay Area wetland communities.
References
Cockburn, A. and W.Z. Lidicker, Jr. 1983. Microhabitat heterogeneity and population ecology of an herbivorous rodent, Microtus californicus. Oecologia
59: 167-177.
DeLong, K.T. 1966. Population ecology of feral house
mice: interference by Microtus. Ecology 47: 481484.
230
Baylands Ecosystem Species and Community Profiles
Geissel, W.H., H. Shellhammer and H.T. Harvey. 1988.
The ecology of the salt marsh harvest mouse (Reithrodontomys raviventris) in a diked salt marsh. J.
Mammology 69: 696-703.
Heske, E.J., R.S. Ostfeld and W.Z. Lidicker, Jr. 1984.
Competitive interactions between Microtus californicus and Reithrodontomys megalotis during two
peaks of Microtus abundance. J. Mammology 65:
271-280.
Krebs, C.J. 1966. Demographic changes in fluctuating
populations of Microtus californicus. Ecol. Monogr.
36: 239-273.
Lidicker, W.Z., Jr. 1966. Ecological observations on a
feral house mouse population declining to extinction. Ecol. Monogr. 36: 27-50.
Salt Marsh Wandering Shrew
Sorex vagrans haliocoetes
Howard S. Shellhammer
Life History
The salt marsh wandering shrew (SMWS) appears to
have some of the most restrictive food and habitat requirements of any mammal inhabiting the marshes of
the greater San Francisco Bay Region—far more, for example, than the endangered salt marsh harvest mouse
(Reithrodontomys raviventris). While little is known of the
SMWS subspecies, shrews in general are insectivores
which are born in the spring and become sexually mature the following winter. SMWSs have gestation periods of about 21 days (Owen and Hoffman 1983). Many
shrew species have only one litter, and adults die after
the young are weaned (Jameson and Peeters 1988).
Historical and Modern Distribution
Chapter 5 —
Mammals
231
Mammals
The historical range of the SMWS extended from the
northern end of the San Francisco Peninsula, down
through the marshes of the South San Francisco Bay,
and up through the marshes of western Contra Costa
County to about the Benicia Straits.
Johnston and Rudd (1957) suggested that between
1951 and 1955 shrews represented about 10% of the
small mammals of the marshes. They were far less numerous in the 1970s and 80s, at least in the southern
part of its range (Shellhammer, pers. obs.). Known or
suspected populations as of 1986 included marshes south
of Foster City and Hayward and in the San Pablo
marshes of the San Pablo Bay (WESCO 1986). This
subspecies of vagrant shrew is currently confined to the
salt marshes of the South San Francisco Bay (Figure
5.3). It exists in a narrow band of tidal salt marsh and
does not seem to be present in diked marshes.
Dr. Richard B. Forbes
______. 1973. Regulation of numbers in an island population of the California vole, a problem in community dynamics. Ecol. Monogr. 43: 271-302.
______. 1989. Impacts of non-domesticated vertebrates
on California grasslands. In: L.F. Huenneke and
H. Mooney (eds). Grassland structure and function: California annual grassland. Kluwer Acad.
Pub., Dordrecht, The Netherlands; pp. 135-150
Pearson, O.P. 1966. The prey of carnivores during one
cycle of mouse abundance. Jour. Anim. Ecol. 35:
217-233.
______. 1971. Additional measurements of the impact
of carnivores on California voles (Microtus californicus). J. Mammology. 52: 41-49.
______. 1985. Predation. Pp. 536-566. In: R. H. Tamarin (ed). Biology of new world Microtus. Amer. Soc.
Mammalogists Spec. Pub. No. 8; 893 pp.
Salvioni, M. and W.Z. Lidicker, Jr. 1995. Social organization and space use in California voles: seasonal,
sexual, and age-specific strategies. Oecologia 101:
426-438.
Thaeler, C.S., Jr. 1961. Variation in some salt-marsh
populations of Microtus californicus. Univ. Calif.
Pub. Zool. 60: 67-94.
Williams, D. F. 1986. Mammalian species of special concern in California. Wildlife Mgt. Div. Admin. Report 86-1, Calif. Dept. of Fish and Game, Sacramento, CA. 112 pp.
Figure 5.3 Salt Marsh
Wandering Shrew –
Some Current Locations
and Suitable Habitat
Mammals
Note: Shrews are likely
present in areas identified
as “ suitable” habitat based
on current information
regarding habitat types.
Shrews may also be present
in other areas.
Suitable Habitat
The SMWS’s habitat is wet, medium high salt marshes.
It is best described by D. Williams (1983) in a draft report for the California Department of Fish and Game
using material primarily from Johnston and Rudd
(1957): “ [Salt marsh wandering shrews] frequent areas
in the tidal marshes providing dense cover, abundant
food (invertebrates), suitable nesting sites, and fairly
continuous ground moisture. Their center of activity is
in the ‘medium high marsh,’ about 6 to 8 feet above sea
level, and in lower marsh areas not regularly inundated.
Suitable sites are characterized by abundant driftwood
and other debris scattered among pickleweed (Salicornia). The pickleweed is usually one to two feet in height.
The detritus preserves moisture and offers refuge in dry
232
Baylands Ecosystem Species and Community Profiles
period to amphipods, isopods and other invertebrates,
and resting sites for shrews. Nesting material consists of
plant material, primarily Salicornia duff. The higher
marsh, 8 to 9 feet in elevation, is too dry and offers only
minimal cover—few to no shrews occupy this zone. The
lower cordgrass (Spartina) zone is subjected to daily tidal
floods and has cover too sparse for shrews.”
Some potential suitable habitat locations are shown
in Figure 5.3.
Conservation and Management
Johnston and Rudd’s 1957 paper represents the last scientific work on the subspecies, per se. The rest of the
reports (Williams 1983; WESCO 1986; this present effort) are all based on that study and that of Rudd 1955.
Many changes have taken place since the early 1950s and
little to nothing is known as to how such changes have
affected the prey or habitat requirements of this shrew.
The southern part of the San Francisco Bay has been
greatly freshened by hundreds of millions of gallons of
treated sewage outflows per day, and this freshening has
brought about changes in plant species composition. Until point source reductions were placed on industrial sewage in the 1980s, large amounts of heavy metals, as well
as polychlorinated biphenyls and petroleum hydrocarbons were poured into the Bay. In addition, the storm
runoff and inflows of creeks and small rivers carried
unknown amounts of pesticides, petroleum compounds,
and other toxic substances. It is not known how decreased salinity and increased toxicity in the South Bay
may have impacted the shrews, either directly, or indirectly, through changes in the amount and diversity of
their prey. In addition to salinity, vegetation changes,
and toxics, many of the marshes of the South Bay have
subsided, and the Salicornia bands have become more
degraded and more heavily inundated. Again little is
known as to the effects on this shrew of such changes.
The SMWS is currently listed as “ Mammalian Species of Special Concern” by the California Deptartment
of Fish and Game and as a candidate species for listing
in Category 2 by the U.S. Fish and Wildlife Service. Neither classification offers legal protection to its habitat.
Little recent biological information is available to support its classification as a protected species, a status it
merits.
References
Sorex ornatus sinuosis
Kevin MacKay
Life History
The Suisun shrew is a small (95-105 mm in total length),
dark, insectivorous mammal with a long, pointed nose,
and a well-developed scaly tail (37-41 mm). Suisun
shrews are carnivores and predators feeding primarily
upon amphipods, isopods, and other invertebrate species (WESCO 1986, Hays 1990). The shrews may also
occasionally serve as prey for several large predators such
as the short-eared owl (Asio flammeus), northern harrier
(Circus cyaneus), and black-shouldered kite (Elanus
caeruleus) (WESCO 1986).
The reproductive period of the Suisun shrew extends from late February through September, with the
majority of breeding occurring from early spring through
May. A second breeding period occurs in late summer
when the young born the previous spring are mature and
able to mate for the first time.
Shrews typically construct domed, cup-like nests
composed of small paper scraps and dead material from
plants such as pacific cordgrass (Spartina foliosa), pickleweed (Salicornia virginica), and salt grass (Distichlis
spicata). The nests are usually placed directly on the soil
surface under driftwood, planks, or wood blocks, and are
situated above the high tide line to escape flooding
(WESCO 1986). Runways enter from the sides and from
beneath, and are not opened until two to three weeks
after the birth of the young (Johnston and Rudd 1957).
After the young have dispersed, the nests may be used
by other small mammals such as the endangered salt
marsh harvest mouse (Reithrodontomys raviventris)
(WESCO 1986).
There are no published data on the gestation period of the Suisun shrew, but the salt marsh wandering
shrew and other small shrews have a gestation period of
about 21 days (Owen and Hoffmann 1983, WESCO
1986). Litter size ranges from four to six individuals,
with a survival rate of 55 to 60 percent from near birth
to just after weaning (Johnston and Rudd 1957). Causes
of mortality include drowning from high tides, death of the
mother, starvation, exposure, and predation (WESCO
1986). The young remain in the nest for up to five weeks
and then move into adjacent areas (Rudd 1955).
Suisun shrews seldom reach their maximum life expectancy of 16 months, and populations turn over on
an annual basis. Populations in the early spring typically
consist of adults born the previous year. These individuals gradually die off during the summer months, and by
fall have been almost completely replaced by young born
the previous spring (Owen and Hoffmann 1983).
Chapter 5 —
Mammals
233
Mammals
Jameson, E.W., Jr. and H.J. Peeters. 1988. California
Mammals. Univ. of Ca. Press, Berkeley, CA.
Johnston, R.F. and R.L. Rudd. 1957. Breeding of the
salt marsh shrew. J. Mammalogy 38: 157-163.
Owen, J.G. and R.S. Hoffmann. 1983. Sorex ornatus.
Mammalian Species No. 212: 1-5. The American
Society of Mammalogists.
Rudd, R. L. 1955. Age, sex, and weight comparisons in
three species of shrews. J. Mammalogy 36: 323-339.
Western Ecological Services Company (WESCO). 1986.
A review of the population status of the salt marsh
wandering shrew, Sorex vagrans haliocoetes, Final
Report.
Williams, D.F. 1983. Mammalian species of special concern in California. Ca. Dept. Fish and Game,
Nongame Wildl. Investigation, E-W-4, IV-14.1,
Draft Final Report. 184 pp.
Suisun Shrew
Activity patterns vary according to season and reproductive condition in the Suisun shrew, but the subspecies is predominately nocturnal, especially during the
breeding season. Sexually mature shrews are very active
in the spring, concurrent with the breeding season, but
are less active during the early summer. Young-of-theyear born in early spring become sexually mature by late
summer, and their activity patterns peak during this second breeding season. Others, born later in the season are
still sexually immature by late summer and remain comparatively inactive during this period (Owen and Hoffmann 1983).
Hays (1990) found that during the non-breeding
season, shrews lived in loose social groups of 10 to 15
individuals. These groups contained only one adult male,
and one such group occupied 0.07 ha. In the spring
other adult males invade these groups, disrupting the
stable structure by competing among themselves.
Territorial behavior in shrews has not been well
documented in the field. However, Rust (1978) noted
territorial patrolling in observations of breeding captive
Suisun shrews.
Mammals
Historical and Modern Distribution
One of the nine subspecies of ornate shrew that occur
in California, the Suisun shrew is a relatively rare inhabitant of the salt marsh ecosystem of San Pablo and Suisun Bays (WESCO 1986). Johnston and Rudd (1957)
estimated that the shrews represent approximately 10
percent of the mammalian fauna present in marsh habitats, and were less abundant than mice (Mus sp.), rats
(Rattus sp.), voles (Microtus sp.), and harvest mice (Reithrodontomys sp.).
The historical extent of the Suisun shrew distribution is unknown (WESCO 1986) (Figure 5.4). According to Rudd (1955) the subspecies historically inhabited
the tidal saline and brackish salt marsh communities of
northern San Pablo and Suisun bays, ranging from the
mouth of the Petaluma River, Sonoma County on the
west, eastward through Southampton and Grizzly Island
to approximately Collingsville, Solano County (WESCO
1986, Rudd 1955, Williams 1983). The western extent
of the range was redefined by Brown and Rudd (1981)
as they identified the shrews inhabiting the marshes west
of Sonoma Creek and Tubbs Island as S. o. californicus
(WESCO 1986, Williams 1983).
However, surveys completed by Grinnell (1913)
discovered Suisun shrews only at Grizzly Island. Researchers (WESCO 1986) have speculated that, at that
time, the shrew was restricted to the greater Grizzly Island area because of the lack of suitable habitat throughout the rest of the historic range. The 1914 soil survey
of the San Francisco Bay Area identifies most of the Napa
Marsh as low tidal mud flats, a habitat that would be
consistently inundated by tidal waters and thus uninhab-
234
Baylands Ecosystem Species and Community Profiles
itable by Suisun shrews or other small mammals. Once
these areas were diked, and suitable habitat created, the
shrew may have expanded its historic range into these
adjacent areas (WESCO 1986).
There are no data available which directly measures
the current densities of Suisun shrew populations. The
number of individuals within a population appears to
vary with season and habitat type. Newman (1970) estimated that the most favorable habitat supported shrew
densities of as many as 111 individuals per hectare. A
related species, the dusky shrew (Sorex obscurus), has
overlapping home ranges averaging 0.037 ha in size, with
a density of 37 to 42 individuals per hectare. These latter figures are probably a more accurate depiction of
Suisun shrew populations as the amount of favorable
habitat is limited throughout most of its range (WESCO
1986).
The Suisun shrew is currently limited in its distribution to the scattered, isolated remnants of natural tidal
salt and brackish marshes surrounding the northern borders of Suisun and San Pablo bays (WESCO 1986).
Rudd (1955) identified four distinct populations
of Suisun shrews: the Grizzly Island population, found
throughout the marshlands east of Suisun Slough; a peripheral population found west of Suisun Slough and on
Morrow Island; the Southampton population, restricted
to the Benicia State Recreation Area; and the Sears Point
Road population located in the Napa marshes.
No Suisun shrews were captured in either of the
two most recent population studies (Williams 1983,
WESCO 1986) that attempted to assess the current distribution of the shrew. This lack of trapping success can
possibly be attributed to the extremely high rainfall in
1982 and 1986. Most of the low-lying marshes were
flooded for extended periods of time, adversely affecting
the small mammal populations. Additional trapping efforts for salt marsh harvest mice occasionally yielded
Sorex captures; however, only one capture, at Cullinan
Ranch on South Slough, was identified as S. o. sinuosis
(WESCO 1986).
WESCO (1986) plotted all known S. o. sinuosis
captures to delineate extant populations. Only two individual areas were identified that support populations
of Suisun shrews: Grizzly Island and Solano Island Number 1. Nine additional marsh areas were also identified
as having a high probability of supporting Suisun shrew
populations: Skaggs Island, Appleby Bay/Coon Island,
Steamboat Slough, Vallejo, Morrow Island, Cordelia
Slough South, Hammond Island, Simmons/Wheeler Islands, and Collingsville (WESCO 1986).
Suitable Habitat
Suisun shrews typically inhabit saline and brackish tidal
marshes characterized by pacific cordgrass (Spartina
foliosa), pickleweed (Salicornia virginica), gumplant
Figure 5.4 Suisun Shrew –
Some Current Locations
and Suitable Habitat
Note: Shrews are likely
present in areas identified
as “ suitable” habitat based
on current information
regarding habitat types.
Shrews may also be present
in other areas.
Mammals
(Grindelia humulis), California bulrush (Scirpus californicus), and common cattail (Typha latifolia). However,
shrew occurrence appears to be more strongly associated
with vegetation structure rather than species composition. Suisun shrews prefer dense, low-lying vegetation
which provides protective cover and suitable nesting
sites, as well as abundant invertebrate prey species (Owen
and Hoffmann 1983). Driftwood, planks, and other
debris found above the high-tide line also affords the
shrew with valuable foraging and nesting sites. In addition, adjacent upland habitats provide essential refuge
areas for Suisun shrews and other terrestrial animals
during periods of prolonged flooding (Williams 1986).
Some areas of potentially suitable habitat for the Suisun
Shrew are shown in Figure 5.4.
Conservation and Management
Williams (1986) identified the lack of an adequate
elevational gradient of marsh vegetation and adjacent
upland habitats as the principal obstacles to the recovery of Suisun shew populations in San Pablo and
Suisun bays. However, as the Suisun shrew does not
seem to make use of upland grasslands (Hays 1990),
and because of evidence of interbreeding with S. o.
californicus, future marsh management practices
should include the provision of elevated sites that
flood only occasionally, but not include upland grassland, which would encourage contact with californicus.
Chapter 5 —
Mammals
235
Mammals
References
Brown, R.J. and R.L. Rudd. 1981. Chromosomal comparisons within the Sorex ornatus-S. vagrens complex. Wassman J. Bio. 39:30-35.
Grinnell, J. 1913. The species of the mammalian genus
Sorex of west-central California. Univ. Ca. Publ.
Zool. 10:179-195.
Hays, W.S. 1990. Population ecology of ornate shre