ISSN 0038-3872

SOuMie RN CALIFORNIA? ACADEMY OF SCIENCES

BULLETIN

Volume 92 Number 3

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BCAS-A92(3) 101-172 (1993) DECEMBER 1993

Southern California Academy of Sciences
Founded 6 November 1891, incorporated 17 May 1907

© Southern California Academy of Sciences, 1993

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Date of this issue 22 December 1993

© This paper meeis the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

Bull. Southern California Acad. Sci.
92(3), 1993, pp. 101-167
© Southern California Academy of Sciences, 1993

The Status and Distribution of the Freshwater Fishes of
Southern California

Camm C. Swift,! Thomas R. Haglund,?
Mario Ruiz,! and Robert N. Fisher?

! Section of Fishes, Natural History Museum, 900 Exposition Boulevard,
Los Angeles, California 90007
?Department of Biology, University of California,
Los Angeles, California 90024-1606
3 Section of Zoology and Center for Population Biology,
University of California, Davis, California 95616

Abstract.—The fresh and low salinity waters of southern California include the
Owens, Mohave, Colorado, and coastal drainages south of Monterey Bay to the
Mexican border. The youthful topography presents a strong dichotomy between
steep rocky streams abruptly meeting relatively flat deserts or coastal plains. Little
or no intermediate, foothill habitat exists. Thirty-eight native freshwater and 23
estuarine fishes have been recorded from this area. In addition, at least 100 species
have been introduced, with widely varying success. Since the late 1940s and 1950s
the native fishes of the Owens, Colorado, and Mohave drainages have been in
jeopardy or extirpated in California. At the same time, the lowland fishes in coastal
drainages, particularly on the Los Angeles Basin, also disappeared. Upland species
of the coastal drainages still remain in a few isolated areas but are so reduced that
special protection is needed. Only one estuarine species, Eucyclogobius newberryi,
is threatened. Some tropical estuarine species of extreme southern California were
last collected 50 to 80 years ago, and are very rare or extirpated here. If the
remaining elements of the fish fauna are to survive, immediate action is needed
to preserve the remaining habitat and to restore areas within the native range.

Coastal and inland southern California has a distinctive, endemic native fresh-
water fish fauna. This fauna is also one of the smallest in the United States,
apparently due to the high topographic relief, long geological isolation from other
continental fish faunas, and the aridity of inland areas (Jordan 1895; Culver and
Hubbs 1917; Miller 1958; Follett 1961; Soltz and Naiman 1978; Smith 1981;
Miller and Smith 1986; Minckley et al. 1986; Swift 1989). Today all of the native
freshwater and some of the euryhaline species are extirpated or severely reduced
in numbers within their native range, and most have been recommended for
special conservation status in California (Miller et al. 1989; Williams et al. 1989;
Moyle and Williams 1990; Moyle et al. 1989; Lufkin 1991; Nehlsen et al. 1991;
Minckley and Deacon 1991). Thirty-eight native freshwater taxa (species, sub-
species, and geographic groups recognized for taxonomic and/or conservation
purposes) are known from this area (Moyle and Williams 1990; this paper). Twen-
ty-three more brackish or estuarine species depend on lower salinity water for at

101

102 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

least part of their life cycle. Some of these are also extinct or much reduced in
range. One brackish species, the tidewater goby, Eucyclogobius newberryi (included
as a freshwater species by Moyle and Williams 1990), is a “‘species of special
concern’ in California, and has been petitioned for Federal Threatened status
(U.S. Fish and Wildlife Service 1991). Most of the native species in the Colorado
River drainages (of California) declined severely or were extirpated many years
ago. Since the last survey of the coastal fauna (Wells et al. 1975), it became apparent
that the species of the coastal drainages were also in jeopardy. Here we document
the status of this fauna and recommend measures to preserve them for the future.
The status and distribution of all the native and introduced fishes known to us
through August 1992 is reviewed.

With at least 100 non-native species now recorded from southern California,
the area has the dubious honor of exceeding all other areas of the state in numbers
of aliens established. California has been a leading recipient of non-native fishes
in the nation (Moyle 1976b; Shapavalov et al. 1981; Courtenay et al. 1991; Baltz
1991; Williams and Jennings 1991). Many of these introductions first arrived in
southern California; others came secondarily after being introduced farther north
(Smith 1896; Shebley 1917a; Evermann and Clark 1931; Curtis 1949; Moyle
1976b; Courtenay et al. 1991; this paper). The success and effects of some of these
introductions is largely due to the highly modified nature of local aquatic habitats.
Moyle (1976b), Baltz (1991), and Minckley and Deacon (1991) reviewed the many
reasons for the arrival of these fishes, including accidental ones.

The natural environment for freshwater fishes in southern California can be
defined in a series of strong contrasts. The relatively well-watered coastal drainages
have a mild, Mediterranean climate that has fluctuated little for the last few million
years due to the ameliorating effect of the ocean (Johnson 1977). In the same
time, the inland deserts have ranged from well-watered lake country with glaciers
in the mountains to the extremely arid climate of today with only a few permanent
streams and springs flowing into saline lakes, many lacking fish (Smith 1981;
Miller 1981; Minckley et al. 1986; Norris and Webb 1990; Jannik et al. 1991;
Enzel et al. 1992). The highest (Mount Whitney, 4430 m) and lowest (Death
Valley, 86 m below sea level) points in the co-terminus United States fall within
this area.

A few million years ago (late Miocene-early Pliocene) the area had a much
flatter, rolling topography, one or more large rivers drained westward or south-
westward to the Pacific, and large lakes and brackish coastal embayments existed
both on the Pacific coast and lower Colorado River (Norris and Webb 1990;
Woodburne et al. 1991). The compression of the local terrain by the Pacific plate
subducting under the North American plate created the high, youthful mountains
separating the coastal areas from the Colorado drainage. Many coastal streams
that formerly flowed to the south or southwest turned west or northwest. Today
most major streams in both coastal and inland areas lie in or are closely aligned
with major faults (Norris and Webb 1990). Faults, often the interface between
different rock types, force groundwater to the surface in the springs that comprise
the habitat of many desert fishes (Soltz and Naiman 1978). They function this
way in wetter areas also, and provide areas of permanent surface water even
during dry periods (Miller 1961). The strongly elevated, still rising, youthful terrain
also erodes rapidly. In most of the area, steep mountains are flanked by relatively

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 103

fiat coastal plains or desert alluvial fans. Foothills or intermediate areas are rare
and mostly restricted to narrow bands at the interface of the mountains and plains.

Although commonly included with the inland deserts, the coastal areas were
originally well-watered and the last few hundred years have seen this environment
greatly altered (Willet 1941; Miller 1961; Gordon 1985; Queenan 1986). Increased
human population has led to drastic modification of water courses, and a dramatic
lowering of the water table, to well below the ground surface in the lowlands.
Even in the mountains where permanent water remains, logging, mining, flood
control, and water storage projects have changed the character of the streams.
The recent drought (1986—present) further reduced the available habitat. Thus,
the coastal habitat is converging artificially toward the natural hydrological con-
ditions of the inland deserts. The increasing threat to the remaining coastal fresh-
water fishes and the increasing appearance of introduced species has prompted
this reassessment of the local fauna.

Methods

For this paper southern California drainages include Pacific coastal streams
south of the Carmel River to the Mexican border, and Colorado River drainage
of California which includes the Owens, Death Valley, Mohave, and Salton Sea
systems (Figs. 1-5). Records are from museum collections, files of the California
Department of Fish and Game and U.S. Forest Service, and a few from a variety
of other sources. Museum collections examined include: Natural History Museum
of Los Angeles County (including material formerly at California State University,
Fullerton; Fullerton College; and California State University, Long Beach), Cal-
ifornia Academy of Sciences, Museum of Zoology of the University of Michigan,
National Museum of Natural History, Field Museum in Chicago, University of
California, Los Angeles, and Scripps Institution of Oceanography. Institutional
abbreviations follow Leviton et al. (1985). Records without voucher specimens
are accepted when by known authorities and/or with adequate descriptions. Rec-
ords from the California Department of Fish and Game, Region 5, Long Beach
are identified with initials (CFG LB), Willis A. Evans (WAE), Phillip A. Douglas
(PAD), Robert R. Bell (RB), James A. St. Amant (JAS), Franklin G. Hoover
(FGH), David Drake (DD), Linda Pardy (LP), and Unarmored Threespine Stick-
leback Recovery Team (UTSRT). Notes made by Ralph G. Miller are noted as
RGM. Copies of all non-LACM sources for data are filed in the Section of Fishes
(LACM). Collections from 1965 onward were entered into the California Fish
Data Base and Preserve Design Project established by Peter Moyle, Jack Williams,
and Roland White, Department of Fisheries and Wildlife, U.C. Davis, and into
the California Department of Fish and Game’s Natural Diversity Data Base (Betsy
Bolster, pers. comm.).

The 38 native freshwater taxa listed are based on Moyle and Williams (1990)
except that the San Antonio Creek stickleback (Vandenberg Air Force Base, Santa
Barbara County) is considered a distinct taxon also. Subspecific names are used
only if more than one subspecies occur in southern California. The conservation
status of each taxon is based on Williams et al. (1989), Miller et al. (1989), Moyle
et al. (1989), Moyle and Williams (1990), and Nehlsen et al. (1991). Moyle and
Williams (1990) relied on preliminary data from this study for their determination
of status of coastal southern California taxa.

104 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Monterey Co

Fig. |. Drainage map of coastal California from Carmel River to Santa Maria River (abbreviations
in Methods; see Fig. 2 for stream names along southeastern edge). 1. San Jose Cr, 2. Malpaso Cr, 3.
Soberanes Cr, 4. Garrapata Cr, 5. Palo Colorado Cr, 6. Bixby Cr, 7. Little Sur R, 8. Big Cr, 9. Vincente
Cr, 10. Limekiln Cr, 11. Mill Cr, 12. Prewitt Cr, 13. Plaskett Cr, 14. Willow Cr, 15. Alder Cr, 16.
Villa Cr, 17. Salmon Cr, 18. San Carpoforo Cr, 19. Ar del Oso, 20. Ar de Corral, 21. Oak Knoll Cr

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 105

We include as freshwater species those that spend all or a distinct portion of
their life cycle in freshwater, thus including the primary and secondary freshwater
and diadromous categories of previous authors (McDowall 1989). These are usu-
ally called inland fishes in the western United States (Moyle 1976a; Wydowski
and Whitney 1979). Estuarine taxa are based on Moyle (1976a), Miller and Lea
(1976), Shapavalov et al. (1981), and our interpretation of the degree of depen-
dence on lowered salinity. Many other species known from bays and estuaries in
California (Horn and Allen 1976), but not as limited to low salinity, are omitted.
Also excluded (as native) are anadromous and euryhaline species recorded from
marine waters in southern California but not native in freshwater in this area
such as Acipenser transmontanus and A. medirostris, species of Pacific salmons
(Oncorhynchus spp., in part), and Thaleichthys pacificus (Miller and Lea 1976;
Eschmeyer et al. 1983). Some of these have been introduced into fresh waters in
southern California. Most marine species introduced into the Salton Sea are not
native there; some are native in marine coastal waters of southern California.
Only freshwater records are listed. Following Shafland and Lewis (1984) and
Allendorf (1991), introductions refer to any movement of fish from one place to
another by humans whether the ultimate origin is local, regional, national, or
international; examples of all these exist in southern California.

Native fishes were greatly reduced in the Colorado River in California/Arizona
by the time Minckley (1973) published his maps, and these are not repeated here.
Also not mapped are the endemics of the Mohave and Owens drainages that have
had historically very limited distribution (except Catostomus fumeiventris), and
are mapped in Lee et al. (1980) and Moyle et al. (1989). The distribution of
Eucyclogobius was mapped by Swift et al. (1989). Maps are redrawn with some
modifications from the U.S. Geological Survey map, State of California,
1:1,000,000, Lambert conformal conic projection, 1970 edition. Abbreviations
on maps and in captions are: Ar, Arroyo (de); Ca, Canada (de); Cn, Canyon; Co,
County; Cr, Creek; L, Lake; R, River; Re, Reservoir; Sl, slough; Spr, spring (s);
trib, tributary(s) (of); W, Wash.

The term Los Angeles Basin is used here to include the drainage areas of the
Los Angeles, San Gabriel, and Santa Ana rivers (the Santa Ana system of Culver
and Hubbs 1917). The spelling of Mohave follows Miller et al. (1991, p. 32) as
originally coined from the native American name rather than as subsequently
changed to a Spanish word (Mojave). Common and scientific names of genera
and species, including those required by the U.S. Food and Drug Administration
for a few commercially important hybrids, usually follow Robins et al. (1991);
some generic and family group names follow Page and Burr (1991; see Mayden
1992). Names for some subspecific groups based on taxonomic, biological, and
conservation criteria follow Moyle and Williams (1990).

—

(Ar Laguna), 22. Ar Tortuga, 23. Ar del Puerto 24. Little Pico Cr, 25. Pico Cr, 26. San Simeon Cr,
27. Leffingwell Cr, 28. Villa Cr, 29. Cayucos Cr, 30. Old Cr and Whale Rock Re, 31. Willow Cr, 32.
Toro Cr, 33. Morro Cr, 34. Chorro Cr, 35. Los Osos Cr, 36. Islay Cr, 37. Coon Cr, 38. San Luis
Obispo Cr, 39. Pismo (Price) Cr, 40. Arroyo Grande Cr, 41. Oso Flaco Cr, 42. Orcutt Cr, 43. Shuman
Cn, 44. San Antonio R and Re, 45. Nacimiento R and Re, 46. Santa Margarita L, 47. San Juan Cr.

SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

106

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FRESHWATER FISHES OF SOUTHERN CALIFORNIA 107

Freshwater Taxa
Petromyzontidae

1. Lampetra cf. pacifica Viadykov. Pacific brook lamprey. Extirpated in
southern California. Hubbs (1967) documented the records of this small, non-
parasitic species in the San Gabriel and Santa Ana drainages. Subsequent ex-
amination of the material of Ewy (1945) at Whittier College (now LACM 32593-
1) verified that she had both this species and the much larger migratory one which
follows. One lot (LACM 31615-1) taken from the Los Angeles River verifies its
existence in that drainage. Very little preserved material of the fish survives, and
it will be difficult to decide ifit was a taxon distinct from the Pacific brook lamprey
farther north. Robins et al. (1991), following Bond and Kan (1985), placed L.
pacifica in the synonymy of L. richardsoni, the former including southern Cali-
fornia populations. Carl L. Hubbs (in litt.) related that it was common in low
gradient portions of Los Angeles Basin streams but disappeared before its dis-
tinction from the anadromous Pacific lamprey was appreciated. The last Pacific
brook lampreys were taken in 1930 from the Los Angeles River near Griffith
Park, in 1943 or 1944 from the San Gabriel River near the mouth of San Jose
Creek, and in 1955 from the Santa Ana River just above the Prado Flood Control
Basin (SIO 61-478).

2. Lampetra tridentata (Gairdner). Pacific lamprey. This anadromous lam-
prey still maintains runs in several creeks in Monterey and San Luis Obispo
County, parts of the Santa Maria and Santa Ynez rivers, parts of the Ventura
River, the Sespe Creek portion of the Santa Clara River drainage, and the lower,
unimpounded reach of Malibu Creek (Fig. 6). Its habitat requirements are similar
to those of steelhead, Oncorhynchus mykiss, and it reaches an elevation of about
900 m in Sespe Creek (Sam Sweet, pers. comm., LACM 45700-1). Shapavalov
and Taft (1954) reported Pacific lamprey absent from all but the largest rivers on
the central California coast; runs in small streams probably were not detected by

—

Fig. 2. Drainage map of coastal California from Santa Maria River to Malibu Creek (abbreviations
in Methods; arrows indicate direction of flow in aqueducts). 1. Arroyo Grande Cr, 2. Oso Flaco Cr,
3. Orcutt Cr, 4. Shuman Cn, 5. Nipomo Cr, 6. Los Berros Cn, 7. Stony Cr, 8. Arroyo Seco, 9. Carrie
Cr, 10. Tepusquet Cr, 11. La Brea Cr, N Fk, 12. La Brea Cr, S Fk, 13. Foxen Cn, 14. Davy Brown
Cr, 15. Manazana Cr, 16. Santa Barbara Cn, 17. Quatal Cn, 18. Apache Cn, 19. Rancho Nuevo Cr,
20. Alder Cr, 21. Honda Cr, 22. Salsipuedes Cr, 23. El Jaro Cr, 24: Jalama Cr, 25. Ca Cojo, 26. Ca
St. Augustine, 27. Ca Bulito, 28. Ca Santa Anita, 29. Ca Alegria, 30. Ca Agua Caliente, 31. Gaviota
Cr, 32. Ca Hondo, 33. Ar Quemado, 34. Refugio Cn, 35. Dos Pueblos Cn, 36. Tecolote Cn, 37.
Winchester (Bell) Cn, 38. Devereaux SI, 39. Goleta S] and tribs. west to east, Tecolito, Carneros, San
Pedro, San Jose, Maria Ignacio, and San Antonio cr, 40. Ar Burro, 41. Mission Cr, 42. Carpinteria
Cr, 43. Rincon Cr, 44. Zaca Cr, 45. Alamo Pintado Cr, 46. Santa Agueda Cr, 47. Cachuma Cr and
L, 48. Santa Cruz Cr, 49. Indian Cr, 50. Mono Cr (trib to Gibraltar Re), 51. Agua Caliente Cn, 52.
Juncal Cr (trib to Jameson L), 53. Padre Juan Cn, 54. Matilija Cr and L, 55. San Antonio Cr, 56.
Santa Paula Cr, 57. Agua Blanca Cr, 58. Lockwood Cr, 59. Gorman Cr, 60. Castac L, 61. Quail L,
62. Munz L, 63. Elizabeth L, 64. Castaic Cr and Re, 65. San Francisquito Cn, 66. Bouquet Cn and
Re, 67. Agua Dulce Cn and trib Escondido Cn, 68. Kentucky Spr Cn, 69. Aliso Cn, 70. Revlon Sl
trib Mugu Lagoon, 71. Ar Conejo, 72. Big Sycamore Cn, 73. Ar Sequit, 74. Zuma Cn, 75. Topanga
Cn, 76. Ballona Marsh and Cr, 77. L Casitas, 78. L Piru, 79. Pyramid Re, 80. California Aq, 81. Los
Angeles Aq.

108 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

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v 118 119

tee

San Bernardino Co

28... {CON )
GOSAA Diego \ONK

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Fig. 3. Drainage map of coastal California from Malibu Creek (Los Angeles County) to the Tijuana
River (Mexican border). (Abbreviations in Methods; arrows indicate direction of flow in aqueducts.)
Malibu Cr, 2. Topanga Cr, 3. Ballona Marsh and Cr, formerly receiving the Los Angeles River (dotted
line), 4. Rio Hondo and Whittier Flood Control Basin, 5. Bull Cn, trib Sepulveda Flood Control Basin,
6. Pacoima Cr, 7. Soledad Cn (upper Santa Clara R), 8. Little Tujunga Cn, 9. Big Tujunga Cr and Re,
trib Hansen Dam Flood Control basin; 10. Arroyo Seco and upper trib Bear Cr, 11. Eaton Cn, 12.
Santa Anita Cn, 13. Monrovia Cn, 14. Fish Cn, 15. W Fk San Gabriel R, trib Cogswell Re, 16. Devils
Cn, 17. Bear Cr, 18. N Fk San Gabriel R and Crystal L, 19. E Fk San Gabriel R trib to San Gabriel
and Morris re, 20. Cattle Cn, 21. Big Dalton Cn, 22. San Dimas Cn, 23. Walnut Cr draining Pud-
dingstone Re, 24. San Jose Cr, 25. Little Rock Cr and Re, 26. Big Rock Cr, 27. San Antonio Cr, and

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 109

Shapavalov and Taft (1954), or some factor other than stream size accounted for
their absence.

It spawned in Los Angeles Basin streams until about 1955 (Hubbs 1967), and
was not recorded again until March 1991 when an adult was observed about one
kilometer above the mouth of the Santa Ana River after heavy rains (Dave Raetz,
pers. comm.). No freshwater records exist for the species south of the Santa Ana
River. The recognition of a southern subspecies was based on the low myomere
counts of the then-unrecognized freshwater non-parasitic form, and subspecies
were not recognized later (Hubbs 1967; Hubbs et al. 1979).

Salmonidae

3. Oncorhynchus mykiss (Walbaum). Rainbow trout. Hubbs (1946) reported
on possible native coarse-scaled trout from the headwaters of the San Luis Rey
drainage, San Diego County. Otherwise southern California native populations
of resident, non-migratory, rainbow trout are only known from coastal drainages
and have not been recognized as taxonomically distinctive (Miller 1950; Hubbs
1946; Needham and Gard 1963; Berg and Gall 1988; Gall et al. 1991; Behnke
1992). Because all other freshwater fishes from this area are endemic or otherwise
diagnostic, a distinctive native trout also would be expected in Los Angeles Basin
streams. Careful examination of some isolated headwater populations may dis-
close such a form, namely upper tributaries of the San Luis Rey and Santa Mar-
garita rivers; San Jacinto River, Santiago, Lytle, and Cajon creeks, all tributaries
of the Santa Ana River; tributaries of San Juan Creek in the Santa Ana Mountains;
upper tributaries of the San Gabriel River; Arroyo Seco (above Pasadena) and
upper Big Tujunga Creek, both tributary to the Los Angeles River; upper tribu-
taries of Piru and Sespe creeks, Santa Clara drainage; and possibly upper tributaries

—_—

upper trib Icehouse Cn; 28. Chino Cr trib Prado Flood Control Area; 29. Cucamonga Cr, 30. Day Cr,
31. Lytle Cr, 32. Cajon Cr, 33. Strawberry Cr, 34. City Cr, 35. Bear Cr outlet of Bear L, 36. S. Fk
Santa Ana R and Dollar L, 37. Mill Cr, 38. Yucaipa Cr, 39. San Timoteo Cn, 40. Ar Tepsequet, 41.
Temescal W, 42. Santiago Cr and Re (Irvine L), 43. Silverado Cr, 44. San Diego Cr trib Newport
Bay, 45. Aliso Cr, 46. Oso Creek, 47. Trabuco Cr, 48. Bell Cn, 49. San Mateo Cr, 50. San Onofre Cn,
51. Las Pulgas (Las Flores) Cn, 52. De Luz Cr, 53. Murietta Cr, 54. Warm Springs Cr, 55. Tucalota
Cr, L Skinner inlet and outlet, 56. Santa Gertrudis Cr, 57. Temucula Cr outlet of Vail L, 58. Wilson
Cr, 59. Tule Cr, 60. Chihuahua Cr, 61. Long Cn, 62. Arroyo Seco, 63. Pechanga Cr, 64. Pala Cr, 65.
Pauma Cr, 66. W Fk San Luis Rey R, trib L Henshaw, 67. Agua Caliente Cr, 68. Buena Vista Cr, 69.
Matagual Cr, 70. Carrizo Cr, 71. Keys Cr, 72. Moosa Cn, 73. Buena Vista Cr, 74. Agua Hedionda
Lagoon and Cr, 75. Batiquitos Lagoon, 76. Escondido Cr, outlet L Wohlford, 77. San Dieguito R, 78.
San Vincente Cr, 79. Wildcat Cr, 80. Temescal Cr, 81. Cedar Cr, 82. Boulder Cr, outlet of Cuyamaca
L, trib El Capitan Re, 83. Conejos Cr, 84. Chocolate Cn, 85. Harbison Cr, 86. Sweetwater R trib
Loveland Re, 87. Lawson Cr, 88. Sweetwater R and Re, 89. Otay R outlet of upper and lower Otay
Re, 90. Deluzra Cr, 91. Hollenbeck Cn, 92. Cottonwood Cr outlet Barrett L, 93. Pine Valley Cr, 94.
Morena Cr trib Morena Re, 95. Cottonwood Cr, 96. Long Cn, 97. La Posta Cr, 98. Potero Cr, 99.
Campo Cr, 100. San Felipe Cr, 101. First San Diego Aq, 102. Second San Diego Aq, 103. Deep Cr,
104. Holcomb Cr, 105. Silverwood L, 106. Lake Gregory, 107. Lake Arrowhead, 108. Baldwin L and
trib Shay Cr, 109. L Elsinore, downstream of Railroad Cn Re, 110. N Fk San Jacinto R, 111. Strawberry
Cr, 112. S Fk San Jacinto R outlet L Hemet, 113. L Matthews and Colorado R Aq, 114. L Perris and
California Aq, 115. Colorado R Aq, 116. L Hodges, 117. L Sutherland, 118. California Aq, 119. Los
Angeles Aq.

110 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

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alee
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Fig. 4. Drainage map of the Colorado River and Salton Sea areas of California (abbreviations in
Methods; arrows indicate direction of flow in aqueducts). 1. Mission Cr, 2. N and S Fk Whitewater
R, 3. San Gorgonio R, 4. Palm Cn, 5. Dry W, 6. Coyote Cr, 7. San Felipe Cr, 8. Fish Cr W, 9.
Deguynas Cn, 10. Vallecitos Cr, 11. Carizo Cn, 12. Coyote W, 13. Westside Main Canal, 14. East
Highline Canal, 15. Ramer L, 16. Senator W, 17. Palo Verde Lagoon, 18. L Cahuilla, 19. Town of
Mecca.

of the Ventura, Santa Inez, or Santa Maria drainages. Recent collections or reports
of self-sustaining populations are known from these areas (Fig. 1-3).

Cooper (1874) reported native trout in San Diego County only in Warner’s Pass
at the head of the San Luis Rey River. Eigenmann and Eigenmann (1890) reported
them in Pala Creek, tributary to the San Luis Rey farther downstream. Hubbs
(1946), referring to Smith (1880, unpublished), reported them abundant in fresh-
water streams near Smiths Mountain and Pala in the headwaters of the San Luis
Rey system. Much of the Warner Ranch area was flooded by Lake Henshaw.

-—3

Fig. 5. Drainage map of the Owens and Mohave drainages in California; Mono to San Bernardino
counties (abbreviations in Methods; arrows indicate direction of flow in aqueducts). 1. Adobe Cr, 2.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 111

FES

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EY 250 \

River Spr, 3. Hot Cr, 4. Crowley L, 5. Pleasant Valley Re, 6. Tinemaha Re, 7. Williamson L, 8.
Haiwee Re, 9. Little L, 10. Salt Cr, 11. Cottonball Marsh, 12. Saratoga Spr, 13. Tecopa Hot Spr, 14.
Shoshone Spr, 15. Salt Cr (fishless), 16. Zyzyx or Soda Spr, 17. Camp Cady L, 18. Deep Cr, trib
Mohave Flood Control Basin; 19. California Aq.

112 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 6. Distribution of migratory (parasitic), Lampetra tridentata, and freshwater (non-parasitic),
L. cf. pacifica, lampreys in coastal southern California. Black circles: Lampetra tridentata from 1970
onward; half circles: before 1970. Open circles: Lampetra cf. pacifica, all 1955 or before. See Figs. 1-
3 for stream names.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 113

Hubbs (1946) reported his examination of specimens of these fish in the San Diego
Society of Natural History, but thought it unlikely that any stocks uncontaminated
by introductions remained. Beginning in the 1890s and extending through the
late 1930s fingerling rainbow trout were planted into almost all possible waters
in southern California (CFG LB). Included were stocks identified at the time as
both rainbow trout and steelhead (Busack and Gall 1980). Field notes made by
State Fish and Game biologists, 1915 to 1945, document three other streams
perennial enough to be strong candidates for permanent trout waters in San Diego
County, south of the San Luis Rey drainage: Boulder Creek, tributary to San Diego
River; Cold Creek, tributary to Green Valley Creek, Sweetwater River drainage;
and Pine Valley Creek, tributary to the Otay River. Many southern California
streams have one or more barrier falls and the extent to which, if any, native
trout occurred above many of these is not known. Today such records can only
be found by searching historical documents or prehistorical river deposits for
remains of these fish.

Rainbow trout were first planted in the lower Colorado River of California in
1935 after the construction of Boulder Dam (Moffett 1942), and much earlier
farther upstream in Arizona and Nevada (Miller and Alcorn 1946; Minckley
1973).

4. Oncorhynchus mykiss (Walbaum). Southern steelhead. Today steelhead
have nearly the same distribution as the Pacific lamprey (Fig. 7) in southern
California. Virtually every coastal stream in Monterey, San Luis Obispo, and
Santa Barbara counties north of Point Conception has had at least a few fish
entering it within the last ten years. South of Point Conception the following
streams have recent records: Gaviota, Mission, and Atascadero creeks, Santa
Barbara County; Ventura and Santa Clara rivers, Ventura County; and Mullhol-
land, Big Sycamore, Malibu, and Topanga canyons, Los Angeles County. His-
torically, fish also entered streams farther south, including Los Angeles and San
Gabriel rivers, Los Angeles County; and San Onofre and San Mateo creeks, Santa
Margarita, San Luis Rey, San Diego and Tijuana rivers, San Diego County (Hubbs
1946). Spawning success south of the Los Angeles basin may have been sporadic.
Large series of juvenile fish were taken in lower San Juan Creek (Miller 1939),
lower San Mateo Creek near San Clemente (UMMZ 132964), and the lower Santa
Margarita River (UMMZ 132968) in the summer of 1939, indicating successful
reproduction at that time. Anglers referred to juvenile fish in coastal lagoons as
sundowners, and often large numbers could be caught in coastal lagoons in the
1930s and earlier (R. Croker, pers. comm.).

Cyprinidae

5. Gila bicolor snyderi Miller. Owens tuichub. State and federally endangered.
Once widespread in the Owens drainage, including Owens Lake (Gilbert 1893;
Miller 1973), it is known only from Fish Slough, Owens Valley Native Fishes
Sanctuary, Inyo County; a small spring and stream, Cabin Bar Ranch, west side
of Owens Lake; about 13 km of the Owens River below Crowley Lake; two springs
at the Hot Creek Fish Hatchery; Little Hot Creek Pond; and an artificial habitat
on Los Angeles Department of Water and Power Land (Bolster 1990b). Many
populations are introgressed with Lahontan tui chubs, Gila bicolor obesa; others
were eliminated by introduced trout (three species) and largemouth bass.

114 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 7. Distribution of steelhead, Oncorhynchus mykiss, in coastal southern California. Black circles
from 1970 onward; half circles before 1970. See Figs. 1-3 for stream names.

6. Gila bicolor mohavensis Snyder. Mohave tui chub, State and federally
endangered. Once widespread in the Mohave drainage, it remains only in artificial
habitats at Soda Springs near Baker; Desert Studies Center Pond at Hinkley;
California Desert Information Center in Barstow; Camp Cady Wildlife Area near
Newberry Springs, San Bernardino County; and China Lake Naval Weapons

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 115

Center, San Bernardino and Kern counties (Hoover and St. Amant 1983; Bolster
1990a). It hybridizes with the introduced G. orcutti (Hubbs and Miller 1943) in
headwater portions of the Mohave drainage, particularly lower Deep Creek and
Moyle (1976a) and Castleberry and Cech (1986) believed arroyo chubs eliminated
the Mohave chub. However, a hybrid taken in the summer of 1991 (LACM 45542-
1) indicates parental influence of both species still exists in the area. Miller (1968)
and Hoover and St. Amant (1983) noted several unsuccessful introductions in
southern California and northern Baja California.

7. Gila orcutti (Eigenmann and Eigenmann). Arroyo chub. State special con-
cern. It is common at three localities within its native range, namely the upper
Santa Margarita River and its tributary, De Luz Creek, Trabuco Creek below
O’Neill Park and San Juan Creek (San Juan Creek drainage), and Malibu Creek.
The Malibu Creek population may be introduced because elsewhere arroyo chubs
always occur with Gasterosteus aculeatus, which is absent from this stream (Culver
and Hubbs 1917). Miller (1968) considered it native in Malibu Creek, and this
is confirmed by the finding of prehistoric remains of arroyo chub in middens
along upper Malibu Creek (Gobalet 1990). It is present but scarce in Big Tujunga
Canyon, Pacoima Creek above Pacoima Reservoir, and in the Sepulveda Flood
Control Basin, Los Angeles River drainage; upper San Gabriel River drainage;
and middle Santa Ana River tributaries between Riverside and the Orange County
line. In the San Gabriel system it is most abundant in the west fork below Cogswell
Reservoir where the gradient is low compared to the north and east forks (Richards
and Soltz 1986). It is usually confined to backwater pools or pools created for
recreational activities. It is scarce within its native range, because it does best in
the lower gradient streams that have largely disappeared. Low water conditions
(1986-present) made conditions more suitable to it in the West Fork of the San
Gabriel River and Big Tujunga Creek where it became abundant. After the rains
of 1991-1992 it again became scarce relative to Catostomus santaanae and Rhin-
ichthys osculus in the San Gabriel. It remains common in Big Tujunga where
Catostomus and Rhinichthys have become scarce and perhaps extirpated. Native
populations of Gila orcutti have been reduced enough to deserve close monitoring
to maintain or improve their status (Moyle and Williams 1990; Tres 1992).

Its rapid artificial range expansion in the 1930s and 1940s, as bait (Vestal 1942),
with trout plants (Miller and Hubbs 1969; CFG WAGE), or with mosquitofish
(Miller 1945a), established it in many extralimital areas (Fig. 8). Shebley’s (1922)
reference to abundant minnows in Big Bear Lake probably refers to introduced
arroyo chubs; they often became numerous and were eradicated as pests in south-
ern California reservoirs (Vestal 1942; CFG LB). It was introduced into tributaries
of the Mohave River, particularly Deep Creek, in the 1930s and has hybridized
there with Gila bicolor mohavensis (Hubbs and Miller 1943; Miller 1946a, 1968).
Fish introduced into the Cuyama drainage hybridize with Hesperoleucas sym-
metricus (Greenfield and Greenfield 1972; Greenfield and Deckert 1973). Miller
(1968) documented unsuccessful introductions of arroyo chubs into the San Felipe
Creek drainage (San Diego County) and the Rio San Tomas (northern Baja Cal-
ifornia) in the 1950s.

8. Gila elegans Baird and Girard. Bonytail chub. Extirpated in California,
state and federally endangered. Originally known only from the lower mainstream
Colorado River (and probably the Salton Sea) in California (Dill 1944; Minckley
1973). Occasional individuals possibly still occur in Lake Havasu (Tyus 199 1a).

116 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

ae

\}

Ee

ww

Fig. 8. Distribution of arroyo chub, Gila orcutti, in southern California. Black circles from 1975
onward; half circles before 1975; black triangles viable introduced populations; open triangles extir-
pated introductions; black square hybrids with roach, Hesperoleucas symmetricus;, open squares hy-
brids with Mohave chub, Gila bicolor mohavensis. Recent reports of this species from Pine Valley
Creek (Tijuana River drainage) occur off the map to the south. See Figs. 1-3 for stream names.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 117

Early observers did not separate G. cypha Miller, not described until 1946, and
it may have also occurred in the lower Colorado River in California (Gobalet
1992).

9. Gila robusta Baird and Girard. Roundtail chub. Extirpated in California.
A few early records of this species from the lower Colorado River date from the
early 1900s, and the species still occurs farther upstream (Minckley 1973; Tyus
199 1a).

10. Plagopterus argentissimus Cope. Woundfin. Extirpated in California and
federally endangered. The woundfin also was collected in the lower Colorado
River in Arizona before 1900, and undoubtedly occurred in the main river in
California, although extant specimens do not exist (Minckley 1973; Hubbs et al.
1979; Gobalet 1992).

11. Ptychocheilus lucius Girard. Colorado squawfish. Extirpated in California,
state and federally endangered. Originally known only from the mainstream Col-
orado River in California (Minckley 1973), and abundant in irrigation canals near
the river and the Salton Sea from 1904-1905 until about 1930 (Dill 1944). Fish
have been introduced into the lower river by the Arizona Department of Fish
and Game since 1985 with uncertain results (Tyus 1991b).

12. Rhinichthys osculus (Girard). Colorado speckled dace. One larval speckled
dace was taken in the lower Colorado River (Minckley 1973); otherwise this species
has not been taken in the lower river.

13. Rhinichthys osculus (Girard). Amargosa Canyon speckled dace. State tax-
on of special concern. This fish occurs only in the Armagosa River near Tecopa.
Other populations are found in Nevada (Moyle et al. 1989). Fish were unsuc-
cessfully introduced into the Borax Works on the west side of Death Valley in
1940 (Miller 1968).

14. Rhinichthys osculus (Girard). Owens speckled dace. State taxon of special
concern. This taxon occurs only in Fish Slough east of Bishop and in a few other
localities in the Owens drainage (Moyle et al. 1989). It was unsuccessfully intro-
duced into Saline Valley in 1939 (Miller 1968).

15. Rhinichthys osculus (Giarard). Santa Ana speckled dace. State taxon of
special concern. This is one of the rarest native freshwater fish in coastal southern
California (Fig. 9). It is abundant only in the lower parts of the East, North, and
West forks of the San Gabriel River. Small populations existed (sightings of one
or a few individuals in the last five years or so) in Fish Canyon (a small tributary
of the San Gabriel River); and Lytle, Cajon, City, Strawberry, Mill, and Silverado
creeks, tributaries of the Santa Ana River system. Fish could not be found in
1990-1992 in Big Tujunga and Santiago creeks despite thorough search and may
be extirpated in them. The fish in Big Tujunga are the last known in the Los
Angeles River drainage (Moyle and Williams 1990).

Miller (1968) noted records of Santa Ana speckled dace in the foothills of the
Montebello Oil fields just northwest of Whittier in 1940. Unpublished records
from the lowlands, Chino Creek and the nearby Santa Ana River (RGM) in 1939,
Santa Ana River, Riverside (LACM 9436) in 1951, and Los Angeles River in
North Hollywood (RGM) in 1939 indicate that relict populations existed across
the lowlands of the Los Angeles basin. Miller (1968) noted the successful estab-
lishment of the Montebello population in River Springs on the east side of Adobe
Valley, Mono County, in 1940.

118 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 9. Distribution of the two forms of speckled dace, Rhinichthys osculus, in coastal southern
California. Black circles, Santa Ana speckled dace from 1970 onward; half circles before 1970; black
triangles introduced. Open circles San Luis Obispo speckled dace; open triangles possibly introduced.
See Figs. 1-3 for stream names.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 119

Miller (1946b) reported speckled dace thought to be this taxon in Castaic Creek
and Elizabeth Lake Canyon, both tributaries of the Santa Clara River. These have
never been collected again and this supports Miller’s suspicion that these may
have been introduced.

16. Rhinichthys osculus (Girard). San Luis Obispo Speckled dace. The San
Luis Obispo Creek population, first noted by Jordan (1895), is probably related
(or identical) to those in the Salinas River, rather than to southern California fish
(Cornelius 1969). Salinas River fish were combined with Sacramento Valley fish
by Moyle and Williams (1990) and Hubbs et al. (1979). We have not found this
taxon in streams north of San Luis Obispo Creek as intimated by Miller (1968),
but rather in two streams to the south, Pismo and Arroyo Grande creeks. Miller
(1968) considered this fish introduced into the next stream to the south, the
Cuyama River of the Santa Maria drainage. Records from this drainage also
include Davy Brown and Tepusquet creeks and the Sisquoc River.

Catostomidae

17. Catostomus /atipinnis Baird and Girard. Flannelmouth sucker. Extirpated
in California. Originally only in the lower Colorado River in California (Minckley
1973), the note of it by Dawson (1923) in the Salton Sea in 1913 is the basis for
its inclusion in the California fauna (Dill 1944; Hubbs et al. 1979). Fish were
introduced below Davis Dam, and specimens were taken in Imperial Reservoir
as late as the 1970s (W. L. Minckley, pers. comm.).

18. Catostomus fumeiventris Miller. Owens sucker. The Owens sucker is com-
mon to abundant in much of the main Owens River including Crowley Lake
(Buth and Murphy 1980). This species apparently entered the Santa Clara system
with Owens River water in the 1930s (Hubbs et al. 1943; Miller 1973; Bell 1978).
Records are from lower Sespe Creek, the outlet of Fillmore Hatchery into and
including the nearby Santa Clara River, and Piru Creek above Piru Lake. It
hybridizes with Catostomus santaanae (also introduced) in the Santa Clara River
(Hubbs et al. 1943; Buth and Crabtree 1982). During winter and early spring high
flows Owens suckers and hybrids with Santa Ana suckers are common in the
Fillmore hatchery area, but are difficult to find at other times of the year when
water flow is much reduced or absent. Formerly large numbers of spawning adults
(100-150) gathered below the outlet of Fillmore hatchery for a few weeks in March,
but only about half this number or less have been observed in the last three or
four years. The artificial permanence of this flow may sustain the population
during the rest of the year. Large suckers observed in the spring of 1990 and 1991
in Piru Creek above Piru Lake are probably this species and possibly hybrids as
well. This species may have been part of the commercial catch in southern Cal-
ifornia reservoirs in the 1950s and 1960s (see C. occidentalis below).

19. Catostomus santaanae (Snyder). Santa Ana sucker. State species of special
concern, Federal candidate 2 species. Native populations still exist in the East,
North, and West forks of the San Gabriel River and in the lower Santa Ana River
from about Mt. Roubidoux downstream to a few miles below Imperial Highway.
In southern California, the Santa Ana sucker is distributed very much like Rhin-
ichthys osculus, but in larger streams (Fig. 10). Thus it is absent from Fish and
Santiago canyons and Cajon and City creeks of the Santa Ana drainage. It is
common in the San Gabriel River system. Large numbers have been taken recently

120 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 10. Distribution of the Santa Ana sucker, Catostomus santaanae, in coastal southern Cali-
fornia. Black circles native records from 1970 onward; half circles native records before 1970; black
triangles introduced; black squares hybrids with dusky suckers, Catostomus fumeiventris. See Figs. 2-
3 for stream names.

in the lower Santa Ana River from Riverside to a few miles below Imperial
Highway, but at other times few or none can be collected. It formerly was native
in the uplands and lowlands of the Los Angeles Basin streams (Smith 1966; Swift
1980c); now it is restricted to the uplands of the Los Angeles and San Gabriel

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 121

systems, and conversely, to the lowlands of the Santa Ana system. Fish became
very rare in the Big Tujunga drainage in 1990-1992 and the species may soon be
extirpated from the Los Angeles River drainage.

Large introduced populations occur in the Santa Clara River (Hubbs et al. 1943;
Bell 1978). In the Sespe Creek area, some hybridization with dusky suckers, C.
fumeiventris, occurs; a few downstream Santa Ana suckers have allozymes of
dusky suckers (Don Buth, pers. comm.). Genetic contamination does not extend
to the isolated Soledad Canyon area upstream (Buth and Crabtree 1982), where
Greenfield et al. (1970) studied the biology of this species. This area is a possible
refuge for C. santaanae since it is becoming rare in its native range. This population
also became much reduced with the drought. A few Santa Ana suckers were
introduced accidentally into River Springs, Mono County in 1940, but did not
become established (Miller 1968).

20. Xyrauchen texanus (Abbott). Razorback sucker. State and federally en-
dangered. The remaining, rare large individuals are believed to be old members
of a dwindling, non-reproductive, remnant stock (Tyus 1991a; Minckley, Marsh
et al. 1991). Buth et al. (1987) reported fertilized eggs collected from Senator
Wash Reservoir, Imperial County in March, 1983 and raised in aquaria. No
recruitment of wild-spawned fish occurs, probably because of predation by intro-
duced fishes (Tyus 1991a). Razorback suckers (then called humpback sucker)
occurred in commercial quantities in the Salton Sea until about 1930 (Croker
1934).

Cyprinodontidae

21. Cyprinodon macularius Baird and Girard. Desert pupfish. State and fed-
erally endangered. This species has declined precipitously in the last 10-15 years
apparently due to the introduction of tilapias into the lower Colorado River
including the Salton Sea (Bolster 1990c). Schoenherr (1988) believed only two
native populations remained in California, in lower San Felipe and Salt creeks,
Imperial County. Subsequent surveys (Lau and Boehm 1991) disclosed popula-
tions at more than 50 localities in canals on the southern and eastern margins of
the Salton Sea, probably maintained by the habitat heterogeneity described earlier
by Schoenherr (1979). A few other populations exist in Arizona and Mexico
(Minckley Meffe et al. 1991). Fish were unsuccessfully introduced to the Owens
Valley in 1940 (Miller 1968).

22. Cyprinodon nevadensis nevadensis Eigenmann and Eigenmann. Saratoga
Springs pupfish. State subspecies of special concern. Known. only from Saratoga
Springs, southeastern corner Death Valley National Monument, Inyo County,
California (Soltz and Naiman 1978; Minckley, Meffe et al. 1991). Fish were
unsuccessfully introduced into Lucerne Valley Reservoir, San Bernardino County
in 1939 and 1940 (Miller 1968).

23. Cyprinodon nevadensis amargosae Miller. Amargosa pupfish. State spe-
cial concern. Known only in the Amargosa River from Tecopa down through
Amargosa Canyon and for about 1.5 km northwest of Saratoga Springs (Soltz and
Naiman 1978; Minckley et al. 1991). Miller (1968) noted the successful intro-
duction of this species into River Springs, Mono County from 1940 to 1965, as
well as its unsuccessful introduction to six other sites.

122 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

24. Cyprinodon nevadensis shoshone Miller. Shoshone pupfish. State special
concern. Known only from Shoshone Spring at Shoshone, Inyo County, California
according to Castleberry et al. (1990) and Minckley, Meffe et al. (1991), who noted
that mosquitofish, Gambusia affinis, far outnumbered pupfish at this locality.
Since then a captive population has been reintroduced into a refugium pond
constructed at the headspring of this locality (Susan Ellis, pers. comm.). An un-
successful introduction was made into the Old Borax Works, Death Valley in
1939 (Miller 1968).

25. Cyprinodon nevadensis calidae Miller. Tecopa pupfish. Extinct. Formerly
occupied the outlets of North and South Tecopa Hot Springs. Alteration of the
spring for bathing and upstream invasion of Armagosa pupfish led to the extinction
of this form (Soltz and Naiman 1978; Minckley, Meffe et al. 1991).

26. Cyprinodon radiosus Miller. Owens pupfish. State and federally endan-
gered. Fish Slough and surrounding marsh, west of Bishop, Inyo County (Owens
Valley Native Fish Sanctuary), Marvin’s Marsh, and BLM Spring are three rel-
atively natural marshes and springs still holding Owens pupfish. These are aug-
mented by four artificial Owens Valley sites (Bolster 1990d). This fish was con-
sidered effective in controlling mosquitos (Kennedy 1916) until habitat modification
and introduced fishes reduced its numbers. Believed to be extinct, it was redis-
covered in the 1940s and with great effort is now maintained in the refuges noted
above (Miller and Pister 1971; Minckley, Meffe et al. 1991).

27. Cyprinodon salinus salinus Miller. Salt Creek pupfish. State subspecies
of special concern. Known only from Salt Creek and associated marshes for about
i.5 to 6 km below McLean Springs, Inyo County, depending on rainfall (Soltz
and Naiman 1978; Minckley, Meffe et al. 1991). It was successfully introduced
into River Spring in Adobe Valley, Inyo County from 1945 to at least 1965 (Miller
1968).

28. Cyprinodon salinus milleri La Bounty and Deacon. Cottonball Marsh pup-
fish. State threatened. Known only from Cottonball Marsh, Inyo County, Cali-
fornia (Soltz and Naiman 1978; Minckley, Meffe et al. 1991).

Fundulidae

29. Fundulus parvipinnis Girard. California killifish. Fish are common in
most larger bays and marshes north to Goleta Slough, and occur disjunctly in
Morro Bay where they are also common. None are known from Gaviota, Jalama,
San Antonio or San Luis Obispo creeks, or the Santa Maria and Santa Ynez rivers,
the most likely places for them to occur between Morro Bay and Goleta Slough.
The record from Monterey Bay (Moyle 1976a) is considered an introduction that
did not survive or a misidentification (Swift 1980d). Subsequent surveys did not
find them in the Elkhorn Slough area (Yoklavich et al. 1991). Eigenmann’s (1892)
assertion that this species was abundant in Lake Elsinore is otherwise unsubstan-
tiated. Freshwater records are numerous (Hubbs 1916; Miller 1939, 1943; Moyle
1976a); new freshwater records (LACM) are from Ventura River, Malibu Creek,
Santa Clara River, and Ballona Marsh.

California killifish were sold for bait on the lower Colorado River in 1951
(Miller 1952). About 150 California killifish were placed in Lake Elsinore in April
1954 (CFG PAD) and did not survive. One or possibly two specimens were

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 123

collected from the Salton Sea in March and April 1968, reportedly a release of
fish being sold for bait four years previously (CFG JAS).

Poecilidae

30. Poeciliopsis occidentalis (Baird and Girard). Gila topminnow. Extinct in
California, federally endangered. Hubbs et al. (1979) record this species from
California, presumably along the Colorado River, but it has not been collected
in California since before 1900. Moyle (1976a) and Shapavalov et al. (1981) did
not include it as part of the California fauna.

Gasterosteidae

31. Gasterosteus aculeatus aculeatus Linnaeus. Fully armored threespine stick-
leback. Fully armored, non-anadromous threespine stickleback have been re-
corded south to San Luis Obispo Creek, San Luis Obispo County by Baumgartner
and Bell (1984). They found the frequency of fully armored fish to increase north-
ward, associated with decreasing mean January temperatures and increasing pre-
cipitation. Baumgartner and Bell (1984) and Miller and Hubbs (1969) thought
anadromous sticklebacks did not occur in our area. Miller and Hubbs (1969)
reported a collection from the Big Sur River that might represent the marine
“trachurus” form.

32. Gasterosteus aculeatus microcephalus Girard. Partially armored threespine
stickleback. This subspecies appears to be widespread north of Point Conception,
but to the south has been declining rapidly during the last five years. Many local
populations no longer exist, and it is possible many of these are gone permanently.
South of the Los Angeles Basin, the only recent records (1991 onward) are from
Trabuco Creek, in and below O’Neill Park; upper San Juan Creek near the mouths
of Hot Spring and Cold Spring canyons; upper reaches of Bell Canyon on Starr
Ranch (all in the San Juan Creek drainage); South Fork of the San Jacinto River
below Lake Hemet, San Diego County, and a one kilometer reach of the Rio San
Domingo in Baja California. Miller and Hubbs (1969) and Ross (1973) found
slight morphological distinctions for San Juan Creek fish when compared to others
south of the Los Angeles Basin.

Gasterosteus a. microcephalus apparently was absent from the Los Angeles Basin
where G. a. williamsoni was present, but the lack of early coastal collections make
any conclusion equivocal (Ross 1973; Miller and Hubbs 1969; Bell 1978; Swift
1989; Swift et al. 1989). Except for the steep Big Sur area, sticklebacks are con-
tinuously distributed in lowland areas north of Point Conception. South of the
Santa Ynez drainage (which lies just north of Pt. Conception), localities are dis-
continuous (north to south); Gaviota Creek, lagoon in Carpenteria, Ventura River,
Santa Clara River, Calleguas Creek, San Juan Creek, San Mateo Creek, Santa
Margarita River, San Luis Rey River, and four localities in northern Baja Cali-
fornia (Miller and Hubbs 1969). In addition fish occurred in Wildcat Canyon a
few miles south of a locality at Tijuana Hot Springs (Smith 1883). These two
records may be the same as those noted from San Diego by Eigenmann (1887);
no other authors have reported them there. Thus several distributional gaps have
always existed south of Point Conception (Fig. 11), in steep small streams lacking
low gradient habitat, and in all but extreme northern and southern San Diego
County.

124 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 11. Distribution of fully and partially armored stickleback, Gasterosteus aculeatus, in southern
California. Open circles native fully armored fish; black circles native partially armored fish from
1970 onward; half circles native records of partially armored fish before 1970; black triangles viable
introduced populations of partially armored fish; open triangles extirpated introduced populations of
partially armored fish. See figures 1-3, 5 for stream names.

y.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 125

This taxon has been introduced widely, and native sticklebacks were promoted
for mosquito control in the early 1900s (Scofield 1915; Hubbs 1919). In the late
1930s and 1940s sticklebacks from the Fillmore area became established in Fill-
more and Mohave River hatchery ponds, and were introduced inadvertently with
trout plants. Established populations have been recorded in Dos Pueblos Canyon,
Santa Barbara County; Big and Little Rock creeks, Holcomb Creek, and Lake
Arrowhead, Mohave River drainage; Big Bear Lake, San Bernardino County;
Sweetwater River, and Pine Creek (tributary to Otay River), San Diego County
(Fig. 11); June Lake, Mono County; and upper and lower Owens River down-
stream to Bishop, Mono and Inyo counties. A few were found among fishes
eradicated from Crystal Lake, San Gabriel River drainage in the 1940s and 1950s
(CFG LB). These sites had no previous records of sticklebacks. Bell (1982) believed
the Holcomb Creek fish were native, but Buth (1984) and Haglund and Buth
(1988) showed they were electrophoretically similar to fish from the lower Sespe
Creek (near Fillmore Hatchery). Fish undoubtedly were placed in other localities
already containing native stickleback populations. The extralimital distribution
of arroyo chubs (Fig. 8), distributed by trout fishermen, may serve as a rough
guide to the introductions of sticklebacks. Some introductions were documented
previously by Ross (1973), Miller and Hubbs (1969), Bell (1982), Haglund and
Buth (1988), and Hendrickson and Brooks (1991).

33. Gasterosteus aculeatus williamsoni Girard. Unarmored threespine stick-
leback. State and federally endangered. This subspecies, originally widespread
and abundant in the Los Angeles Basin (Culver and Hubbs 1917; George 1927),
is restricted to a 14 km stretch of the Soledad Canyon portion of the upper Santa
Clara River and upper San Francisquito Canyon (U.S. Fish and Wildlife Service
1985; Buth 1984; Haglund and Buth 1988; Fig. 12). It is distinctive morpholog-
ically and electrophoretically (Miller and Hubbs 1969; Bell and Richkind 1981;
Buth 1984; Buth et al. 1984). Unplated sticklebacks apparently native in the upper
Huasna River and its tributary Alamo Creek (Santa Maria River drainage) and
introduced into the Mohave River drainage (both based on pre-1940 collections)
were eliminated by subsequent introductions of partially armored fish (Miller and
Hubbs 1969; Buth 1984). Those from upper Piru Creek, Elizabeth Lake Canyon,
and Castaic Creek (Miller and Hubbs 1969) have disappeared (U.S. Fish and
Wildlife Service 1985), and a population was recently discovered in Escondido
Canyon tributary to Agua Dulce Canyon in September 1992 (Linda Pardy, Jon-
athan Baskin, pers. comm.). This population is only one to two kilometers up-
stream of the main extant population in Soledad Canyon.

Gasterosteus a. williamsoni has been absent from the Los Angeles Basin since
the early 1950s. The last records were in 1925 (Los Angeles River, LACM 982,
987), 1951 (San Gabriel River, Mission Creek, Whittier Narrows, CFG RB, PAD),
and 1947 (Santa Ana River, Lake Norco, DFG WAE). In 1972 and 1973, 458
fish from upper Soledad Canyon were placed in upper San Felipe Creek and
Sentenac Canyon, Salton Sea drainage, San Diego County. These did well until
the winter of 1979-1980, after which they disappeared. In 1981, 180 more fish
were placed in the stream, and these have survived through September of 1990.
Unsuccessful transplants have been made of fish from upper Soledad Canyon: in
July 1975 to Malibu Creek in the vicinity of Tapia Park, in September 1973 to
the West Fork of the San Gabriel River just below Cogswell Reservoir, in 1982

126 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Distribution of the forms of unplated stickleback, Gasterosteus aculeatus, in southern

Fig. 12.
California. Black circles native records of Gasterosteus aculeatus williamsoni from 1960 onward; half

circles before 1960; black triangles viable introduced populations; half blackened triangles extirpated
introductions. Black squares San Antonio Creek stickleback; open squares introduced viable popu-

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 127

about 300 fish to a small artificial stream in the Botanical Garden on the campus
of the University of California, Los Angeles (LACM W82-13), and in 1972 into
Piru Pond, Ventura County (CFG LB UTSRT; Wells et al. 1975).

34. Gasterosteus aculeatus. San Antonio Creek stickleback. Status undeter-
mined. This taxon is known only from the lower 6 to 8 km of San Antonio Creek,
Vandenberg Air Force Base, Santa Barbara County (Fig. 12). It is also low-plated
and originally was considered G. a. williamsoni (Irwin and Soltz 1982; U.S. Fish
and Wildlife Service 1985), based on morphology. Electrophoretically it does not
differ significantly from local populations of the subspecies G. a. microcephalus
(Buth 1984; Haglund and Buth 1988). It is distinct morphologically, and such
evolutionary novelties are the reason the population is important to preserve.
Fish from San Antonio Creek (about 850) were planted into Honda Canyon on
Vandenberg Air Force Base in 1984 and survive today (CFG UTSRT).

35. Gasterosteus aculeatus. Shay Creek stickleback. This population is low-
plated and distinctive electrophoretically (Haglund and Buth 1988). It was recently
discovered (1984) in the Baldwin Lake drainage just east of Big Bear Lake (Fig.
12) and is restricted to two or three large pools and interconnecting meadow (CFG
UTSRT). It has been transplanted to one other site, Sugarloaf Meadows, 7 miles
airline southeast of Baldwin Lake in the San Bernardino Mountains. This form
is certainly one of the strongest candidates for immediate protection among the
taxa covered here. :

Because all native southern California sticklebacks known are from lower el-
evations, this may be an introduced population. Shebley (1927) participated in
the first introductions of fish into the Big Bear area in 1891-1892, transporting
rainbow trout by muleback on the old trail up the Santa Ana River. Although
most southern California mountains are highly eroded, Norris and Webb (1990)
note that the San Bernardino Mountains uncharacteristically retain a significant
rolling upland plateau. Pre-historical continuity of this feature possibly provided
habitat for the survival of native sticklebacks as the mountains were uplifted.

Moyle et al. (1989) and Moyle and Williams (1990) use the subspecific com-
bination G. a. santaanae Regan (1909) for this fish Regan described from fish
taken from the Santa Ana River, Riverside. Shay Creek is not in the Santa Ana
River drainage today, and it is premature to assign this name until adequate
comparisons are made. Thus we only use the vernacular, Shay Creek stickleback.

Cottidae

36. Cottus asper Richardson. Coastal prickly sculpin. This sculpin is wide-
spread in coastal streams north of Point Conception where juveniles are common
in coastal lagoons (Fig. 13). Farther southward only four localities are known:
Santa Anita Canyon (LACM 36657-2), Gaviota Creek (LACM 35423-1), Arroyo
Quemado (SBMNH), and lower Ventura River (UMMZ 132891). The first three
date from 1975, and the last from 1938. The few, rarely collected, large individuals
indicate that larvae stray southward and colonize, but do not reproduce. Larval
Cottus asper occur in near-shore plankton farther north (Matarese et al. 1989).

—

lations. Open circle Shay Creek stickleback; open triangle viable introduced population. See Figs. 1—
3 for stream names.

128 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Fig. 13. Distribution of ihe species of Cofttus in coastal southern California. Black circles native
records of coastal prickly sculpin, C. asper from 1970 onward; half circle records before 1970; black
triangles introduced inland prickly sculpin, Cottus asper; circles coastrange sculpin, C. aleuticus. See
Figs. 1-3 for stream names.

.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 129

Recent single collections from reservoirs, Lake Cachuma in 1975 and Lake Casitas
in 1989, probably are introduced, but could possibly be native coastal forms
trapped on the upstream side of dams.

Hubbs (1921) and Kresja (1967a) note variation in the degree of prickling in
the vicinity of San Luis Obispo. Juveniles of the coastal form can have half or
more of the lateral sides of the body with prickles. These are lost with growth in
coastal prickly sculpin. Adults of the inland form (discussed below) retain prickling
on the body. This is the only catadromous taxon in California, namely that
descends from streams into tidal or lagoonal waters to spawn (Kresja 1967b).
Prickly sculpin (and threespine stickleback) have been recorded from only a few
localities between the Carmel River and Arroyo de la Cruz (Fig. 13); these larger
streams have lagoon development, apparently a requirement for C. asper. Hubbs
(1921) noted that Jordan’s (1895) record of C. gulosus from San Luis Obispo
Creek was based on C. asper (see account of C. aleuticus below).

37. Cottus aleuticus Gilbert. Coastrange sculpin. This sculpin has been re-
corded south to Oso Flaco Creek, Santa Barbara County (LACM 35163-1), ex-
tending the range southward from “‘a stream between San Simeon and Piedras
Blancas” (Hubbs 1921) (Fig. 13). Despite the fact Matarese et al. (1989) thought
the larvae may not occur in marine plankton, it occurs in many coastal streams,
even several steep ones that lack C. asper. No specimens have been found to
verify various records of riffle sculpin, Cottus gulosus, from coastal streams of
San Luis Obispo County (Barclay 1975; CFG records); we believe all such records
were based on misidentifications of coastrange sculpins. The latter keys out with
C. gulosus in Robins and Miller (1957).

Gobiidae

38. Eucyclogobius newberryi (Girard). Tidewater goby. Petitioned for federal
threatened status. In our area 28 localities remain and more of these may disappear
with the continuing drought. Extant localities are (north to south): Arroyo Laguna
(Oak Knoll Creek), Tortuga Canyon, Arroyo del Puerto, Little Pico, Pico, San
Simeon, Santa Rosa, Villa, San Geronimo, Cayucos, Pismo creeks, and Santa
Maria River (San Luis Obispo County), Shuman Canyon, San Antonio Creek,
Santa Ynez River, Jalama Creek, Canadas de Cojo, Las Agujas, Santa Anita, and
Alegria, Gaviota Creek, and Refugio and Bell canyons (Santa Barbara County),
Ventura and Santa Clara rivers (Ventura County), Malibu Creek, re-established
1991 (Los Angeles County), San Onofre Creek, Las Pulgas Canyon, and Santa
Margarita River (San Diego County). In September 1992 a few were seen in
Cockleburr Canyon (between Santa Margarita and Las Pulgas), a site that lacked
them or was dry on many previous visits since 1980. Only fourteen localities exist
south of Point Conception.

Tidewater gobies have disappeared from many localities, and rarely recolonize
(Swift et al. 1989). Small lagoons often have only a few adult fish, thus lacking
continuously established populations, such as Arroyo de Corral and Leffingwell
and Willow creeks in San Luis Obispo County. In 1989 fish were newly collected
from Refugio Creek (Tom Taylor, in litt.) and lower San Luis Obispo Creek (Tim
Pafford, pers. comm., LACM 44824-1). None had been previously taken at these
localities since 1916 despite collecting by many individuals in the intervening
years.

130 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Swift et al. (1989) suggested re-introducing fish to localities where they formerly
occurred, and in June 1991, 52 adults from the mouth of the Ventura River were
placed in Malibu Lagoon by the Topanga—Las Virgines Resources Conservation
District and California Department of Parks and Recreation. In August about 75
mostly smaller fish were collected. In April 1992 and April 1993, several adults
that had overwintered were collected and in August 1992 many fish were seen in
the lagoon, indicating successful establishment.

Estuarine Fishes
Carcharhinidae

1. Carcharhinus leucas (Valenciennes). Bull shark. Miller and Lea (1976)
report this species from southern Caifornia, based on Fry and Roedel (1945) and
Rosenblatt and Baldwin (1958). Compagno (1984) did not record it north of
southern Baja California, but Eschmeyer et al. (1983) state that it possibly reached
southern California. Robins et al. (1991) accept it as recorded from the Pacific
coast of the United States. Elsewhere in the world this species is closely tied to
fresh and low salinity water, and the early demise of this habitat in southern
California may have led to its disappearance here.

Elopidae

2. Elops affinis Regan. Machete. Minckley (1973) and Swift (1980a) sum-
marized the marine and freshwater records from southern California. A fish (43
gm) taken in October 1980 (Colorado River below the confluence of the Gila
River) was listed as an angling size record for the state of Arizona (Sport Fishing
Institute Bulletin, Oct.-Nov., 1980, p. 8). Thus this species is a regular, if rare
(or at least rarely collected), member of the fauna of the lower Colorado River in
California. Fish have not been recorded upstream of Laguna Dam since the 1940s.
All freshwater records are from the Colorado River and the Salton Sea, although
marine records exist for coastal Ventura (Swift 1980a; Fitch and Schultz 1978)
and San Diego (R. N. Lea, pers. comm.) counties.

Clupeidae

3. Clupea pallasi Valenciennes. Pacific herring. Pacific herring spawn in low
salinity water in mid-winter in southern California; populations have been re-
corded from Morro Bay, San Luis Obispo Creek, Santa Ynez River lagoon, Goleta
Slough, San Diego Bay, and are suspected in Los Angeles Harbor (Miller and
Schmidtke 1956; Spratt 1981; Milton Love, pers. comm.). Miller and Schmidtke
(1956) note that Morro Bay and San Luis Obispo Creek fish are larger than others
in California.

Engraulidae

4. Engraulis mordax Girard. Northern anchovy. Common in bays and es-
tuaries, the northern anchovy only rarely enters fresh water. Two records known
to us are those of Hubbs (in Lane 1977a) and one juvenile taken in upper Santa
Margarita lagoon on 23 December 1983 (UT 30.8).

5. Anchoa delicatissima (Girard). Slough anchovy. This species has often been
reported from low salinity and fresh water in southern California (Allen 1982),
and occurs northward to Los Angeles Harbor.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 131

6. Anchoa compressa (Girard). Deepbody anchovy. This species is also abun-
dant in bays and harbors north to Morro Bay and also from fresh water (Horn
1981; Allen 1982). A hiatus in distribution occurs between Mugu Lagoon and
Morro Bay.

Syngnathidae

7. Syngnathus auliscus (Swain). Barred pipefish. This tropical species occurs
north to the Santa Barbara channel but is frequent only in southern San Diego
Bay (Fritzsche 1980) and Newport Bay (LACM, 7 collections). Starks and Morris
(1906) note that it favors more upstream, lower salinity locations than S. lepto-
rhynchus.

8. Syngnathus leptorhynchus Girard. Bay pipefish. This is the only other
species of pipefish that can be taken in brackish or fresh water in our area.
Freshwater records include Morro Bay and Malibu lagoon. This species was once
common in many bays but is much more scarce in Anaheim Bay than formerly
(Lane 1977a) and possibly is declining.

Atherinidae

9. Atherinops affinis (Ayres). Topsmelt. Topsmelt enter fresh water in the
upper portions of coastal lagoons and estuaries, particularly in spring when spawn-
ing in waters of lowered salinity. They dominate southern California estuaries
numerically and in biomass (Horn 1981, 1988; Allen 1982; Horn and Allen 1985),
but possibly were much less common originally (Lane 1977a). At many localities
north of Point Conception topsmelt are found primarily near the mouths of
estuaries and lagoons except in spring when they move inland to spawn in lower
salinity waters (Schultz 1934).

10. Leuresthes tenuis (Ayres). Grunion. Occasional fish are washed over the
barrier sand bars into coastal lagoons where they can be collected in brackish and
fresh water. Some early observers felt they tended to spawn near stream mouths,
but this was discounted by Hubbs (1916), Thompson (1919) and Walker (1952).
Most beaches occur, however, where stream and river mouths deposit sand and
where originally much more freshwater influence existed.

Hemirhamphidae

11. Hyporhamphus rosae (Jordan and Gilbert). California halfbeak. This spe-
cies also was recorded as common in San Diego and Mission bays near the turn
of the century (Eigenmann 1892), and has been rarely collected since (Miller and
Lea 1976). It is known from low salinity and fresh water farther south (Miller
1945b; Castro-Aguirre 1978) as H. patris, now a synonym of H. rosae (Collette
etal 1992):

Embiotocidae

12. Cymatogaster aggregata Gibbons. Shiner surfperch. This species is com-
mon in bays and estuaries in southern California. During the breeding season in
spring, it enters the warmer estuaries and lagoons, often of lowered salinities
(Odenweller 1977; Moyle 1976a).

132 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Cottidae

13. Leptocottus armatus Girard. Staghorn sculpin. This species is ubiquitous
in the mouths of streams throughout our area, particularly in mid-winter when
the young-of-the-year invade and coastal lagoons are mostly or completely dom-
inated by freshwater (Tasto 1977; Moyle 1976a; Horn 1981; LACM records).

Mugulidae

14. Mugil cephalus Linnaeus. Striped mullet. Silvery young-of-the-year in-
dividuals are often taken from December to March in coastal streams tributary
to lagoons and bays (Eigenmann 1892; Horn and Allen 1985). We know of such
fish taken as far north as the Santa Clara River (LACM 45110-1), and adults have
been taken in marine waters north to San Francisco Bay (Eschmeyer et al. 1983).
In December 1972 twelve juveniles were taken in the Santa Ana River at Featherly
Park about 35 km upstream from the mouth in the ocean (LACM 35223-2; 22-
26 mm SL). Individuals of all sizes occur today in the lower Colorado River
(Minckley 1973); commercial quantities occurred in the Salton Sea until about
1930 (Croker 1934), and adults were taken through the 1970s (CFG JAS). In
March 1966 about 50 “‘half-pound”’ striped mullet were placed in Puddingstone
Reservoir, Los Angeles County, and a few were caught up to two years later (CFG
LB).

Paralichthyidae

15. Paralichthys californicus (Ayres). California halibut. This species depends
on estuarine conditions for the first few years of life (Haaker 1977; Allen 1988;
Kramer 199 1a, b, 1992). It occurs primarily in larger estuaries with tidal openings,
but we have records of individual juveniles from Ballona Marsh and Malibu
Lagoon (LACM).

Pleuronectidae

16. Hypsopsetta guttulata (Girard). Diamond turbot. Young and juveniles of
this species are mostly taken in estuaries which it uses as nursery areas (Lane
1977b; Kramer 1991b). It is more numerous than starry flounder in southern
California estuaries; all of our records of young-of-the-year and juveniles in low
salinity water are south of Point Conception. .

17. Platichthys stellatus (Pallas). Starry flounder. Juveniles of this species
invade fresh or low salinity water in winter and early spring. Most such records
are north of Point Conception, but the southernmost is from Goleta Slough, Santa
Barbara County in the late 1960s (Milton Love, pers. comm.).

Sciaenidae

18. Cynoscion parvipinnis Ayres. Shortfin corvina. This species was collected
in San Diego and Mission bays until about 1925 (Follett 1976), and again in 1991
(R. N. Lea, pers. comm., SIO material). Eigenmann (1892) noted its probable
occurrence in the lower Sweetwater River in San Diego County. It is known from
low salinity and fresh water farther south in Mexico (Castro-Aguirre 1978). Cy-
noscion parvipinnis was introduced into the Salton Sea from the Gulf of California
from 1953 to 1955 (Walker et al. 1961), where it established a breeding population
for a number of years (Shapavalov et al. 1981).

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 133

Eleotridae

19. Eleotris picta Kner and Steindachner. Spotted sleeper. Recorded once in
1953 in the lower Colorado River and not taken since (Hubbs 1953; Minckley
1973).

Gobiidae

20. Gillichthys mirabilis Cooper. Mudsucker. This goby is often taken in fresh
water in the upper parts of bays and sloughs and can tolerate a wide range of
salinity. One juvenile (LACM 38569-1) was taken from the stomach of a garter
snake, Thamnophis c. atratus, from Arroyo de Corral in northern San Luis Obispo
County (Marc Hayes, pers. comm.).

Mudsuckers have long been used as live bait both in fresh and salt water. Dealers
obtain fish from coastal southern California and the outer coast of Baja California
near San Quintin (Miller 1952; Walker et al. 1961; Turner and Sexsmith 1967).
Fish from San Diego Bay were introduced to the Salton Sea by 1931 (California
Department of Fish and Game 1931). Miller (1952) recognized two subspecies
of G. mirabilis. He noted the only California record of the Gulf of California
form, G. m. detrusus Gilbert and Scofield, was based on dead specimens from a
pelican rookery near the Salton Sea. Later Barlow (1963) did not recognize sub-
species because of wide variation in many characters. Probably both Gulf of
California fish imported to California and southern California fish have been
released in the lower Colorado River, and into some southern California bays
and estuaries. Fish in lower Salt Creek near the Salton Sea successfully reproduce
(CFG JAS). Gillichthys seta Ginsburg, endemic to the upper Gulf of California,
was also introduced once into the Salton Sea (Walker et al. 1961).

21. Clevelandia ios (Jordan and Gilbert). Arrow goby. Lane (1977a) recorded
the early collections of Carl Hubbs (1916) from fresh or low salinity water in
upper Anaheim Bay, and other southern California freshwater records are from
the mouths of the Santa Clara (LACM 45110-1) and Santa Margarita (UT 122.36)
rivers and Malibu Creek.

22. Gobionellus sagittula (Gunther). Longtail goby. Only a few early records
exist for this species in San Diego Bay, as G. /ongicauda (Starks and Morris 1906),
and it apparently is extirpated in California.

Scombridae

23. Scomberomorus concolor (Lockington). Gulf sierra. Extirpated in Cali-
fornia. This species was last recorded in the early 1950s and disappeared soon
thereafter (Fitch and Flechsig 1949; Fitch 1952). Now only known from the upper
Gulf of California (Collette and Nauen 1983). Other species in the genus are
estuarine and its disappearance may be due to alteration of local estuaries.

Introduced Species
Acipenseridae

1. Acipenser transmontanus Richardson. White sturgeon. In March 1967, 39
white sturgeon were planted in Lake Havasu, Colorado River from San Pablo
Bay, California (CFG LB). Several more were released in 1968 (Minckley 1973).
In June 1967 and March 1969 single fish were taken downstream at Imperial

134 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Dam. Another was caught at the mouth of the Bill Williams River, Arizona in
1976 (Los Angeles Times, 27 March 1981, Pt. III, p. 16). No evidence of repro-
duction has been found. Over 200 white sturgeon (from the Arrowhead Fishery
in Red Bluff) were planted in Irvine Lake and the Santa Ana River lakes in
September 1987 (Los Angeles Times, 18 September 1987, Pt. III, p. 12) and
several have been captured since.

Osmeridae

2. Hypomesus nipponensis McAllister. Wakasagi. This species was introduced
into Big Bear Lake in 1959, and several hundred were recorded during chemical
treatment of the lake in the fall of 1960 (Wales 1962). None have been recorded
since.

Esocidae

3. Esox lucius Linnaeus. Northern pike. Shebley (1917a) and Smith (1896)
note that in 1891, 400 yearling northern pike were placed in Lake Cuyamaca, San
Diego County, and were still common in January 1896. Smith (1896) and Sha-
pavalov et al. (1959) provide evidence that some (or possibly all) of these fish
were E. americanus americanus Gmelin, redfin pickerel.

Salmonidae

4. Oncorhynchus clarki (Richardson). Cutthroat trout. Salmo evermanni Jor-
dan and Grinnell from the San Gorgonio Mountains was based on introduced
Lahontan cutthroat trout, O. c. henshawi (Gill and Jordan), that were placed in
several tributaries of the Santa Ana River before 1900 (Benson and Behnke 1961).
This population survived into the 1930s in the upper south fork of the Santa Ana
River below Dollar Lake (R. Croker, in litt.; Miller 1950; Needham and Gard
1963). Gold et al. (1978) noted the survival of pure Colorado cutthroat trout, O.
c. pleuriticus (Cope), in the Williamson Lakes, Owens River drainage after their
introduction in 1931. Most likely, several early literature records of this subspecies
from the lower Colorado River and the Salton Sea were misidentified rainbow
trout introduced earlier; straying downstream of cutthroat trout was considered
almost impossible (Dill 1944; Miller 1950; Minckley 1973).

5. Oncorhynchus aguabonita (Jordan). Golden trout. Widely introduced into
high lakes in the eastern Sierra Nevada Mountains (Moyle 1976a), golden trout
were also placed into the upper Santa Ana River in 1918 (Shebley 1919), but were
not recorded subsequently.

6. Oncorhynchus kisutch (Walbaum). Silver or coho salmon. Hatchery raised
young and juvenile coho salmon have been introduced into southern California
reservoirs, estuaries, and the Salton Sea from as early as 1935 (Walker et al. 1961;
CFG LB). Some have been caught in coastal waters, but no returns to freshwater
streams are known.

We have several verbal reports of a “‘salmon,” in addition to steelhead trout,
that entered southern California coastal streams prior to the 1950s. However, no
well-authenticated records of wild fish entering fresh water south of Monterey
Bay are known to us. Only this salmon species commonly enters spawning streams
south of San Francisco Bay. All five species of Pacific salmon have been recorded
southward at sea to between Los Angeles and northern Baja California (Hubbs

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 135

1946; Miller and Lea 1976; Eschmeyer et al. 1983). Given the large numbers at
sea locally and the hatchery fish introduced into local estuaries it is surprising
some have not been taken in local streams.

7. Oncorhynchus tshawytscha (Walbaum). Chinook salmon. Since 1984 young
chinook salmon from outside California have been raised and released in lower
San Luis Obispo Creek. This effort has created some returns of adult fish from
the sea, but to date depends on artificial propagation (CFG; San Luis Obispo
County Telegram Tribune, 12 February 1991, p. A-3).

Jordan’s (1895) record for king (now called chinook) salmon in San Luis Obispo
Creek was probably based on steelhead (Hubbs 1946). Jordan and Gilbert’s (1883)
early contention that the southern limit of king salmon was the Ventura River
was probably based on Jordan and Henshaw (1878) (quoted by Stone 1884) who
visited fish markets, talked to local fishermen, and did not actually secure spec-
imens from fresh water.

8. Oncorhynchus nerka (Walbaum). Kokanee salmon. This species was in-
troduced into Lake Arrowhead in the late 1950s and supported a fishery through
the 1960s (CFG LB; R. Croker, pers. comm.).

9. Salmo trutta Linnaeus. Brown trout. Although as widely introduced as the
other trout, and well established and famous in the Owens River since before
1900, brown trout have not fared as well in desert streams and coastal drainages
farther south. Fish were even placed in reservoirs on Catalina Island (Shebley
1925). Since 1970, authentic verbal and/or specimen records of this introduced
fish exist for Piru Creek, the Arroyo Seco-Bear Creek drainage, Deep Creek-
Mohave drainage, Ice House Canyon (San Antonio Creek drainage, Los Angeles-
San Bernardino county line), and the upper Santa Ana River including Bear Creek.
Reproduction by brown trout has been documented for Deep Creek and the Santa
Ana River and tributaries, where rainbow trout are very rare or non-existent
(Deinstadt et al. 1988). These authors demonstrate the virtual absence of brown
trout where rainbows are abundant in Piru Creek and the West Fork of the San
Gabriel River. Before 1938 more streams supported reproducing populations of
brown trout, such as the upper San Gabriel River, Santa Anita Canyon, Fish
Canyon, and Holy Jim Canyon (Trabuco Creek drainage). After the severe floods
of 1938-1939 many of these populations diminished or disappeared.

10. Salvelinus fontinalis (Walbaum). Brook trout. Thousands of fingerling
brook trout were introduced in many mountainous streams in the 1877-1940
period (Smith 1896; CFG LB). These only became established in high lakes and
creeks of the eastern Sierra Nevada mountains tributary to the Owens River.

Clupeidae

11. Alosa sapidissima (Wilson). American shad. This species was introduced
into the lower Colorado River from 1884-1886 (Smith 1896) but did not survive
(Minckley 1973). Marine records from coastal southern California date from 1880
(Los Angeles County, Smith 1896), and are known south to northern Baja Cali-
fornia (Eschmeyer et al. 1983). American shad also should be expected with
aqueduct water from central California.

12. Dorosoma petenense (Gunther). Threadfin shad. The original introduc-
tion of this species from the Tennessee River into southern California was recorded
by Kimsey (1954) and Shapavalov et al. (1959) (SIO H53-180 and 183). It also

136 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

is imported with Central Valley and Colorado River water. It occurs in freshwater
lakes, reservoirs, larger rivers, and coastal bays and estuaries (Moyle 1976a), the
lower Colorado River and, formerly, the Salton Sea (Walker et al. 1961; Minckley
1973).

Anguillidae

13. Anguilla australis Richardson. Shortfinned eel. Three specimens (LACM)
of about three dozen noted by biologists between 1978 and 1986 came from
Puddingstone Reservoir, Legg Lake, and lower Los Angeles River, Los Angeles
County. The origin of these is not known, but many live eels were imported during
this period (McCosker 1989; Williamson and Tabeta 1991). Williamson and
Tabeta found no evidence for natural straying by A. japonica into the northeastern
Pacific Ocean. McCosker reviewed California anguillid records, provided a key
to the species most likely to be imported, and pointed out that freshwater eels
probably would not successfully reproduce here. They may significantly prey on
native fishes; A. australis greater than 50 cm are largely piscivorous in New Zealand
(Ryan 1986).

Characidae

14. Astyanax mexicanus (Fillippi). Mexican tetra. This species was collected
a few times in the lower Colorado River in 1950 (Minckley 1973) as releases of
fishes sold for bait (Miller 1952).

Cyprinidae

15. Carassius auratus (Linnaeus). Goldfish. Goldfish have been known from
California fresh waters since before 1900 (Moyle 1976a, b), and were widely used
for bait (Dill 1944). They occur in reservoirs, ponds and lowland streams. The
sources and times of arrival are not well documented, but Gill (1862) reported a
collection from California taken a year or two earlier. They appeared in San Diego
County reservoirs with the importation of Colorado River water (CFG LB).

16. Cyprinus carpio Linnaeus. Commoncarp. This species was imported from
Germany in 1872 and Japan in 1877 (Shebley 1917a). It was abundant in the
Owens River by 1891 (Gilbert 1893), and occurs in most major reservoirs and
lowland portions of the larger rivers. It was commercially fished in many of the
a lakes and reservoirs until the 1960s (Davis 1963).

. Ctenopharyngodon idella (Valenciennes). Grass carp. This species, well
aUere in the central United States (Courtenay et al. 1984, 1986), was im-
ported, discovered, and eradicated at one coastal southern California locality,
Padre Juan Canyon, Ventura County in November 1975 (LACM 35498-1, 2).
Since the late 1970s they have been widely introduced in the Imperial Valley
(CFG LB, JAS; Thiery 1991). In April 1980 hybrids between this species and
Hypophthalmichthys nobilis, bighead carp, were planted in several waterways in
the Cochella Valley, and two were taken from Lake Cahuilla, Riverside County
in April 1981 (CFG LB). In 1985 18,000 triploid grass carp were introduced in
the Coachella Canal (Thiery 1991).

18. Gila bicolor obesa (Cope). Lahontan tui chub. Native to the Lahontan
Basin, this chub has been widely introduced into the Owens drainage, where it
has introgressed with the Owens chub (Miller 1973).

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 137

19. Hesperoleucas symmetricus (Baird and Girard). California roach. This
species, possibly introduced into the Cuyama River (Miller 1945c), hybridizes
with Gila orcutti, definitely introduced there (Miller 1945a; Greenfield and Green-
field 1972; Greenfield and Deckert 1973). The occurrence of Rhinichthys in this
drainage lends support to Greenfield and Greenfield’s (1972) contention that the
dace and roach are native.

20. Ptychocheilus grandis (Ayres). Sacramento squawfish. This species, es-
tablished before 1975 in Chorro and Los Osos creeks, tributaries to Morro Bay
(Barclay 1975; Moyle 1980), occurs down to upper tidal areas of the bay. One
specimen was taken in Lake Arrowhead in June 1966 (SIO66-75), and in 1988
from Pyramid Lake (CFB LB). The latter probably arrived with central California
water. In the Eel River, introduced squawfish restrict habitat use by other native
fishes (Brown and Moyle 1991).

21. Lavinia exilicauda Baird and Girard. Hitch. This species was reported
from Pyramid Lake in 1984 (CFG DD), Lake Silverwood in 1988 (CFG FGH,
LP), and one was taken in June 1992 at Pyramid Lake (LACM 45701-1). It is
possibly established or at least a regular immigrant from central California. Miller
(1952) reported hitch sold as bait in the lower Colorado River, and Kimsey and
Fisk (1960) reported one collected in Ramer Lake, Imperial County. Hitch were
collected in the spring of 1992 from isolated pools of Aliso Canyon, tributary to
Santa Clara River in Soledad Canyon (LACM 45699-1), and thought to have
been there for about 40 years (Tom Hale, pers. comm.).

22. Orthodon microlepidotus (Ayres). Blackfish. Blackfish were in Tweedy
Lake in 1918 and by 1938 had spread into Munz and Elizabeth lakes and into
Lake Hughes in northern Los Angeles County (CFG LB). These were eradicated
in the early 1940s. One was taken in Echo Park Lake, Los Angeles in February
1984. The species has been established in the lower Santa Ana River below Prado
Reservoir since about 1986. It is one of the few native Central Valley species to
thrive in highly modified aquatic habitats (Jennings and Saiki 1990).

A specimen from Pyramid Lake taken in June 1992 has intermediate features
and apparently is a hybrid with Lavinia exilicauda (LACM 45701-2), a combi-
nation first reported by Hopkirk (1974), and found to be common in San Luis
Reservoir in central California by Moyle and Massingill (1981).

23. Pogonichthys microlepidotus Ayres. Splittail. One fish was reported taken
in Lake Silverwood in February 1977 (CFG LB). The specimen was not kept, but
the species would be expected to come in with aqueduct water.

24. Notemigonus crysoleucas (Mitchill). Golden shiner. This well known bait
fish can be taken almost anywhere that permanent fresh water exists that is also
utilized by fishermen. It arrived in 1891 in San Diego County and quickly became
widespread (Hubbs 1919; Kimsey and Fisk 1964; Miller 1952). Since about 1980,
bait farms outside California have raised rudd, Scardinus erythrophthalmus (Lin-
naeus), a minnow that closely resembles golden shiner. It has been distributed to
14 states, and wild captures have occurred in eight states (Burkhead and Williams
1991). Burkhead and Williams (1991) document hybridization between golden
shiners and rudd. Since rudd are popular bait for striped bass, they will probably
turn up in California.

25. Cyprinella lutrensis (Baird and Girard). Red shiner. Between 1948 and
1951 red shiners cultured near Blythe became established in the lower Colorado

138 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

River and Salton Sea areas (Hubbs 1954). About 100 were unsuccessfully intro-
duced to Puddingstone Reservoir in 1955. Red shiners appeared in Big Tujunga
Creek in 1983 and were established by August 1985 (DFG DD; Jennings and
Saiki 1990). They were still present in the lower creek below Tujunga Reservoir
in June 1992. Red shiners occurred in uppermost Newport Bay in September
1986 at the mouth of San Diego Creek (LACM 44508-1, 44509-1), and further
upstream at Peters Canyon Wash in 1983. Fish were abundant in the lagoon at
the mouth of San Juan Creek in this same period, could not be found there in
August 1992, but were recorded farther upstream at the crossing of Ortega Hwy
in July 1992. They occur with arroyo chubs in Big Tujunga Canyon, but so far
occur downstream of arroyo chub populations in San Juan Creek.

26. Pimephales promelas Rafinesque. Fathead minnow. This bait fish is com-
mon and established in most if not all the larger low gradient streams in the area,
as well as in some lakes and reservoirs. It first appeared along the lower Colorado
River with red shiners near Blythe in 1950 and quickly became widespread in
southern California (Miller 1952; Shapavalov et al. 1959; Kimsey and Fisk 1964).

Catostomidae

27. Catostomus occidentalis Ayres. California sucker. This species, estab-
lished in Cayucos Creek and Whale Rock Reservoir, San Luis Obispo County
from 1986 to 1989, was absent after low water conditions in 1990 (Dan Holland,
pers. comm.). Fish were present in Chorro and Morro creeks (Barclay 1975) until
1978 (CFG SLO); all of these probably were introduced from the Salinas drainage.
Although Davis (1963) noted “‘western suckers’? (often used in reference to C.
occidentalis) in the commercial catch from southern California reservoirs, these
most likely were Owens suckers (see above). Since 1972 California suckers are
potential immigrants with central Valley water.

28. Ictiobus cyprinellus (Valenciennes). Bigmouth buffalo. The bigmouth buf-
falo was introduced (probably from Arizona) into the Los Angeles Aqueduct
system in Inyo and Los Angeles counties, and supported a commercial fishery
through the 1960s in southern California reservoirs (Evans 1950; Davis 1963).
Minckley (1973) noted that Arizona fish rarely occurred in the lower Colorado
River, and that two other species existed in Arizona reservoirs, J. niger (Rafin-
esque) and J. bubalus (Rafinesque). Shapavalov et al. (1981) reported a fish from
a southern California waterway (1969) identified as J. bubalus by C. Hubbs and
J. St. Amant. None have been collected since and they probably no longer occur
outside the Colorado River (Shapavalov et al. 1981).

Ictaluridae

29. Ictalurus punctatus Rafinesque. Channel catfish. This species is wide-
spread in rivers and reservoirs in this area as well as the Colorado River (Minckley
1973; Moyle 1976a). It was first introduced into Lake Cuyamaca, San Diego
County in 1891 (Smith 1896) with fish originating from the northeastern United
States. Later introductions have come from Texas (CFG LB).

30. Ictalurus furcatus (Lesueur). Blue catfish. Richardson et al. (1970) doc-
ument the introduction of this species into Lake Jennings, San Diego River drain-
age, San Diego County, in October 1969 (the 1966 date of Shapavalov et al. [1981]

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 139

undoubtedly a misprint). It has since been placed in many southern California
reservoirs, the lower Colorado River, and has appeared in the Sacramento-San
Joaquin Delta (Taylor 1980; Young and Marsh 1991).

31. Ameiurus natalis (Lesueur). Yellow bullhead. This species is rare in south-
ern California, being first introduced in California in 1874, along with the black
and brown bullheads and white catfish (Smith 1896; Curtis 1949). We have records
from Evans Lake, Riverside County; and Santa Ana River, Orange County; and
another (UCLA) from Cochella Valley irrigation canals. Minckley (1973) noted
that they were widespread but rare in collections from Arizona, including the
Colorado River. The original plants of yellow and brown bullheads came from
the Schuylkill River, Pennsylvania (Neale 1915), as did channel and white catfish.
One or more species of bullheads also came from Lake Champlain, Vermont and
the Mississippi Valley (Shebley 1917a).

32. Ameiurus nebulosus (Lesueur). Brown bullhead. Definite records of brown
bullhead are from Zaca Lake in the Santa Ynez drainage near Buellton; lower
Owens River, Inyo County; Irvine Lake and the Santa Ana River in Orange
County; Evans Lake and Lake Hemet, Riverside County; and San Luis Rey River
just below Lake Henshaw, San Diego County. Brown bullheads were abundant
in the Owens River near Lone Pine by 1891 (Gilbert 1893). In August 1963 in
Lake Hemet, Riverside County numerous (but not counted) bullheads thought to
be brown turned out to be four black to one brown (L. K. Puckett, CFG LB).
These two species are often confused. Minckley (1973) expected them to occur
in the lower Colorado River but they had not actually been collected.

33. Ameiurus melas (Rafinesque). Black bullhead. Black bullheads are the
most common bullhead in the streams and rocky areas of reservoirs, and are the
basis for many unpublished reports of brown bullheads (see above). Minckley
(1973) reported black bullheads numerous but stunted in the lower Colorado
River.

34. Ameiurus catus (Linnaeus). White catfish. In 1933 white catfish were
noted from upper Otay Lake, Tia Juana (sic) River drainage, San Diego county
(CFG LB); these may have been channel catfishes, but A. catus was reported (as
Schuylkill catfish) introduced to Los Angeles and San Diego counties in 1874
(Smith, 1896). Specimens were “‘verified from San Diego County” by Curtis (1949)
and taken from the Los Angeles Aqueduct system in 1951 (SIO H51-337). In
1988 they were recorded in Pyramid and Castaic lakes, probably recent arrivals
with central California water (CFG LB).

35. Pylodictis olivaris (Rafinesque). Flathead catfish. Bottroffet al. (1969) and
Minckley (1973) reported the introduction of this species into the lower Colorado
River in Arizona in March 1962. It was first taken in California in the Highline
Canal, Imperial County, in 1968 and subsequently spread west into canals of the
Salton Sea area (Shapavalov et al. 1981).

Cobitidae

36. Misgurnus anguillicaudatus (Cantor). Oriental weatherfish. First discov-
ered in 1968 in Westminster flood control channel, Orange County (St. Amant
and Hoover 1969), oriental weatherfish survived at least through June 1979 (LACM
collections; Shapavalov et al. 1981). Courtenay (in litt., January 1990 to P. Moyle)
believed it remained in flood drains in Westminster.

140 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Atherinidae

37. Menidia beryllina (Cope). Inland silverside. This species was reported in
Lake Silverwood, Pyramid and Castaic lakes, and Skinner Reservoir in 1988
(CFG LP, FGH), and young-of-the-year and adults were taken in June 1992 in
Pyramid Lake (LACM 45698-1), indicating an established population. Inland
silversides have been in the Central Valley since 1967 (Moyle 1976a) and spread
rapidly (Shapavalov et al. 1981).

Fundulidae

38. Lucania parva (Baird and Girard). Rainwater killifish. McCoid and St.
Amant (1980) reported fish collected in Irvine Lake (1963 and 1964) and Arroyo
Seco Creek, tributary to Vail Lake (1976). Cyprinodon sp. was on a list of fishes
introduced to Vail Lake from Dexter, New Mexico in November 1948 (CFG
RRB); undoubtedly these were rainwater killifish as suspected by McCoid and St.
Amant (1980). In the early 1980s a population was established in the campus
lagoon near Goleta Point, University of California, Santa Barbara (Sam Sweet,
pers. comm.).

Rivulidae

39. Rivulus harti (Boulenger). Trinidad rivulus. St. Amant (1970) and Moyle
(1976b) report the existence of this fish in drains near the Salton Sea in the late
1960s. Thought to have disappeared after 1969 (Shapavalov et al. 1981), fish were
collected again in mid-1990 (CFG JAS).

' Poeciliidae

40. Gambusia affinis (Baird and Girard). Western mosquitofish. Mosquitofish
are found in almost all fresh and low salinity waters, and are continually being
reintroduced by county mosquito abatement districts. It was introduced to south-
ern California in December 1922 (John D. Lynch, in litt.) and was noted frequently
by field biologists by the late 1930s (CFG LB). Shapavalov et al. (1981) report
frequent California introductions of eastern mosquitofish, Gambusia holbrooki
Girard. John D. Lynch (pers. comm.) believes only western mosquitofish have
been placed in California waters, and all fish we have identified have been western
mosquitofish. Although implicated in the demise of native southwestern fishes
(Minckley 1973; Meffe 1985), including sticklebacks in the Los Angeles Basin
(Miller 1946b), Bell (1978) questioned this latter conclusion because both species
coexist in many other areas of southern California today. In captivity mosquitofish
feed much less on larval fishes than formerly believed (Hubbs 1991). The species
is less prevalent at cooler coastal localities north of Point Conception (Swift et
al. 1989) because it has slow growth and a low reproductive rate at 20°C or lower
(Vondracek et al. 1988).

41. Poecilia latipinna (Lesueur). Sailfin molly. This species has been estab-
lished in the Salton Sea, lower Colorado River, and environs since the 1950s
(Minckley 1981; Schoenherr 1979). A population existed in a coastal, tidal slough
in Oxnard from April 1977 (LACM 36402-3) to April 1983 and probably still
lives there. Fish were found in Ballona Marsh in July 1990 (David Soltz, pers.
comm.), and in the Santa Ana River near Prado reservoir in 1991 (Paul Winkel,

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 141

pers. comm.). This herbivorous, euryhaline species survives well in degraded
environments (Felley and Daniels 1992), and may disperse (or be transported by
man) to other coastal and inland localities.

42. Poecilia mexicana (Steindachner). Shortfin molly. This species is restrict-
ed to a small area south of Mecca near the Salton Sea, and to a few areas of the
lower Colorado River (Schoenherr 1979; Lau and Boehm 1991).

43. Poeciliopsis gracilis Heckel. Porthole livebearer. First collected in 1965
from a drain south of Mecca near the Salton Sea (Mearns 1975), this species
remains locally established (Schoenherr 1979; Courtenay et al. 1986).

44. Xiphophorus helleri Heckel. Green swordtail. This species was collected
from a drain in Westminster and near Mecca (St. Amant and Hoover 1969; Mearns
1975) in the late 1960s and early 1970s but disappeared soon thereafter (Courtenay
et al. 1986).

45. Xiphophorus maculatus (Gunther). Southern platyfish. This species was
present for a year or so in Westminster flood control drain in the late 1960s (St.
Amant and Hoover 1969) but has since disappeared (Courtenay et al. 1986).

46. Xiphophorus variatus (Meek). Variable platyfish. This fish was collected
at a few localities in Orange and Riverside counties in the late 1960s (St. Amant
and Hoover 1969; St. Amant and Sharp 1971). Courtenay et al. (1986) thought
they had disappeared, but a few were caught in the spring of 1991 in the vicinity
of the Salton Sea (Lau and Boehm 1991).

Moronidae

47. Morone saxatilis (Walbaum). Striped bass. The introduction of this well
known sport fish into northern California and the lower Colorado River is well
documented (Minckley 1973; Moyle 1976a; Baltz 1991). Introductions into the
Salton Sea in 1929 and 1930 were unsuccessful (Walker et al. 1961). Colorado
River fish, introduced in 1959, migrate from Lake Havasu down into the Imperial
Valley and the river delta in Mexico and back up through the New and Alamo
rivers (Minckley 1973, pers. comm.). Although natural reproduction occurred,
repeated plantings once were necessary (Minckley 1973), but the species is self-
sustaining today. From at least 1972 to 1982 small numbers of striped bass strayed
into Lake Cahuilla near Indio at the end of the Coachella Canal that originates
in the Colorado River (CFG LB).

Fish may stray southward in marine waters from central California. This has
not been definitely documented, and is confused by the unsuccessful introduction
of fish from Cambria to San Diego since 1903 (Shebley 1917b; Scofield and Bryant
1926; Horn etal. 1984). Since the early 1970s fish entered Pyramid and Silverwood
lakes via the California Aqueduct, and large individuals caught when dewatering
the aqueduct were put in Pyramid Lake also (CFG LB).

Hybrids between striped and white basses have been raised by fish farms in
Riverside and San Diego counties for several years to supply marketable food
fish. In August 1992, some of these hybrids were introduced into Irvine and
Anaheim lakes, Orange County, as sport fish and probably will be placed in other
waters as well. Both possible hybrids have separate common names; those reared
from striped bass eggs are palmetto bass and those from white bass eggs are
sunshine bass (Robins et al. 1991).

48. Morone chrysops (Rafinesque). White bass. This species was introduced

142 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

unsuccessfully into the lower Colorado River near Yuma in the late 1960s (Minck-
ley 1973; Shapavalov et al. 1981). White bass were introduced into Nacimiento
Reservoir, Salinas drainage (von Geldern 1966) rather than into south-central
coastal drainages as Moyle (1976b) indicated.

Percidae

49. Percina macrolepida Stevenson. Bigscale logperch. This logperch has been
established in the Central Valley of California since 1953 (Moyle 1976a). Its pelagic
larvae and adaptation to large turbid rivers (Simon and Kaskey 1992) facilitated
its transport southward to Silverwood, Pyramid, and Castaic lakes in 1988 (CFG
LP), and to Irvine Lake in July 1990 (LACM 45147-1, 45701-3). It is apparently
established in these lakes.

50. Perca flavescens (Mitchill). Yellow perch. This species was introduced
into Lake Cuyamaca in 1891 (Smith 1896; Curtis 1949) and into the lower Col-
orado River (Dill 1944; Minckley 1973) at an unknown date, both unsuccessfully.

51. Stizostedion vitreum (Mitchill). Walleye. Walleye were unsuccessfully in-
troduced into Puddingstone Reservoir, Los Angeles County (1959-1962), and San
Vicente (1959-1960), and El Capitan (1962-1963) reservoirs, San Diego County
(CFG LB); these fish grew and entered the sport catch, but did not reproduce
(Shapavalov et al. 1981).

Centrarchidae

52. Archoplites interruptus (Girard). Sacramento perch. One specimen was
collected in Ramer Lake, near the Alamo River, Imperial County in May 1956
(SIO 56-53). Introduced into Crowley Lake, Owens drainage in the late 1960s, it
now occurs downstream in Pleasant Valley and Hiawee reservoirs and in Little
Lake in the Owens Valley (Aceituno and Nicola 1976), and may be further dis-
tributed in southern California.

53. Micropterus salmoides (Lacepede). Largemouth bass. This species is wide-
spread in southern California, first arriving between 1888 and 1891 (Smith 1896;
Shebley 1917a; Glidden 1931). It was later introduced to the Colorado River
(Minckley 1973). These fish all came from the north central United States. In
1959 and 1960, fish came from Texas, and the southern subspecies, M. s. floridanus
(Lesueur), was brought in to many southern California waters (Shapavalov et al.
1981). Largemouth bass are established also in reservoirs on Santa Catalina Island,
Los Angeles County (LACM 42309-1).

54. Micropterus dolomieui Lacepede. Smallmouth bass. This species is much
less common than the largemouth in southern California. It is known from the
Santa Ynez River (Moyle 1976a) and a few other southern California reservoirs;
Morris Dam, San Gabriel River and Pyramid Lake, Los Angeles County; Lake
Hemet, San Diego County; and Lake Arrowhead and Big Bear Lake, San Ber-
nardino County (CFG LB). Smith (1896) reported smallmouth bass planted in
San Diego and Los Angeles counties in 1889 and 1894, respectively. They were
brought to lower Otay and Chimney lakes, San Diego County from Friant in
central California in 1933 (CFG LB). They can be locally abundant in the lower
Colorado River (Minckley 1981) where they were introduced in 1950 (Beland
1953).

55. Micropterus coosae Hubbs and Bailey. Redeye bass. Redeye bass were

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 143

introduced into the Santa Ana River, Riverside County, Santa Margarita River,
San Diego County and the Sisquoc River, Santa Barbara County in 1962 and
1963 and again in 1968 and 1969 (Moyle 1976a; Shapavalov et al. 1981). The
Santa Margarita fish survived at least into 1987, the Santa Ana fish have not been
recollected, and the status of the Sisquoc fish is not known.

56. Micropterus punctulatus (Rafinesque). Spotted bass. Several San Diego
County reservoirs and Lake Perris, Riverside County have established populations
of spotted bass dating from 1974 (Moyle 1976a). The stocks originated in Alabama
(CFG LB).

57. Lepomis cyanellus Rafinesque. Green sunfish. This sunfish is found in
many freshwater streams, ponds, and reservoirs in coastal southern California.
They are usually common and self-sustaining. It is also found throughout the
lower Colorado River (Minckley 1973). The first arrived in southern California
from Illinois in 1891 for placement in Lake Cuyamaca, San Diego County (Shebley
1917a).

58. Lepomis macrochirus Rafinesque. Bluegill. Bluegill are common to abun-
dant in many ponds and reservoirs in the area, including the lower Colorado
River. They may have arrived in 1891 with the green sunfish, but were not
definitely recorded until 1908 (Smith 1896; Curtis 1949). The original stocks were
from the northeastern United States, but from the late 1940s onward stocks from
Texas and Florida were planted as well (Miller 1952; Shapavalov et al. 1981;
CFG LB). Fish have been taken from reservoirs on Santa Catalina Island, Los
Angeles County also.

59. Lepomis gibbosus (Linnaeus). Pumpkinseed. This species is occasionally
found in lakes and ponds. It was first collected in southern California near Mecca,
Riverside County in 1939 after arriving about 25 years earlier (Dill et al. 1955).
Miller (1968) reported them in Lucerne Valley, San Bernardino County in 1940.
Minckley (1973) found most, if not all, records of pumpkinseed from the Colorado
River were based on redear. Pumpkinseed often hybridize with bluegills adding
to the confusion in identification.

60. Lepomis microlophus (Gunther). Redear. This species was first introduced
into the Headgate Rock Dam area of the lower Colorado River in 1948 (Beland
1953; California Department of Fish and Game 1973), and Minckley (1973)
reported them common there though not often collected. Redear were brought
into coastal southern California in 1954 (Shapavalov et al. 1959), and were in-
troduced into San Diego County reservoirs beginning in 1962 to prey on snails
known to be vectors of bass and sunfish parasites (CFG LB).

61. Lepomis gulosus (Cuvier). Warmouth. Fish placed in Lake Cuyamaca in
1891 did not survive (Smith 1896). This species has been abundant in the lower
Colorado River since being introduced in 1961 (Lanse 1965; Minckley 1973;
Moyle 1976b). It should be expected in reservoirs receiving water from both the
Colorado River and the Central Valley.

62. Pomoxis annularis Rafinesque. White crappie. In both the lower Colorado
River (Minckley 1973) and coastal southern California, this species is less common
than the black crappie, Pomoxis nigromaculatus. Both crappies arrived in Cali-
fornia in 1891 from Illinois and were placed in Lake Cuyamaca, San Diego County
(Smith 1896; Curtis 1949). These died out and other transplants in 1908 to
Ventura, Los Angeles, Riverside, and Orange counties (Vogelsang 1931) and in

144 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

1917 to San Diego County (Glidden 1931) survived and became widespread
(California Department of Fish and Game 1973). Some fish probably also rep-
resent immigrants from both central California and the Colorado River.

63. Pomoxis nigromaculatus (Lesueur). Black crappie. Most, if not all, of the
larger reservoirs hold this species, which, as noted above, arrived in southern
California in 1891 and later. In 1981 fish from the St. John’s River, Florida were
brought to Buena Vida Farms, Lakeview, Riverside County and the Chino Fish
and Wildlife Base in December 1981 (CFG LB) for distribution in southern
California.

Sciaenidae

64. Cynoscion xanthulus Jordan and Gilbert. Orangemouth corvina. This
species has been established in the Salton Sea since the 1950s (Walker et al. 1961).
In September 1966 fish were taken about three-fourths of a kilometer up the
Whitewater River from the Salton Sea (CFG LB). They also invade freshwater in
Mexico (Castro-Aguirre 1978); in March 1961, 37 fingerlings were introduced to
Puddingstone Reservoir and not recorded subsequently. Horn and Allen (1978)
erroneously list them from coastal southern California.

65. Bairdiella icistia Jordan and Gilbert. Bairdiella. One specimen of this
species was taken in the fresh water of the lower Whitewater River, with the
orangemouth corvina noted above. Otherwise biardiellas are restricted to the
saline waters of the Salton Sea (Walker et al. 1961), but invade fresh water in
Mexico (Castro-Aguirre 1978).

- Gobiidae

66. Acanthogobius flavimanus (Temminck and Schlegel). Yellowfin goby.
Haaker (1979) noted the first records of this species in southern California (Los
Angeles Harbor) and it has been subsequently taken in Newport Bay (Allen 1982)
and upstream in lower San Diego Creek, Ballona Marsh (LACM 39764-6), and
Mugu Lagoon (44205-1). It is rare or absent in smaller coastal estuaries: Malibu
Lagoon (Manion and Dillingham 1989), San Onofre, San Mateo, Las Pulgas, and
Santa Margarita lagoons, and Morro Bay.

67. Tridentiger trigonocephalus (Gill). Chameleon goby. Established in the
1960s in San Francisco Bay and Los Angeles Harbor (Haaker 1979), it was col-
lected in Pyramid Lake, Los Angeles County (LACM 45016-1) in October 1990.
Locally abundant in some rocky areas of the lake, it was also taken downstream
in Piru Creek on 25 June 1992 (LACM 45688-1). It presumably wiil move down-
stream throughout the Santa Clara River drainage. A record upstream invasion
of salt water into the Sacramento-San Joaquin delta enabled chameleon gobies
to be entrained with water diverted south (Kathy Heib, pers. comm.). Establish-
ment of the species in fresh water was predicted by Moyle (1976a).

Cichlidae

68. Oreochromis aurea (Steindachner). Blue tilapia. The blue tilapia has be-
come established in several areas of the lower Colorado River (Courtenay et al.
1986, 1991), and some have turned up in the Salton Sea area as well (Courtenay,
in litt.). In 1992 O. aurea have been collected from the Salton Sea, as well as fish

-_

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 145

that appear to be hybrids between O. aurea and O. mossambica (W. L. Minckley,
W. Courtenay, in litt.).

69. Oreochromis mossambica (Peters). Mozambique tilapia. This species be-
came established in drains of the lower Colorado River near Yuma in 1963, the
Salton Sea area in January 1964, and tidal channels of the Santa Ana, San Gabriel,
and Los Angeles rivers by 1974 (St. Amant 1966; Shapavalov et al. 1981). A few
unidentified tilapias taken from the Sepulveda Flood Control Basin on the Los
Angeles River and the Prado Flood Control Basin on the Santa Ana River are
believed to be this species. Courtenay et al. (1984, 1986, 1991) note that some
of these introductions may be of O. honorum Trewavas or of hybrids between O.
mossambica and O. honorum, the last sometimes treated as a subspecies of O.
urolepis Norman. The hybrid combination is called the red tilapia (Robins et al.
1991). Oreochromis mossambica largely has replaced T. zi/li in the Salton Sea
and possibly in coastal southern California as well, having been taken in upper
Newport Bay (LACM 44375-2). Tilapias obviously have been a major factor in
a decline of the desert pupfish in the Salton Sea area, but their impact on native
coastal estuarine species is not known (Horn 1988). By 1983 five pure species,
O. mossambica, O. honorum, O. aurea, T. zilli, and T. nilotica, and ten possible
hybrid combinations of these were tentatively identified in the lower Colorado
River area of California and Arizona (Barrett 1983; Lopez and Ulmer 1983).

70. Tilapia zilli (Gervais). Redbelly tilapia. Introductions of this species for
weed control (Salton Sea area) and mosquito abatement (Orange County drains)
led to established populations (Knaggs 1977; Moyle 1976a, b; Courtenay et al.
1984, 1986), and even to records from the open coast off Huntington Beach after
heavy winter rains in December 1974 (LACM 36203-1). The only other estuarine
collections are those of O. mossambica from Newport Bay noted above. Despite
their salinity tolerance, tilapias have invaded local estuaries only in highly mod-
ified channels where power plant effluent keeps the water warm. Tilapia zilli has
been partially and possibly completely replaced by other tilapias locally (Courtenay
et al. 1984, 1986, 1991).

Embiotocidae

71. Hysterocarpus traski Gibbons. Tule perch. This species was collected in
Lake Silverwood in 1988 or 1989 (CFG FGH, LP), and about twenty adults were
taken from widely scattered localities in Pyramid Lake in June 1992 (LACM
45701-7), indicating establishment there or continued immigration with imported
water.

Cottidae

72. Cottus asper Richardson. Interior prickly sculpin. Bell (1978) recorded
presence of this species in the Santa Clara River. They have remained common
in most of the drainage, but have never been taken in the Soledad Canyon section
upstream. A specimen of Cottus sp. was noted in June 1972 in the Gorman Canal,
tributary to Pyramid Lake (CFG LB), the first year this water began to flow into
the Santa Clara Basin. In 1987 they were taken in the lower Santa Ana River
(LACM 44384-1, 44385-1) and Irvine Lake (LACM 45146-1), Orange County,
and upstream in Big Bear Lake and its outlet Bear Creek, San Bernardino County

146 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

(Deinstadt et al. 1988). Prickly sculpin are known from Skinner, Castaic, Silver-
wood (including the west fork of the Mohave River downstream), Cachuma, and
Casitas reservoirs also (CFG LB). Casitas and Cachuma do not receive imported
water, and are in drainages originally inhabited by the native coastal form noted
above. The recency of these two collections in otherwise previously well collected
drainages indicates they are introduced. Mixing of coastal and interior forms in
southern California seems inevitable (Fig. 13).

Other Introductions

About twenty-five additional freshwater fishes have been placed in southern
California waters, caught once and never seen again. A specimen of Pimephales
notatus, bluntnose minnow, was taken in Victorville in a shipment of channel
catfish from Arkansas in 1968 (SIO-164). A specimen of Notropis atherinoides,
emerald shiner, was collected from a fish pond in Oak Glen in the summer of
1975 (LACM 35569-1). Several urban aquatic sites in Los Angeles received Asian
milfoil from Florida in late 1980; viable eggs on these plants hatched into bluefin
killifish, Lucania goodei, that survived several months in an outdoor pond (LACM
39884-1). Marine, estuarine, and anadromous species, both northern and south-
ern, were released into southern California waters with live bait discarded by
returning tuna boats in the 1930s and 1940s (Clark 1932, 1940). Additional species
documented include twenty-eight marine and estuarine fishes introduced into the
Salton Sea (Walker et al. 1961), 16 freshwater fishes taken from bait dealers along
the lower Colorado River (Miller 1952), and aquarium species captured once near
fish farms or in warm springs (St. Amant 1966; St. Amant and Hoover 1969;
Naiman and Pister 1974; Shapavalov et al. 1981). These bring the total number
of introductions to at least 100 and probably more for southern California, not
including local transfers of native species and the recent plantings of hybrids.

Discussion

The native freshwater fish fauna of coastal drainages of southern California
comprises a meagre southwestern extreme of an already depauperate temperate
freshwater fish fauna in western North America. South of the Carmel River to
Point Conception, two species almost continuously inhabit freshwater streams,
namely Oncorhynchus mykiss and Cottus aleuticus. Cottus aleuticus does not quite
range southward to Point Conception. A third, Lampetra tridentata, may be
continuous also, but is rarely collected and data are not complete enough to
establish a pattern. Lampetra tridentata extends southward to the Los Angeles
Basin, and Oncorhynchus mykiss ranges to northern Baja California in drainages
of the outer Pacific Coast. Cottus asper and Gasterosteus aculeatus are continuously
distributed in lower gradient areas, but are absent from most of the steeper streams.
Cottus asper ranges southward to the mouth of the Ventura River, and Gasterosteus
aculeatus occurs south to northern Baja California. All of these species, except C.
aleuticus, are polytypic in California and all but C. aleuticus and C. asper have
one or more California populations that have special conservation status. None
is continuously distributed south of Point Conception, and C. asper may only
accidentally stray south of this point. Fish collections before 1950 were not nu-
merous and it is difficult to separate the effects of natural causes from those due

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 147

to human alteration of the environment since the mid-1700s. Holocene sediments
hold remains of some of this fauna and may help document the historical distri-
bution of these taxa (Swift 1989).

Only one area of coastal southern California, the Los Angeles Basin, contains
endemic freshwater fishes, namely Rhinichthys osculus, Gila orcutti, Catostomus
santaanae, Lampetra cf. pacifica, and one or two taxa of threespine stickleback.
These taxa are distributed in one or more of the four physiographically defined
freshwater fish communities that can be defined from the region, namely 1) low-
land (and low gradient), 2) upland (and high gradient), 3) anadromous, and 4)
coastal lagoon. These correspond somewhat to those formulated by Hopkirk
(1974) and Moyle and Nichols (1974) in the Clear Lake region and the southern
Central Valley, respectively, of California. These latter two classifications also
include a foothill fish community defined as an intermediate one between the
upland and lowland communities. This foothill component is not well developed
in southern California; steep mountains rather abruptly meet the relatively flat
alluvial coastal plains leaving little or no foothill habitat to intervene between the
upland trout-dace habitat and the lowland stickleback-chub habitat. In steep,
coastal streams, upland (high gradient) habitat extends directly to the sea without
either lowland or foothill habitat intervening.

Upland habitat is dominated by rainbow trout (and/or steelhead) and speckled
dace, with trout extending farther upstream than the dace. In some larger rivers
(Los Angeles, San Gabriel, Santa Ana), Santa Ana suckers also inhabit the upland
zone. Arroyo chubs are infrequent in this zone and may be mostly introduced,
maintaining themselves only in the scarce low gradient areas or in artificial im-
poundments. Today, steelhead and Pacific lamprey use this zone for spawning in
mid-winter, and probably also utilized the lowlands formerly. Farther north Cottus
aleuticus inhabits high gradient streams of upland character where they flow
directly into the ocean without intervening foothills or lowlands.

The lowland habitat was occupied primarily by Pacific brook lamprey, arroyo
chub, and unarmored threespine stickleback. Arroyo chubs and sticklebacks were
particularly widespread (Culver and Hubbs 1917; George 1927). Santa Ana suck-
ers inhabited large streams in the lowlands, and speckled dace occurred in a few,
isolated, often spring-fed, foothill localities. Steelhead and Pacific lamprey tra-
versed this area to spawn. Lowland and upland streams and coastal lagoons served
as nursery areas for juveniles of these migratory fishes before they returned to the
sea. Moore (1980) observed steelhead spawning in the lowlands of the Ventura
River. Lowland spawning was probably more prevalent before dams flooded the
major rivers where they meet the lowlands. These areas of alluvial gravels and
strong spring flow undoubtedly were significant for the spawning of both anad-
romous species. In lowland streams north and south of the Los Angeles Basin,
partially armored sticklebacks replaced the unarmored forms, and Pacific brook
lampreys were absent. North of Point Conception, prickly sculpins occupy lowland
areas as well.

The foothill zone in southern California consists of large alluvial fans at the
base of steep mountains. True foothills are small isolated hills between the moun-
tains and the coast. During the dry season, streams might dry up in the alluvial
fan areas, and return to the surface farther downstream (Miller 1961; Pourade
1968). Originally, during hot weather, evapotranspiration by the extensive riparian

148 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

gallery forests caused streams to dry up during the day and begin flowing again
in the evening, a phenomenon often noted by early explorers (Pourade 1968). It
seldom occurs today in the lowlands since artificial lowering of the water table
led to the loss of much of the gallery forests. The few, widely scattered, apparently
relict, populations of Rhinichthys osculus on the Los Angeles Basin indicate that
the foothill habitat was attenuated and almost non-existent.

Coastal streams often were perennial in the mountains, but disappeared un-
derground in the large alluvial fans at the bases of the mountains, and reemerged
at places like Whittier Narrows (San Gabriel River), the Riverside-Mt. Roubidoux
area (Santa Ana River), and along the north side of the present day Hollywood
Hills (Los Angeles River) (Miller 1961). Thus, coastal and inland populations of
freshwater species were probably often isolated from each other seasonally. South
of the San Luis Rey drainage, the mountains are farther from the coast, rainfall
is less, and native freshwater fishes consisted of a few widely scattered populations
of threespine sticklebacks, rainbow trout, and steelhead (Cooper 1874; Eigenmann
and Eigenmann 1890; Hubbs 1946; Miller and Hubbs 1969).

Farther north in Cuyama River tributaries, Huasna River and Alamo Creek,
Rhinichthys osculus, Hesperoleucas symmetricus, Gila orcutti, and unplated Gas-
terosteus aculeatus are considered introduced (Miller 1945a, c, 1968; Miller and
Hubbs 1969; Greenfield and Greenfield 1972). Gila orcutti is an obvious intro-
duction, but the other three are not as certain. These fish were restricted to upland
areas of the Cuyama River and could have been missed by early collectors. Low
divides separate the upper Huasna drainage from the Salinas system where these
species occur. Rhinichthys particularly can fluctuate widely in numbers from year
to year, and 1s restricted to spring-fed rocky and riffle areas that may not be in
good supply in the Cuyama River. The unplated stickleback were lost with the
introduction of partially plated fish (Miller and Hubbs 1969), and Hesperoleucas
and Rhinichthys still occur in the drainage. These two species should be studied
further to clarify their relationships.

The coastal lagoon habitat, largely absent in southern California today, contains
a mixture of lowland freshwater, estuarine, and a few marine species dependent
on low salinity during part of’their life cycle. Five estuarine species, California
killifish, tidewater goby, longjaw mudsucker, cheekspot goby, arrow goby (Swift
et al. 1989; Allen 1991) would live almost exclusively in such habitats, although
the larvae of the latter three occur along the open coast. The lagoons provided
nursery areas for juvenile steelhead and, farther north, coastrange and prickly
sculpins. There are three groups of estuarine fishes: 1) widespread tropical species
reaching their extreme northern extent, 2) northern temperate species reaching
their southern limit, and 3) a group endemic to this area, often called the Cali-
fornian province of authors (Ricketts et al. 1985; Horn and Allen 1978). Cali-
fornian species are typically distributed between Point Conception on the north
and Magdalena Bay, outer Baja California on the south, often with an isolated
disjunct population in the northern Gulf of California. Two tropical species occur
inland only in the Colorado River, Elops affinis and Eleotris picta. Five of the
tropical coastal species are rare or extirpated in California, namely Carcharhinus
leucas, Hyporhamphus rosae, Cynoscion parvipinnis, Scomberomorus concolor,
and Gobionellus sagittula. Some of these are listed as coastal but non-bay inhab-
iting by Horn and Allen (1978), but are so scarce now as to be undeterminable.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 149

Definite bay records are known for Gobionellus sagittula (as G. longicauda) and
Cynoscion parvipinnis, and strongly suspected for Carcharhinus leucas, Eucinos-
tomus argenteus, Oligoplites saurus, Chaetodipterus zonatus, Myrophis vafer,
Trichiurus nitens, Mugil curema, and Bagre panamensis. Other tropical species,
rare in coastal southern California, fall in this same category, but are not as
exclusively associated with low salinity water, such as Galeocerdo cuveri, Poly-
dactylus opercularis, and P. approximans. Some of these species were caught for
live bait by tuna boats from the 1930s to the 1950s (Alverson and Shimada 1957).
Clarke (1932, 1940) noted that both northern and southern estuarine species were
inadvertently introduced into southern California coastal waters by these vessels.
Four tropical species still occur in southern California, Anchoa delicatissima, A.
compressa, Syngnathus auliscus, and Paralichthys californicus. Although from
tropical genera, three of these (excluding S. auliscus) are Californian in distri-
bution. The tropical species represent a combination of: extirpation of taxa re-
stricted to brackish waters; occasional colonization by taxa with more marine life
stages but dependent on warmer waters that infrequently invade southern Cali-
fornia; introductions of bait species by tuna vessels; and taxa that have contin-
uously inhabited the coastal estuaries of southern California.

Eight temperate estuarine species are all common, except Eucyclogobius new-
berryi, which has been rapidly disappearing. Some of the temperate species, like
Atherinops affinis and Fundulus parvipinnis, are probably much more common
than in the past because of the changed character of California estuaries (Lane
1977a; Swift et al. 1989) particularly in southern California. The California en-
demic, Eucyclogobius newberryi, only survives in a few coastal lagoons and river
mouths in southern California that are at least partially in a natural state. Some
of these localities went from virtual disappearance in February 1991 to a tentative
reprieve for another year due to heavy rains in March 1991.

The history of droughts and the aridity of the spectacular deserts of southeastern
California have masked awareness of the relatively wet conditions that existed in
much of coastal southern California until the arrival of Europeans. The high
mountain ranges stopped considerable moisture that accumulated in deep sedi-
mentary basins fronting the coast. Early observers and paleontological records
document extensive surface water, springs, fresh and brackish water marshes, and
artesian springs in the lowlands and near the coast in many areas (Willett 1941;
Emery et al. 1957; Gorsline 1967; Miller 1961; Warme 1971; Gordon 1985; Swift
et al. 1989). Gorsline (1967) noted:

‘“‘The southern California rivers are seasonal streams and in their natural state
are sealed off at their mouths by beach deposits during the dry season. A few
southern California estuaries (Mugu Lagoon, Newport Bay) are abandoned
mouths of the major streams which have been closed off by longshore drift
and then opened to the sea by artificial channels.”

The original coastal lagoons were largely brackish or fresh and were cooler in
summer and warmer in winter than the nearby ocean. The need for harbors during
the last 150 years led to these habitats being replaced by much smaller estuaries
with regular tidal action artificially maintained. Temperature and salinity now
closely parallel that of the ocean. These systems harbor a much larger and more
marine fauna (Lane 1977b; Horn and Allen 1976; Zedler 1984). The original

150 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

coastal lagoon habitat is important to preserve as a unique physiographic feature
indispensable for certain life stages of many aquatic and terrestrial organisms
(Barnes 1991), but only rarely are aquatic taxa exclusively restricted to them (i.e.,
tidewater goby, Swift et al. 1989).

The native freshwater fishes of the lower Colorado River drainage in California
have been extirpated or very rare since the 1950s. Most are riverine forms with
at least small populations farther upstream in tributaries outside California. Two
of these, Oncorhynchus clarki and Rhinichthys osculus, are at best strays in the
lower river. Poeciliopsis occidentalis exists only by virtue of great effort to manage
the few remaining populations in Arizona and Mexico, and may not have lived
along the main river in California. Cyprinodon macularius still exists in many
small populations around the Salton Sea; with increasing salinity it may be the
only fish to survive (Lau and Boehm 1991), a situation that probably has repeated
itself many times in the Salton Sea area for thousands of years (Miller 1946a;
Norris and Webb 1990). Lower Colorado River freshwater fishes invaded the
Salton Sea with river floodwaters in 1904-1907 (Gobalet 1992), a phenomenon
that occurred six times (on a smaller scale) from 1840 to 1891 (Carpelan 1961).
Since 1907 only the controlled release of irrigation water comes into the Salton
Sea area. Farther north in the Mohave, Armagosa, and Owens drainages, native
fishes only occurred in the lowland areas originally flooded by Pleistocene lakes;
many tributaries at higher elevations with perennial water originally lacked native
fish as far as is known (Miller 1948). The efforts to protect the native fishes of
these areas are discussed by Minckley and Rinne (1991) and in Minckley and
Deacon (1991).

The accelerating rate of habitat reduction, partially drought related, means all
the native coastal freshwater fishes require protection to prevent their disappear-
ance. The reductions in habitat make it less likely that native fishes will be able
to expand their ranges with the return of wetter conditions. Low water concentrates
native fishes in the larger pools favored by introduced centrarchids, particularly
green sunfish. Local residents build small dams to store this water for supression
of potential fires, and stock these ponds with predatory warm water fishes. Farther
north low water levels in streams made stickleback vulnerable to predation by
introduced bullfrogs (Dan Holland, pers. comm.). Introduced clawed frogs prey
on stickleback in southern California (CFG UTSRT). Brown trout predation may
explain the scarcity of native fishes today (including trout?) in the mountainous
portions of the Santa Ana River and tributaries. They have been implicated in
the reduction of native fishes in northern California (Moyle 1976a) and elsewhere
(Taylor et al. 1984).

In some areas, secondary water sources, usually sewer treatment plant effluent,
provide more surface water than was available historically. Although potentially
useful to preserve riparian habitats, this water is often detrimental. It is much
warmer than natural waters emerging from underground sources. Its high nutrient
load encourages a different suite of species and can put the native fauna (and flora)
at a competitive disadvantage (Morris 1991). These conditions favor introduced
aquatic vertebrates like red shiners, grass carp, goldfish, common carp, sailfin
mollies, tilapia, and clawed frogs. These are all known to impact native species,
and the severity of impact seems related to degree of habitat alteration (Ross
1991). Several other introduced species known to have been established in the

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 151

late 1970s and early 1980s for a few years or more may still exist, but have not
been reassessed recently (Courtenay et al. 1984, 1986, 1991).

The established, successful introduced stream fishes in coastal southern Cali-
fornia comprise an assemblage of five species from the central United States
already known for survival in small plains streams, often turbid and fluctuating
widely in flow. They are both pioneer and colonizing species elsewhere (Capone
and Kushlan 1991; Fausch and Bramblett 1991; Meador and Matthews 1991).
These species thrive in similar conditions in southern California, and include
fathead minnows, black bullheads, mosquitofish, green sunfish (Moyle 1976a, b)
and, recently, the red shiner. All typically become self-sustaining. Mosquitofish
are actively distributed by local mosquito abatement districts, despite early evi-
dence that native sticklebacks and pupfishes effectively controlled mosquitos (Sco-
field 1915; Kennedy 1916; Hubbs 1919). In addition, sailfin mollies and tilapias,
long established in the lower Colorado River and Salton Sea, turn up in a few
more coastal drainage localities each year.

The community of reservoir fishes is much more dependent on continual in-
troductions and highly modified (impounded) habitat for survival (Minckley 1973;
Moyle 1976a, b). This assemblage of introduced species characterizes ponds, lakes,
and reservoirs, namely threadfin shad, golden shiner, goldfish, carp, channel cat-
fish, mosquitofish, bluegill, redear, pumpkinseed, black crappie, and largemouth
bass. Most of these species would not be self-sustaining in the natural streams of
southern California. Since the early 1970s, additional native and introduced fishes
of the Central Valley have been carried into southern California reservoirs with
imported water. The following are known or suspected to be established in one
or more of these reservoirs: inland silverside, white catfish, inland prickly sculpin,
tule perch, striped bass, bigscale logperch, and chameleon goby. Most are restricted
to lake and reservoir environments. Some, like the inland prickly sculpin and
chameleon goby (Santa Clara River and some tributaries) and blackfish (lowland
Santa Ana River), are established in outlet streams. Earlier efforts to prevent a
similar invasion of carp.and goldfish from the Colorado River into the San Diego
reservoir system in the early 1900s were unsuccessful (CFG LB), and the prognosis
is no better now. For sculpins, sticklebacks, minnows, and suckers, the possibility
exists of introducing closely related taxa and mixing natural gene pools. This is
in addition to the usual problems of competition and predation. Non-native
genetic additions are almost always detrimental to the integrity of the native stock
(Hindar et al. 1991), as well as being undesirable for a variety of other ecological
and economic reasons (Allendorf 1991). Such genetic additions are justifiable only
when no alternative exists to maintain genetic diversity (Echelle 1991). A recent
trend, introducing sterile hybrids among the grass carps, tilapias, and temperate
basses, is still being evaluated. Such fish theoretically will not become seif-sus-
taining or reproductive, but have many of the same effects attributed to the
parental species while living.

As expected from the mixture of flowing and impounded areas on the lower
Colorado River bordering California, it and the larger water distribution canals
have an introduced fish fauna that is a mixture of the stream and reservoir species,
with a few taxa restricted to the Colorado River area (Schoenherr 1979; Minckley
1981; Lau and Boehm 1991). In addition the cool, hypolimnic water released
from some of the dams creates thermal conditions that segregate the species. Cool

152 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

sections are dominated by rainbow trout and striped bass. Farther downstream
carp, red shiners, channel catfish, green sunfish, bluegill, green sunfish, and black
bullhead predominate. Only in the warmest lower river are tilapias and sailfin
mollies abundant. As pointed out by Minckley (1981), the instability of the river
temperature and water levels means that the species live in transitory habitats
where “‘populations explode, stunt, then stabilize and often disappear.”

The irrigation canals in and around the Salton Sea are dominated by tilapias,
mosquitofish, porthole livebearer, shortfin, and sailfin mollies, and red shiners
(Schoenherr 1979, 1988; Lau and Boehm 1991). Several other species occur in
small numbers, including the only native species in these habitats, the desert
pupfish, Cyprinodon macularius. Usually most of these species will occur at any
given locality, but in areas of natural or artificially induced thermal gradients,
considerable species segregation can occur. Individual species often segregate from
each other linearly along a canal (Schoenherr 1979). Such conditions allow the
desert pupfish to survive by taking advantage of such scattered optimal micro-
habitats. Only the tilapias and sailfin mollies occur extensively in the main Salton
Sea, along with the few marine and estuarine species acclimated there (Walker et
al. 1961; see above).

Introduced species from eastern North America are usually of mixed genetic
stocks with more than one species or subspecies having been stocked in California
(Miller and Alcorn 1946; Miller 1952; Shapavalov et al. 1959, 1981). Moyle
(1976b) noted two eras of introductions into California, namely 1871-1890 and
1960-1976. Superimposed on these are three periods representing changing sources
for stocks of the fishes involved. Early introductions came from the northeastern
United States, including the Great Lakes and upper Mississippi Valley. In the
1940s and 1950s stocks of the same species originated in Texas and arrived via
the Dexter National Fish Hatchery, New Mexico. In the late 1950s and 1960s
stocks from Florida (or other southern localities) were brought in. The species
known or suspected of being polytypic are Notemigonus crysoleucas, Micropterus
salmoides, M. punctulatus, Lepomis macreochirus, Ictalurus punctatus, Pomoxis
nigromaculatus, and Morone saxatilis (Hubbs and Lagler 1964; Miller 1952; Swift
et al. 1986; Page and Burr 1991). The sources for several other species are less
certain but may be equally diverse, particularly for Ameiurus nebulosus, A. natalis,
A. melas, Gambusia affinis, Micropterus dolomieui, and Pimephales promelas.
Cultured varieties also complicate the picture, and Minckley (1973) contended
that distinctive forms will be swamped out by the established one. Pelzman (1980)
documented the mixing of genes of northern and southern largemouth bass in
California reservoirs.

Conservation Recommendations

As noted above, the native fish fauna of the greater Colorado River drainage
covered in this paper has been threatened, endangered or extirpated for a long
time. With a few exceptions the conservation of remaining populations primarily
or exclusively involves stocks outside southern California (Minckley and Deacon
1991).

Today, natural habitats for the freshwater fishes of coastal southern California
exist in hilly or mountainous headwater areas and in a few coastal localities that

_

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 153

have remained protected. The broad lowland areas between are highly modified
and largely uninhabitable for resident species and those that migrate between the
headwaters and the coast. Thus the priorities for the preservation of the native
fauna are, 1) protection of the remaining coastal and interior habitats still con-
taining elements of the native fauna, and 2) restoration and/or rehabilitation of
some portion of the now unsuitable intervening areas. These objectives are similar
to those proposed for the recovery of the federally endangered unarmored three-
spine stickleback (U.S. Fish and Wildlife Service 1985). To succeed the whole
ecosystem must be considered and a variety of natural stocks must be preserved
to re-establish populations (Nehlsen et al. 1991).

The areas between the mountains and the coast suffer primarily from extreme
channel modification and degraded water quality. Many are completely concrete
lined. In addition, introduced species dominate these sites. Man-made structures
encroach so closely on the water courses that they represent a major obstacle to
returning the areas to more natural conditions. In other areas, particularly flood
control basins, semi-natural habitat exists. Often the stream bottom could not be
covered over with concrete since groundwater upwelling had to be accommodated.
Maintenance of stable surface flow and improvement of water quality would return
the habitat to conditions that could support native fishes. However, non-native
fishes also have to be controlled or removed for the native fishes to maintain
themselves. :

The habitats critical for preserving the native freshwater fish fauna of coastal
southern California fall into the same four categories noted above as habitat types.
One or a few localities are critical in the Los Angeles Basin for the three endemic
cypriniform fishes. A mutually exclusive five to eight streams south of Morro Bay
to Malibu Creek still hold anadromous steelhead and lamprey. About twenty
coastal lagoons and marshes contain populations of Eucyclogobius newberryi.
Several additional freshwater and estuarine localities formerly held native fishes
and must be improved to restore populations of the native fish involved.

Recently, the remaining native species endemic to the Los Angeles Basin existed
only in the Big Tujunga Creek drainage (Los Angeles River drainage), the com-
bined east, west and north forks of the San Gabriel River above San Gabriel
Reservoir, and widely scattered and isolated portions of the Santa Ana River
drainage. The Los Angeles and Santa Ana river populations of these species are
restricted in distribution and rare where they occur. Only the San Gabriel system
appears large, stable, and unimpacted enough to maintain the three native species,
Gila orcutti, Rhinichthys osculus, and Catostomus santaanae (and possibly native
Oncorhynchus mykiss). Conditions in the other two systems have to be improved
so the native stocks can increase in numbers, and in the Santa Ana drainage this
may require the removal of brown trout. Gila orcutti also naturally still occurs in
parts of the San Juan Creek and Santa Margarita River drainages farther south.
However, these populations are also restricted in distribution and need formal
protection (Moyle et al. 1989). Particularly San Juan Creek and its tributary
Trabuco Creek also hold one of two extant native populations of partially armored
sticklebacks south of the Los Angeles Basin in the United States, the other in the
South Fork of the San Jacinto River below Lake Hemet. Only one other population
is known today from northern Baja California.

The two anadromous species, Lampetra tridentata and Oncorhynchus mykiss,

154 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

occur in several drainages where relatively natural habitat exists continuously
from ocean to headwaters, noted above. Their maintenance and increase will
depend on restoration of lowland areas, both as migratory routes to spawning
areas and as coastal lagoons for nursery grounds, the latter probably important
for steelhead only. Dams and debris basins block migration and obliterate large
areas that were undoubtedly used for spawning in the past. Even if access is
provided to headwaters above such impoundments, they may not provide as
much habitat as expected unless adequate beds of spawning gravel are available.
Intermittent streams can be important for spawning habitat in spring, and lower
reaches, including coastal lagoons, equally so for summer and fall habitation after
the smaller headwater areas lose adequate flow (Erman and Leidy 1975). Such
transitory habitat should be relatively more important in southern California and
is often not included in fish habitat listings that concentrate on permanent wa-
tercourses. Southern California populations may be particularly important as
genetic stocks that are better adapted to warmer water conditions than more
northerly populations. The occurrence of adult Pacific lamprey at the mouth of
the Santa Ana River in March 1991 is an encouraging sign that stocks are present
to colonize Los Angeles Basin streams that recover enough for these fish to com-
plete their life cycle.

The heavily impacted lowland areas where streams are cement-lined channels
must be improved to assure continued survival of native fishes. Two taxa have
been extirpated from Los Angeles Basin streams, namely Lampetra cf. pacifica
and Gasterosteus aculeatus williamsoni. Both taxa exist elsewhere and reintro-
duction is possible. A high conservation priority must be the existing habitat of
the three taxa of unplated threespine sticklebacks, the federally endangered form
in the upper Santa Clara River, the Shay Creek stickleback in the Baldwin Lake
area, and the San Antonio Creek stickleback. Both the maintenance of the existing
populations and designation of areas suitable for their expansion are of highest
priority. Unplated stickleback and Pacific brook lamprey formerly occurred in
lowlands of the Sepulveda Basin (Los Angeles River) system, of the Whittier
Narrows area (San Gabriel River), and the middle Santa Ana River from Mt.
Rubidoux (City of Riverside) downstream to a few kilometers below Prado Flood
Control Basin. Gila orcutti and Catostomus santaanae still occur in a few areas
on the Santa Ana and Los Angeles rivers. These areas have artificially permanent
water flow, albeit of low quality water, and are the best candidates for restoration
for management of native fishes. They are also the intermediate areas needed for
anadromous fishes and could be managed as spawning areas as well.

No brackish marshes or coastal lagoons exist in the conditions that were present
at first European contact. A few small systems have remained relatively unchanged
by human activity, namely several small lagoons in San Luis Obispo and western
Santa Barbara counties, Morro Bay, the mouths of the Ventura and Santa Clara
rivers, Mugu Lagoon, and four lagoons on Camp Pendleton. The two largest of
these, Santa Margarita Lagoon (on Camp Pendleton) and Morro Bay, are partic-
ularly important as the only two remaining with a full complement of bay gobies
(Swift et al. 1989). Most of these have artificially maintained entrances that make
them more like bays and estuaries than the coastal lagoons they once were. Having
irreparably lost most of the larger systems, it is important to maintain all that
remain, only two of which are large. The tidewater goby is the only coastal fish

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 155

in jeopardy, but a whole suite of other threatened organisms requires this habitat
for survival (Smith et al. 1989, Barnes 1991).

The current drought accelerated the reduction of natural habitat for native fishes
previously due to increases in human population, demand for water, and a general
lack of appreciation for scarce aquatic habitats in the southwestern United States
(Moyle and Williams 1990; Moyle and Ellison 1991). During the last surge in
human population after World War II, all the species of the lower Colorado River
drainage in California became threatened, endangered or extirpated (Minckley
1973). At this same time the large flood control projects on the Los Angeles basin
were completed (Van Wormer 1991), and about 40% of the freshwater fishes of
the Los Angeles Basin were extirpated. As the originally well-watered habitat of
coastal southern California has become artificially desertified, the fishes became
subjected to many of the same impacts affecting the desert regions (Minckley and
Deacon 1991). Today all of the remaining coastal species are, to some degree, in
jeopardy for long term survival as wild populations. As noted above, a variety of
threats impinge on the habitat, and a unified effort is needed to maintain the
integrity of the native aquatic systems and the riparian habitat associated with
them to allow these fishes, and a host of other riparian dependent species, to
survive (Nehlsen et al. 1991; Gregory et al. 1991; Johnson 1991; Moyle and Leidy
1992).

To organize the effort to save the freshwater fishes of California, Moyle and
Ellison (1991) classified the aquatic habitats of California, and defined criteria
important for the survival of native fish populations, namely Aquatic Diversity
Management Areas (ADMA). One hundred ADMAs are proposed for the southern
California area covered here. They are heavily biased towards sites where native
fishes still occur, or sites perceived to be recoverable for native fishes. The pro-
tection of these habitats must be incorporated into long term management plans
for the land areas involved. If the distinctive freshwater fish fauna of southern
California is to survive,.protection must be implemented immediately.

Acknowledgments

The Natural History Museum of Los Angeles County has supported work on
southern California freshwater fishes for many years. Additional support came
from the California Department of Fish and Game, contracts AB-23 (1975) and
FG7455 (1989), facilitated by Steve Nicola and Betsy Bolster, respectively. Terry
Garrett largely set up the HABIT computerized data base program at LACM.
California Department of Fish and Game biologists provided specimens and
access to department records, particularly James St. Amant, Frank Hoover, Sho-
ken Sasaki, Chuck Marshall, Karen Worcester (and her predecessor, Paul Chap-
pell), and Linda Pardy. United States Forest Service biologists provided data on
their respective forests: Helen Hamilton, Cleveland National Forest, Steve Loe
and Rose Robinson, San Bernardino National Forest, William Brown, Angeles
National Forest, and Maeton Freel, Los Padres National Forest. Other biologists
have contributed records and information over the years, namely Michael Bell,
Jonathan Baskin, Willis Evans, Dave Drake, Peter Moyle, James Diana, Alan
Wells, the late Carl L. Hubbs, Robert R. Miller, David Greenfield, Stephen Ross,
Mark Capelli, and Thomas Hablett. In addition the following persons have helped
document this fauna: L. Allen, R. Lavenberg, R. Thompson Swift, S. Manion, J.

156 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Dillingham, R. Feldmuth, G. Arita, M. Capelli, C. B. Crabtree, J. Johnson, A.
Naydol, S. Buck, J. Wolfe, B. Harper, P. Rice, J. Seigel, S. Goldberg, W. Tippets,
J. Malcolm, D. and R. Soltz, J. Irwin, J. Edmondson, R. Brownell, M. Horn, D.
Woefel, T. Taylor, A. Schoenherr, N. Villa, R. Feeney, the late W. I. Follett, M.
Love, G. and K. Marsh, M. Georgiu, E. Lichwardt, M. Fuller, A. Horne, J. Roth,
G. Kerns, C. Nagano, P. Winkel, W. Sinnen, D. Kresja, H. Fierstine, M. White,
R. Nakamura, S. Sweet, K. Howe, S. Imsand, T. Mack, and W. Mongey.

Museum collections were examined courtesy of: Reeve M. Bailey, Robert R.
Miller, and Douglas Nelson (UMMZ), Donald Stewart and Hyman Marx (FMNH),
William N. Eschmeyer, Tomio Iwamoto, Lillian Dempster, and Pearl Sonoda
(CAS), Richard Rosenblatt and the late Carl L. Hubbs (SIO), Richard Vari and
Susan Jewett (USNM), Karsten Hartel (MCZ), Boyd W. Walker, Don Buth, and
C. Ben Crabtree (UCLA), David Greenfield (formerly CSUF) and Michael Horn
(CSUP), David Soltz and Richard Bray (CSULB), and Eric Hochberg and Kathryn
Rindlaub (SBMNBH).

Earlier versions of the manuscript benefitted from the comments of Donald
Buth, Robert R. Miller, Michael A. Bell, Susan Ellis, W. L. Minckley, Willis A.
Evans, Richard Coker, Peter Moyle, Michael Horn, and Robert N. Lea.

Richard Meier and John De Leon provided photographic reproductions of the
maps which were largely drafted by the first author.

Partial funding for publication provided by California-Nevada Chapter of the
American Fisheries Society and the California Department of Fish and Game.

Literature Cited

Aceituno, M. E., and S. J. Nicola. 1976. Distribution and status of the Sacramento perch, Archoplites
interruptus (Girard), in California. Calif. Fish Game, 62:246-254.

Allen, L.G. 1982. Seasonal abundance, composition, and productivity of the littoral fish assemblage

in upper Newport Bay, California. U.S. Nat. Mar. Fish. Ser., Fish. Bull., 80:769-790.

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. S. Calif. Acad. Sci., 87:19-30.

Allen, M. J. 1991. Beam-trawl survey of bay and nearshore fishes of the soft-bottom habitat of
southern California in 1989. Calif. Coop. Oceanic Fish. Invest. Rept., 32:112-127.

Allendorf, F. W. 1991. Ecological and genetic effects of fish introductions: synthesis and recom-
mendations. Canad. J. Fish. Aquat. Sci., 48(Suppl. 1):178-181. ;

Alverson, F. G., and B. M. Shimada. 1957. A study of the eastern Pacific fishery for tuna baitfishes,
with particular reference to the anchoveta (Cetengraulis mysticetus). Inter-Amer. Trop. Tuna
Comm. Bull., 2:25-61.

Baltz, D. M. 1991. Introduced fishes in marine systems and inland seas. Biol. Conser., 56:151-177.

Barclay, L. 1975. Fishery survey of six coastal streams and the Salinas River drainage, San Luis
Obispo County. Contract, Calif. Dept. Fish and Game, 32 pp.

Barlow, G. W. 1963. Species structure of the gobiid fish Gillichthys mirabilis from coastal sloughs
of the eastern Pacific. Pac. Sci., 17:47-72.

Barnes, R.S. K. 1991. Dilemmas in the theory and practice of biological conservation as exemplified
by British coastal lagoons. Biol. Conser., 55:315-328.

Barrett, P. J. 1983. Systematics of fishes of the genus Tilapia (Perciformes; Cichlidae) in the lower
Colorado River basin. MS thesis, Arizona State University, Tempe, Arizona.

Baumgartner, J. V., and M. A. Bell. 1984. Lateral plate morph variation in California populations
of the threespine stickleback, Gasterosteus aculeatus. Evolution, 38:665-674.

Behnke, R. J. 1992. Native trout of western North America. Amer. Fish. Soc. Mono. 6, 275 pp.

Beland, R. D. 1953. The occurrence of two additional centrarchids in the lower Colorado River.
Calif. Fish Game, 39:149-151.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 157

Bell, M. A. 1978. Fishes of the Santa Clara system, southern California. Contrib. Sci., Nat. Hist.

Mus. L.A. Co., No. 295, 20 pp.

1982. Melanism in a high elevation population of Gasterosteus aculeatus. Copeia, 1982:
829-835.

, and K. E. Richkind. 1981. Clinal variation of lateral plates in threespine stickleback fish.

Amer. Nat., 117:113-132.

Benson, S. B., and R. J. Behnke. 1961. Salmo evermanni a synonym of Salmo clarki henshawi. Calif.
Fish Game, 47:257-259.

Berg, W. J., and G. A. Gall. 1988. Gene flow and genetic differentiation among California coastal
rainbow trout populations. Canad. J. Fish. Aquat. Sci., 45:122-132.

Bolster, B. B. 1990a. Mohave tui chub, five year status report. Inland Fisheries Division, Calif. Dept.

Fish Game, Rancho Cordova, 10 pp. (mimeo)

1990b. Owens tui chub, five year status report. Inland Fisheries Division, Calif. Dept. Fish
Game, Rancho Cordova, 10 pp. (mimeo)

1990c. Desert pupfish, five year status report. Inland Fisheries Division, Calif. Dept. Fish
Game, Rancho Cordova, 12 pp. (mimeo)

1990d. Owens pupfish, five year status report. Inland Fisheries Division, Calif. Dept. Fish
Game, Rancho Cordova, 10 pp. (mimeo)

Bond, C. E., and T. T. Kan. 1985. Systematics and evolution of the lampreys of Oregon. P. 919
(abst.) in Proceedings Second International Conference on Indo-Pacific Fishes, Ichthyological
Society of Japan, Tokyo.

Bottroff, L., J. A. St. Amant, and W. Parker. 1969. Addition of Pylodictis olivaris to the California
fauna. Calif. Fish Game, 55:90.

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). Canad. J. Fish. Aquat. Sci., 48:849-856.

Burkhead, N. M., and J. D. Williams. 1991. An intergeneric hybrid of a native minnow, the golden
shiner, and an exotic minnow, the rudd. Trans. Amer. Fish. Soc., 120:781-795.

Busack, C. A., and G. A. E. Gall. 1980. Ancestry of artificially propagated California rainbow trout
strains. Calif. Fish Game, 66:17-24.

Buth, D.G. 1984. Genetic affinities of freshwater populations of Gasterosteus aculeatus with special

reference to Gasterosteus aculeatus williamsoni. Final Report, U.S. Forest Service Job Order

40-91T5-0-974. Pasadena, CA.

, and C. B. Crabtree. 1982. Genetic variability and population structure of Catostomus san-

taanae in the Santa Clara River drainage. Copeia, 1982:439-444.

: , R. D. Orton; and W. J. Rainboth. 1984. Genetic differentiation between freshwater

subspecies of Gasterosteus aculeatus in southern California. Biochem. Syst. Ecol., 12:423-432.

, and R. W. Murphy. 1980. Catostomus fumeiventris. P. 379 in Atlas of North American

freshwater fishes. (D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. Mc. Allister, and

J. R. Stauffer, Jr., eds.), North Carolina State Museum, Raleigh, NC.

—, , and L. Ulmer. 1987. Population differentiation and introgressive hybridization of
the flannelmouth sucker and of hatchery and native stocks of the razorback sucker. Trans.
Amer. Fish. Soc., 116:103-110.

California Department of Fish and Game. 1931. [Report of] Bureau of Fish Culture. Calif. Fish

Game, 17:97-98.

1973. Warmwater game fishes of California. 3rd Edition, revised. Calif. Dept. Fish Game,
Sacramento, 39 pp. (noted on last page as the reorganization by R. L. Johnson and L. Shapavalov
of an earlier leaflet by B. Curtis).

Capone, T. A., and J. A. Kushlan. 1991. Fish community structure in dry-season stream pools.
Ecology, 72:983-992.

Carpelan, L. 1961. History of the Salton Sea. Pp. 9-15 in The ecology of the Salton Sea, California,
in relation to the sportfishery. (B. W. Walker, ed.), Calif. Dept. Fish Game, Fish Bull. No. 113.

Castleberry, D. T., and J. J. Cech, Jr. 1986. Physiological responses of a native and an introduced
desert fish to environmental stressors. Ecology, 67:912-918.

——., J. E. Williams, G. M. Sato, T. E. Hopkins, A. M. Brasher, and M. S. Parker. 1990. Status
and management of Shoshone Pupfish, Cyprinodon nevadensis shoshone (Cyprinodontidae), at
Shoshone Spring, Inyo County, California. Bull. S. Calif. Acad. Sci., 89:19-25.

Castro-Aguirre, J. L. 1978. Catalogo sistematico de los peces marinos que penetran a las aguas

158 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

continentales de Mexico con aspectos zoogeograficos y ecologicos. Dir. Gral. Inst. Nac. Pesca,
Mexico, Ser. Cient., 19, 298 pp.
Clark, F.N. 1932. Introduction of Mexican fishes into southern California waters. Calif. Fish Game,
18:61-62.
1940. True smelt found near Los Angeles Harbor. Calif. Fish Game, 26:180.

Collette, B. B.,and C. E. Nauen. 1983. Scombrids of the world. An annotated and illustrated catalogue
of tunas, mackerels, bonitos and related species known to date. U.N. Food Agri. Org., Fish.
Synopsis No. 125, vol. 2, vit + 137 pp.

——.,N. V. Parin, and M. S. Nizinski. 1992. Catalog of type specimens of recent fishes in the
National Museum of Natural History, Smithsonian Institution, 3: Beloniformes (Teleostei).
Smiths. Contrib. Zool., 525:1-16.

Compagno, L. J. V. 1984. Sharks of the world. United Nations, FAO Fisheries Synopsis No. 124,
vol. 4, Pt. 2, x + 251-655.

Cooper, J. G. 1874. Animal life of the Cuyamaca Mountains. Amer. Nat. (Peabody Acad. Sci.,
Salem, Mass.), 8:14—-18.

Cornelius, R.H. 1969. The systematics and zoogeography of Rhinichthys osculus (Girard) in southern
California. MS thesis, California State University, Fullerton.

Courtenay, W. R., Jr., D. A. Hensley, J. N. Taylor, and J. A. McCann. 1984. Distribution of exotic
fishes in the continental United States. Pp. 41-77 in Distribution, biology, and management
of exotic fishes. (W. R. Courtenay and J. R. Stauffer, eds.), John Hopkins Press, Baltimore.

—— : , and 1986. Distribution of exotic fishes in North America. Pp. 675-
698 in The F OOueOR ADH of North American freshwater fishes. (C. H. Hocutt and E. O. Wiley,
eds.), John Wiley and Sons, New York.

—., D. P. Jennings, and J. D. Williams. 1991. Appendix 2: exotic fishes. Pp. 97-107 in Common
and scientific names of fishes from the United States and Canada. Fifth Edition. (C. R. Robins,
R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott, eds.),
Amer. Fish. Soc. Spec. Publ. 20, Bethesda.

Crabtree, C. B. 1985. Allozyme variability in the tidewater goby, Eucyclogobius newberryi (Pisces:
Gobiidae). Isozyme Bull., 18:70.

Croker, R. S. 1934. The freshwater commercial fisheries of California. Calif. Fish Game, 20:375-
384.

Culver, G. B., and C. L. Hubbs. 1917. The fishes of the Santa Ana system streams in southern
California. Lorquinia, 1:82-83.

Curtis, B. 1949. The warm-water game fishes of California. Calif. Fish Game, 35:255-273.

Davis, S. P. 1963. Commercial freshwater fisheries of California. Calif. Fish Game, 49:84-94.

Dawson, W. L. 1923. The birds of California, vol. 3. South Moulton Company, San Diego, xiii +
1433-2121. ;

Deinstadt, J. M., E. J. Pert, F.G. Hoover, and S. Sasaki. 1988. Survey of fish populations in southern
California streams: 1987. Calif. Fish and Game, Inland Fisheries Division, Administrative
Rept. No. 88-5, 85 pp.

Dill, W. A. 1944. The fishery of the lower Colorado River. Calif. Fish Game, 30:109=211.

, M. Coots, and P. A. Douglas. 1955. The status of the pumpkinseed, Lepomis gibbosus

(Linne), in California. Calif. Fish Game, 41:157-159.

Echelle, A. A. 1991. Conservation genetics and genic diversity in freshwater fishes.of western North
America. Pp. 141-153 in Battle against extinction. Native Fish management in the American
west. (W. L. Minckley and J. E. Deacon, eds.), Univ. Arizona Press, Tucson.

Eigenmann, C. H. 1887. A review of the American Gasterosteidae. Proc. Acad. Nat. Sci. Phila.,

1886 (1887):233-252.

1892. The fishes of San Diego. Proc. U.S. Nat. Mus., 15:123-178.

, and R. S. Eigenmann. 1890. Additions to the fauna of San Diego. Proc. Calif. Acad. Sci.,

Ser. 2, 3:1-24.

Emery, K. O., R. E. Stevenson, and J. W. Hedgpeth. 1957. Estuaries and lagoons. Pp. 673-749 in
Treatise on marine ecology and paleoecology. Vol. 1, Ecology. (J. W. Hedgpeth), Geol. Soc.
Amer. Mem. 67.

Enzel, Y., W. J. Brown, R. Y. Anderson, L. D. McFadden, and S. G. Wells. 1992. Short-duration
Holocene lakes in the Mojave River drainage basin, southern California. Quat. Res.,
38:60-73.

Erman, D. C., and G. R. Leidy. 1975. Downstream movement of rainbow trout fry in a tributary

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 159

of Sagehen Creek, under permanent and intermittent flow. Trans. Amer. Fish. Soc., 104:467-
473.

Eschmeyer, W. N., E. S. Herald, and H. Hamann. 1983. A field guide to Pacific coast fishes of North
America. Houghton Mifflin Co., Boston, xii + 336 pp.

Evans, W. A. 1950. Notes on the occurrence of the bigmouth buffalo in southern California. Calif.
Fish Game, 36:332-333.

Evermann, B. W., and H. W. Clark. 1931. A distributional list of the species of freshwater fishes
known to occur in California. Calif. Dept. Fish Game, Fish Bull. 35, 67 pp.

Ewy, E. E. 1945. An ecological study of certain portions of the San Gabriel River basin. MS thesis,
Whittier College, Whittier, CA.

Fausch, K. D., and R. G. Bramblett. 1991. Disturbance and fish communities in intermittent trib-
utaries of a western Great Plains river. Copeia, 1991:659-674.

Felley, J. D., and G. L. Daniels. 1992. Life history of the sailfin molly (Poecilia latipinna) in two
degraded waterways of southwestern Louisiana. Southwest. Nat., 37:16—21.

Fitch, J. E. 1952. Distributional notes on some Pacific coast marine fishes. Calif. Fish Game, 38:

557-564.

,and A. O. Flechsig. 1949. A brief account of the Monterey spanish mackerel (Scomberomorus

concolor). Calif. Fish Game, 35:275—280.

, and S. A. Schultz. 1978. Some rare and unusual occurrences of fishes off California and Baja

California. Calif. Fish Game, 64:74-92.

Follett, W. I. 1961. The freshwater fishes—their origin and affinities. Pp. 212—232 in Symposium:

the biogeography of Baja California and adjacent seas. Syst. Zool., vol. 9.

. 1976. Fish remains from an archaeological site at Rancho Carrillo on the Silver Strand, San

Diego County, California. Bull. S. Calif. Acad. Sci., 75:131-137. ‘

Fritzsche, R. A. 1980. Revision of the eastern Pacific Syngnathidae (Pisces: Syngnathiformes), in-
cluding both recent and fossil forms. Proc. Calif. Acad. Sci., 42:181-227.

Fry, D. H., Jr., and P. M. Roedel. 1945. The shark, Carcharhinus azureus, in southern California
waters. Calif. Fish Game, 31:209.

Gall, G. A. E., B. Bently, and R. C. Nuzum. 1991. Genetic isolation of steelhead rainbow trout in
Kaiser and Redwood creeks, California. Calif. Fish Game, 76:216-223.

George, F. A. 1927. The eastern bullfrog in southern California. Calif. Fish Game, 13:222.

Gilbert, C. H. 1893. Report on the fishes of the Death Valley Expedition, collected in southern
California and Nevada in 1891, with descriptions of new species. North Amer. Fauna, 7:229-
234.

Gill, T. 1862. Notice ofa collection of the fishes of California presented to the Smithsonian Institution
by Mr. Samuel Hubbard. Proc. Acad. Nat. Sci., Phila., 14:274—282.

Glidden, D. 1931. Spiny-rayed fish, San Diego County lakes. Calif. Fish Game, 17:89-90.

Gobalet, K. W. 1990. Fish remains from archaeological site LAN-229: Freshwater and marine fishes

exploited. Jn Archaeological studies at site CA-LAN-229: an experiment in inference justifi-

cation. (M. Rabb, ed.), Northridge Center for Public Archaeology, Calif. St. University, North-

ridge.

. 1992. Colorado River fishes of Lake Cahuilla, Salton Basin, southern California: a cautionary

tale for zooarchaeologists. Bull. S. Calif. Acad. Sci., 91:70-83.

Gold, J. R., G. A. E. Gall, and S. J. Nicola. 1978. Taxonomy of the Colorado cutthroat trout (Salmo
clarki pleuriticus) of the Williamson Lakes, California. Calif. Fish Game, 64:98-103.

Gordon, B. L. 1985. Monterey Bay area: natural history and cultural imprints. 2nd Ed. Boxwood
Press, Pacific Grove, CA.

Gorsline, D. S. 1967. Contrasts in coastal bay sediments on the Gulf and Pacific coasts. Pp. 219-
225 in Estuaries. (G. H. Lauff, ed.), Amer. Assoc. Adv. Sci., Publ. 83, Washington, D.C.

Greenfield, D. W., and G. D. Deckert. 1973. Introgressive hybridization between Gila orcutti and

Hesperoleucas symmetricus (Pisces: Cyprinidae) in the Cuyama River Basin, California: II.

Ecological Aspects. Copeia, 1973:417—427.

, and T. Greenfield. 1972. Introgressive hybridization between Gila orcutti and Hesperoleucas

symmetricus (Pisces: Cyprinidae) in the Cuyama River Basin, California: I. Meristics, mor-

phometrics and breeding. Copeia, 1972:849-859.

, S. T. Ross, and G. D. Deckert. 1970. Some aspects of the life history of the Santa Ana

sucker, Catostomus santaanae (Snyder). Calif. Fish and Game, 56:166-179.

Gregory, S. V., F.G. Swanson, W. A. McKee, and D. W. Cummins. 1991. An ecosystem perspective
of riparian zones. Focus on links between land and water. Bioscience, 41:540-551.

160 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Haaker, P. 1977. The biology of the California halibut, Paralichthys californicus (Ayres) in Anaheim
Bay, California. Pp. 137-151 in The marine resources of Anaheim Bay. (E. D. Lane and C. W.
Hill, eds.), Calif. Dept. Fish Game, Fish Bull. 165. (1975)

1979. Two asiatic gobiid fishes, Tridentiger trigonocephalus and Acanthogobius flavimanus,

in southern California. Bull. S. Calif. Acad. Sci., 78:56-61.

Haglund, T. R., and D. G. Buth. 1988. Allozymes of the unarmored threespine stickleback (Gas-
terosteus aculeatus williamsoni) and identification of the Shay Creek population. Isozyme Bull.,
21:196.

Hendrickson, D. A., and J. E. Brooks. 1991. Transplanting short-lived fishes in North American
deserts: review, assessment, and recommendations. Pp. 283-298 in Battle against extinction.
Native fish management in the American West. (W. L. Minckley and J. E. Deacon, eds.), Univ.
Arizona Press, Tucson.

Hindar, K., N. Ryman, and F. Utter. 1991. Genetic effects of cultured fish on natural fish populations.
Canad. J. Fish. Aquat. Sci., 43:945-957.

Hoover, F., and J. A. St. Amant. 1983. Results of Mohave chub, Gila bicolor mohavensis, relocations
in California and Nevada. Calif. Fish Game, 69:54—56.

Hopkirk, J. D. 1974. Endemism in the fishes of the Clear Lake region. Univ. Calif. Publ. Zool., 96,
ii + 135 pp., 4 pls., publ. 1974 according to Shapavalov et al. (1981).

Horn, M. L. 1981. Diet and seasonal variation in abundance and diversity of shallow water fish

populations in Morro Bay, California. U.S. Nat. Mar. Fish Ser., Fish. Bull., 78:759-770.

1988. The fish community of the upper Newport Bay Ecological Reserve. Pp. 80-92 in The
natural and social sciences of Orange County. (H. C. Koerper, ed.), Mem. Nat. Hist. Found.

Orange Co., vol. 2.

, and L. G. Allen. 1976. Numbers of species and faunal resemblance of marine fishes in

California bays and estuaries. Bull. S. Calif. Acad. Sci., 75:159-170.

, and 1978. A distributional analysis of California coastal marine fishes. J. Biogeogr.,
5:23-42.
, and 1985. Fish community ecology in southern California bays and estuaries. Ch.

8, pp. 169-190 in Ecology in estuaries and coastal lagoons: towards an ecosystem integration.
(A. Yanez-Arancibia, ed.), DR (R) UNAM UNAM Press, Mexico.

, and F. D. Hagner. 1984. Ecological status of striped bass, Morone saxatilis, in upper

Newport Bay, California. Calif. Fish Game, 70:180-182.

Hubbs, C. L. 1916. Notes on the marine fishes of southern California. Univ. Calif. Publ. Zool., 16:
153-169.

. 1919. The stickleback: a fish eminently fitted by nature as a fish destroyer. Calif. Fish Game,
5:21-24.

1921. Notes on Cottus asper and Cottus aleuticus. Copeia, 90:7-8.

1946. Wandering of pink salmon and other salmonid fishes into southern California. Calif.
Fish Game, 32:81-86.

1953. Eleotris picta added to the fish fauna of California. Calif. Fish Game, 39:69-76.

. 1954. Establishment ofa forage fish, the red shiner (Notropis lutrensis), in the lower Colorado
River system. Calif. Fish Game, 40:287-294.

1967. Occurrence of the Pacific lamprey, Entosphenus tridentatus, off Baja California and
in streams of southern California, with remarks on its nomenclature. Trans. San Diego Nat.
Hist. Soc., 14:301-312.

—, W. I. Follett, and L. C. Hubbs. 1979. List of the fishes of California. Occ. Pap. Calif. Acad.

Sci. No. 133, 51 pp.

, L.C. Hubbs, and R. E. Johnson. 1943. Hybridization in nature between species of catostomid

fishes. Contr. Lab. Vert. Biol. Univ. Mich. No. 22, 76 pp.

,and K. F. Lagler. 1964. Fishes of the Great Lakes Region, with a new preface. Univ. Michigan

Press, Ann Arbor, MI, xv + 213 pp., 251 figs., 44 pls.

, and R. R. Miller. 1943. Mass hybridization between two genera of cyprinid fishes in the

Mohave desert, California. Pap. Mich. Acad. Sci. Arts Lett., 28:343-378.

Hubbs, C. 1991. Intrageneric “cannibalism” in Gambusia. Southwest. Nat., 36:153-157.

Irwin, J. F., and D. L. Soltz. 1982. The distribution and natural history of the unarmored threespine
stickleback, Gasterosteus aculeatus williamsoni (Girard), in San Antonio Creek, California. U.S.
Fish Wild. Ser., Sacramento, CA. Contract No. 11310-0289-80, 90 pp.

3

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 161

Jannik, N. O., F. M. Phillips, G. I. Smith, and D. Elmore. 1991. A 2°Cl chronology of lacustrine
sedimentation in the Pleistocene Owens River system. Geol. Soc. Amer. Bull., 103:1146-1159.

Jennings, M. R., and M. K. Saiki. 1990. Establishment of red shiner, Notropis lutrensis, in the San
Joaquin Valley, California. Calif. Fish Game, 76:46—57.

Johnson, D. L. 1977. The late Quaternary climate of coastal California: evidence for an Ice Age
refugium. Quat. Res., 8:154-179.

Johnson, J. E. 1991. Management concerns for endangered and threatened species of fish in western
warmwater springs and streams. Pp. 160-168 in Warmwater fisheries symposium I. (J. L. Cooper
and R. H. Hamre, Coordinators), USDA Forest Service, Gen. Tec. Rept. RM-207.

Jordan, D. S. 1895. Notes on the fresh-water fishes of San Luis Obispo County, California. Bull.

U.S. Fish Comm., 14:141-142.

, and C. H. Gilbert. 1883. Synopsis of the fishes of North America. Bull. U.S. Nat. Mus., 3:

lvi + 1018 pp.

, and W. W. Henshaw. 1878. Report on fishes collected during the years 1875, 1876, and

1877 in California and Nevada. U.S. Geol. Geogr. Surv. Terr. Rept., 1878:187—200.

Kennedy, C. H. 1916. A possible enemy of the mosquito. Calif. Fish Game, 2:179-182.

Kimsey, J. B. 1954. The introduction of the redeye bass and the threadfin shad into California. Calif.

Fish Game, 40:203-204.

, and L. O. Fisk. 1960. Keys to the freshwater and anadromous fishes of California. Calif.

Fish Game, 46:453-479.

, and L. O. Fisk. 1964. Freshwater nongame fishes of California. Calif. Dept. Fish Game,

Sacramento, CA, 54 pp.

Knaggs, E. H. 1977. Status of the genus 7i/apia in California’s estuarine and marine waters. Cal-
Neva Wildl. Trans., 1977:60-67.

Kramer, S. H. 1991a. Growth, mortality, and movements of juvenile California halibut Paralichthys

californicus in shallow coastal and bay habitats of San Diego County, California. U.S. Nat.

Mar. Fish. Ser. Fish. Bull., 89:195-—207.

. 1991b. The shallow water flatfishes of San Diego County. Calif. Coop. Oceanic Fish. Invest.

Rep., 32:128-142.

1992. Distribution and abundance of juvenile California halibut, Paralichthys californicus,
in shallow waters of San Diego County. Pp. 99-126 in The California halibut, Paralichthys
californicus, resource and fisheries. (C. H. Haugen, ed.), Calif. Dept. Fish Game Fish Bull. 174.
(1990)

Kresja, R. J. 1967a. The systematics of the prickly sculpin, Cottus asper Richardson, a polytypic

species. Part I. Synonymy, nomenclatural history and distribution. Pac. Sci., 21:241-251.

1967b. The systematics of the 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.

Lane, E.D. 1977a. Early collections of fishes from Anaheim Bay made between 1919 and 1928. Pp.

13-15 in The marine resources of Anaheim Bay. (E. D. Lane and C. W. Hill, eds.), Calif. Dept.

Fish Game, Fish Bull. 165. (1975)

. 1977b. Quantitative aspects of the life history of the diamond turbot, Hypsopsetta guttulata

(Girard), in Anaheim Bay. Pp. 153-174 in The marine resources of Anaheim Bay. (E. D. Lane

and C. W. Hill, eds.), Calif. Dept. Fish Game, Fish Bull. 165. (1975)

Lanse, R. 1965. The occurrence of warmouth, Chaenobryttus gulosus (Cuvier), in the lower Colorado
River. Calif. Fish Game, 51:123.

Lau, S., and C. Boehm. 1991. A distribution survey of desert pupfish (Cyprinodon macularius)
around the Salton Sea, California. Dept. Fish Game, Region 5, Final Rept. Proj. EF90XII-1,
21 pp.

Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, jr. 1980.
Atlas of North American freshwater fishes. North Carolina State Museum, Raleigh, NC, xiii
+ 867 pp.

Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and
ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology
and ichthyology. Copeia, 1985:802-832.

Lopez, J., and L. Ulmer. 1983. Preliminary distribution abundance survey of Tilapia found in
irrigation drains in the Palo Verde Valley, Riverside County, California. Calif. Dept. Fish Game,
Inland Fish., Region 5, unpublished report.

162 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Lufkin, A. 1991. California’s salmon and steelhead: the struggles to restore an imperiled resource.
Univ. Calif. Press, Berkeley, xxiii + 305 pp.

Manion, B. S., and J. H. Dillingham. 1989. Malibu Lagoon: a baseline ecological survey. Topanga-
Las Virgenes Res. Cons. Dist., Topanga, CA, vii + 180 pp.

Matarese, A. C., A. W. Kendall, Jr., D. M. Blood, and B. M. Vinter. 1989. Laboratory guide to early
life history stages of northeast Pacific fishes. NOAA Tech. Rept. (U.S. Nat. Mar. Fish. Ser.),
80, iv + 652 pp.

Mayden, R. L. 1992. Review of: A field guide to freshwater fishes of North America north of Mexico,
by L. M. Page and B. B. Burr, 1991. (Houghton Miffin Co., Boston, xii + 432 pp.) Copeia,
1992:920-925.

McCosker, J. E. 1989. Freshwater eels (Family Anguillidae) in California: current conditions and
future scenarios. Calif. Fish Game, 75:4—10.

McCoid, M. J., and J. A. St. Amant. 1980. Notes on the establishment of the rainwater killifish,
Lucania parva, in California. Calif. Fish Game, 66:124—125.

McDowali, R. M. 1989. Diadromy in fishes. Migrations between freshwater and marine environ-
ments. Timber Press, Portland, OR, ix + 308 pp.

Mearns, A. J. 1975. Poeciliopsis gracilis (Heckel), a newly introduced poeciliid fish in California.
Calif. Fish Game, 61:251-253.

Meffe, G. K. 1985. Predation and species replacement in American southwestern fishes: a case study.
Southwest. Nat., 30:173-—187.

Meador, M. R., and W. J. Matthews. 1991. Spatial and temporal patterns in fish assemblage structure
of an intermittent Texas stream. Amer. Mid. Nat., 127:106—-114.

Miller, D. J., and R. N. Lea. 1976. Guide to the coastal marine fishes of California. Calif. Dept.

Fish Game, Fish Bull. 157, 235 pp.

, and J. Schmidtke. 1956. Report on the distribution and abundance of Pacific herring (C/upea
pallasi) along the coast of central and southern California. Calif. Fish Game, 42:163-—187.

Miller, R. R. 1939. Occurrence of the cyprinodont fish, Fundulus parvipinnis, in fresh water in San

Juan Creek, southern California. Copeia, 1939:168.

1943. Further data on freshwater populations of the Pacific killifish, Fundulus parvipinnis.

Copeia, 1943:51-52.

. 1945a. Anewcyprinid fish from southern Arizona, and Sonora, Mexico, with the description

of a new subgenus of Gila and a review of related species. Copeia, 1945:104—110.

1945b. Hyporhamphus patris, a new species of hemiramphid fish from Sinaloa, Mexico,
with an analysis of the generic characters of Hyporhamphus and Hemiramphus. Proc. U.S. Nat.

Mus., 96:185-193.

—. 1945c. The status of Lavinia ardesiaca, a cyprinid fish from the Pajaro-Salinas River basin,

California. Copeia, 1945:197-204.

1946a. Correlation between fish distribution and Pleistocene hydrography in eastern Cali-

fornia and southwestern Nevada, with a map of the Pleistocene waters. J. Geol., 54:43-53.

1946b. Distributional records for North American fishes, with nomenclatorial notes on the

genus Psenes. J. Wash. Acad. Sci., 36:206—212.

———. 1948. The cyprinodont fishes of the Death Valley system of eastern Califouna and south-
western Nevada. Univ. Mich. Mus. Zool. Misc. Publ. 68, 155 pp.

1950. Notes on the cutthroat and rainbow trouts with the description of a new species from
the Gila River, New Mexico. Occ. Pap. Mus. Zool. Univ. Mich. No. 529, 42 pp.
1952. Bait fishes of the lower Colorado River from Lake Mead, Nevada, to Yuma, Arizona,

with a key for their identification. Calif. Fish Game, 38:7-42.

1958. Origin and affinities of the freshwater fish fauna of western North America. Pp. 187-

222 in Zoogeography. (C. L. Hubbs, ed.), Amer. Assoc. Adv. Sci., Washington, D.C.

1961. Man and the changing fish fauna of the American southwest. Pap. Mich. Acad. Sci.,

Arts Lett., 46:365-404.

. 1968. Records of some native freshwater fishes transplanted into various waters of California,

Baja California, and Nevada. Calif. Fish Game, 54:170—-179.

——. 1973. Two new fishes, Gila bicolor snyderi and Catostomus fumeiventris, from the Owens
River basin, California. Occ. Pap. Mus. Zool. Univ. Mich., No. 667, 19 pp.

——. 1981. Coevolution of deserts and pupfishes (genus Cyprinodon) in the American southwest.
Pp. 39-94 in Fishes in North American deserts. (R. J. Naiman and D. L. Soltz, eds.), John
Wiley and Sons, New York.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 163

,andJ.R. Alcorn. 1946. The introduced fishes of Nevada, with a history of their introduction.

Trans. Amer. Fish. Soc., 73:173-193.

, and C. L. Hubbs. 1969. Systematics of Gasterosteus aculeatus, with special reference to

intergradation and introgression along the Pacific Coast of North America: a commentary on

a recent contribution. Copeia, 1969:52-69.

, C. Hubbs, and F. H. Miller. 1991. Ichthyological explorations of the American west: the

Hubs-Miller era, 1915-1950. Pp. 19-40 in Battle against extinction. Native fish management

in the American west. (W. L. Minckley and J. E. Deacon, eds.), Univ. Arizona Press, Tucson.

, and E. Pister. 1971. Management of the Owens pupfish, Cyprinodon radiosus, in Mono

County, California. Trans. Amer. Fish. Soc., 100:502-509.

, and M. L. Smith. 1986. Origin and geography of the fishes of central Mexico. Pp. 487-517

in The zoogeography of North American freshwater fishes. (C. H. Hocutt and E. O. Wiley,
eds.), John Wiley and Sons, New York.

—, J. D. Williams, and J. E. Williams. 1989. Extinctions of North American fishes during the
past century. Fisheries, 14:22-38.

Minckley, W. L. 1973. Fishes of Arizona. Arizona Game Fish Dept., Phoenix, xv + 293 pp.

. 1981. Trophic interrelations among introduced fishes in lower Colorado River, southwestern

United States. Calif. Fish Game, 68:78-89.

, and J. E. Deacon. (eds.). 1991. Battle against extinction. Native fish management in the

American west. Univ. Ariz. Press, Tucson, xviii + 517 pp.

—,, D. A. Hendrickson, and C. E. Bond. 1986. Geography of western North American freshwater

fishes: description and relationships to intracontinental tectonism. Pp. 519-613 in The zoo-

geography of North American freshwater fishes. (C. H. Hocutt and E. O. Wiley, eds.), John

Wiley and Sons, New York. ‘

, P. C. Marsh, J. E. Brooks, J. E. Johnson, and B. L. Jensen. 1991. Management toward

recovery of the razorback sucker. Pp. 303-358 in Battle against extinction. Native fish man-

agement in the American west. (W. L. Minckley and J. E. Deacon, eds.), Univ. Arizona Press,

Tucson.

, G. K. Meffe, and D. L. Soltz. 1991. Conservation and management of short-lived fishes:

the cyprinodontoids. Pp. 247-282 in Battle against extinction. Native fish management in the

American west. (W. L. Minckley and J. E. Deacon, eds.), Univ. Arizona Press, Tucson.

, and J. Rinne. 1991. Native fishes of arid lands: a dwindling resource of the desert southwest.

U.S. Forest Ser., Rocky Mtn. Forest Range Exp. Sta., Gen. Tech. Rept., RM-206, ii + 45 pp.

Moffett, J. W. 1942. A fishery survey of the Colorado River below Boulder Dam. Calif. Fish Game,
28:76-86. i

Moore, M. R. 1980. Factors influencing the survival of juvenile steelhead rainbow trout (Salmo
gairdneri gairdneri) in the Ventura River. MS thesis, Humbolt State University, Arcata, CA.

Morris, J. T. 1991. Effects of nitrogen loading on wetland ecosystems with particular reference to
atmospheric deposition. Annu. Rev. Ecol. Syst., 22:257-79.

Moyle, P. B. 1976a. Inland fishes of California. Univ. California Press, Berkeley, viii + 405 pp.

1976b. Fish introductions in California: history and impact on native fishes. Biol. Conser.,

9:101-118.

1980. Ptychocheilus grandis (Ayres). P. 347 in Atlas of North American freshwater fishes.
(D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr.,
eds.), North Carolina State Museum, Raleigh.

, and J. P. Ellison. 1991. A conservation-oriented classification system for the inland waters
of California. Calif. Fish Game, 77:161-—180.

,and R.A. Leidy. 1992. Loss of biodiversity in aquatic ecosystems: evidence from fish faunas.
Pp. 127-169 in Conservation biology. The theory and practice of nature conservation, pres-
ervation, and management. (P. L. Fiedler and S. K. Jain, eds.), Chapman and Hall, New York.
, and M. Massingill. 1981. Hybridization between hitch, Lavinia exilicauda, and Sacramento
blackfish, Orthodon microlepidotus, in San Luis Reservoir, California. Calif. Fish Game, 67:
196-198.

, and R. Nichols. 1973. Ecology of some native and introduced fishes of the Sierra-Nevada
foothills, central California. Copeia, 1973:478-490.

, and J. E. Williams. 1990. Biodiversity loss in the temperate zone: decline of the native fish
fauna of California. Conser. Biol., 4:275—-284.

: ,and R. K. White. 1990. Draft Final Report: an inventory of California’s inland fishes

164 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

with recommendations for candidate preserves to protect biodiversity of fishes. Rept. Inland

Fish. Div., Calif. Dept. Fish Game, Rancho Cordova, 220 pp. (mimeo)

, and E. D. Wikramanayake. 1989. Fish species of special concern of California. Final
Rept., Inland Fish. Div., Calif. Dept. Fish Game, Rancho Cordova, Contract No. 7337, ii +
222 pp.

Naiman, R. J., and E. P. Pister. 1974. Occurrence of the tiger barb, Barbus tetrazona, in the Owens
Valley, California. Calif. Fish Game, 60:100-101.

Neale, G. 1915. The catfish in California. Calif. Fish and Game, 1(2):62—64.

Needham, P. R., and R. Gard. 1963. Rainbow trout in Mexico and California with notes on the
cutthroat series. Univ. Calif. Publ. Zool., 67, 124 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, 16:4—21.

Norris, R. M., and R. W. Webb. 1990. Geology of California. John Wiley and Sons, New York, xiii
+ 541 pp.

Odenweller, D. B. 1977. The life history of the shiner surfperch, Cymatogaster aggregata Gibbons,
in Anaheim Bay, California. Pp. 107-115 in The marine resources of Anaheim Bay. (E. D.
Lane and C. W. Hill, eds.), Calif. Dept. Fish Game, Fish Bull. 165 (1975).

Page, L. M., and B. B. Burr. 1991. A field guide to freshwater fishes of North America north of
Mexico. Houghton Miffin Co., Boston, MA, xii + 432 pp.

Pelzman, R. J. 1980. Impact of Florida largemouth bass, Micropterus salmoides floridanus, intro-
ductions at selected northern California waters with a discussion of the use of meristics for
detecting introgression and for classifying individual fish of intergraded populations. Calif. Fish
Game, 66:133-162.

Pourade, R. F. 1968. The call to California. Union-Tribune Publishing Co., San Diego, xiv + 194 pp.

Queenan, C. F. 1986. Long Beach and Los Angeles. A tale of two ports. Windsor Publications Inc.,
Northridge, CA, 205 pp.

Regan, C. T. 1909. The species of three-spined stickleback. Ann. Mag. Nat. Hist., Ser. 8, 4:435-
437.

Richards, C., and D. L. Soltz. 1986. Feeding of rainbow trout (Salmo gairdneri) and arroyo chubs
(Gila orcutti) in a California mountain stream. Southwest. Nat., 31:250-253.

Richardson, W. M., J. A. St. Amant, L. J. Bottroff, and W. L. Parker. 1970. Introduction of blue
catfish into California. Calif. Fish Game, 70:311-312.

Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W. Phillips. 1985. Between Pacific tides. Fifth
Edition. Stanford Univ. Press, Stanford, CA, xvi + 652 pp.

Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott.

1991. Common and scientific names of fishes from the United States and Canada. Sth ed.

Amer. Fish. Soc. Spec. Publ. 20, Bethesda, MD, 183 pp.

, and R. R. Miller. 1957. Classification, variation, and distribution of the sculpins, genus

Cottus, inhabiting Pacific slope waters in California and southern Oregon, with a key to the

species. Calif. Fish Game, 43:213-233.

Rosenblatt, R. H., and W. J. Baldwin. 1958. A review of the eastern Pacific sharks of the genus
Carcharhinus, with a redescription of C. malpeloensis (Fowler) and California records of C.
remotus (Dumeril). Calif. Fish Game, 44:137-159.

Ross, S. T. 1973. The systematics of Gasterosteus aculeatus (Pisces: Gasterosteidae) in central and

southern California. Contrib. Sci. Nat. Hist. Mus. Los Angeles Co., 243, 20:pp.

1991. Mechanisms structuring stream fish assemblages: are there lessons from introduced

species? Environ. Biol Fish., 30:359-368.

Ryan, P. A. 1986. Seasonal and size related changes in the food of the short-finned eel, Anguilla
australis, in Lake Ellesmere, Canterbury, New Zealand. Environ. Biol. Fish., 15:47-58.
Schoenherr, A. A. 1979. Niche separation within a population of freshwater fishes in an irrigation

drain near the Salton Sea, California. Bull. S. Calif. Acad. Sci., 78:46—-55.

—. 1988. A review of the life history and status of the desert pupfish, Cyprinodon macularius.
Bull. S. Calif. Acad. Sci., 87:104—134.

Schultz, L. P. 1934. The age and growth of Atherinops affinis oregonia Jordan and Snyder and of
other subspecies of baysmelt along the Pacific Coast of the United States. Univ. Wash. Pubi.
Biol., 2:45-102.

Scofield, N. B. 1915. Fish and the mosquito problem. Calif. Fish Game, 1:230-231.

, and H. C. Bryant. 1926. The striped bass in California. Calif. Fish Game, 12:55-74.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 165

Shafland, P. L., and W. M. Lewis. 1984. Terminology associated with introduced organisms. Fish-
eries, 9(4):17-18.

Shapavalov, L., A. J. Cordone, and W. A. Dill. 1981. A list of the freshwater and anadromous fishes
of California. Calif. Fish Game, 67:4—38.

——,, W.A. Dill, and A. J. Cordone. 1959. A revised check list of the freshwater and anadromous

fishes of California. Calif. Fish Game, 45:159-180.

, and A. C. Taft. 1954. The life histories of the steelhead rainbow trout (Salmo gairdneri

gairdneri) and silver salmon (Oncorhynchus kisutch). Calif. Dept. Fish Game, Fish Bull. 98,

575 pp.

Shebley, W. H. 1917a. History of introduction of food and game fishes into the waters of California.
Calif. Fish Game, 3:3-12.

1917b. Hatchery notes. Calif. Fish Game, 3:29-34.

1919. Hatchery notes. Calif. Fish Game, 5:37-39.

1922. A history of fish cultural operations in California. Calif. Fish Game, 8:62-98.

1925. Hatchery notes. Calif. Fish Game, 11:83-84.

1927. Report of the Department of Fish Culture. Pp. 33-61 in Twentyninth Annual Report
of the California Fish and Game Commission for the years 1924-1926. Calif. Dept. Fish Game,
Sacramento.

Simon, T. P., and J. B. Kaskey. 1992. Description of eggs, larvae, and early juveniles of the bigscale
logperch, Percina macrolepida Stevenson, from the west fork of the Trinity River Basin, Texas.
Southwest. Nat., 37:28-34.

Smith, G. R. 1966. Distribution and evolution of the North American catostomid fishes of the
subgenus Pantosteus, genus Catostomus. Misc. Publ. Mus. Zool. Univ. Mich., No. 129, 132
pp., 1 pl.

Smith, H. M. 1896. A review of the history and results of the attempts to acclimatize fish and other
water animals in the Pacific states. Bull. U.S. Fish Comm., (1895) 15:379-472.

Smith, M. L. 1981. Late Cenozoic fishes in the warm deserts of North America: a reinterpretation
of desert adaptations. Pp. 11-38 in Fishes in North American deserts. (R. J. Naiman and D.
L. Soltz, eds.), John Wiley and Sons, New York.

Smith, R. 1883. Notes on the fishes of Todos Santos Bay, lower California. Proc. U.S. Nat. Mus.,
6:232-236.

Soltz, D. L., and R.J. Naiman. 1978. The natural history of native fishes in the Death Valley system.
Nat. Hist. Mus. Los Angeles Co., Sci. Ser. No. 30, ix + 76 pp.

Spratt, J.D. 1981. Status of the Pacific herring, Clupea harengus pallasii, resource in California 1972
to 1980 Calif. Dept. Fish Game, Fish Bull. 171, 99 pp.

St. Amant, J. A. 1966. Addition of Tilapia mossambica Peters to the California fauna. Calif. Fish
Game, 52:54—-55.

—-. 1970. Addition of Hart’s rivulus, Rivulus harti (Boulenger), to the California fauna. Calif.

Fish Game, 56:138.

, and F. G. Hoover. 1969. Addition of Misgurnus anguillicaudatus (Cantor) to the California

fauna. Calif. Fish Game, 55:330-331.

, and I. Sharp. 1971. Addition of Xiphophorus variatus (Meek) to the California fauna. Calif.

Fish Game, 57:128-129.

Starks, E. C., and E. L. Morris. 1906. The marine fishes of southern California. Univ. Calif. Publ.
Zool., 3:159-251.

Stone, L. 1884. The quinnat or California salmon-Oncorhynchus chouicha. Pp. 479-548 in The
fisheries and fisheries industries of the United States. Sec. I, Natural history of aquatic animals.
(G. Goode, ed.), Washington, D.C.

Swift, C. C. 1980a. Elops affinis. P. 55 in Atlas of North American freshwater fishes. (D. S. Lee, C.

R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr., eds.), North

Carolina State. Mus., Raleigh, NC.

1980b. Gila orcutti. P. 169 in Atlas of North American freshwater fishes. (D. S. Lee, C. R.
Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr., eds.), North
Carolina State Mus., Raleigh, NC.

1980c. Catostomus santaanae. P. 389 in Atlas of North American freshwater fishes. (D. S.
Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr., eds.),
North Carolina State Museum, Raleigh, NC.

1980d. Fundulus parvipinnis. P. 524 in Atlas of North American freshwater fishes. (D. S.

166 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES

Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr., eds.),

North Carolina State Museum, Raleigh, NC.

1989. Late Pleistocene freshwater fishes from the Rancho La Brea deposit, southern Cali-
fornia. Bull. S. Calif. Acad. Sci., 83:93-102.

, C. R. Gilbert, S. A. Bortone, G. H. Burgess, and R. W. Yerger. 1986. Zoogeography of the

freshwater fishes of the southeastern United States: Savanah River to Lake Pontchartrain. Pp.

213-265 in The zoogeography of North American freshwater fishes. (C. H. Hocutt and E. O.
Wiley, eds.).

———., J. L. Nelson, C. Maslow, and T. Stein. 1989. Biology and distribution of the tidewater goby,
Eucyclogobius newberryi (Pisces: Gobiidae) of California. Cont. Sci. Los Angeles Co. Mus. Nat.
Hist., 404, 19 pp.

Tasto, R. N. 1977. Aspects of the biology of Pacific staghorn sculpin, Leptocottus armatus Girard,
in Anaheim Bay. Pp. 123-135 in The marine resources of Anaheim Bay. (E. D. Lane and C.
W. Hill, eds.), Calif. Dept. Fish Game, Fish Bull. 165. (1975)

Taylor, J. N., W. R. Courtenay, Jr., and J. A. McCann. 1984. Known impacts of exotic fishes in the
contintenal United States. Pp. 322-373 in Distribution, biology, and management of exotic
fishes. (W. R. Courtenay and J. R. Stauffer, eds.), p. 58. John Hopkins Press, Baltimore.

Taylor, T. L. 1980. A blue catfish from the Sacramento-San Joaquin delta. Calif. Fish Game, 66:
120-121.

Thiery, R. G. 1991. Grass carp in the Coachella Canal: 1980-1990. Pp. 103-112 in Warmwater
fisheries symposium I. (J. L. Cooper and R. H. Hamre, Coordinators), USDA Forest Service,
Gen. Tech. Rept. RM-207.

Thompson, W. F. 1919. The spawning of the grunion (Leuresthes tenuis). Calif. Fish Game, Fish
Bull. 3, 27 pp.

Tres, J. 1992. Breeding biology of the Arroyo chub, Gila orcutti (Pisces: Cypridae) Ms Thesis,
California Polytechnic Univ., Pomona. 73 pp.

Turner, C. H., and J. C. Sexsmith. 1967. Marine baits of California. (1st rev. ed.) Calif. Dept. Fish
Game, Sacramento, CA, 71 pp.

Tyus, H. M. 1991la. Management of Colorado River fishes. Pp. 175-182 in Warmwater fisheries

symposium I. (J. L. Cooper and R. H. Hamre, Coordinators), USDA Forest Service, Gen. Tech.

Rept. RM-207.

1991b. Ecology and management of the Colorado squawfish. Pp. 379-402 in Battle against
extinction: native fish management in the American west. (W. L. Minckley and J. E. Deacon,
eds.), Univ. Arizona Press, Tucson.

U.S. Fish and Wildlife Service. 1985. Unarmored threespine stickleback recovery plan (revised).

U.S. Fish Wild. Ser., Portland, OR, 80 pp.

1991. Endangered and threatened wildlife and plants; 90-day findings and commencement
of status reviews for seven petitions to list five species as threatened or endangered. Federal
Register, 56:12146-12148.

Van Wormer, S. R. 1991. A history of flood control in the Los Angeles County drainage area. S.
Calif. Quart., 73:55—-94. ;

Vestal, E. H. 1942. Reclamation with rotenone of Crystal Lake, Los Angeles County, California.
Calif. Fish Game, 28:136-142.

Vogelsang, C. A. 1931. Early history of introduction and distribution of spiny-rayed fishes in Cal-
ifornia waters. Calif. Fish Game, 17:238—240. ;

Vondracek, B., W. A. Wurtsbaugh, and J. J. Cech., Jr. 1988. Growth and reproduction of the
mosquitofish, Gambusia affinis, in relation to temperature and ration level: consequences for
life history. Environ. Biol. Fish., 21:45-57.

von Geldern, C. E. 1966. The introduction of white bass (Roccus chrysops) into California. Calif.
Fish Game, 52:303.

Wales, J. B. 1962. Introduction of pond smelt from Japan into California. Calif. Fish Game, 48:
141-142.

Walker, B. W. 1952. A guide to the grunion. Calif. Fish Game, 38:409-420.

, R. R. Whitney, and G. W. Barlow. 1961. The fishes of the Salton Sea. Pp. 77-91 in The

ecology of the Salton Sea, California, in relation to the sport fishery. (B. W. Walker, ed.), Calif.

Dept. Fish Game, Fish Bull. 113.

Warme, J. E. 1971. Paleoecological aspects of a modern coastal lagoon. Univ. Calif. Publ. Geol.
Sci., 87, vi + 110 pp.

FRESHWATER FISHES OF SOUTHERN CALIFORNIA 167

Wells, A. W., J. S. Diana, and C. C. Swift. 1975. Survey of the freshwater fishes and their habitats
in the coastal drainages of southern California. Final Rep., Calif. Fish Game, Inland Fish.,
Sacramento. Contract AB-26, vi +364 pp.

Willett, G. 1941. Faunal changes in Los Angeles County. Quart. Mus. Patrons Assoc. Los Angeles
Co. Mus., 1:19-23.

Williams, J. D.,and D. Jennings. 1991. Computerized data base for exotic fishes: the western United
States. Calif. Fish Game, 77:86-93.

Williams, J. E., J. E. Johnson, D. A. Hendrickson, S. Contreras-Balderas, J. D. Williams, M. Navarro-
Mendoza, D. E. McAllister, and J. E. Deacon. 1989. Fishes of North America, endangered,
threatened, or of special concern. Fisheries, 14:2—20.

Williamson, G. R., and O. Tabeta. 1991. Search of Anguilla eels on the west coast of North America
and on the Aleutian and Hawaiian Islands. Jap. J. Ichthyol., 38:315-317.

Woodburne, M. O., R. E. Reynolds, and D. P. Whistler (eds.). 1991. Inland southern California:
the last 70 million years. Quart. San Bernardino Co. Mus. Assoc., 38, 120 pp.

Wydowski, R. S., and R. R. Whitney. 1979. Inland fishes of Washington. Univ. Washington Press,
Seattle, xxxii + 220 pp.

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, Cali-
fornia. Estuaries, 14:465—480.

Young, K. L., and P. C. Marsh. 1991. Age and growth of flathead catfish in four southwestern rivers.
Calif. Fish Game, 76:224—233.

Zedler, J. B. 1984. The ecology of southern California coastal salt marshes: a community profile.
U.S. Fish Wild. Ser., FWS/OBS-81-54, 110 pp. (2nd corrected printing)

Accepted for publication 18 December 1991.

Bull. Southern California Acad. Sci.
92(3), 1993, pp. 168-169
© Southern California Academy of Sciences, 1993

INDEX TO VOLUME 92
Abraham, Sue, see Leonard R. Brand

Bicak, Charles J., and Daniel Steinberg: Water Relations of an Annual Grass,
Bromus diandrus, in the Central Valley of California, 54

Boardman, Constance J., and Charles T. Collins: Lead in the Feathers of the
California Least Tern (Sterna antillarum browni), 95

Brand, Leonard R., Andrew Reese, Sue Abraham, and Ivan Rouse: Laboratory
Tests of Species Discrimination in Three Species of Peromyscus, 89

Bursey, Charles R., see Stephen R. Goldberg

Cash-Clark, Cora E., see Joel W. Martin
Chang, L., see P. Wells
Collins, Charles T., see Constance J. Boardman

Delaney, Paul M.: X-ray Microanalysis of the Cuticle Surface of Two Southern
California Marine Isopod Species, 64

Fisher, Robert N., see Camm C. Swift

Goldberg, Stephen R., Charles R. Bursey and Rana Tawil: Gastrointestinal Hel-
minths of the Western Brush Lizard, Urosaurus graciosus (Phrynosomatidae),
43

Grismer, L. Lee, and Jimmy A. McGuire: The Oases of Central Baja California,
Mexico. Part I. A Preliminary Account of the Relict Mesophilic Herpetofauna
and the Status of the Oases, 2

Haglund, Thomas R., see Camm C. Swift
Han, K., see P. Wells
Han, R., see P. Wells

Laughlin, Jimmy, see Bruce Thompson
ee, ©. see Ps Wells

Martin, Joel E., and Cora E. Cash-Clark: The Spinicaudatan Clam Shrimp Leptes-
theria Sars, 1898 (Crustacea, Brachiopoda) in California, 78
McGuire, Jimmy A., see L. Lee Grismer

Reese, Andrew, see Leonard R. Brand
Rouse, Ivan, see Leonard R. Brand
Ruiz, Mario, see Camm C. Swift

Sternberg, Daniel, see Charles J. Bicak
Swift, Camm C., Thomas R. Haglund, Mario Ruiz, and Robert N. Fisher: The
Status and Distribution of the Freshwater Fishes of Southern California, 101

168

INDEX TO VOLUME 92 169

Tawil, Rana, see Stephen R. Goldberg

Thompson, Bruce, David Tsukada, and Jimmy Laughlin: Megabenthic Assem-
blages of Coastal Shelves, Slopes, and Basins off Southern California, 25

Tsukada, David, see Bruce Thompson

Vadehra, N., see P. Wells
Vu, V., see P. Wells

Wells, H., see P. Wells
Wells, P., H. Wells, V. Vu, N. Vadehra, C. Lee, R. Han, K. Han, and L. Chang:
Does Honey Bee Nasanov Pheromone Attract Foragers?, 70

Proceedings of the Symposium

Interface Between Ecology and Land Development
in California

Edited by Jon E. Keeley

An outstanding compilation of timely papers by researchers, conser-
vationists, consultants, and policymakers, including:

Peter H. Raven Ecology and Species Extinction

Jonathan L. Atwood California Gnatcatcher and Coastal Sage Scrub

Martin L. Cody Theoretical and Empirical Aspects of Habitat Fragmen-
tation

Jim A. Bartel Endangered Species Act: Land Development, Politics, and
Reauthorization

Cheryl Swift et al. Habitat Linkages in an Urban Mountain Chain

Wayne R. Ferren et al. Rare and Threatened Wetlands of California

G. Ledyard Stebbins Conservation of California’s Rare Habitats

Todd Keeler-Wolf Rare Community Conservation in California

Joy B. Zedler Restoring Biodiversity to Coastal Marshes

Ted V. St. John et al. Use of Mycorrhizal Plants in Revegetation and Restora-
tion

Marylee Guinon Habitat Valuation and Restoration Costing

Jim Jokerst An Alternative Approach to Vernal Pool Mitigation Plan-
ning

and twenty-nine other papers dealing with land mangement, biodiversity,
habitat corridors, buffer zones, rare species, and community restoration.

Price, $26.00 (soft cover, includes tax and shipping)

Available from:
Southern California Academy of Sciences
900 Exposition Blvd. Los Angeles, CA 90007

Symposium

NATURAL RESOURCES
IN THE PUENTE HILLS—CHINO HILLS
CORRIDOR

IMPLICATIONS FOR LAND USE AND PLANNING

WHITTIER
MARCH 18-19, 1994
WHITTIER COLLEGE

CALL FOR PAPERS: Professional and student papers on topics pertaining to existing natural
resources, conservation, and land use policy in this geographic region are solicited. The purpose
of the symposium is to provide a forum for discussion of completed or continuing research on
these issues within this geographic location. The Puente Hills—Chino Hills Corridor encompasses
a large amount of both private and public open space under the jurisdiction of three counties and
numerous municipalities. Professionals and students engaged in research on land use policy in
this region are encouraged to attend.

Direct abstracts and correspondence to: Dr. Cheryl Swift
Department of Biology
Whittier College
Whittier, CA 90602

Please see other side for instructions regarding abstract preparation and submission. ABSTRACTS
DUE FEBRUARY 1, 1994.

Sponsored By: The Santa Monica Mountains Conservancy « Whittier College « The Whittier
Conservancy * The Whittier Audubon Society

ABSTRACT INSTRUCTIONS

WITH YOUR ABSTRACTS, PLEASE SUBMIT THE
FOLLOWING ON A 3x5 FILE CARD

Your name and affiliation.

Your mailing address and telephone number.
Title of your paper.

Audio-visual equipment required.

Ne

Abstracts will appear in the program exactly as submitted, so a clean, camera-ready copy is required. Abstracts
must be typed and submitted on an 812 x 11” sheet. Abstracts prepared on a word processor whould be printed
on a laser printer if possible. Typewritten abstracts should be prepared clean and have to erasures. Arrange
in a space 6” wide and 4” high (see the example below). Do not show the enclosing box. Please follow these
instructions.

1. TITLE must be in all capital letters. Use italics only for species names.

2. List the author(s) name(s) with the presenter’s name given first.

3. List the institution and department or affiliation for each author as indicated in the sample abstract.
4. Skip a single line and begin text. Do not indent. Text should be single spaced.

Sample Abstract

AGE STRUCTURE OF SYCAMORES, Platanus Racemosa, IN THE WHITTIER HILLS.
C. Quirarte, C. Allen, and C. Swift, Department of Biology, Whittier College, Whittier, CA
90608.

The age structure of a population of sycamores, Platanus racemosa, in Sycamore Canyon
in the Whittier Hills was determined, etc... .

MAIL ABSTRACT AND CARD TO: Dr. Cheryl Swift
Department of Biology
Whittier College
Whittier, CA 90602

ABSTRACTS DUE FEBRUARY I, 1994

Qn

1647

INSTRUCTIONS FOR AUTHORS

The BULLETIN is published three times each year (April, August, and December) and includes articles in English
in any field of science with an emphasis on the southern California area. Manuscripts submitted for publication
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The author should submit at least two additional copies with the original, on 82 x 11 opaque, nonerasable paper,
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A feature article comprises approximately five to thirty typewritten pages. Papers should usually be divided into
the following sections: abstract, introduction, methods, results, discussion and conclusions, acknowledgments,
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A research note is usually one to six typewritten pages and rarely utilizes subheadings. Consult a recent issue of
the BULLETIN for the format of notes. Abstracts are not used for notes.

Abbreviations: Use of abbreviations and symbols can be determined by inspection of a recent issue of the
BULLETIN. Omit periods after standard abbreviations: 1.2 mm, 2 km, 30 cm, but Figs. 1-2. Use numerals before
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Taxonomic procedures: Authors are advised to adhere to the taxonomic procedures as outlined in the International
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Viruses (Buchanan et al. 1958), and the International Code of Zoological Nomenclature (Stoll et al. 1961). Special
attention should be given to the description of new taxa, designation of holotype, etc. Reference to new taxa in
titles and abstracts should be avoided.

The literature cited: Entries for books and articles should take these forms.

McWilliams, K. L. 1970. Insect mimicry. Academic Press, vii + 326 pp.

Holmes, T. Jr., and S. Speak. 1971. Reproductive biology of Myotis lucifugus. J. Mamm., 54:452-458.

Brattstrom, B. H. 1969. The Condor in California. Pp. 369-382 in Vertebrates of California. (S. E. Payne, ed.),

Univ. California Press, xii + 635 pp.

Tables should not repeat data in figures (line drawings, graphs, or black and white photographs) or contained in
the text. The author must provide numbers and short legends for tables and figures and place reference to each of
them in the text. Each table with legend must be on a separate sheet of paper. All figure legends should be placed
together on a separate sheet. Illustrations and lettering thereon should be of sufficient size and clarity to permit
reduction to standard page size; ordinarily they should not exceed 8'2 by 11 inches in size and after final reduction
lettering must equal or exceed ‘the size of the typeset. All half-tone illustrations will have light screen (grey)
backgrounds. Special handling such as dropout half-tones, special screens, etc., must be requested by and wili be
charged to authors. As changes may be required after review, the authors should retain the original figures in their
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Assemble the manuscript as follows: cover page (with title, authors’ names and addresses), abstract, introduction,
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A cover illustration pertaining to an article in the issue or one of general scientific interest will be printed on the
cover of each issue. Such illustrations along with a brief caption should be sent to the Editor for review.

PROCEDURE

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College, 1600 Campus Road, Los Angeles, California 90041. Authors are requested to submit the names, addresses
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Proof: The galley proof and manuscript, as well as reprint order blanks, will be sent to the author. He or she
should promptly and carefully read the proof sheets for errors and omissions in text, tables, illustrations, legends,
and bibliographical references. He or she marks corrections on the galley (copy editing and proof procedures in
Style Manual) and promptly returns both galley and manuscript to the Editor. Manuscripts and original illustrations
will not be returned unless requested at this time. All changes in galley proof attributable to the author (misspellings,
inconsistent abbreviations, deviations from style, etc.) will be charged to the author. Reprint orders are placed with
the printer, not the Editor.

CONTENTS

The Status and Distribution of the Freshwater Fishes of Southern Califor-
nia. By Camm C. Swift, Thomas R. Haglund, Mario Ruiz, and Robert
IN FESHGT 2 i IN a eae REN GD LTA NSE AD SEU 101

Index tO: Vole D2). oii ENN a El APNE ee Ae 168

COVER: Leonard G. Hogue: ‘‘Steelhead (Salmon Trout) caught in Los Angeles River in City of Los
Angeles. June 1940. 25 inches long.” This was a locality near Glendale. Photo provided by Charles
L. Hogue.