123456789 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 A B C D E F G H I J K L M N O FINAL REPORT BIOLOGICAL CHARACTERISTICS OF NEARSHORE FISHES OF CALIFORNIA:

A REVIEW OF EXISTING KNOWLEDGE AND PROPOSED ADDITIONAL STUDIES FOR THE PACIFIC OCEAN INTERJURISDICTIONAL FISHERIES MANAGEMENT PLAN COORDINATION AND DEVELOPMENT PROJECT SUBMITTED TO AL DIDIER PACIFIC STATES MARINE FISHERIES COMMISSION AUGUST 31, 2000 PRINCIPAL INVESTIGATOR:

Dr. Gregor M. Cailliet, Professor Moss Landing Marine Laboratories 8272 Moss Landing Road Moss Landing, California 95039 ( 831) 632 4432; fax ( 831) 632 4403 cailliet@ mlml. calstate. edu MLML RESEARCH STAFF:

Erica J. Burton, Project Leader; burton@ mlml. calstate. edu Jason M. Cope, Graduate Research Assistant; jcope@ mlml. calstate. edu Lisa A. Kerr, Graduate Research Assistant; lkerr@ mlml. calstate. edu Moss Landing Marine Laboratories 8272 Moss Landing Road, Moss Landing, California 95039 ( 831) 632 4419; fax ( 831) 632 4403 CO INVESTIGATORS:

Dr. Ralph J. Larson, Professor Dr. Robert N. Lea Department of Biology California Department of Fish and Game San Francisco State University 20 Lower Ragsdale Drive, Suite 100 San Francisco, California 94132 Monterey, California 93940 ( 415) 338 1027 ( 831) 649 2835; fax ( 831) 649 2894 rlars@ sfsu. edu rlea@ dfg. ca. gov Mr. David VenTresca Mr. Eric Knaggs California Department of Fish and Game California Department of Fish and Game 20 Lower Ragsdale Drive, Suite 100 20 Lower Ragsdale Drive, Suite 100 Monterey, California 93940 Monterey, California 93940 ( 831) 649 2881; fax ( 831) 649 2894 ( 831) 649 7193; fax ( 831) 649 2894 dventres@ dfg. ca. gov ehknaggs@ dfg. ca. gov Page 1 of 1167 5 2220e  Investigators and Affiliations 56 57 58 59 60 61 A B C D E F G H I J K L M N O Page 2 of 1167 5 2220e  Investigators and Affiliations 123456789 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 A B C D E F G H I J K L M PROJECT SUMMARY As part of the recent Marine Life Management Act, the California Department of Fish and Game is required to produce a Fishery Management Plan for the nearshore region as described in the Nearshore Fisheries Management Act. This requires the compilation of a species list that includes the species of fishes for which size limits are prescribed in the Act and other vulnerable species, and a detailed analysis of existing knowledge of the life history ( age, growth, age at maturity, age composition, habitat utilization,

reproduction, recruitment, trophic relationships, population health, and stock structure, including genetics) of nearshore fish species,

especially those subject to the newly developing live fish ( premium) fishery. During this project, we compiled a list of species currently taken in the nearshore fishery and others that could be particularly vulnerable ( n= 124). In addition, we created a life history parameter questionnaire to guide our literature search and compilation of life history parameters, surveyed the existing literature,

constructed a life history database of the nearshore fish species list, and compiled a literature cited document. Gaps in the existing knowledge are identified, and specific research projects are proposed on such subjects as age at maturity, age validation, reproduction,

trophic interactions, genetic variance, and mobility of fish stocks subject to the nearshore fisheries of California. It is hoped that this project will result in additional years of support to answer questions that will provide management agencies with sufficient information to effectively manage this fishery.

BACKGROUND The 1998 Keeley Bill ( AB 1241), which is now law as the Marine Life Management Act ( MLMA), has several requirements,

including size limits for selected species, and status reports and Fishery Management Plans ( FMPs) to be prepared by the California Department of Fish and Game ( CDFG) for important commercial and sport fishery species in California. Funding has been made available from the Pacific States Marine Fisheries Commission ( PSMFC) to support a research institution ( Moss Landing Marine Labs, MLML) to complement the work CDFG is doing on this subject.

Nearshore fisheries have existed in California for decades ( Lea et al. 1999), but a recent fishery for live fish that are used in the restaurant business and shipped overseas, started in southern California in the late 1980s, spread to northern California in the early 1990s, and now has become common in central California ( McKee Lewis 1996). The fishery uses gear ranging from hook and line to pole fishing and traps. The landings of live finfish in California have exhibited an exponential increase from ~ 20,000 to 1,200,000 pounds in the past 7 years, and were worth from ~ $ 20,000 to $ 1,300,000. The number of species landed in this fishery range from ~ 5 to > 50, depending upon location, season, and year. Even though some reports of landings and the status of this fishery exist ( CDFG 1998, Starr et al. 1998) much information is still needed.

The role of life histories in this nearshore fishery has not yet been evaluated. Could the live fish fishery be doomed to failure because it targets mainly on small, immature specimens of shallow, nearshore fishes, some of which are not targeted in any other way?

Certainly, fisheries for juvenile fishes, especially if they target all areas where these juveniles live, cannot last long. The large and increasing numbers of fishes being harvested before most get a chance to reproduce for the first time is of particular concern regarding the live fish fishery. Since rockfish are typically long lived, often take many years to mature ( Echeverria 1987, Love and Johnson 1998, Lea et al. 1999), and are known to have highly variable, successful recruitment ( Karpov et al. 1995), harvesting most individuals before they mature and reproduce can be disastrous for their populations and ultimately the ecosystem.

The live fish fishery targets on sizes and species of fishes that were not heavily fished prior to now ( Lea et al. 1999). It is known that many of these nearshore fishes are very site intensive. Thus, any heavy fishing on their populations, most of which have either home sites or territories and do not move much after settling, could be very deleterious to the local population densities.

In June 1999, the University of California Sea Grant Marine Advisory Program and CDFG sponsored a Workshop on Assessing and Managing Nearshore Fisheries in Santa Barbara. Many fishery biologists attended this useful workshop and produced numerous recommendations, including the need to obtain the best possible data on catch, effort, species and size composition, reproduction,

maturity, location, site specificity, larval dispersal and recruitment, and socio economic variables to enable CDFG to evaluate the trends in this increasing fishery. Page 3 of 1167 5 2220e  Project Summary & Background 123456789 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 A B C D E F G H I J K L 1) Species List The first objective was to produce a list of the species now taken in the nearshore fisheries of California. This list includes the species listed in the MLMA, but also includes species frequently caught that are not on this size limit list and those which could be particularly vulnerable. Among personnel at MLML, and Bob Lea from CDFG, we decided on a list of species having some or one of the following criteria: 1) occurring in 40 fathoms or less; 2) listed in the MLMA; 3) listed in McKee Lewis ( 1996); 4)

common in the sport fishery; 5) common as bait fish; 6) common in public aquaria or the aquaria trade; 7) common as bycatch and prone to overfishing; or 8) marketed incorrectly and possibly prone to overfishing. These criteria could, of course, include almost all of the nearshore fishes off the coast of California. Therefore, our species list was subjectively honed down from > 239 to 124 species ( 42 families); a relatively reasonable number of species to survey during this 8 month project ( Table 1).

2) Literature Survey The second objective was to survey the existing published literature on those subjects recently included in the Fisheries Management Plan ( FMP) and Biological Information Catalog system being developed by David VenTresca ( CDFG, Monterey).

Life history subjects in our survey include, but are not limited to, age and growth ( length/ weight/ age relationships, growth parameters, longevity, age composition); spawning and reproductive status ( parturition period, size/ age at sexual maturation,

fecundity); recruitment ( size at settlement, recruitment to fisheries); population or stock genetics; mortality estimates ( natural,

fishing, total); species habitat associations; trophic interactions ( feeding, predation); and behavior ( competition, movements,

home ranges). We constructed a LIFE HISTORY QUESTIONNAIRE ( APPENDIX A) to guide our literature search, and used this questionnaire to compile life history parameters into a FLAT DATABASE ( MICROSOFT   EXCEL WORKBOOK:

LIFE_ HISTORIES. XLS).

3) Database Format To read, understand, and utilize the database and its format, the LIFE HISTORY QUESTIONNAIRE ( APPENDIX A) should be used while viewing the database. The FLAT DATABASE is composed of 32 MICROSOFT   EXCEL WORKSHEETS within a SINGLE WORKBOOK ( LIFE_ HISTORIES. XLS). Worksheets are organized by family name or species group ( elasmobranchs and flatfishes only). Within a spreadsheet, species are listed in rows within column B ( Scientific name). Life history parameter categories are listed at the top of the spreadsheet( s) from left to right ( following the order of the LIFE HISTORY QUESTIONNAIRE, APPENDIX A), columns A thru DG ( or DH, in some cases). Life history information is then contained within spreadsheet cells. Abbreviations used in the database are listed at the end of APPENDIX A. The literature source for each life history parameter is listed at the bottom of each column. Therefore, the citations are column specific, and are footnoted accordingly ( i. e. renumbered for each column). The corresponding literature can be found in the LITERATURE CITED document ( APPENDIX B), where citations are alphabetized.

In some cases, life history categories are related; therefore, information is listed across a single row ( encompassing several columns). For example, the " von Bertalanffy Growth Parameters " section within " Age and Growth" encompasses several columns ( from " Geographic Area" to " Verification"); and data within one row ( across 16 columns) will refer to one study. Other similar formats are found in the " Maturity," " Fertilization and Spawning Period," " Habitat Association & Trophic Interactions,"

" Genetic Variance," " Recruitment Into Fishery," and " Mortality" sections. Other columns stand alone, and footnoted citations may not correspond across a row, from column to column. Page 4 of 1167 5 2220e  Database Format 123456789 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 A B C D Family Name Scientific Name Common Name Anarhichadidae Anarrhichthys ocellatus eel, wolf Atherinopsidae Leuresthes tenuis grunion, California Atherinopsis californiensis jacksmelt Atherinops affinis topsmelt Batrachoididae Porichthys notatus midshipman, plainfin Bothidae Paralichthys californicus halibut, California Citharichthys sordidus sanddab, Pacific Citharichthys stigmaeus sanddab, speckled Carangidae Seriola dorsalis yellowtail Chimaeridae Hydrolagus colliei ratfish Clinidae Heterostichus rostratus kelpfish, giant Cottidae Scorpaenichthys marmoratus cabezon Hemilepidotus hemilepidotus Irish lord, red Leptocottus armatus sculpin, staghorn Cynoglossidae Symphurus atricauda tonguefish, California Dasyatididae Urolophus halleri stingray, round Embiotocidae Amphistichus argenteus surfperch, barred Embiotoca jacksoni surfperch, black Amphistichus koelzi surfperch, calico Damalichthys vacca surfperch, pile Amphistichus rhodoterus surfperch, redtail Rhacochilus toxotes surfperch, rubberlip Cymatogaster aggregata surfperch, shiner Embiotoca lateralis surfperch, striped Hyperprosopon argenteum surfperch, walleye Phanerodon furcatus surfperch, white Girellidae Girella nigricans opaleye Gobiidae Lythrypnus dalli goby, bluebanded Lythrypnus zebra goby, zebra Gillichthys mirabilis mudsucker, longjaw Haemulidae Anisotremus davidsoni sargo Xenistius californiensis salema Heterodontidae Heterodontus francisci shark, horn Hexagrammidae Hexagrammos decagrammus greenling, kelp Hexagrammos superciliosus greenling, rock Ophiodon elongatus lingcod Table 1. Nearshore fishes of California included in the life history survey ( 124 species, 42 families). List is alphabetized first by Family name, then by common name. Page 5 of 1167 5 2220e  Species List ( Table 1) 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 A B C D Hexanchidae Notorynchus maculatus shark, sevengill Hexanchus griseus shark, sixgill Kyphosidae Hermosilla azurea zebra perch Labridae Oxyjulis californica senorita Semicossyphus pulcher sheephead, California Halichoeres semicinctus wrasse, rock Malacanthidae Caulolatilus princeps whitefish, ocean Mugilidae Mugil cephalus mullet, striped Muraenidae Gymnothorax mordax eel, California moray Myliobatididae Myliobatis californica ray, bat Osmeridae Thaleichthys pacificus eulachon Spirinchus thaleichthys smelt, longfin Spirinchus starksi smelt, night Hypomesus pretiosus smelt, surf Platyrhinidae Platyrhinoidis triseriata thornback Pleuronectidae Platichthys stellatus flounder, starry Parophrys vetulus sole, English Eopsetta jordani sole, petrale Lepidopsetta bilineata sole, rock Psettichthys melanostictus sole, sand Lyopsetta exilis sole, slender Pleuronichthys coenosus turbot, C O Pleuronichthys decurrens turbot, curlfin Hypsopsetta guttulata turbot, diamond Pleuronichthys verticalis turbot, hornyhead Pleuronichthys ritteri turbot, spotted Pomacentridae Chromis punctipinnis blacksmith Hypsypops rubicundus garibaldi Rajidae Raja binoculata skate, big Raja inornata skate, California Raja rhina skate, longnose Raja kincaidii skate, sandpaper Raja stellulata skate, starry Rhinobatidae Rhinobatos productus guitarfish, shovelnose Sciaenidae Menticirrhus undulatus corbina, California Genyonemus lineatus croaker, white Roncador stearnsii croaker, spotfin Umbrina roncador croaker, yellowfin Seriphus politus queenfish Atractoscion nobilis white sea bass Page 6 of 1167 5 2220e  Species List ( Table 1) 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 A B C D Scorpaenidae Sebastes melanops rockfish, black Sebastes chrysomelas rockfish, black and yellow Sebastes mystinus rockfish, blue Sebastes auriculatus rockfish, brown Sebastes dallii rockfish, calico Sebastes nebulosus rockfish, China Sebastes caurinus rockfish, copper Sebastes rubrivinctus rockfish, flag Sebastes carnatus rockfish, gopher Sebastes rastrelliger rockfish, grass Sebastes chlorostictus rockfish, greenspotted Sebastes elongatus rockfish, greenstriped Sebastes semicinctus rockfish, halfbanded Sebastes atrovirens rockfish, kelp Sebastes serranoides rockfish, olive Sebastes maliger rockfish, quillback Sebastes rosaceus rockfish, rosy Sebastes ovalis rockfish, speckled Sebastes hopkinsi rockfish, squarespot Sebastes constellatus rockfish, starry Sebastes saxicola rockfish, stripetail Sebastes nigrocinctus rockfish, tiger Sebastes serriceps rockfish, treefish Sebastes miniatus rockfish, vermilion Sebastes ruberrimus rockfish, yelloweye Scorpaena guttata scorpionfish, spotted Sebastolobus altivelis thornyhead, longspine Sebastolobus alascanus thornyhead, shortspine Scorpididae Medialuna californiensis halfmoon Scyliorhinidae Cephaloscyllium ventriosum shark, swell Serranidae Paralabrax nebulifer bass, barred sand Stereolepis gigas bass, giant sea Paralabrax clathratus bass, kelp Paralabrax maculatofasciatus bass, spotted sand Sphyraenidae Sphyraena argentea barracuda, California Squalidae Squalus acanthias shark, spiny dogfish Squatinidae Squatina californica shark, Pacific angel Stichaeidae Cebidichthys violaceus eel, monkeyface Xiphister mucosus prickleback, rock Stromateidae Peprilus simillimus butterfish, Pacific Syngnathidae Syngnathus leptorhynchus pipefish, bay Syngnathus californiensis pipefish, kelp Hippocampus ingens seahorse, Pacific Torpedinidae Torpedo californica ray, Pacific electric Page 7 of 1167 5 2220e  Species List ( Table 1) 166 167 168 169 A B C D Triakidae Mustelus henlei shark, brown smoothhound Mustelus californicus shark, grey smoothhound Triakis semifasciata shark, leopard Galeorhinus zyopterus shark, soupfin Page 8 of 1167 5 2220e  Species List ( Table 1) 123456789 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 A B C D E F G H I ABBREVIATIONS USED IN DATABASE SE: Standard Error SL: Standard Length FL: Fork Length TL: Total Length BL: Body Length DL: Dressed Length NL: Notocord Length ? L: measurement not defined m: meters cm: centimeters mm: millimeters If information is not provided, the following abbreviations should be used to explain why NA: Not Applicable ( Refers to a life history parameter that is not valid for a species; e. g.

egg stage not applicable for embiotocids).

NG: Not Given ( Refers to a study where similar parameters are provided but not for the specified parameter; e. g. a maturity study with age at first maturity but not age at 50% maturity; an age and growth study where von Bertalanffy growth parameters are provided for males and females, but not for the sexes combined.)

NF: Not Found ( Refers to a parameter where no information was found; either it does not exist, or we could not find the information.) Page 9 of 1167 5 2220e  ABBREVIATIONS 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 A B C D E Family name Scientific name ( Synonym) Common name( s) Geographic range Observed depth range ( m)

Anarhichadidae1 Anarrhichthys ocellatus 1 wolf eel1 Sea of Japan and Aleutian Is. to s. CA ( Imperial Beach) 1 1subtidal to 355 1 Robins et al. 1991 1 Eschmeyer 1998 1 Robins et al. 1991 1 Love et al 2000 1 Love et al 2000 TAXONOMY RANGE Page 10 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 F G H I JKLMNOPQ Length weight equation Growth Parameters Max. reported Geographic area Linf in cm ( SE) k ( SE) to ( SE)

Length conversion equations length ( cm) ( length measurement) Male Female Both Male Female Both Male Female NF NF 1 240 NF NF NF NF NF NF NF NF NF 1 Love et al 2000 AGE & GROWTH Length ( TL= cm & FL, SL= mm) to weight ( g)

equation. * denotes TL = mm or FL, SL= cm Page 11 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 R S T U V W X Y Z AA AB AC Age composition Both Male Female Both Ageing structure / Method Validation Verification Other refs. Longevity ( yrs.) information Sexuality Mating system NF NF NF NF NF NF NF NF 1 4 NF Dioecious 1 Matarese et al. 1989 1 Parra et al. 2000 Promiscuous, but within season mate fidelity1 r2 Page 12 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 AD AE AF AG AH AI AJ AK Mode of reproduction Parental Care M vs. F length Secondary Characteristics Male Female Male Female Oviparous1,2,3 Yes1,2 1Males> Females NF NF NF NF 1 Watson 1996a 1 Hart 1973 1 Garrison & Miller 1982 1 Garrison & Miller 1982 2 Barsukov 1972 2 Watson 1996a 2 Cruys & Cantera 1980 2 Friese 1966 3 Garrison & Miller 1982 3 Fitch & Lavenberg 1975 4 Garrison & Miller 1982 5 Cruys & Cantera 1980 6 Love 1996 TL, FL*, or SL^ in cm ( age in yrs) at Breeding Behavior Sexual dimorphism 1st maturity 50% maturity nest guarded by both sexes3,4,5,6 males have thicker lips, more powerful head1 females show bulging abdominal cavity during breeding season2 Males with lumpy head and oversized jaw2 Page 13 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 AL AM AN AO AP AQ AR AS AT AU AV AW AX AY AZ BA BB Fertilized females present Spawning females present Both reproductive states p Ageing structure / Method Validation Verification Other refs. Geographic Spawning Area Jan. Feb. March April May June July Aug. Sept. Oct. Nov. Dec.

NF NF NF NF California1,2 1 Watson 1996a 2 Love 1996 Fertilization and Spawning Period (^^^^^= peak fertilization;*****= peak spawn; ##### = doubtful spawning records REPRODUCTION Page 14 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 BC BD BE BF BG Spawning Frequency Size specific fecundity equation ( per year) ( TL= cm, SL= mm; * denotes TL= mm; SL= cm) Egg duration NF 1NG ( 3,500 to 5,000) 1,4 10,000 ( NG) NF 1,2 3 to 4 months 27,000 ( NG) 3 2 months 3 5,000 ( NG)

1 Garrison & Miller 1982 1 Matarese et al. 1989 1 Watson 1996a 2 Fitch & Lavenberg 1971 2 Matarese et al. 1989 3 Cruys & Cantera 1980 3 Cruys & Cantera 1980 4 Love 1996 Mean ( range) fecundity in eggs/ g female wt. Fecundity Fecundity range in # eggs (@ cm, grams, or age of female) Page 15 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 BH BI BJ BK BL BM BN BO Size at settlement Size Fish Size Larval duration ( SL in mm) Annual cohort success ( diameter in mm) Habitat Predators ( mm) Habitat 1 2 months 1150 to 610 NF 5.2 to 5.51 demersal, adhesive1,2,3,4,5 118.6 to 47.6 pelagic1,2,3,4 2~ 200 5.2 to 5.42 233.5 to 330 to 35 to 4 30.5 to 1 Garrison & Miller 1982 1 Watson 1996a 1 Watson 1996a 1 Watson 1996a 1 Love 1996 1 Watson 1996a 1 Watson 1996a 2 Marliave 1978 2 Matarese et al. 1989 2 Fitch & Lavenberg 1971 2 Matarese et al 1989 2 Barsukov 1972 3 Fitch & Lavenberg 1975 3 Friese 1966 3 Garrison & Miller 1982 4 Garrison & Miller 1982 4 Cruys & Cantera 1980 4 Cruys & Cantera 1980 5 Cruys & Cantera 1980 rockfish; kelp greenlings1 Eggs DISPERSAL & RECRUITMENT Page 16 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 BP BQ BR BS BT BU BV BW BX Fish Size Food habits Foraging guild Foraging method When / Frequency Predators ( SL mm) Habitat Foraging guild Foraging method When / Frequency carnivore NF NF NF 147.6 to maturity NF carnivore NF NF ( fish, such as walleye pollock) 1 1 Garrison & Miller 1982 1 Watson 1996a 1 Barraclough 1967b ( zooplankton, such as copepods;

larval walleye pollock) 1 Larvae HABITAT ASSOCIATION & TROPHIC INTERACTIONS Food habits Life Stage Juveniles Page 17 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 BY BZ CA CB CC CD CE Adults Food habits Predators Habitat Foraging guild Foraging method When / Frequency Predators Substocks 1lincod; rockfish; cabezon carnivore NF nocturnal1,2 salmon1 NF river otters2 harbor seals3 1 Cruys & Cantera 1980 1 Burge & Schultz 1973 1 Hart 1973 1 Hulberg & Graber 1980 1 Fitch & Lavenberg 1971 2 Barsukov 1972 2 Hulberg & Graber 1980 2 Barsukov 1972 2 Stevens et al. 1984 3 Walton 1979 3 Barsukov 1972 3 Olesiuk et al. 1990 4 Dewees & Gotshall 1974 4 Fitch & Lavenberg 1971 5 Fitch & Lavenberg 1975 5 Fitch & Lavenberg 1975 6 Bodkin 1986 6 Hallacher & Roberts 1985 7 Friese 1966 7 Friese 1966 8 Cruys & Cantera 1980 8 Hobson 1994 9 Love 1996 9 Cruys & Cantera 1980 10 Love 1996 ( crustaceans; echinoderms, mostly sea urchins,

mollusks; fish, such as juvenile rockfishes)

1,2,3,4,5,6,7,8,9,10 rocky reefs in crevices; stony bottoms;

artificial reefs; kelp forests 1,2,3,4,5,6,7,8,9 Page 18 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 CF CG CH CI CJ CK CL CM CN CO Genetic variance Haplotype diversity Mean number of % Polymorphic First Recruitment Full Recruitment ( Geographic Area) alleles per locus ( SE) loci Expected ( SE) Observed ( SE) Approach Effective population size Total length ( age) Total length ( age)

NF NF NF NF NF NF NF NF NF NF Nucleotide diversity (#) Mean heterozygosity GENETICS Page 19 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 CP CQ CR CS CT CU CV CW CX CY CZ DA RECRUITMENT INTO FISHERY Mortality Estimates Rate of Exploitation Method of ge range use Rate of Exploitation Sampling gear Ageing structure / Method Validation Verification Natural ( M) Fishing ( F) Total ( Z) mortality estimation alculate mort exploitation ( u) ratio ( F/ Z) Competition NF NF NF NF NF NF NF NF NF NF NF NF MORTALITY Page 20 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 DB DC DD DE DF DG Interactions / associations with other organisms Tagging studies Resident or Transient Homing ability ( Home range) Schooling Territoriality NF Residential1,2,3 Yes1,2 NF Yes1,2,3,5 Yes, even when juveniles4 1 Parra et al. 2000 1 Parra et al. 2000 1 Miller 1982 1 Parra et al. 2000 2 Hulber & Graber 1980 2 Quast 1968c 2 Miller 1982 3 Gascon & Miller 1981 3 Hulber & Graber 1980 4 Garrison & Miller 1982 5 Cruys & Cantera 1980 symbiotic relationships with rockfish,

lingcod, and sailfin sculpin; compete with Pacific giant octopus1 infested by parasites from the family Gyrodactylidae2 BEHAVIOR Movement Page 21 of 1167 5 2220e  Anarhichadidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 A B C D Family name Scientific name ( Synonym) Common name( s) Geographic range Atherinidae1,2 Leuresthes tenuis 1,2 California grunion Atherinopsidae 3 Atherinidae1,2 Atherinopsis californiensis 1,2 jacksmelt Atherinopsidae 3 Atherinidae1,2 Atherinops affinis 1,2 topsmelt Vancouver Island, nearshore, also in estuaries 1 Atherinopsidae 3 TAXONOMY RANGE silversides, California smelt, horse smelt, blue smelt,

pescado del rey, Peixe rey, Pesce rey 1,2 smelt, little smelt, least smelt, grunion, silversides 1,2,5 San Francisco, northern California to Bahia Magdelena,

southern Baja California 1 Yaquina Bay, Oregon to at least Bahia Magdelena,

southern Baja California 1 bay smelt, rainbow smelt, panzarotto, little smelt, least smelt,

silverside, capron, jack, pescadillo, panzarotti 1,2,3,4 Page 22 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 E F G H I JK Total Length ( cm) Max. reported Geographic area Observed depth range ( m) to weight ( g) equation Length conversion equations length ( cm) ( length measurment) Male Female 1 inshore to 18 m 1Both: W= 0.0089L3 ( immature fall fish and winter fish) ( L= Body lengthNF 119 cm TL NF NF NF 1Both: W= 0.0096L3 ( adult fall fish) ( L= Body length)

1nearshore NF 1SL= 1.167 x TL( 0.832); SE: 2.366 148.6 cm TL NF NF NF 1TL= 1.646 x SL( 1.200); SE: 2.841 1TL= 2.645 x FL( 1.076); SE: 5.453 1FL= 1.509 x TL( 0.925); SE: 5.057 2Both: W= 0.00000886L3.03574 ( L= SL ( mm)) 1SL= 6.217 x TL( 0.863); SE: 1.491 137.6 cm TL NF NF NF 3Both: W= 0.0000992L2.59 ( L= SL ( mm)) ( specimens over 135 mm) 1TL= 7.565 x SL( 1.156); SE: 1.725 1TL= 0.228 x FL( 1.092); SE: 1.272 1FL= 0.399 x TL( 0.914); SE: 1.164 Linf in cm ( SE 1 intertidal and nearshore,

also in estuaries Page 23 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 L M N O P Q R S T U V W X r2 Both Male Female Both Male Female Both Male Female Both Ageing structure / Method Validation Verification NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF AGE and GROWTH Growth Parameters to ( SE)

E) k ( SE) Page 24 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Y Z AA AB AC AD AE Age composition Other refs. Longevity ( yrs.) information Sexuality Mating system Mode of reproduction Parental Care 1,2( larvae), 5 2,3,54 yrs. NF dioecious polygamous1 oviparous2,3 No 83 yrs polyandrous1,2 4,6 3,5,6,1011 yrs. NF dioecious1 NF oviparous1,2,3,4 No 79 10 yrs. ( 30 cm)

911 yrs ( 40.6 cm)

3,7 16 7 yrs. NF dioecious1 polyandrous3 oviparous1,2,3,4 No 5,6,97 8 yrs.

108 yrs. Breeding Behavior Page 25 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 AF AG AH AI AJ AK AL Male : Female SL ratio Secondary Characteristics Male Female Male Female Ageing structure / Method females larger2,3 NF 51 yr. 1 yr. NG NG scales/ surface reading 92 yr. 2 yr. NG NG NG NF NF 32 3 yrs 2 3 yrs NG NG NG 4> 14 cm ( 2 3 yrs) > 14 cm ( 2 3 yrs) NG NG NG 7> 20 cm ( 2 yrs) > 20 cm ( 2 yrs) NG NG NG females larger1,4,5 NF 1,2NG 1,22 3 yrs. NG NG NG 31 2 yrs 2 3 yrs NG NG NG 610 11 cm ( 1 yr.) 10 11 cm ( 1 yr.) NG NG NG 712 cm ( 2 yrs) 12 cm ( 2 yrs) NG NG NG 82 yrs. 2 yrs. NG NG NG 102 yr. 2yr. NG NG scales Sexual dimorphism 1st maturity 50% maturity Total length ( age) at REPRODUCTION Page 26 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 AM AN AO AP AQ AR AS AT AU AV AW AX AY Fertilized eggs present Spawning females present Validation Verification Other refs. Geographic Spawning Area Jan. Feb. March April May June July Aug. Sept.

No between scale comparison NF southern California4,5,7,8,9,10,13 NG NG CalCOFI area2 ***** *****

southern CA to northern Baja18 NG NG NF Tomales Bay, CA5,17 NG NG San Pablo Bay, CA5,11,17 NG NG San Francisco Bay, CA5,15,17 central California6 Moss Landing Harbor, CA15 ***** ***** ***** ***** *****

southern California11,12,15,19,21 ***** *****

California18,20 northeast Pacific1 CalCOFI area2 ***** ***** *****

Pacific coast10 NG NG NF Coos Bay, Oregon3,22 NG NG San Francisco, CA6,11,17 ***** *****

NG NG Moss Landing Harbor, CA15 ***** *****

NG NG central California6 NG NG Anaheim Bay, CA22 No No San Diego Bay, CA22 Newport Bay, CA5,14,16,17 ***** *****

CalCOFI area2 ***** ***** ***** *****

California18 northeast Pacific1 Fertilization and Spawning Period (^^^^^= peak fertilization;*****= peak spawn; ##### = doubtful spawning r Page 27 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 AZ BA BB BC BD BE BF BG Mean Fecundity Size specific Both reproductive states present eggs/ g Number of eggs fecundity eq Oct. Nov. Dec. Spawning Frequency ( weight range) ( at cm, grams, or age) ( length or weight range) Egg duration Multiple spawners1,4,5,9 1,5up to 3,000 eggs every two weeks NF 2,3,5,7,8,9,10,1410 days 2,4,91,000 3,000 eggs 410.5 days( 23 C), 20.5days ( 18 C 31,500 2,200 eggs 67.5 days 1314 days 62,200 ( avg.)

81,600 to 3,600 eggs per spawn Multiple spawners6,7,8,11,13 7over 2,000 eggs per female NF NF 37 19 days ***** *****

***** ***** Multiple spawners2,7,10,11 2200 1,000 eggs per season NF 19 days 411.3 days ( 21 C)

records 33,000 eggs per spawn/ spawning every two weeks, 12,000 24,000 per 11,475 eggs ( 13.7 cm TL) 2,528 eggs ( 17.1 cm TL) 3,11,1235 days ( 55 F) to less than 9 days ( 81 F)

several hundred ( 11 12 cm)

to1,000 (> 16 cm) 2,3 females spawn 4 8 times a season1,3,5 Fecundity Page 28 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 BH BI BJ BK BL Eggs Size at settlement Size Larval duration & location Annual cohort success ( diameter in mm) Habitat 140 days NF NF 2,4,51.5 1.6 mm 230 40 days 31.5 2.2 6,71.75 2.20 viable limit for hatching 14.0 28.5 C5,8 8< 2.0 mm viable limit for hatching 16.0 26.5 C21 eurythermal27 3> 24 days 125 mm NF 1,41.9 2.5 mm 22.0 2.5 mm 31.9 2.5 mm 90.88 2.4 mm 100.9 2.2 mm 4> 18 days NF NF 1,4,101.5 1.7 mm 21.62 mm ( avg.)

31.4 1.7 x 1.5 1.8 mm benthic in estuaries, bays, and lagoons25 found on vegetation in shallow water estuaries and bays25 eggs attached to algae, eel grass hydroids, and submerged aquatic plants4,12,26 eggs are deposited in the sand at night just below the high spring tide line1,2,4,6,9,10,11,12,13,15,17,18,19,22 attached to algae, grass, and submerged aquatic plants4,12,14,16,20,23,24,29 eggs attached to spawning substrate and to one another via adhesive filaments1,12,20,30 eggs spawned in estuaries and along open coast; primarily nighttime spawners3 embryonic development occurs from 12.8 to 26.8 C21 eggs are laid primarily on eelgrass and adhere to macroalgae on tidal flats25,26 eggs attached to spawning substrate and to one another via adhesive filaments1,20,29,30 eggs attached to estuarine and marine plants and other floating objects3,20 eggs develop in sand and hatch when uncovered and agitated by surf of next high tide15,17,18,22,28 duration of in beach sand incubation is dictated by tidally driven wave activity6,8,9 grunion spawn only on three or four nights following each full or new moon6,7,17,26,28 DISPERSAL & RECRUITMENT Page 29 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 BM BN BO BP BQ Predators Fish Size Habitat Foraging guild Foraging method 26.5 20.0 mm SL planktonic1 planktivores1 NF ( larvae retain yolk sac for 4 6 days) 1 reduced salinity tolerance of larvae with age8 isopods, flies, and beetles6 NF 17.5 19.5 mm SL planktonic1 planktivores NF 26.0 19.8 mm SL neuston of bays and shallow coastal waters2,

38.1 mm TL ( size at hatching) found in mesohaline salinities4,

57.5 8.6 mm ( at hatching) 25 mm abundant inshore, common in surface canopy13 shiner surfperch and adult topsmelt514.3 15 mm SL planktonic1 planktivores NF 24.3 21 mm SL ( small crustaceans) 4 35.4 mm TL ( size at hatching)

44.3 18.5 mm tolerates euryhaline conditions5 abundant in upper few inches of kelp13 eel grass beds 14 found in estuarine, bay, and kelp bed habitats and actively school near the 11 surface waters in harbors, along sandy beaches, and in the kelp canopy6,12 ( Stenosomella spp, Labidocera trispinosa ( nauplii), Euterpina acutifrons , Euterpina acutifrons ( nauplii), cirriped nauplii, bivalve veligers, Coscinodiscus spp., Paracalanus parvus , and Oncea spp. ) 2 54.3 4.9 mm ( at hatching)  18.5 mm found over soft, unconsolidated sediments and other substrates11 planktonic, school near the surface, found in embayments11 ( copepod nauplii, tintinnid protozoans, cyclopoid copepods, and harpactacoid copepods) 1,3 beetles, possibly annelids,

amphipods, and isopods4 larvae showed reduced viability below and above 25.2 C9 Larvae Food habits ( Coscinodiscus spp., Peridinium spp.,

Stenosomella spp., Euterpina acutifrons,

E. acutifrons ( nauplii), Acartia tonsa , unidentified copepods) 2 found in neuston and upper water column in coastal waters2,3,7 sand worms, shore birds, isopods,

flies, beetles, and ground squirrels1,3 sand worms, shore birds, and ground squirrels2 found in neuston and upper water column in bays and coastal waters2 collected at surface during the day with a shift to lower depths occurring at night10 Page 30 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 BR BS BT BU BV When / Frequency Predators Fish Size Habitat Foraging guild fed during early evening1 131.0 38.8 mm SL found in shallow waters nearshore6 planktivores1 fed during early evening1 kelp bass and yellowtail2 125.8 37.0 mm SL planktivores4 NF NF 130.1 39.3 mm SL planktivores4,6 218.5 120.0 mm ( algae and kelpfly larvae) 2,5 ( algae and copepods) 3 abundant in surf grass beds5 ( insects, amphipods, and copepods ( 51 75 mm))

resident populations in lagoons10 found in neritic, estuarine, and bay environments9 large schools of juveniles reside in estuaries and bays in summer; in fall,

they migrate to coastal waters3 surface waters in harbors, along sandy beaches, and in the kelp canopy2,11 occur along sandy beaches, in kelp beds, over rocky reefs, and around 9 halibut, bass, white croakers,

and other large predators1 Juveniles preference for modal temperature of 25 C7 shallow inshore waters in bays and along coast1 mostly in shallow polyhaline waters of bays and along coast1 shallow inshore waters used as nursery grounds4 found by night and day in surface waters close to kelp forests8 ( foraminiferans, protozoans, cladocerans,

ostracods, copepods, cirrepedian larvae,

gammarid amphipods, caprellid amphipods,

Callianassa larvae, decapod zoea, dipteran adults, dipteran larvae, algae, and detritus) 6 HABITAT ASSOCIATION & TROPHIC INTERACTIONS Life Stage Page 31 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 BW BX BY BZ CA Foraging method When / Frequency Predators Habitat Foraging guild NF NF planktivores1,2 omnivores8 found in midkelp environment13 NF kelp bass and yellowtail4 omnivores8,17 ( plankton and small fishes) 1 ( Engraulis mordax ) 3,9 ( small fish and crustaceans) 5 ( small crustaceans) 7,19 ( green algae and diatoms) 11 ( zooplankton and flora) 11 NF kelp bass3 omnivores8,17 ( plankton, algae, and insect larvae) 1,5 usually found near surface2 ( Engraulis mordax ) 3,9 abundant in surf grass beds8 ( planktonic crustaceans) 4,5,7,14 found in euryhaline conditions4,7,10,12 ( crustaceans, amphipods, and copepods) 8 ( plant material, algae, diatoms, and small crustacea) 8,17 ) 7 kelp column and canopy11 ( zooplankton and flora) 11 ( detritus, diatoms, and plant material) 12 found in neritic, estuarine, and bay environments12 ( diatoms, filamentous algae, detritus, chironomic midge larvae, and amphipods) 6 found in kelp beds, harbors, and sandy beaches3 opportunistic feeder, characterized as a herbivore,

detrivore, and low level carnivore10 ( calanoid copepods, anchovies, polychaete worms,

fish eggs, zoea, nauplei, cypris larvae, and 13 ( Enteromorpha spp ., and other filamentous algae,

benthic diatoms, small crustaceans, and detritus) 17 occurs in estuaries, around kelp beds,

in bays, and in open water2,6,7 often found in lower reaches of coastal streams5 Food habits Fo California halibut, sand bass, white croakers, and other large predators,

including man1,2 primarily filter feeders,

occasionally more predatory1 schools nearshore, usually at or near surface along open coast and in bays1 primarily filter feeders,

occasionally more predatory1 schools nearshore along open coast and in bays1 schools nearshore, especially in bays and estuaries and near kelp beds1,6 feed near water surface, but feed on bottom when in shallow water5 found in sand bottom kelp holdfast,

midkelp, and rocky bottom kelp bed habitat13 aggregates by day in surface waters close to kelp forests, but at night larger Page 32 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 CB CC CD CE Foraging method When / Frequency Predators Substocks NF Diurnal feeders1 least terns and doublecrested cormorants1 NF larger fish and humans12 Diurnal feeders2 NF larger fish, such as kelp bass and yellowtail3 western gull and Caspian and Fosters' terns8 harbor seal9,10 Caspian terns11 sooty shearwaters13 Diurnal feeders2,3,4,5 several subspecies1 larger fish, such as kelp bass and yellowtail3 olive rockfish4 birds, such as sea gulls5 California gull chicks7 western gull and Caspian and Fosters' terns8 harbor seals9 Caspian terns11 typically feed within 10 15 feet of the surface over shallow rocky areas or in kelp beds4 planktonic feeders ( ocean dwellers),

bottom feeder ( estuarine dwellers) 2,1 filter feeders and opportunistic particulate predators3 feed throughout the water column, but especially on or near the bottom3 Atherinops affinis affinis, subspecies in salt ponds of San Francisco2 feed throughout the water column, but especially on or near the bottom3 piscivorous birds and fishes, including yellowtail6,14 Adults ood habits marine fishes ( yellowtail), marine mammals ( harbor seal,

northern fur seals), and sea birds ( brown pelicans, western gulls,

least terns Brandt's and doublecrested cormorants) 1 yellowtail, kelp bass, sharks, other piscivorous fishes,

piscivorous birds ( brown pelicans and gulls), and marine mammals6 halibut, bass, white croakers, and other large predators2 California sea lions, harbor seals, least terns, and Brandt's and double crested cormorants1 feed at bottom, swallowing sand and shell fragments along with plant and animal material ( bay dwellers) 1 filter feeders and opportunistic particulate predators3 Atherinops affinis oregonia ( ranges from Oregon to Humbolt Bay), A. affinis affinis ( ranges from San Francisco and surrounding areas to Monterey), A. affinis littoralis ( integrades with A. affinis affinis from Monterey to Santa Barbara and extends south to San Diego), and A. affinis insularum ( Channel Islands form) 3,4 Page 33 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 CF CG CH CI CJ CK CL Genetic variance Haplotype diversity Mean number of % Polymorphic ( Geographic Area) alleles per locus ( SE) loci Expected ( SE) Observed ( SE) Approach NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF GENETICS Nucleotide diversity (#) Mean heterozygosity Page 34 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 CM CN CO CP CQ CR CS RECRUITMENT INTO FISHERY First Recruitment Full Recruitment Effective population size Fishery / Total Length ( age) Fishery / Total Length ( age) Sampling gear Ageing structure / Method Validation Verification NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF Page 35 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 CT CU CV CW CX CY CZ DA Mortality Estimates Rate of Exploitation Method of Age range used Rate of Exploitation Natural ( M) Fishing ( F) Total ( Z) mortality estimation to calculate mortality exploitation ( u) ratio ( F/ Z) Competition NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF MORTALITY Page 36 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 DB DC DD DE DF DG Interactions / associations with other organisms Tagging studies Resident or Transient Homing ability ( Home range) Schooling Territoriality NF NF Yes2,3,12 NF ectoparasite: Argulus melanstictus 4 parasite: trematode ( Asymphylodora atherinopsidis) 7 parasites: 15 families, 16 spp. ( see reference) 12 parasite: trematode ( Asymphylodora atherinopsidis) 7 NF NF Yes2,3,4,6,7,12 NF parasite: nematode10 often schools with topsmelt3,8,11 larvae school near surface waters7 cleaning symbiosis between topsmelt, Atherinops affinis ,

cleaners and gray whale, Eschrichtius robustus 1,2 NF NF Yes1,2,3,4,6,11 NF schools near surface waters5,7,9,10,14 larvae school near surface waters7,11 often found schooling with jacksmelt3,8 often associated with schools of shiner perch5 school by size10 this species is commonly impinged on power plant intake screens10 moderate to dense schools above sand or mud bottoms13 resident; limited marking experiments idicate the grunion are nonmigratory3 move inshore and into bays and estuaries to spawn during late winter and early spring2 this species is seldom found far from shore2 adult Oxyjulus californica and juvenile Herosilla azurea,

Girella nigricans, and Medialuna californiensis have been observed cleaning adult Atherinops affinis8 highly susceptible to adverse effects from pollution and habitat modification10,11 may school in separate sexes,

particularly during spawning season8 may school in separate sexes,

particularly during spawning season8 topsmelt and grunion occur together in larval and juvenile stages9 unique spawning behavior makes them vulnerable to any reduction or change in spawning habitat11 parasites: copepods( Bomolachus pectinatus , Caligus olsoni , and Clauellopsis ), brachiuran ( Argulus melanostictus ), isopods( Lironeca californica and Nerocila californica ), trematodes ( Asymphylodora atherinosidis and Lepocredium manteri ) 1 parasites: Lepocreadium manteri, hyperparasite ( Nosema lepocreadii) 6 associated with Fundulus parvipinnis, Clevandia ios,

Gillichthys mirabilis, and Gambusia affinis 3 school in murky waters, preferable over sand14 school separately from jacksmelt, but in the same area10 topsmelt and grunion occur together in larval and juvenile stages9 many move into estuaries during spring and summer and into shallow, open coasts in fall and winter1,2 BEHAVIOR Movement Page 37 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 DH DI DJ DK DL DM DN DO Page 38 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 A B C D 1 Eschmeyer 1998 1 Robins et al 1991 1 Eschmeyer and Herald 1983 1 Love et al 2000 2 Robins et al 1991 2 Eschmeyer 1998 2 Gates and Frey 1974 3 Pers comm R. N. Lea, CDFG, Monterey 3 Hart 1973 4 Love 1996 5 Thompson 1919 Page 39 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 E F G H I JK 1 Love et al 2000 1 Clark 1925 1 Karpov and Kwiecien 1988 1 Love et al 2000 2 Quast 1968a 3 Schultz 1933 Page 40 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 L M N O P Q R S T U V W X Page 41 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 Y Z AA AB AC AD AE 1 Fritzsche et al 1985 1 Matarese et al 1980 1 Emmett et al 1991 1 Gregory 1992 1 Matarese et al 1980 2 Spratt 1986 2 Breder and Rosen 1966 2 Moser 1996a 3 Love 1996 3 Turner 1960 3 Love 1996 3 Fronk 1969 4 Fitch and Lavenberg 1971 4 Emmett et al 1991 4 Clark 1929 5 Gregory 1992 5 Clark 1925 6 DeLeon 1999a 6 Emmett et al 1991 7 Clark 1929 7 Schultz 1933 8 Clark 1925 9 Ruagh 1976 10 Emmett et al 1991 2 Vasquez Yeomans and Sosa Cordero 1992 Page 42 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 AF AG AH AI AJ AK AL 1 Hart 1973 1 Fitch and Lavenberg 1975 2 Fitch and Lavenberg 1971 2 Gregory 1992 3 Thompson 1919 3 DeLeon 1999a 4 Feder et al 1974 4 Clark 1929 5 Schultz 1933 5 Clark 1925 6 Carpelan 1955 7 Emmett et al 1991 8 Turner 1960 9 Fritzsche et al 1985 10 Schultz 1933 Page 43 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 AM AN AO AP AQ AR AS AT AU AV AW AX AY 1 Matarese et al 1989 2 Moser 1996a 3 Hart 1973 4 Spratt 1986 5 Love 1996 6 Wang 1981 7 Walker 1952 8 Fritzsche et al 1985 9 Fitch and Lavenberg 1971 10 Gregory 1992 11 DeLeon 1999a 12 Clark 1929 13 Clark 1925 14 Horn and Allen 1985 15 Ruagh 1976 16 Fronk 1969 17 Emmett et al 1991 18 C. D. F. G. 1982 19 Limbaugh 1955 20 Turner 1960 21 Feder et al 1974 22 Scultz 1933 Page 44 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 AZ BA BB BC BD BE BF BG 1 Spratt 1986 1 Spratt 1986 1 Fritzsche et al 1985 1 Matarese et al 1989 2 Love 1996 2 Love 1996 2 Fronk 1969 2 Spratt 1986 3 Walker 1952 3 Wang 1981 3 Emmett et al 1991 3 Love 1996 4 Fritzsche et al 1985 4 Walker 1952 4 Hubbs 1965 5 Fitch and Lavenberg 1971 5 Fitch and Lavenberg 1971 5 Ehrlich and Farris 1971 6 Breder and Rosen 1966 6 Thompson 1919 6 David 1939 7 DeLeon 1999a 7 Emmett et al 1991 7 Walker 1952 8 Clark 1929 8 Daugherty 1960a 8 Fritzsche et al 1985 9 Clark 1925 9 Moffatt and Thomson 1978b 9 Fitch and Lavenberg 1971 10 Fronk 1969 10 Gregory 1992 11 Emmett et al 1991 11 DeLeon 1999a 12 Daugherty 1960a 12 Emmett et al 1991 13 Turner 1960 13 Limbaugh 1955 14 Daugherty 1960a Page 45 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 BH BI BJ BK BL 1 Vasquez Yeomans and Sosa Cordero 1992 1 Emmett et al 1991 1 Matarese et al 1989 1 Moser 1996a 2 Reynolds et al 1976 2 Ahlstrom et al 1984 2 Spratt 1986 3 Clark 1929 3 Moser 1996a 3 Love 1996 4 Ehrlich et al 1978 4 Wang 1981 5 Ehrlich and Farris 1971 5 David 1939 6 Moffatt and Thomson 1978b 6 Moffatt and Thomson 1978b 7 Walker 1952 7 Fritzsche et al 1985 8 Fritzsche et al 1985 8 Breder and Rosen 1966 9 Thompson et al 1936 9 Clark 1929 10 Fitch and Lavenberg 1971 10 Emmett et al 1991 11 Middaugh et al 1983 12 Gregory 1992 13 Breder and Rosen 1966 14 DeLeon 1999a 15 Moffatt and Thomson 1978a 16 Horn1980 17 Clark 1925 18 Spratt 1981 19 Thomson and Muench 1976 20 Ruagh 1976 21 Ehrlich and Muszynski 1981 22 Thompson 1919 23 Fitch and Lavenberg 1975 24 Fronk 1969 25 Emmett et al 1991 26 C. D. F. G. 1982 27 Ehrlich et al 1979 28 Daugherty 1960a 29 Turner 1960 30 Feder et al 1974 Page 46 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 BM BN BO BP BQ 1 Fritzsche et al 1985 1 Matarese et al 1980 1 Moser 1996a 1 Fritzsche et al 1985 2 Fitch and Lavenberg 1971 2 Moser 1996a 2 Wang 1981 2 Watson and Davis 1989 3 Gregory 1992 3 Middaugh et al 1990 3 Fritzsche et al 1985 3 Kauffman et al 1981 4 Thompson 1919 4 DeLeon 1999a 4 Middaugh et al 1990 4 Feder et al 1974 5 Fronk 1969 5 Emmett et al 1991 5 Middaugh and Shenker 1988 6 Daugherty 1960aa 6 Greogory 1992 7 Watson and Davis 1989 8 Reynolds et al 1976 9 Ehrlich and Muszynski 1981 10 Kauffman et al 1981 11 Emmett et al 1991 12 Turner 1960 13 Feder et al 1974 14 Hoffman 1986 Page 47 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 BR BS BT BU BV 1 Watson and Davis 1989 1 Fitch and Lavenberg 1971 1 Moser 1996a 1 Wang 1981 1 Fritzsche et al 1985 2 Turner 1960 2 Emmett et al 1991 2 Gregory 1992 2 Fitch and Lavenberg 1975 3 DeLeon 1999a 3 Horn and Allen 1985 4 Horn 1980 4 Ruagh 1976 5 DeMartini 1981 5 Quast 1968d 6 Reynolds et al 1977 6 Klingbeil 1972 7 Reynolds et al 1977 7 Mitchell 1953 8 Hobson et al 1981 9 Emmett et al 1991 10 Allen and Horn 1975 11 Turner 1960 Page 48 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 BW BX BY BZ CA microcarnivore15 sensitive to low and high temperatures15 1 Ruagh 1976 1 Fritzsche et al 1985 1 Moser 1996a 1 Love 1996 2 Fitch and Lavenberg 1971 2 Love 1996 2 Fritzsche et al 1985 3 Love and Ebeling 1978 3 Gregory 1992 3 Coyer and Hall 1993 4 Turner 1960 4 Fronk 1969 4 Fitch and Lavenberg 1975 5 Moyle 1976 5 Gregory 1992 6 Carlisle et al 1964 6 Moyle 1976 7 DeLeon 1999a 7 Carlisle et al 1964 8 DeMartini 1981 8 DeLeon 1999a 9 Hobson et al 1981 9 Coyer and Hall 1993 10 Carpelan 1955 10 Allen, L. G. 1982 11 Quast 1968b 11 Barry et al 1996 12 Emmett et al 1991 12 Horn and Allen 1985 13 Limbaugh 1955 13 Ruagh 1976 14 Feder et al 1974 14 Fitch and Lavenberg 1975 15 Doudoroff 1945 15 Quast 1968b 16 Quast 1968d 17 Emmett et al 1991 18 Klingbeil 1972 19 Turner 1960 occur along sandy beaches, in kelp beds, over rocky reefs, and around found in sand bottom kelp holdfast,

midkelp, and rocky bottom kelp bed habitat13 ( planktonic crustaceans, larval bryozoans, algae, and insect larvae) 16 ( diatoms, algae, mysids, jacksmelt eggs, and calanoid copepods) 13 ( diatoms, copepods, ostracods, mysids, algae,

amphipods, and zoea larvae) 13 estuary/ bay inhabitants ( algae, diatoms, small crustaceans, benthic invertebrates); oceanic inhabitants ( gammarid and caprellid amphipods,

mysids, ostracods, copepods, and crustacean larvae) 17 ( detritus, foraminiferans, gastropods, ostrocods,

copepods, cumaceans, tanaidaceans, gammarid amphiods, caprellid amphipods, Callianassa larva,

dipterans, A. affinis eggs, and algae) 18 primarily a marine fish that prefers estuaries, bays, sloughs, and lagoons12 individuals disperse close beneath the water surface over deeper water9 Page 49 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 CB CC CD CE 4 subspecies in California: San Francisco topsmelt, Atherinops affinis affinis ; bay topsmelt, A. affinis littoralis ; kelp topsmelt, A. affinis cedroscensis ; and the island topsmelt, A. affinis insularum 5 1 Fronk 1969 1 Fritzsche et al 1985 1 Love 1996 1 Fronk1969 2 DeLeon 1999a 2 Coyer and Hall 1993 2 Fitch and Lavenberg 1971 2 Carpelan 1955 3 Ruagh 1976 3 Hobson et al 1981 3 Carlisle et al 1964 3 Fronk 1969 4 Quast 1968d 4 Emmett et al 1991 4 Love and Ebeling 1978 4 Turner 1960 5 Emmett et al 1991 5 Hobson et al 1981 5 Fronk 1969 5 Feder et al 1974 6 Emmett et al 1991 6 Hubbs 1918 7 Dierks 1990 8 Harvey 1982 9 Trumble 1995 10 Torok 1994 11 Parkin 1998 12 Daugherty 1960a 13 Krasnow 1978 14 Feder et al 1974 6 subspecies: Atherinops affinis littoralis,

A. affinis magdalenae, A. affinis cedroscensis, A. affinis guadalupae, A.

affinis affinis , and A. affinis insularum 6 feed near water surface, but feed on bottom when in shallow water5 Page 50 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 CF CG CH CI CJ CK CL Page 51 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 CM CN CO CP CQ CR CS Page 52 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 CT CU CV CW CX CY CZ DA Page 53 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 DB DC DD DE DF DG due to spawning habitat any pollution or alteration of bays and harbors may have a deterious effect on the species 11 parasites: 18 families, 19 spp. ( see reference) 12 parasite: isopod ( Lironeca californica ) 13 1 Fritzsche et al 1985 1 Love 1996 1 Matarese et al 1980 2 Swartz 1981 2 Emmett et al 1991 2 Moser 1996a 3 Allen, L. G. 1982 3 C. D. F. G. 1982 3 Love 1996 4 Benz et al 1995 4 Coyer and Hall 1993 5 Moyle 1976 5 Fitch and Lavenberg 1975 6 Canning and Olson 1980 6 Gregory 1992 7 Olson 1977 7 DeLeon 1999a 8 Sikkel 1986 8 Ruagh 1976 9 Ehrlich et al 1978 9 Fitch and Lavenberg 1975 10 Emmett et al 1991 10 Fronk 1969 11 C. D. F. G. 1982 11 Emmett et al 1991 12 Love and Moser 1983 12 Limbaugh 1955 13 Bennett 1993 13 Turner 1960 14 Feder et al 1974 Page 54 of 1167 5 2220e  Atherinopsidae 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 DH DI DJ DK DL DM DN DO Page 55 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 A B C D Page 56 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 E F G H I JK Page 57 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 L M N O P Q R S T U V W X Page 58 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 Y Z AA AB AC AD AE Page 59 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 AF AG AH AI AJ AK AL Page 60 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 AM AN AO AP AQ AR AS AT AU AV AW AX AY Page 61 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 AZ BA BB BC BD BE BF BG Page 62 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 BH BI BJ BK BL Page 63 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 BM BN BO BP BQ Page 64 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 BR BS BT BU BV Page 65 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 BW BX BY BZ CA Page 66 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 CB CC CD CE Page 67 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 CF CG CH CI CJ CK CL Page 68 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 CM CN CO CP CQ CR CS Page 69 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 CT CU CV CW CX CY CZ DA Page 70 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 DB DC DD DE DF DG Page 71 of 1167 5 2220e  Atherinopsidae 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 DH DI DJ DK DL DM DN DO Page 72 of 1167 5 2220e  Atherinopsidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 A B C D E Family name Scientific name ( Synonym) Common name( s) Geographic range Observed depth range ( m)

Batrachoiddidae1,2 Porichthys notatus 1,2 plainfin midshipman 1 intertidal to 383 m 1 Robins et al 1991 1 Robins et al 1991 1 Eschmeyer and Herald 1983 1 Love et al 2000 1 Love et al 2000 2 Eschmeyer 1998 2 Eschmeyer 1998 2 Gates and Frey 1974 3 Hart 1973 southeastern Alaska ( Sitka)? British Columbia ( Smith Sound) to Bahia Magdelena, southern Baja California 1 northern midshipman, bullhead,

singing fish, toad fish, grunter,

midshipman 1,2,3 TAXONOMY RANGE Page 73 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 F G H I J K L M Total Length ( cm) Max. reported Geographic area to weight ( g) equation Length conversion equations length ( cm) ( length measurment) Male Female Both Male 1W= 0.0274 L0.605 ( sessile larvae; L= SL) 1TL= 1.091 SL + 4.8 138 cm TL NF NF NF NF NF 2W= 0.0087L3.04 1 Crane 1981 1 Demartini 1990 1 Love et al 2000 2 Phillips 1973 Linf in cm ( SE)

Page 74 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 N O P Q R S T U V W X r2 Female Both Male Female Both Male Female Both Ageing structure / Method Validation Verification NF NF NF NF NF NF NF NF NF NF NF k ( SE) AGE and GROWTH to ( SE) Growth Parameters Page 75 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 Y Z AA AB AC AD AE Age composition Other refs. Longevity ( yrs.) information Sexuality Mating system Mode of reproduction Parental Care 1,2,3 17 yrs. NF dioecious polygamous1,3,5 oviparous1,2 nest guarded by male1,2,3,5,6,7,8,9,11,12 polygynous2,3,4 1 Campana 1984b 1 Love 1996 1 Hart 1973 1 Moser 1996a 1 Moser 1996a 2 Phillips 1973 2 Demartini 1988 2 Garrison and Miller 1982 2 Love 1996 3 Ibara 1967 3 Arora 1948 3 Arora 1948 4 Knapp et al 1999 4 Knapp et al 1999 5 Garrison and Miller 1982 5 Breder and Rosen 1966 6 Baxter 1999c 7 Phillips 1973 8 Garrison and Miller 1982 9 Hubbs 1920a 10 Turner et al 1969 11 Allen, M. J. 1982 12 Feder et al 1974 male clears area for egg laying and guards nest10 males guard nests with clutches at very different stages of development4,8 androgens play an important role in parental care behavior of males4 Breeding Behavior Page 76 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 AF AG AH AI AJ AK AL Male : Female SL ratio Secondary Characteristics Male Female Male Female Ageing structure / Method males larger1,2 13 4 yrs 3 4 yrs NG NG NG 218 19 cm 14 15 cm NG NG NG large sound producing muscles in males2 1 Fitch and Lavenberg 1971 1 Brantley and Bass 1994 1 Phillips 1973 2 Ibara 1967 2 Ibara 1967 2 Ibara 1967 1st maturity 50% maturity Sexual dimorphism two male reproductive morphs,

dimorphic body and vocal muscle traits 1 Total length ( age) at REPRODUCTION Page 77 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 AM AN AO AP AQ AR AS AT AU AV AW AX AY AZ Fertilized eggs present Spawning females present Both reprod Validation Verification Other refs. Geographic Spawning Area Jan. Feb. March April May June July Aug. Sept. Oct.

NG NG NF CalCOFI area1 NG NG San Franscisco Bay, CA3 central California2 ***** *****

Santa Barbara, CA2 California4 1 Moser 1996a 2 Love 1996 3 Baxter 1999c 4 Feder et al 1974 Fertilization and Spawning Period (^^^^^= peak fertilization;*****= peak spawn; ##### = doubtful spawning records Page 78 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 BA BB BC BD BE BF BG Mean Fecundity Size specific ductive states present eggs/ g Number of eggs fecundity eq Nov. Dec. Spawning Frequency ( weight range) ( at cm, grams, or age) ( length or weight range) Egg duration 1,5200 500 eggs per nest NF 1F= 14+ 5.13W, r= 0.698 ( 1978) 1,3,4,5,616 20 days 2up to 1,600 eggs per nest 1F= 40+ 3.91W, r= 0.652 ( 1979) 215 days 3,6200 eggs( avg. female) 1F= 29+ 4.92W, r= 0.741 ( 1984)

1F= 17+ 5.95W, r= 0.698 ( 1985)

1F= 37+ 4.28W, r= 0.565 ( 1986)

480 200 eggs per season 720 800 eggs per nest 1 Fitch and Lavenberg 1971 1 Hart 1973 1 DeMartini 1990 1 Hart 1973 2 Baxter 1999c 2 Crane 1986 2 Love 1996 3 Phillips 1973 3 DeMartini 1988 3 Crane 1981 4 Love 1996 4 Phillips 1973 5 Arora 1948 5 Garrison and Miller 1982 6 Fitch and Lavenberg 1971 6 Ibara 1967 7 Garrison and Miller 1982 31,000 1,200 eggs per nest ( represents production of 5 6 females)

single spawner, both sexes die after spawning1,2 females usually die after spawning, males may spawn more than once3 Fecundity Page 79 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 BH BI BJ BK BL BM Eggs Size at settlement Size Larval duration & location Annual cohort success ( diameter in mm) Habitat Predators NF NF 14.0 6.0 mm x 4.0 8.0 mm eggs attached to nest wall1 crabs and snails1 24.0 7.5 mm 36.0 mm 44.0 6.0 mm 1 Crane 1981 1 Moser 1996a 1 Moser 1996a 1 Crane 1986 2 Wang 1981 2 Hart 1973 3 Breder and Rosen 1966 3 Wang 1981 4 Hubbs 1920a 4 Crane 1981 2 5 Arora 1948 6 Fitch and Lavenberg 1971 7 Garrison and Miller 1982 8 Turner et al 1969 9 Hubbs and Schultz 1939 10 Limbaugh 1955 11 Allen, M. J. 1982 12 Feder et al 1974 demersal, adhesive eggs spawned in shallow intertidal coastal waters3,5 14 5 weeks larvae break away from disc and swim free of the eggs laid on the underside of intertidal and subtidal rocks4,7,8,10,12 eggs spawned in shallow water or intertidal zone in a nest2,9,11 DISPERSAL & RECRUITMENT Page 80 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 BN BO BP BQ BR BS Fish Size Habitat Foraging guild Foraging method When / Frequency Predators 17.0 19 mm larvae attached to nest wall1 planktivore3 NF NF 27.0 30 mm TL ( crustacean larvae) 1,2 ( copepods) 4 ( crustacean larvae and zooplankton) 5 1 Moser 1996a 1 Moser 1996a 1 Hart 1973 1 Arora 1948 2 Wang 1981 2 Wang 1981 2 Crane 1986 3 Arora 1948 3 Love 1996 4 Crane 1981 5 Allen, M. J. 1982 attached to substrate in shallow intertidal coastal waters2,3 Food habits attached larvae feed on yolk reserves, free swimming larvae feed on small crustacean larvae and other l kt 5 Larvae Page 81 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 BT BU BV BW BX BY Fish Size Habitat Foraging guild Foraging method When / Frequency Predators NF planktivore1 NF NF harbor seals1 ( copepods) 1 young gradually migrate to deeper waters2 ( copepods and amphipods) 2 ( small crustaceans and zooplankton) 3,4 seek cover by burrowing in the mud or sand4 1 Wang 1981 1 Ibara 1968 1 Torok 1994 2 Phillips 1973 2 Phillips 1973 3 Garrison and Miller 1982 3 Garrison and Miller 1982 4 Limbaugh 1955 avoid light and remain buried in the sand during the day, coming out at night to feed 3 living on bottom of shallow and deeper coastal waters1 Juveniles Food habits HABITAT ASSOCIATION & TROPHIC INTERACTIONS Life Stage Page 82 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 BZ CA CB CC Habitat Foraging guild Foraging method When / Frequency carnivore detect prey by motion rather than sight1 nocturnal1,2,3,4,5,6 ( fishes and crustaceans) 1,6 feeds in midwater2 ( krill, mysid shrimps, amphipods, and squids) 2 ( shrimplike crustaceans and small fish( anchovies)) 3 ( euphausiids and mysid shrimp) 5 exhibit diel vertical migration4,5 ( euphausiids, mysids, and gammarids) 7 found in rocky bottom kelp bed habitat7 males fast while guarding eggs6 1 Moser 1996a 1 Hart 1973 1 Ibara 1968 1 Love 1996 2 Love 1996 2 Love 1996 2 Phillips 1973 2 Fitch and Lavenberg 1971 3 Fitch and Lavenberg 1971 3 Fitch and Lavenberg 1971 3 Ibara 1967 3 Phillips 1973 4 Ibara 1968 4 Phillips 1973 4 Cailliet et al 1979 4 Ibara 1967 5 Phillips 1973 5 Ibara 1967 5 Allen, M. J. 1982 5 Cailliet et al 1979 6 Hubbs and Schultz1939 6 Hubbs 1920a 6 Garrison and Miller 1982 6 Allen, M. J. 1982 7 Limbaugh 1955 7 Allen, M. J. 1982 8 Allen, M. J. 1982 9 Feder et al 1974 emerges at night, swims upward and snaps at the plankton as it drifts and swims through its midst3 sand and mud bottoms; bury themselves in substrate during the 2 3 8 9 vertical migrator, moving off the bottom in search of prey at night4,5 occurs in deep water in the north and south of range6 ambush prey from a partially buried position on the bottom and drift in the water column and snap up prey that comes near5 demersal, living primarily on soft bottom of the continental shelf; near rocky areas during spawning1 Adults Food habits Page 83 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 CD CE CF CG CH CI Genetic variance Haplotype diversity Mean number of % Polymorphic Predators Substocks ( Geographic Area) alleles per locus ( SE) loci harbor seals1,7,8 NF NF NF NF NF rock cod4 California gull chicks5 western gull, and Caspian and Fosters' terns6 California sea lion9 1 Olesiuk et al 1990 2 Love 1996 3 Fitch and Lavenberg 1971 4 Hubbs 1920a 5 Dierks 1990 6 Harvey 1982 7 Trumble 1995 8 Torok 1994 9 Hawes 1983 kelp bass, northern elephant seals, California sea lions,

northern fur seals, Pacific white sided dolphins, Steller sea lions, pygmy sperm whales, harbor porpoises,

Pacific striped dolphins, cormorants, western gulls,

common murres, arctic loons and pink footed shearwaters2 rockcod, giant seabass, lingcod, sea lions and numerous other large predators3 GENETICS Nucleotide diversity (#)

Page 84 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 CJ CK CL CM CN CO CP CQ RECRUITMENT INTO FISHERY First Recruitment Full Recruitment Expected ( SE) Observed ( SE) Approach Effective population size Fishery / Length ( age) Fishery / Length ( age) Sampling gear eing structure / Meth NF NF NF NF NF NF NF NF Mean heterozygosity Page 85 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 CR CS CT CU CV CW CX CY Mortality Estimates Rate of Ex Method of Age range used Rate of Validation Verification Natural ( M) Fishing ( F) Total ( Z) mortality estimation to calculate mortality exploitation ( u)

NF NF NF NF NF NF NF NF MORTALITY Page 86 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 CZ DA DB DC DD xploitation Exploitation Interactions / associations ratio ( F/ Z) Competition with other organisms Tagging studies Resident or Transient NF NF NF male midshipman humming attracts females 2,5 photophores may play a role in mating behavior6,8 possesses venom apparatus, not toxic to humans 5 possesses venom apparatus, may have toxic action 7 parasites: 5 families, 10 spp. ( see reference) 10 parasites: digenea ( Tubluovesicula lindbergi) 11 1 Demartini 1991b 1 Love 1996 2 Crane 1986 2 Garrison and Miller 1982 3 Ibara et al 1983 3 Ibara 1967 4 Harper 1999 4 Hubbs 1920a 5 Phillips 1973 6 Ibara 1967 7 McAllister 1968 8 Crane 1965 9 Allen, M. J. 1982 10 Love and Moser 1983 11 Arai 1969 P. notatus migrates into intertidal zone for spawning in late spring and early summer3,4 males migrate inshore in winter for spawning, females follow soon after, both sexes return to deep water in fall1,2 disruptive counterillumination used by some adults as an antipredatory strategy4,5 invertebrates compete with plainfin midshipman embryos for attachment space1 possible functions of bioluminescence: mimicry, warning, courtship,

and counterillumination9 BEHAVIOR Moveme Page 87 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 DE DF DG DH DI DJ DK DL DM DN DO Homing ability ( Home range) Schooling Territoriality NF NF Yes( males defending nest) 1 1 Arora 1948 ent Page 88 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 A B C D E Page 89 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 F G H I J K L M Page 90 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 N O P Q R S T U V W X Page 91 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 Y Z AA AB AC AD AE Page 92 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 AF AG AH AI AJ AK AL Page 93 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 AM AN AO AP AQ AR AS AT AU AV AW AX AY AZ Page 94 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 BA BB BC BD BE BF BG Page 95 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 BH BI BJ BK BL BM Page 96 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 BN BO BP BQ BR BS Page 97 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 BT BU BV BW BX BY Page 98 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 BZ CA CB CC Page 99 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 CD CE CF CG CH CI Page 100 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 CJ CK CL CM CN CO CP CQ Page 101 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 CR CS CT CU CV CW CX CY Page 102 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 CZ DA DB DC DD Page 103 of 1167 5 2220e  Batrachoididae 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 DE DF DG DH DI DJ DK DL DM DN DO Page 104 of 1167 5 2220e  Batrachoididae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 A B C D E F Family name Scientific name ( Synonym) Common name( s) Geographic range Observed depth range ( m)

Carangidae1 Seriola lalandi 3 n. B. C. to Chile1 1surface to 69 1Male: W= 0.00000005689FL2.89 S. dorsalis 1 1Female: W= 0.00000007747FL2.84 S. lalandei 2 1Both: W= 0.00000007439FL2.85 1 Robins et al. 1991 1 Love et al 2000 1 Gates & Frey 1974 1 Love et al 2000 1 Love et al 2000 1 Craig 1960b 2 Crooke 1983 2 Robins et al. 1991 3 Eschmeyer 1998 TAXONOMY RANGE Length ( TL= cm & FL, SL= mm) to weight ( g)

equation. * denotes TL = mm or FL, SL= cm yellowtail; California yellowtail; amberjack; amberfish;

white salmon; cavasina; jack1,2 Page 105 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 G H I J K L M N O P Q R S T U Growth Parameters Max. reported Geographic area Linf in cm ( SE) k ( SE) to ( SE)

Length conversion equations length ( cm) ( length measurement) Male Female Both Male Female Both Male Female Both Male Female Both NF 1 193 s. California ( FL) 1 NF NF 12.91 NF NF 0.136 NF NF  1.9 NF NF NF 1 Love et al 2000 1 Smith & Mekjian 1960 AGE & GROWTH r2 Page 106 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 V W X Y Z AA AB AC Age composition Ageing structure / Method Validation Verification Other refs. Longevity ( yrs.) information Sexuality Mating system scales tag & recapture No NF 1,2,3 12 NF Dioecious1 NF 1 Strategic Assessment Branch 1990 1 Strategic Assessment Branch 1990 2 Baxter 1960b 3 Love 1996 Page 107 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 AD AE AF AG AH AI AJ AK AL Mode of reproduction Parental Care Male : Female SL ratio Secondary Characteristics Male Female Male Female Ageing structure / Method Oviparous1 NF NF NF 1NG ( NG) 1 to 2 ( 50.6) NG ( NG) NG ( NG) NG 2,3,4 NG ( NG) 55.9 ( 2) NG ( NG) 71.1 ( 3) NG 1 Strategic Assessment Branch 1990 1 Smith & Paul 1960 2 Crooke 1992b 3 Fitch & Lavenberg 1971 4 Love 1996 Sexual dimorphism 1st maturity TL, FL*, or SL^ in cm ( age in yrs) at 50% maturity Breeding Behavior Page 108 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 AM AN AO AP AQ AR AS AT AU AV AW AX AY AZ BA Fertilized eggs present Spawning females present Both reproductive state Validation Verification Other refs. Geographic Spawning Area Jan. Feb. March April May June July Aug. Sept. Oct. Nov.

NG NG NF WA to CA2 NG NG s. California1 California3,4,6,7 ***** *****

Mexico2 1 Smith & Paul 1960 2 Strategic Assessment Branch 1990 3 Watson et al. 1996 4 Fitch & Lavenberg 1971 5 Kramer & Smith 1973 6 Baxter 1960b 7 Love 1996 REPRODUCTION Fertilization and Spawning Period (^^^^^= peak fertilization;*****= peak spawn; ##### = doubtful spawning records Page 109 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 BB BC BD BE BF BG BH es present Spawning Frequency Size specific fecundity equation Dec. ( per year) ( TL= cm, SL= mm; * denotes TL= mm; SL= cm) Egg duration Larval duration NF NF NF NF Multiple4 2450,000 ( 3 yrs.)

3,4 450,000 ( NG) to 700,000 ( NG)

1 Smith & Paul 1960 1 Smith & Paul 1960 2 Strategic Assessment Branch 1990 2 Fitch & Lavenberg 1971 3 Crooke 1992b 3 Baxter 1960b 4 Baxter 1960b 4 Love 1996 1,4 729,000 ( 72.8 FL) to 3,914,000 ( 105.3 FL) eggs Fecundity range in # eggs (@

cm, grams, or age)

Mean ( range) fecundity in eggs/ g female wt.

Once ( prolonged over a 3 month period);

older capable of mulitple1,2,3 DISPERS Fecundity Page 110 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 BI BJ BK BL BM BN BO BP BQ Size at settlement Size Fish Size ( SL in mm) Annual cohort success ( diameter in mm) Habitat Predators ( mm) Habitat Foraging guild Foraging method NF NF 1 0.8 to 1.6 planktonic1,2,3 NF 1,33.2 to 24.5 pelagic1 NF NF 2 1.4 23 to 4 to 3 1.27 4 1.69 1 Smith & Paul 1960 1 Feder et al 1974 1 Strategic Assessment Branch 1990 1 Watson et al. 1996 2 Watson et al. 1996 2 Watson et al. 1996 2 Watson et al. 1996 3 Brownell 1979 3 Limbaugh 1955 3 Sumida et al. 1985 4 Baxter 1960b Food habits Eggs Larvae SAL & RECRUITMENT Page 111 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 BR BS BT BU BV BW BX BY Fish Size Food habits When / Frequency Predators ( SL mm) Habitat Foraging guild Foraging method When / Frequency Predators NF NF 124.5 to maturity drifitng kelp1 carnivore NF NF Least terns1 ( pelagic crustacans; juvenile fish) 1 1 Sumida et al. 1985 1 Mitchell & Hunter 1970 1 Strategic Assessment Branch 1990 1 Love 1996 Juveniles HABITAT ASSOCIATION & TROPHIC INTERACTIONS Life Stage Page 112 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 BZ CA CB CC Adults Food habits Habitat Foraging guild Foraging method When / Frequency carnivore NF diurnal1,2,3,4,5 1 Feder et al 1974 1 Feder et al 1974 1 Strategic Assessment Branch 1990 2 Strategic Assessment Branch 1990 2 Strategic Assessment Branch 1990 2 Crooke 1992b 3 Carlisle et al. 1964 3 Crooke 1983 3 Crooke 1983 4 Turner et al. 1969 4 Carlisle et al. 1964 4 Fitch & Lavenberg 1971 5 Mitchell & Hunter 1970 5 Craig 1960a 5 Schmitt & Strand 1982 6 Limbaugh 1955 6 Fitch & Lavenberg 1971 7 CA Dept of Fish & Game 1982 7 Schmitt & Strand 1982 8 Love 1996 8 Limbaugh 1955 9 Love 1996 pelagic/ neritic; kelp beds; rocky peaks; drifting kelp; artificial reefs 1,2,3,4,5,6,7,8 ( squid; small schooling fishes, such as sardine, mackerel, and anchovy; pelagic crabs and other crustaceans) 1,2,3,4,5,6,7,8,9 Page 113 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 CD CE CF CG CH CI CJ CK CL Genetic variance Haplotype diversity Mean number of % Polymorphic Predators Substocks ( Geographic Area) alleles per locus ( SE) loci Expected ( SE) Observed ( SE) Approach NF NF NF NF NF NF NF 1 Strategic Assessment Branch 1990 1 Crooke 1992b 2 Fitch & Lavenberg 1971 2 Crooke 1983 3 Brian 2000 None from s. CA to Baja CA3 Mean heterozygosity Baja California ( south of Cedros Is.) and s.

California ( north of Cedrose Is.) 1,2 Nucleotide diversity (#)

pinnipeds; large fish, such as giant sea bass 1,2 GENETICS Page 114 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 CM CN CO CP CQ CR CS CT CU CV CW RECRUITMENT INTO FISHERY Mortality Estimates First Recruitment Full Recruitment Method of Effective population size Total length ( age) Total length ( age) Sampling gear Ageing structure / Method Validation Verification Natural ( M) Fishing ( F) Total ( Z) mortality estimation NF NF NF NF NF NF NF NF NF NF NF MORTALITY Page 115 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 CX CY CZ DA DB DC DD DE Rate of Exploitation ge range use Rate of Exploitation Interactions / associations alculate mort exploitation ( u) ratio ( F/ Z) Competition with other organisms Tagging studies Resident or Transient Homing ability ( Home range)

NF NF NF NF sometimes schools with S. argentea 1,6 1to 404 miles some residents, but mostly transient 1 NF cleaned by Oxyjulis californica 2,3 may clean parasites from blue sharks4 1 Strategic Assessment Branch 1990 1 Baxter & Collyer 1960 1 Baxter & Collyer 1960 2 Hobson 1971 2 Collyer 1954 3 Turner et al. 1969 4 Fitch & Lavenberg 1971 5 Schmitt & Strand 1982 6 Limbaugh 1955 7 Hose et al. 1983 8 Love & Moser 1983 infected by parasites from the families Dasyrhynchidae, Caligidae, Pennellidae,

Acanthocolpidae, Bucephalidae,

Hemiuridae, and Heteraxinidae8 displays intraspecific cooperative foraging5 may be negatively affected by chlorine discharge plums from power plants7 BEHAVIOR Movement 2up to 240 miles; but most < 10 miles Page 116 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 DF DG Schooling Territoriality Yes1,2,3,4 NF 1 Feder et al 1974 2 Strategic Assessment Branch 1990 3 CDF & G 1982 4 Love 1996 Page 117 of 1167 5 2220e  Carangidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 A B C D E Family name Scientific name ( Synonym) Common name( s) Geographic range Observed depth range ( m)

Clinidae1,2 Heterostichus rostratus 1,2 giant kelpfish 1intertidal to 40 m 1 Robins et al 1991 1 Robins et al 1991 1 Eschmeyer and Herald 1983 1 Love et al 2000 1 Love et al 2000 2 Eschmeyer 1998 2 Eschmeyer 1998 2 Gates and Frey 1974 British Columbia to Cabo San Lucas, southern Baja California, including Isla Guadalupe1 kelpfish, eel, kelp blenny, butterfish 1,2 TAXONOMY RANGE Page 118 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 F G H I J K L M N Total Length ( cm) Length conversion equations Max. reported Geographic area to weight ( g) equation length ( cm) ( length measurment) Male Female Both Male Female 1Both: W= 0.00310456L3.243 1SL= 0.598+ 0.819( TL) ( larvae) 161 cm TL NF NF NF NF NF NF 1SL= 0.580+ 0.906( TL) ( juveniles and adults)

1 Stepien 1986 1 Stepien 1986 1 Love et al 2000 Linf in cm ( SE) k ( SE)

Page 119 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 O P Q R S T U V W X r2 Both Male Female Both Male Female Both Ageing structure / Method Validation Verification NF NF NF NF NF NF NF NF NF NF Growth Parameters to ( SE)

AGE and GROWTH Page 120 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 Y Z AA AB AC AD AE Age composition Other refs. Longevity ( Length) information Sexuality Mating system Mode of reproduction Parental Care 1 15 yrs ( females) 1 dioecious seasonal monogamy1 oviparous1,2,3,5 male guards nest1,2,3,4,5 13 yrs ( males)

25 yrs.

37 yrs.

1 Stepien 1986 1 Matarese et al 1980 1 Stepien 1986 1 Coyer 1982 1 Matarese et al 1980 1 Matarese et al 1980 2 Stepien 1986 2 Moser 1996a 2 Moser 1996a 3 Tenera Environmental Services 2000 3 Love 1996 3 Coyer 1982 4 Tenera Environmental Services 2000 4 Breder and Rosen 1966 5 Coyer 1982 5 Limbaugh 1955 6 Holder 1907 Breeding Behavior Page 121 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 AF AG AH AI AJ AK AL Male : Female SL ratio Secondary Characteristics Male Female Male Female Ageing structure / Method females larger1 118.6 cm ( 1 1.5 yrs.) 18.6 cm ( 1 1.5 yrs.) NG NG otolith/ whole 22 yrs. 2 yrs. NG NG NG 1 Stepien 1986 1 Stepien 1986 1 Stepien 1986 2 Stepien 1986 2 Tenera Environmental Services 2000 adult males and females exhibit different melanin patterns1 1st maturity Total length in cm ( age in yrs) at REPRODU 50% maturity Sexual dimorphism adult males are brown ( infrequently olive green) and adult females exhibit three color morphs; red, brown, and green1 Page 122 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 AM AN AO AP AQ AR AS AT AU AV AW AX AY AZ Fertilized eggs present Spawning females present Both reprod Validation Verification Other refs. Geographic Spawning Area Jan. Feb. March April May June July Aug. Sept. Oct.

No interreader comparison California5 NG NG northeast Pacific1 CalCOFI2 ***** ***** *****

throughout range 3 ***** ***** *****

Santa Catalina Island, CA4 ***** ***** ***** ***** *****

1 Matarese et al 1980 2 Moser 1996a 3 Stepien 1986 4 Love 1996 UCTION Fertilization and Spawning Period (^^^^^= peak fertilization;*****= peak spawn; ##### = doubtful spawning records Page 123 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 BA BB BC BD BE BF BG Fecundity Mean Fecundity Size specific ductive states present eggs/ g Number of eggs fecundity eq Nov. Dec. Spawning Frequency ( weight range) ( at cm, grams, or age) ( length or weight range) Egg duration 1,2,3400 1,200 eggs 13,550 eggs ( 155 mm TL female) NF 1,2,312 17 days 2700 eggs ( avg.) 214 days ( avg.)

1 Matarese et al 1980 1 Matarese et al 1980 1 Wang 1981 1 Matarese et al 1980 2 Stepien 1986 2 Stepien 1986 2 Stepien 1986 3 Tenera Environmental Services 2000 3 Love 1996 3 Love 1996 may spawn several times a year1,2,3 Page 124 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 BH BI BJ BK BL BM Size at settlement Size Larval duration & location Annual cohort success ( diameter in mm) Habitat Predators 1approx. 8 week NF 11.35 mm ( avg.) adhesive eggs1,5,11 22 weeks 1,31.4 mm ( avg.)

21.2 1.4 mm eggs spawned in coastal intertidal to 30 m6 1 Stepien 1986 1 Stepien 1986 1 Matarese et al 1980 1 Matarese et al 1980 1 Coyer 1982 2 Love 1996 2 Moser 1996a 2 Moser 1996a 3 Stepien 1986 3 Stepien 1986 4 Coyer 1982 5 Love 1996 6 Wang 1981 7 Hubbs 1920b 8 Breder and Rosen 1966 9 Orton 1955 10 Limbaugh 1955 11 Tenera Environmental Services 2000 DISPERSAL & RECRUITMENT Eggs 30 50 mm TL ( fronds of giant kelp, brown algae, and green 1 Gibbonsia elegans, Heterostichus rostratus, Semicossyphus pulcher, and Oxyjulis californica 1 eggs are found on sargassum, ribbon kelp, giant kelp, and surfgrass10 two color morphs of eggs; purple yolk with orange oil globules and yellow yolk and yellow oil globules9 eggs attached in algal nest and brooded by male2,3,4,5,7,8 Page 125 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 BN BO BP BQ BR BS BT Fish Size Habitat Foraging guild Foraging method When / Frequency Predators Fish Size 15.5 50.0 mm SL pelagic1,2 planktivore NF NF NF 130 50 mm to maturity 25.1 45.0 mm SL rocky reef species3 near kelp beds in coastal waters2 1 Matarese et al 1980 1 Matarese et al 1980 1 Stepien 1986 1 Stepien 1986 2 Moser 1996a 2 Moser 1996a 2 Wang 1981 3 Pondella and Stephens 1994 HA ( single celled algae, rotifers,

mollusk larvae, and barnacle and copepod larvae) 1 Food habits Larvae newly hatched larvae have large yolk sacs1 Page 126 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 BU BV BW BX BY BZ Habitat Foraging guild Foraging method When / Frequency Predators Habitat NF NF NF 1tunas and lancetfish among kelp beds and algae of coastal waters3 inhabits surf grass beds4 rocky reef species4 1 Stepien 1986 1 Moser et al 1986 1 Moser 1996a 2 Stepein 1987 2 Stepein 1987 3 Wang 1981 3 Hubbs 1920b 4 Pondella and Stephens 1994 4 DeMartini 1981 5 Hobson et al 1981 6 Stepien 1986 7 Quast 1968b 8 Hubbs 1952 9 Limbaugh 1955 10 Holder 1907 BITAT ASSOCIATION & TROPHIC INTERACTIONS Life Stage Food habits kelp forests and rocky bottom with macrophyte cover1 fronds of giant kelp, the brown alga,

Sargassum muticum, and green surfgrass1 majority of juvenile color morphs were collected in plant habitats that closely matched the color of the fish2 found in sand bottom kelp holdfast,

midkelp, and rocky bottom kelp bed habitat9 lies almost motionless along the strands of kelp, usually at about a 45 ° angle, with the head up or down8 common in kelp column and canopy,

also about laminarian algae and eel grass on the bottom7 Juveniles hovers among columns of giant kelp druing both day and night5 majority of adult color morphs were collected in plant habitats that closely matched the color of the fish2,3,6,8,10 Page 127 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 CA CB CC CD CE Foraging guild Foraging method When / Frequency Predators Substocks carnivores NF diurnal feeders1,2 Brandt's cormorant and least terns1 1Santa Catalina Island San Diego Punta Clara San Benito Islands ( mysids, isopods, amphipods, shrimp, and fish) 2 Guadalupe Islands ( small crustaceans, small fish, and mollusks) 3,4 ( isopods and decapods ( 113 283 mm) 5 1 Love 1996 1 Love 1996 1 Love 1996 1 Stepien and Rosenblatt 1991 2 Quast 1968d 2 Hobson et al 1981 2 Hubbs 1952 3 Limbaugh 1955 4 Feder et al 1974 5 Mitchell 1953 Food habits Adults 2subspecies Heterosticus rostratus guadalupensis ( from Guadalupe Island)

( small crustaceans; such as mysids, shrimps, and amphipods, small fish; such as clingfish, kelp perch, and senorita) 1 Page 128 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 CF CG CH CI CJ CK CL CM Genetic variance Haplotype diversity Mean number of % Polymorphic ( Geographic Area) alleles per locus ( SE) loci Expected ( SE) Observed ( SE) Approach Effective population size NG NG 1.25 + 0.07 25.00% NG 0.07 + 0.02 allozymes NG 1.30 + 0.09 25.00% 0.07 + 0.02 1.20 + 0.06 20.00% 0.07 + 0.03 1.23 + 0.07 22.50% 0.05 + 0.02 1.04 + 0.04 3.57% 0.01 + 0.01 Nucleotide diversity (#) Mean heterozygosity GENETICS Page 129 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 CN CO CP CQ CR CS CT CU RECRUITMENT INTO FISHERY First Recruitment Full Recruitment Fishery / Length ( age) Fishery / Length ( age) Sampling gear Ageing structure / Method Validation Verification Natural ( M) Fishing ( F)

NF NF NF NF NF NF NF NF Page 130 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 CV CW CX CY CZ DA DB Mortality Estimates Rate of Exploitation Method of Age range used Rate of Exploitation Interactions / associations Total ( Z) mortality estimation to calculate mortality exploitation ( u) ratio ( F/ Z) Competition with other organisms NF NF NF NF NF NF illicit cleaning behavior from senoritas 1 larvae tend to school with mysid shrimp 2 parasites: 5 families, 5 spp. ( see reference) 3 1 Coyer 1982 2 Love 1996 3 Love and Moser 1983 MORTALITY Page 131 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 DC DD DE DF DG DH DI DJ DK DL Tagging studies Resident or Transient Homing ability ( Home range) Schooling Territoriality NF Residents1 NF Yes ( larvae) 1,2,3 Yes1,2,3,4,5 ( newly recruited juveniles) 2,3 adults solitary3 1 Tenera Environmental Services 2000 1 Stepien 1986 1 Stepien 1986 2 Love 1996 2 Limbaugh 1955 3 Feder et al 1974 3 Feder et al 1974 4 Coyer 1982 5 Holder 1907 BEHAVIOR Movement Page 132 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 DM DN DO Page 133 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 A B C D E Page 134 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 F G H I J K L M N Page 135 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 O P Q R S T U V W X Page 136 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 Y Z AA AB AC AD AE Page 137 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 AF AG AH AI AJ AK AL Page 138 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 AM AN AO AP AQ AR AS AT AU AV AW AX AY AZ Page 139 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 BA BB BC BD BE BF BG Page 140 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 BH BI BJ BK BL BM Page 141 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 BN BO BP BQ BR BS BT Page 142 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 BU BV BW BX BY BZ Page 143 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 CA CB CC CD CE Page 144 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 CF CG CH CI CJ CK CL CM Page 145 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 CN CO CP CQ CR CS CT CU Page 146 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 CV CW CX CY CZ DA DB Page 147 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 DC DD DE DF DG DH DI DJ DK DL Page 148 of 1167 5 2220e  Clinidae 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 DM DN DO Page 149 of 1167 5 2220e  Clinidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 A B C D E Family name Scientific name ( Synonym) Common name( s) Geographic range Observed depth range ( m)

Cottidae Hemilepidotus hemilepidotus 2,3,4 red Irish lord 3,4,5 Sea of Okhotsk to Monterey Bay, California 1 Intertidal to 51 m ( 156 ft) 1 [ spotted Irish lord, bullhead, red sculpin] 3 Kamchatka ( Russia) to Monterey Bay, CA ( common AK, rare CA) 2 San Miquel Is, southern CA 3 intertidal to 275 m ( 908 ft) 4 Eastern Kamchatka, Russia and Commander Is, Bering Sea to St Paul I, Aleutian Is4 Diablo Cove, central CA 5 Cottidae Leptocottus armatus 2,3,4 Pacific staghorn sculpin 3,5 Chignik, Alaska to San Quintin Bay, Baja California 1 Intertidal to 98 m ( 300 ft) 1 [ bullhead, staghorn sculpin 4, cabezon,

buffalo sculpin, smooth cabezon] 3 S Bering Sea to northern central Baja2 max 275 m ( 906 ft) 3 TAXONOMY RANGE Page 150 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 F G H I J KLM AGE and GROWTH Length ( mm) to Max. recorded Geographic Area Linf in mm ( SE)

weight ( g) equation Length conversion equations length ( cm) ( length measurement) Male Female Both Male 3 Male: W = 4.6759 x 10 4TL 2.4063 Female: W = 7.3659 x 10 6TL 3.1510 NF 1 51 cm TL ( 20 in) NF NF NF NF NF 4 W = 0.000341 SL 2.896 2 FL = 1.166( SL) + 1.549 SL = 0.856( FL)  1.116 ( range 56  178 mm SL; n= 10) 1 30.5 cm TL ( 12 in) 3 San Juan Island, WA ( TL) NG NG NG NG 5 W = 5.14 x 10 3 TL 3.31 ( TL = cm) 2 TL = 1.013( FL)  0.083 FL = 0.987( TL) + 0.093 ( range 67  213 mm FL; n= 10) 2 46 cm TL ( 18 in) 4 San Francisco Bay, CA ( TL) NG NG NG NG 6 log W =  5.243 + 3.134 ( log TL)

(< 190 mm TL) 2 SL = 0.846( TL)  1.056 TL = 1.181( SL) + 1.462 ( range 68  215 mm TL; n= 10) 5 Anaheim Bay, CA ( SL) NG NG NG NG 8 W = 3.9 6 x TL 3.2037 Page 151 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 N O P Q R S T U V W X Y Z k ( SE) to ( SE)

Female Both Male Female Both Male Female Both Ageing structure / Method Validation Verification Other refs. Longevity ( Length)

NF NF NF NF NF NF NF NF NF NF NF NF 6 > 6 yr NG NG NG NG NG NG NG NG 3 otoliths/ whole immersed in glycerin ( n= 262) No No NF 3 10 yr ( 228 mm TL)

NG NG NG NG NG NG NG NG 4 Length frequency < 2yr; Otoliths/ Whole &

cleared > 2yr ( n= 146) No No 4 Male: 3 yr ( 193.5 mm TL);

Female: 3 yr ( 197.3 mm TL)

NG NG NG NG NG NG NG NG 5 length frequency; otoliths/ whole (< 1yr) No No 5 NG r2 Growth Parameters Page 152 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 AA AB AC AD AE AF Age composition information Sexuality Mating system Mode of reproduction Parental Care Male : Female SL ratio NF Dioecious Male: Promiscuous and/ or Polygynous 2 Oviparous 1 Both sexes: guard nest3,4 Female: larger 3 NF Dioecious NF Oviparous 1 NF NF external fertilization Breeding Behavior Sexual d Page 153 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 AG AH AI AJ AK AL Secondary Characteristics Male Female Male Female Ageing structure / Method Male: black & white pelvic fins Female: white 4 4 22.0  31.3 cm TL ( 4 6 yr) 4 28.7  38.5 cm TL ( 4 6 yr) NG NG NG NF 3 NG ( SF Bay, CA) 3 12.8  15.3 cm TL ( 1 yr; SF Bay, CA) NG NG Otoliths / whole 3 12.0  12.2 cm TL ( 1 yr; Walker Creek, CA) 3 12.0  12.2 cm TL ( 1 yr; Walker Creek, CA) NG NG Otoliths / whole 50% maturity dimorphism 1st maturity Total length in cm ( age in yrs) at REPROD Page 154 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 AM AN AO AP AQ AR AS AT AU AV AW AX Fertilized eggs present Spawning females present Validation Verification Other refs. Geographic Spawning Area Jan. Feb. March April May June July Aug.

NG NG NF Puget Sound and nearby areas, WA 7 No No NF Walker Creek estuary ( SF Bay, CA) 8 ***** *****

No No San Pablo Bay ( SF Bay, CA) 9 ***** *****

San Francisco Bay 10 Anaheim Bay ( S CA) 11 ***** *****

Fertilization and Spawning Period (^^^^^= peak fertilization;*****= peak spawn; ##### = doubtful s DUCTION Page 155 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 AY AZ BA BB BC BD BE BF Mean Fecundity Size specific Both reproductive states present eggs/ g Number of eggs fecundity eq Sept. Oct. Nov. Dec. Spawning Frequency ( weight range) ( at cm, grams, or age) ( length or weight range)

once annually 4 NF NF 3 Log ( ova) = 2.9145 log( TL)  2.4311 ( range: 326  382 mm TL)

once annually 5 3 5000 eggs/ female NF 4 # maturing eggs = 0.355 TL 1.849 ( range: ~ 135  235 mm TL)

group synchronous ( 2 4 clutches) 6 no evidence of multiple spawning6 pawning records Fecundity Page 156 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 BG BH BI BJ BK Size at settlement Size Egg duration Larval duration & location Annual cohort success ( diameter in mm)

NF NF NF NF 3 1.5  1.6 mm 5 1.5  1.7 mm 11 NG 8 NG 9 NG 12 NG 2,3 9 14 days at 15oC NF NF NF 3 1.4  1.5 mm ( avg 1.43 mm)

6 1.36  1.50 mm DISPERSAL & RECRUITMENT Page 157 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 BL BM BN BO Habitat Predators Fish Size Habitat demersal, adhesive, guarded 3,11 NF 2 ~ 5  20 mm NL pelagic 2  1 to + 1 m 5 3 5  6 to > 19  23 mm SL pelagic 3 egg masses adhered to substrate ( rocks,

inverts) 5,11 7 22 mm FL ( n= 1) pelagic 7 shallow water or intertidal zone 8 8 6.5  10.5 mm pelagic 8 tough, pink conspicuous masses, often above low tide mark 9 11 16.5  22 mm FL pelagic, surface 11 shallow water, among sessile inverts in areas of high currents 12 demersal 3,6,7,10 NF 2 ~ 4 11 mm TL pelagic 2 adhesive 3,6,10 3 3.8 5.0 mm SL ( hatching); 15 20 mm SL ( transformation) pelagic 2 4 4.5 mm SL ( hatching); 15 20 mm SL ( metamorphosis) NG 4 5 3 11 mm SL tidal embayment & seasonal estuary ( Elkhorn Slough,

central CA) 5 6 3.8 5.0 mm TL ( at hatching) planktonic in polyhaline to mesohaline areas ( Sacramento San Joaquin estuary, Moss Landing Harbor and Elkhorn Slough, CA) 6 9 ~ 3.9  20 mm SL shallow water, riverine, estuarine, marine 9 Eggs Page 158 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 BP BQ BR BS BT Foraging guild Foraging method When / Frequency Predators Fish Size NG 2 NF NF NF NF NG 3 copepods 7 NG 8 copepods, amphipods 11 NG 2 NF NF NF 2 < 70 mm SL NG 3 2 > 70 mm SL NG 4 3 45 79 mm TL NG 5 3 80 119 mm TL NG 6 4 NG planktivorous 9 5 66 166 mm TL ( juv & adult)

6 3.3 24.3 cm TL 7 54 140 mm SL? ( juv & adult)

7 59 220 mm SL? ( juv & adult)

7 105 290 mm SL? ( juv & adult)

8 107 160 mm TL ( juv & adult)

9 99 205 mm SL ( juv & adult)

Food habits Larvae Page 159 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 BU BV BW BX Habitat Foraging guild Foraging method When / Frequency NF NF NF NF estuary ( Anaheim Bay, southern CA) 2 carnivore: decapods, amphipods, fishes 2 NF estuary ( Anaheim Bay, southern CA) 2 carnivore: ( decapods, fishes) & opportunistic benthic feeder 2 bay ( Padilla Bay, WA) 3 carnivore: amphipods, isopods 3 bay ( Padilla Bay, WA) 3 carnivore: amphipods, isopods, crabs 3 estuary ( Elkhorn Slough, central CA) 4 epifaunal crustaceans ( amphipods, copepods, polychaetes) 4 marsh ( China Poot, Alaska) 5 generalist: amphipods, diptera larvae, plants, fish, detritus, insects, crustacean, oligochaetes 5 estuary ( Yaquina Bay, OR) 6 NG 6 lagoon ( Big Lagoon, Humboldt Co, CA) 7 amphipods 7 river ( Mad River, Humboldt Co, CA) 7 algae, shrimp, amphipod, Dungeness crab legs, fish 7 bay ( Humboldt Bay, Humboldt Co, CA) 7 Dungeness crab, fish, amphipods, polychaete, isopods, shrimp 7 intertidal 8 Dungeness crab 8 bay/ estuary ( San Francisco Bay, CA) 9 shrimp, crabs, fish, amphipods, Callianassids, bivalve mollusks,

polychaetes, isopods, other 9 Food habits Life Stage Juveniles HABITAT ASSOCIATION & TROPHIC INTERACTIONS Page 160 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 BY BZ CA Predators Fish Size Habitat NF 2 287 385 mm TL ( breeding females) guarding nests 2 2 220 313 mm TL ( breeding males) guarding nests 2 19 NG Nanaimo, B. C. 19 gulls and cormorants 1 3 NG estuary ( SF Bay, northern CA) 3 Great Blue Heron ( Ardea herodias ) 2 3 NG estuary ( Walker estuary, northern CA) 3 Caspian tern ( Hydroprogne caspia ) 3 4   120 mm TL bay ( Padilla Bay, WA) 4 waterfowl, ducks 4 5 NG Mugu Lagoon, southern CA 5 western grebe ( Aechmophorus occidentalis ) 5 9 NG estuary ( Elkhorn Slough, central CA) 9 mammals ( harbor seal, CA sea lion, N. sea lion) 6 6 66 166 mm TL ( juv & adult) marsh ( China Poot, Alaska) 6 size NG ( juv or adult): harbor seal ( Phoca vitulina richardsi ) 9 6 3.3 24.3 cm TL estuary ( Yaquina Bay, OR) 6 size NG ( juv or adult): California gull ( Larus californicus ) 10 7 54 140 mm SL? ( juv & adult) lagoon ( Big Lagoon, Humboldt Co, CA) 7 Caspian tern ( Sterna caspia ) 8,11 7 59 220 mm SL? ( juv & adult) river ( Mad River, Humboldt Co, CA) 7 western gull ( Larus occidentalis ) 11 7 105 290 mm SL? ( juv & adult) bay ( Humboldt Bay, Humboldt Co, CA) 7 harbor seal ( Phoca vitulina richardsi ) 12 8 107 160 mm TL ( juv & adult) intertidal 8 10 > 70 mm SL tideflat ( Coos Bay, OR) 10 Page 161 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 CB CC CD CE CF Foraging guild Foraging method When / Frequency Predators Substocks Female ( breeding): carnivore ( crabs, shrimp, fishes) 2 NF Both sexes: suggested nocturnal 1 NF NF Male ( breeding): carnivore ( crabs, shrimp, fishes,

conspecific eggs) 2 carnivore: crabs ( Hemigrapsus, Cancer productus ),

shrimps, barnacles, mussels 19 carnivore: shrimps, fishes 3 NF NF gulls and cormorants 1 NF carnivore: amphipods, nereid worm, dipteran larvae,

shrimp 3 Great Blue Heron ( Ardea herodias ) 2 carnivore: isopods, crabs, fish 4 Caspian tern ( Hydroprogne caspia ) 3 carnivore: bivalve siphons 5 waterfowl, ducks 4 epifaunal crustaceans ( crabs, amphipods, macroalgae)

9 western grebe ( Aechmophorus occidentalis ) 5 generalist: amphipods, diptera larvae, plants, fish, detritus, insects, crustacean, oligocha etes 6 mammals ( harbor seal, CA sea lion, N. sea lion) 6 NG 6 harbour seal 7 amphipods 7 size NG ( juv or adult): harbor seal ( Phoca vitulina richardsi ) 9 algae, shrimp, amphipod, Dungeness crab legs, fish 7 size NG ( juv or adult): California gull ( Larus californicus ) 10 Dungeness crab, fish, amphipods, polychaete,

isopods, shrimp 7 caspian tern ( Sterna caspia ), western gull ( Larus occidentalis ) 11 Dungeness crab 8 harbor seal ( Phoca vitulina richardsi ) 12 burrowing ghost shrimp ( Callianassa californiensis ) 10 Food habits Adults Page 162 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 CG CH CI CJ CK CL CM CN CO Genetic variance Haplotype diversity Mean number of % Polymorphic First Recruitment ( Geographic Area) alleles per locus ( SE) loci Expected ( SE) Observed ( SE) Approach Effective population size Fishery / Length ( age)

NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF Nucleotide diversity (#) Mean heterozygosity GENETICS Page 163 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 CP CQ CR CS CT CU CV CW CX CY CZ DA RECRUITMENT INTO FISHERY Mortality Estimates Rate of Exploitation Full Recruitment Method of Age range used Rate of Exploitation Fishery / Length ( age) Sampling gear Ageing structure / Method Validation Verification Natural ( M) Fishing ( F) Total ( Z) mortality estimation to calculate mortality exploitation ( u) ratio ( F/ Z)

NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF NF MORTALITY Page 164 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 DB DC DD DE DF Interactions / associations Competition with other organisms Tagging studies Resident or Transient Homing ability ( Home range)

NF Parasite: mature Trematode ( Derogenes varicus ); Bering Sea2 2 < 5 m2 to 500 900 m2 NF NF Parasites: 9 Families; ~ 13 Species; See Refs 11 Parasites: Digenea ( 7 sp), Nematoda ( Ascarophis sebastodis ),

Acanthocephala ( Echinorhynchus gadi ) 12 NF Parasites: nematode Spirocamallanus pererai in gastrointestinal track 3 3 0 800 m ( Anaheim Bay, CA) Resident: Elkhorn Slough, central CA ( tidal embayment & seasonal estuary) 1 NF Parasite: nematode larvae Contracaecum in coelum 3 Resident: Elkhorn Slough, central CA ( tidal embayment & seasonal estuary) 2 Parasite: trematode ( Gyrodactylus sp) on gills 3 Resident: Elkhorn Slough, central CA ( tidal embayment & seasonal estuary) 3 Parasites: trematodes ( Tubulovesicula lindbergi & Stephenostomum sp)

in small intestine 3 Parasite: cestode larvae ( Acanthobothrium) in small intestine 3 Parasite: glossiphonid leech at isthmus 3 Parasite: cyclopoid copepod ( Holobomolochus prolixus) at operculum 3 Parasite: nematode ( Contracaecum magnus) 4 Parasite: Trematode ( Ascocotyle diminuta ) on gills; Pt Mugu, CA 5 Parasite: Haemogregarina leptocotti in blood; Humboldt Bay, CA 6 Parasite: acanthocephalan ( Echinorhynchus gadi ) in intestine; Coos Bay,

OR 8 Parasite: acanthocephalans ( Podocotyle olssoni, P. reflexa ) in intestine;

Coos Bay, OR 8 Movement BEHAVIOR Page 165 of 1167 5 2220e  Cottidae 1234 5 67 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 DG DH Schooling Territoriality NF NF NF NF Page 166 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 A B C D E Cottidae Scorpaenichthys marmoratus 3,4 cabezon3,4,5; giant marbled sculpin1,2,3 Sitka, Alaska to Pt. Abreojos, central Baja California 1,2,3 Intertidal to 82 m ( 250 ft) 1,2 ( Hemitripterus marmoratus) 1, 2 [ bullhead, blue cod, sculpin, giant sculpin,

marble sculpin, bull cod, salpa, scorpion,

biggyhead, scaleless sculpin] 3 Max @ 110 m ( 335 ft) 3 [ cab, caby, cabezony, bullhead] 6 Page 167 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 F G H I J KLM 1 W = 1.85 x 10 5 TL 2.9836 1 SL = 0.83 TL 1 99 cm TL ( 39 in) 1 Puget Sound, WA ( TL) 690.25 ( NG) 740.87 ( NG) NG 0.241 ( NG)

2 W = 6.84 x 10 6 TL 3.16 2 central Cal, S Cal, & Baja ( TL) 604 ( NG) 760 ( NG) NG 0.83 ( NG)

7 Male: W = 2.546 x 10 6 TL 3.307 Female: W = 8.780 x 10 6 TL 3.113 Both: W = 5.498 x 10 6 TL 3.185 1 O'Connell's data ( central Cal, S Cal, Baja) recalculated 536 ( NG) 679 ( NG) NG 0.457 ( NG)

Page 168 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 N O P Q R S T U V W X Y Z 0.354 ( NG) NG  1.23 ( NG) 0.84 ( NG) NG NG NG NG 1 Otoliths/ whole; fin rays/ sectioned ( n= 87) No between structures NF 1 Male: 17 yr ( 648 mm TL)

Female: 16 yr ( 725 mm TL)

0.91 ( NG) NG NG NG NG NG NG NG 2 otoliths/ whole ( n= 105) No intrareader 2 Male: 9+ yr ( 531 mm TL)

Female: 13+ yr ( 720 mm TL)

0.230 ( NG) NG NG NG NG NG NG NG 1 otoliths/ whole ( n= 105) No NG Page 169 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 AA AB AC AD AE AF NF Dioecious Polygynous 1 Oviparous 1 Male: guard nest1,2 Female: larger, older 1 tend eggs laid on Iridaea splendens 5 Female: larger 2 Page 170 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 AG AH AI AJ AK AL Juveniles: silver red 1 1 34.3 cm TL ( 2 yr) 1 44.5 cm TL ( 3 yr) NG NG NG Male: ( adult): red Female: ( adult): green 1,2 2 ~ 33.9 cm TL ( 2+ yr) 2 ~ 44.5 cm TL ( 3+ yr) NG NG Otoliths / whole Both sexes: ( Non breeding adults):

white1,2 Male: ( near spawning females):

brown, red, black Male:

( nesting): white Female ( adult): turquoise green 3 Page 171 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 AM AN AO AP AQ AR AS AT AU AV AW AX NG NG NF British Columbia 1 No between structures British Columbia 2 southern Strait of Georgia, B. C. 3 Puget Sound, WA 4,5 ***** *****

central CA 4,5 central CA 6 *****

Page 172 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 AY AZ BA BB BC BD BE BF batch spawner1,3 1 26.4  14.5 per batch ( 2.5  10.5 kg TW) NF 1 # eggs = 0.29 TL  87.54 ( range: 500  775 mm TL)

at least twice during spawning season2 2 34.8  21.0 per batch ( 1.4  4.6 kg TW) 2 Y = 27.3 + 1.53X ( range: 1.4  4.6 kg TW; Y =

1000' s of eggs; X = 100' s of g TW)

***** Page 173 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 BG BH BI BJ BK 1 2 3 weeks Nov  Mar 1 1 40 mm TL (~ 3 4 months), tidepools NF 1 1.4  1.7 mm 3 4 months 3 2 < 0.40  1.5 mm 3 1.4  1.9 mm 4 1.5 mm 5 1.5  1.6 mm 7 NG 13 nests: 46 cm diam ( 5  10 cm thick)

14 NG Page 174 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 BL BM BN BO NG 1 eggs toxic to man 1 1 10 40 mm TL planktonic 1 NG 2 eggs toxic to lab animals 1,3,5,6,7 2 4 35 mm SL ( hatch at 4 6 mm) pelagic ( Oregon < 65 km offshore) 2 demersal, adhesive, attached eggs 3 eggs avoided by natural predators 2 3 NG pelagic 3 hard surfaces ( rock, iron, steel, wood) 15 Artedius harringtoni, Embiotoca lateralis,

Damalichthys vacca, other male cabezon 4 6 hatch: 3.1 4.8 mm TL NG 6 5 NG 10 8  17 mm FL pelagic, surface 10 demersal, adhesive, clusters 7 11 14  22 mm FL pelagic, surface 11 intertidal, under rocks, deep crevices, subtidal 13 12 3  6 mm SL ( hatching); 14 mm SL ( transformation) pelagic until ~ 35 mm 12 algae ( Iridaea splendens ) 14 13 6 mm TL ( hatching) NG 13 Page 175 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 BP BQ BR BS BT 10,11 60 118 mm TL ( 0+)

12 ~ 26 31.4 mm TL ( juv)

13 45 245 mm TL ( juv & adult)

13 45 79 mm TL ( juv)

13 80 119 mm TL ( juv)

14 NG ( juvenile or adult)

15 20 220 mm TL ( juv & adult)

16 < 100 mm TL 18 33 290 mm TL ( juv & adult)

copepods ( crustaceans) 1 NF NF NF 1 41 200 mm TL ( arbitrary: juvs)

NG 2 1 201 400 mm TL ( arbitrary: pre adults)

NG 3 17 200 499 mm SL ( juv & adult )

NG 6 19 NG barnacle larvae ( Cirripedia: Balanus cypris stage),

copepod ( nauplii, Copepodites, Pseudocalanus minutus ), amphipod ( Parathemisto pacifica ),

decapod ( Brachyura zoea), Euphausiacea ( furcilia),

fish ( Atheresthes stomias ) 10 copepods, fish ( Stichaeidae) 11 NG 12 NG 13 Page 176 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 BU BV BW BX estuary ( Grays Harbor, WA) 10,11 carnivore: nereid polychaetes, ghost shrimp, sand shrimps, juv crabs, bivalve siphon tips, unid crustaceans 10,11 bay ( Skagit Bay, Puget Sound, WA) 12 carnivore: crustaceans, polychaetes, insects 12 bay ( Padilla Bay, WA) 13 generalist: amphipod, isopods, crabs, polychaetes, moluscs,

fishe, algae, wood, rock, shrimp 13 bay ( Padilla Bay, WA) 13 carnivore: amphipods, isopods, polychaetes, molluscs, other 13 bay ( Padilla Bay, WA) 13 carnivore: amphipods, isopods, crabs, polychaetes, other 13 estuary ( west of Vancouver Island, B. C., Canada) 14 amphipods, insects ( Chironamid larvae & pupae), Oligochaetes,

isopods ( Gnorimosphaeroma bregonensis ), eggs, calanoid copepods 14 bay, cove, strait ( Strait of Juan de Fuca) 15 generalist: decapods ( Natantia, Reptantia), mysids, amphipods,

fish ( incl Pleuronectidae, Cottidae), polychaetes, isopods 15 estuaries ( Vancouver Island, B. C. Canada) 16 generalist: amphipods ( Eogammarus confervicolus , Corophium sp ), polychaetes ( Nereis sp ), isopod ( Gnorimosphaeroma oregonensis ), chum salmon fry ( Oncorhynchus keta ) 16 bay ( Everett Bay, WA) 18 carnivore: amphipods ( Corophium , gammarids), fish ( juv staghorn sculpin, sandlance, starry flounder, threespine stickleback), ghost shrimp ( Callianassa ) 18 demersal; hard bottoms; tidepools 1 carnivore: Crustacea, fish 1 NF NF demersal; hard bottoms; tidepools 1 carnivore: Crustacea, Mollusca, fish 1 S Cal rocky inshore and kep beds 17 cephalopods ( octopus, squid), crabs ( cancroid, spider),

crustaceans, shrimp, fish, algae ( red & green), gastropods,

lamellibranchs 17 Diablo Cove, CA 19 fish ( incl juv rockfish), crustaceans, mollusks, Octopus sp, see ref for more ID 19 Page 177 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 BY BZ CA 11 99 205 mm SL ( juv & adult) bay/ estuary ( San Francisco Bay, CA) 11 12,13   118 mm TL ( 1+ and older) estuary ( Grays Harbor, WA) 12,13 14 45 245 mm TL ( juv & adult) bay ( Padilla Bay, WA) 14 14 > 120 mm TL ( adult) bay ( Padilla Bay, WA) 14 15 NG ( juvenile or adult) estuary ( west of Vancouver Island, B. C., Canada) 15 16 20 220 mm TL ( juv & adult) bay, cove, strait ( Strait of Juan de Fuca) 16 18 33 290 mm TL ( juv & adult) bay ( Everett Bay, WA) 18 sea otters, pigeon guillemots, least terns, Brandt's cormorants 7 1 401 720 mm TL ( arbitrary ) demersal; hard bottoms1 1   488 mm tidepools 1 19 NG Nanaimo, B. C. 19 20 NG ( Adult) Kelp bed bottom 20 17 200 499 mm SL ( juv & adult ) S Cal rocky inshore and kep beds 17 Page 178 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 CB CC CD CE CF shrimp, crabs, fish, amphipods, Callianassids, bivalve mollusks, polychaetes, isopods, other 11 carnivore: ghost shrimp, nereid polychaete, mud shrimp, juv crabs, gammarid amphipod, sand shrimp,

fish 12,13 generalist: amphipod, isopods, crabs, polychaetes,

moluscs, fishe, algae, wood, rock, shrimp 14 generalist: crabs, isopods, fishes, algae, wood, rock 14 amphipods, insects ( Chironamid larvae & pupae),

Oligochaetes, isopods ( Gnorimosphaeroma bregonensis ), eggs, calanoid copepods 15 generalist: decapods ( Natantia, Reptantia), mysids,

amphipods, fish ( incl Pleuronectidae, Cottidae),

polychaetes, isopods 16 carnivore: amphipods ( Corophium , gammarids), fish ( juv staghorn sculpin, sandlance, starry flounder,

threespine stickleback), ghost shrimp ( Callianassa ) 18 carnivore: Crustacea, Mollusca, fish, eggs 1 NF NF sea otters, pigeon guillemots, least terns, Brandt's cormorants 8 NF NG 1 carnivore: fish ( Hemilepidotus hemilepidotus, fish eggs, small perch), crabs ( Epialtus , Cancer ,

Hemigrapsus ), shrimp 19 mesocarnivore 20 cephalopods ( octopus, squid), crabs ( cancroid,

spider), crustaceans, shrimp, fish, algae ( red &

green), gastropods, lamellibranchs 17 Page 179 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 CG CH CI CJ CK CL CM CN CO NF NF NF NF NF NF NF NF NF Page 180 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 CP CQ CR CS CT CU CV CW CX CY CZ DA NF NF NF NF NF NF NF NF NF NF NF NF Page 181 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 DB DC DD DE DF Parasite: trematode ( Tubulovesicula sp ) in stomach, intestine, gill region;

Coos Bay, OR 8 Parasite: nematodes ( Contracaecum sp, Anisakis sp ) in intestine and body musculature; Coos Bay OR 8 Parasite: trematode ( Derogenes varicus, Genolinea manteri ) in stomach 9 Parasites: 21 Families; ~ 43 Species; See Refs 11 Parasites: Digenea ( 5 sp), Nematoda ( 2 sp), Acanthocephala ( 2 sp) 12 NF Parasite: Nematode ( Capillaria margolisi ) in pyloric caeca1 1 0 naut mi NF NF Parasite: Protozoa, Sarcomastigophora ( Trypanoplasma beckeri ) in blood and body fluids 7 Parasite and vector for T. beckeri : Leech, Hirudinea, Piscicolidae ( Malmiana diminuta ) 7 Parasite: trematode ( Genolinea robusta ) in stomach 9 Parasite: copepod ( Chondracanthus gracilis ) in gill cavity 10 Parasite: copepod ( Chondracanthus pinguis ) 10 Parasites: 11 Families; ~ 18 Species; See Refs 11 Page 182 of 1167 5 2220e  Cottidae 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 DG DH NF NF Page 183 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 A B C D E 1  Lea and Eschmeyer 1988 1  Clemens and Wilby 1961 1  Miller and Lea 1972 1  Miller and Lea 1972 2  Eschmeyer 1998 2  Pillsbury 1957 2  Eschmeyer and Herald 1983 2  Eschmeyer and Herald 1983 3  Miller and Lea 1972 3  Gates and Frey 1974 3  Love et al. 2000 3  Love et al. 2000 4  Robins et al. 1991 4  Miller and Lea 1972 4  Allen and Smith 1988 4  Allen and Smith 1988 5  Robins et al. 1991 5  Burge and Schultz 1973 6  Reilly et al. 1994 Page 184 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 F G H I J KLM 1  O'Connell 1953 1  O'Connell 1953 1  Miller and Lea 1972 1  Lauth 1987 2  Lauth 1987 2  Karpov and Kwiecien 1988 2  Hart 1973 2  O'Connell 1953 3  DeMartini and Patten 1979 3  Weiss 1969 4  Tasto 1975 4  Jones 1962 5  Bayer 1985 5  Tasto 1975 6  Weiss 1969 7  Lea et al. 1999 8  Armstrong 1991 Page 185 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 N O P Q R S T U V W X Y Z 1  O'Connell 1953 1  Lauth 1987 2  Lauth 1987 2  O'Connell 1953 3  Weiss 1969 3  Weiss 1969 4  Jones 1962 4  Jones 1962 5  Tasto 1975 5  Tasto 1975 6  Love 1996 Page 186 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 AA AB AC AD AE AF 1  Lauth 1987 1  Matarese et al. 1989 1  Feder et al. 1974 1  O'Connell 1953 2  DeMartini and Patten 1979 2  Lauth 1987 2  Lauth 1987 3  DeMartini and Patten 1979 3  DeMartini and Patten 1979 4  Garrison and Miller 1982 5  Burge and Schultz 1973 Page 187 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 AG AH AI AJ AK AL 1  O'Connell 1953 1  Fitch and Lavenberg 1971 1  Fitch and Lavenberg 1971 1  Fitch and Lavenberg 1971 2  Orton 1955 2  O'Connell 1953 2  O'Connell 1953 2  O'Connell 1953 3  Lauth 1987 3  Jones 1962 3  Jones 1962 3  Jones 1962 4  DeMartini and Patten 1979 4  Garrison and Miller 1982 4  Garrison and Miller 1982 Page 188 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 AM AN AO AP AQ AR AS AT AU AV AW AX 1  Fitch and Lavenberg 1971 1  Fitch and Lavenberg 1971 1  Hart 1973 2  O'Connell 1953 2  O'Connell 1953 2  Fraser 1923 3  Jones 1962 3  Jones 1962 3  Pillsbury 1957 4  Lauth 1987 5  Lauth 1988 6  O'Connell 1953 7  DeMartini and Patten 1979 8  Jones 1962 9  DeVlaming et al. 1984 10  Boothe 1967 11  Tasto 1975 Page 189 of 1167 5 2220e  Cottidae 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 AY AZ BA BB BC BD BE BF 1  O'Connell 1953 1  Lauth 1988 ( TW = total weight) 1  Lauth 1988 2  Lauth 1987 2  O'Connell 1953 ( TW = total weight) 2  O'Connell 1953 ( TW = total weight)