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Fishes of Marine Protected Areas Near La Jolla, California PDF

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Bull. SouthernCaliforniaAcad. Sci. 113(3),2014,pp.200-231 ©SouthernCaliforniaAcademyofSciences,2014 Fishes of Marine Protected Areas Near LaJolla, California P. A. Hastings,1* M. T. Craig,2 B. E. Erisman,3 J. R. Hyde,4 and H. J. Walker1 1Marine Biology Research Division, Marine Vertebrate Collection & Centerfor Marine Biodiversity and Conservation, Scripps Institution ofOceanography, University ofCalifornia San Diego, La Jolla, CA 92093-0208 U.S.A. 2Department ofEnvironmental and Ocean Sciences, University ofSan Diego, San Diego, CA 92110 U.S.A. 3University ofTexas, The University ofTexas Marine Science Institute, 750 Channel View Drive, Port Aransas, TX 78373 4Southwest Fisheries Science Center, NOAA/NMFS, 8901 La Jolla Shores Dr., La Jolla, CA 92037, USA — Abstract. The marine waters surrounding La Jolla, California have a diverse array ofhabitats and include several marine protected areas (MPAs). We compiled a list of the fish species occurring in the vicinity based on records of specimens archived in the Marine Vertebrate Collection (MVC) ofthe Scripps Institution ofOceanography MVC (SIO). Collection of fishes from La Jolla in the started in 1905, but greatly accelerated in 1944 when Carl L. Hubbs moved to SIO. By 1964, 90% of the 265 species recorded from the area had been collected and archived in the MVC. The fishes of La Jolla are dominated by species whose center of distribution is north of Point Conception (111 species), or between there and Punta Eugenia (96), with fewer species with southern distributions (57), and one exotic species. Reflecting the diversity ofhabitats in the area, soft-substrate species number 135, pelagic species 63, canyon-dwelling species 123 (including 35 rockfish species of the genus Sebastes ), and hard-bottom species 140. We quantified the abundance of the latter group between 2002 and 2005 by counting visible fishes in transects along the rocky coastline ofLa Jolla, both within and adjacent to one ofthe region’s MPAs. In 500 transects, we counted over 90,000 fishes representing 51 species. The fish MPA communities inside and outside of the were similar and, typical of southern California kelp forests, numerically dominated by Blacksmith, Chromispunctipinnis (Pomacentridae), and Senorita, Oxyjulis californica (Labridae). Natural history MVC collections such as the are important resources for conservation biology for determining the faunal composition of MPAs and surrounding habitats, and documenting both the disappearance and invasion of species. The coastal environment in and around La Jolla, San Diego County, California is notable for its complex and diverse array ofhabitats within a relatively small area. These include kelp forests, rocky reefs, rocky intertidal, sandy beaches, sand and mud subtidal areas, eelgrass and surf-grass stands, pier pilings, and submarine canyons, as well as the pelagic environment. Because of the proximity of the La Jolla and Scripps submarine canyons, depths range to over 500 meters within less than 7 km of the coastline. These diverse habitats support a rich marine community, which has served as the focus of a *Corresponding author: [email protected] 200 FISHES OF LA JOLLA 201 variety of scientific investigations (e.g., Limbaugh 1955; Quast 1968; Craig et al. 2004; Brueggeman 2008). The nearshore environment of La Jolla is especially important within the context of marine conservation because it houses a network ofmarine protected areas (MPAs) of varying size and age and with varying levels ofuse restrictions (Fig. 1). Historically, the first ofthese was the San Diego Marine Life Refuge established in 1957, which included the Scripps Coastal Reserve, a part of the University of California Natural Reserve system (McArdle 1997). Directly to the south is the San Diego-La Jolla Ecological Reserve (SDLJER), a small no-take reserve on the northern end of the La Jolla rocky coastline that was established in 1971. The San Diego-La Jolla Ecological Reserve Areas of Special Biological Significance was established in 1974 and largely overlapped the SDLJER (McArdle 1997). Recently the SDLJER was included in the Matlahuayl State Marine Reserve (CDFG 2013). In addition, the area on the southern part ofthe La Jolla peninsula was recently designated as the South La Jolla State Marine Reserve, and an areadirectlywest ofthatwasdesignated asthe South LaJolla State MarineConservation Area (CDFG 2013). These MPAs have a variety ofuse restrictions, but all recognize and seek to protect the biodiversity ofthis ecologically important region. This study documents the fish fauna in and around this series of marine protected areas. The primary source of information on fishes occurring in the La Jolla area was the Scripps Institution of Oceanography’s Marine Vertebrate Collection (MVC). The MVC important roles ofnatural history collections such as the to conservation biology have been widely documented (Allman 1994; Pyke and Ehrlich 2010; Drew 2011). These includecompiling biotic inventories, documenting theloss or degradation ofhabitats and associated biota, documenting changes in the distribution and occurrence of native species, and documenting species invasions. In addition to compiling an inventory offish species collected in the area and archived in the MVC, we report on diver surveys ofthe abundance and diversity offishes in kelp forests, one ofthe most prominent habitats in the area, in and around the SDLJER (now the Matlahuayl State Marine Reserve) from 2002 to 2005. BriefHistory ofFish Collecting in the La Jolla Area Themarinefishes ofCaliforniahavebeen studied formanydecades andarewell-known (Miller and Lea 1972, 1976; Hubbs et al. 1979; Love et al. 2005; Allen et al. 2006) with at least519speciesknownfromstatewaters(Hornetal. 2006). Inadditiontoearlycollections offishesfromtheSanDiegoareareportedbyJordanandGilbert (1880, 1881), thestudyof fishes in the San Diego region of southern California was begun in earnest with the Albatross surveys (Moring 1999) as reported by C.H. Gilbert (1890, 1896, 1915), aswell as inventories by Eigenmann and Eigenmann (1890) and Eigenmann (1892). The establishment ofthe Scripps Institution ofOceanography (SIO) in the San Diego region in 1903 and its subsequent move to La Jolla in 1905 marked a significant increase in the study ofthe region’s biota (Hastings and Rosenblatt 2003). The on-site aquarium displayed many ofthe common shallow-water fish species ofthe area. In 1918, Percy S. Barnhart (Fig. 2A) was appointed as Collector and Curator of the Aquarium, and in 1926, Barnhart was elevated to the position of Curator of the Biological Collections, a position he held until 1948. Barnhart studied the local fishes leading to a publication on the fishes ofsouthern California (Barnhart 1936), and he assembled a small collection of preserved specimens from the region that ultimately formed the basis ofthe SIO Marine Vertebrate Collection (MVC). 202 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES 117*20OW 117*18'0'W 117*16‘0"W ii7*i-row i SouthernCaliforniaMPAs igiMatlahuaylSlateMarineReserve BBBSanDiego-ScrippsCoastalStateMarineConservationArea SouthLaJoliaMarineConservationArea SouthLaJollaStateMarineReserve Studyarea L.VJStudyarea Fig. 1. Map ofstudy area with MPAs designated. Kelp forestfishes were surveyed at La Jolla Cove and Boomers. Knowledge ofthe ichthyofauna ofthe region greatly increased after 1944when Carl L. Hubbs (Fig. 2B) moved to SIO and began actively archiving collections of fishes in the MVC. In negotiations regarding his forthcoming move from the University ofMichigan to SIO, Hubbs wrote to then SIO Director Harald Sverdrup: “I would no doubt want to put considerable emphasis on systematic and variational studies ofwest coast marine fishes, particularly those in which speciation would be correlated with oceanographical (sic) conditions...I would no doubt be interested in exploratory work, for instance with the fauna of the deep basins off the southern California coast. I will probably be interested too in detailed analyses of the distribution of fishes along the entire west coast, again as correlated with the oceanographic conditions.” (Shor et al. 1987, pp. 226-227). Before his arrival in October 1944, Hubbs convinced Sverdrup to invest in facilities to store his anticipated collection of fishes, leading to the ultimate establishment of the FISHES OF LA JOLLA 203 Fig.2. PhotosofA)PercyS. Barnhart, 1935;B)CarlL.Hubbs, 1973;C)RichardH.Rosenblatt, 1979; D) aviewofLaJolla Shores and LaJollapeninsulalookingsouthwardfrom the SIOcampus, circa 1910 (arrow indicates small embayment at La Jolla Shores). All images are from the Scripps Institution of Oceanography Archives, UC San Diego Library. MVC (Shor et al. 1987). Hubbs wasted little time in collecting fishes from the surroundings ofthe SIO campus, amassing over 200 collections in his first year and over 500 by the end ofthe decade (Fig. 3). Hubbs had another California project in mind when considering the move to La Jolla. In 1944hewrotetoW. I. Follett in Oakland, withwhom hehad beencorrespondingfora decade: “I look forward particularly to cooperating with you in making better known the California fish fauna. I no doubt will have newmaterial published from time to time on the systematics and biology of the fishes but will definitely hope that you will 204 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES Year Fig. 3. Number ofcollections offishes made within the study area and archived in the SIO Marine Vertebrate Collection by year (bars) and cumulative number ofspecies recorded from those collections within the study area (line). maintain your plan to work toward a ‘Fishes ofCalifornia.’ It will be a pleasure to make records and other information available for your project.” (Shor et al. 1987, p. 227). That project was published shortly after Hubb’s death in 1979 (Hubbs et al. 1979). In 1958, Professor and Curator Richard H. Rosenblatt (Fig. 2C) was hired to oversee the growing collection of fishes at SIO. He and other researchers and students at SIO actively collected fishes in and around the La Jolla region. By the end of 1969, over 1,000 MVC collections and well over 3,000 lots offishes from La Jolla had been archived in the (Fig. 3). Local collecting of fishes declined after that time due to diverging research MVC interests, and constraints ofspace in the for storage ofspecimens. Since that time, the MVC has archived specimens offishes from the La Jolla region primarily when new or unusual specimens become available, specimens from focused efforts associated with faunal inventories of the area (e.g., Craig et al., 2004; Craig and Pondella, 2006), and MVC voucher specimens for the growing tissue collection, established by H.J. Walker in 1993 (Hastings and Burton, 2008). Materials and Methods We compiled a list of fishes recorded from La Jolla, California based on specimens MVC collected and archived in the of the Scripps Institution of Oceanography, University ofCalifornia San Diego (Table 1). We included all species collected less than W 10 km from shore (east of 117°20.5' longitude) and from Torrey Pines State Beach southward to Tourmaline Beach (32°54' N - 32°48' N latitude). This area includes the entire rocky headland ofLaJolla, aswell as theprimaryconservation areas in thevicinity (Fig. 1). These include the San Diego-Scripps Coastal State Marine Conservation Area, the Matlahuayl State Marine Reserve, the South La Jolla State Marine Reserve and Marine Conservation Area (CDFG 2013). FISHES OF LA JOLLA 205 Whilesomecollectionsdatetotheearly 1900sandafewweremadeinrecentyears,most weremadebetween 1944and 1969(Fig. 3). Archivedspecimenswerecollectedusingawide variety ofsamplingmethods and were taken from the beach or ocean surface to depths of over500meters. SpeciesarelistedinTable 1 arrangedintaxonomicorder. Commonnames offamilies and species follow those recognized by the American Fisheries Society (Page et al. 2013). Brief information on habitat or habitats occupied and special occurrences is provided for each species. Habitat type was divided into three categories as follows: H — hard substrates including rocky reef, rocky intertidal, kelp forest, sides of pier pilings, , bouldersoranyothertypeofhardsubstrate; S= softbottom includingsand,mud,eelgrass , and surfgrass; P = pelagic including species that always swim well above the bottom, as , well as those that periodically or regularly swim several meters above the bottom. Also indicated in Table 1 are the species collected in the La Jolla and Scripps Canyons from > 30metersdepth (= Cn), andthespeciesobservedintransectsconductedinandadjacentto the SDLJER (= Tr) between 2002 and 2005 (see below). Frequencyofoccurrenceisbasedonthenumberoflots(occurrences)ofeachspeciesthat MVC havebeenarchivedinthe thatwerecollectedwithinthedesignatedarea, regardlessof theabundanceofthespecies. Common (C) speciesarethose representedbyelevenormore collections, uncommon U) specieswerecollectedinthe studyareafrom threeto tentimes, ( and rare (R) species were collected only once or twice in the La Jolla study area. In a few cases, species that are known byus orreported by others to be more common in the study area than indicated by collection records are indicated with an asterisk (*). The biogeographic distribution ofeach species was designated based on the mid-point ofthe entire known range of the species. Range endpoints are from Horn et al. (2006), supplemented as needed based on published distribution records (e.g., Love et al. 2005). Southernspecies(S)arethosewhoserangemidpointissouthofPuntaEugeniaontheouter coast ofBaja California (27°50' N), northern species N) have range midpoints north of ( Point Conception (34°27' N), and central species (C) have range midpoints between these well-established biogeographic barriers (Brusca and Wallerstein 1979; Horn et al. 2006). Between2002 and 2005, theabundance offishes associatedwithkelpforestswasvisually surveyed at La Jolla Cove (within the SDLJER, now called the Matlahuayl State Marine Reserve) and an adjacent site (Boomers) a short distance beyond the reserve boundary m (Fig. 1). Both sites have moderate reliefrocky reefs (1-3 high) scattered throughout the area and are dominated by red algal turfreefs and kelp forests (Parnell et al. 2005, 2006). Survey protocols were modeled after established techniques for assessing abundance and densityofconspicuousfishes(McCormickandChoat 1987;Pondellaetal. 2005). Randomly selected, quantitative belt transects were swum by two SCUBA divers for a period of 5 minutes over a distance of50 m. All fishes (excluding pelagic species) observed within a two-meter window (i.e., one meter on either side of the diver and one meter above and below) along the transect were identified to species where possible, and the number of individuals was counted by each diver. Ifcounts from the divers differed, the average was recorded. Each transect accounted for 100 m2 of bottom area surveyed. Three replicate transects wereconducted along rockyreefsubstrates ateach offourdepths (3 m, 6 m, 9 m, and 12 m) for a total of twelve transects at each site per survey period. Surveys were conducted every two months for a total ofsix periods per year between January 2002 and December2005. Atcertain periods throughoutthe study, persistent foulweatherprevented full surveys at each sample site, especially for the shallowest depth contours. However, at least six transects were conducted at every site during each sample period throughout the studyexceptforJuly/August2004whencontinuoushighsurfprecludedsurveysatBoomers. i 206 SOUTHERN CALIFORNIAACADEMY OF SCIENCES = of o <u area. records; P kelp Xg g Bto uaj ^O Zo Zo Zo Zo Zo U 2 etc);in endpoint'C g1 fsrtoumdy1to (reefs, occurrence h0<J02§£(3 fst (oNo Gr"d fftt f(tN or<onn fctn fCtN mo latitude G record r=are, bottom and o ZII w fmt xO oX cmn min oX OX Xm R <D •*> southern 2 -2 fMiVrsCt records; Hha=rd depth, = —2Gaj1 U<oG9u w00 mCN o(N- o(oN ^m fctn XCN <fNt CfNt meters SE T3 r of 10 30 3). H X i X year3to categories: > = Table at Date see atjo‘GG Cn X X X X 1 X X X (Hab) s sw uncommon, Canyons g w X X X Reserve; 0 G IG X X ffi DC X X X Collection. = Habitat 1 1 IX pu ai cL DC DC U Scripps c g Oc D u U * U U U Ecological oj text). or records; Vertebrate more (seeLJolala Jolla gC oCl QCC^DJ iO1f—nNt1 fO1o—tNo1 OfitNn 1oOf—Nt1 fOX^tN fOitNn fOitNn OfitNn1 indicate T3 Marine or for 11 Diego-La s-h bO SIO records records ii u ft ft ithne Cc=ommon, collection collection (SfoarmnerlyZ 23 T®tGc3oO*rS^C=C0cO230 0QOft0to X1<o!GoGo3—o deposited area: than (aagZ>D U2 X<OD OCO oft f13Gt 0GO§3 and area respectively,Reserve ^3 00 00 ft 00 ft ft study •GS wG California, study Marine from ^ I the indicate, in State ft} 2 Jolla, Tr collections La and TLof1cfoialirslbe.sfhoclettmeesd foOiMreo=VcqccfnuurrCeenncecy*iskbntpndimceocoeocamowtrmienoesensspb“uSCeonotlx=dtafnoeg”mtir,c;fiaatMtoarrtrnaholnenusaesdhencuttdsaylX£-12£GS>o><<IUuno032o8o2a«8§!f0-h)t\mooSo&goGop <£U X'-XstacS(22/oU)5?."£^ScKR§^~g$<^O2§—sui xor2SoGtSo-o* IUG1B0tOGZ3))*^fM^G2<3tu fGfi IgGS tXoOr—8cCcctGs-!oOuoo~o• "XfofrOXac*o^eCto--s,;3>lr Xoif*caGft0ot1jtro1-f^^tQ'5gR:O2 3f0ffGgGG<Gg00tttu0-<.^Cas>^_KS> 3ffXSngo4GG2ZttfI>ii ^^^tQf^VSR|GSgtj_ 1 i I FISHES OF LAJOLLA 207 3X 3X Mo 7o1 £o O U(U cQ/5 O(U UU 0o0 0o0 wo Uu m m m wa oo 38.5 40.5 50 m x m 53 27 28 - - x 00 GO X O0 OO P,H X oT X 00 00 X S C X £ * u X X & u U £ c* P* C* C* R* o o m m 1945 <onn oono iOnN XOXN) oOdoN- odOoN- idO-nn dO-N xOn OixNn oOooNf oO—oNI ooOooN 1949 X TShhreasrhekr 0XXWcI0a) «XX3xaocoa »XXocOp—oia h<fOut3X™Oso2£o) ^-™OcoO&i^Xft _XX0cgooo0 XX0XXcPcO<—0LaaD, XX0XQi>2d0n.XXj3ocdoa sX'oXoX2Ogco5a X3X<X2o<2CuJD Xf£|ocofai PERlaecciatfriiycc oo 00 ffl c/3 00 oo oo OO o sharks 1(Bo7nn8ate8rr)e, "2S OP Mwx•3<0SSo0S,3nD3~ «o,—C^2co<onDr .r,o—s-nlr, oXxOO X^oo XO^d* oXSo S elerctariycsA1y8r5e5s, X0<Cu J£<s-3 cc*a j—1 In o dcar b*§s .tads.J&gs*o O torpedo thresh-er vulpinus 3§Sci1a ^xJ<R©CJ 340d>3 X<?%*5S 1*ClL>> -§^S3 - californica Lamniformes AlopiidaeAlopias 1 ^ o 3 ^ ^a ^a £-a £•2 § X>XNcK^VcCol orpediniformes TorpedinidaeTorpedo 00 O0 T 1 1 208 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES U<D U<D O ZO ugo ^O %O 0 0 O 0 u<D U(U 0 0 u<u go GO GO GO GO GO GO m in in in d•|^-t (0N0 dT NxlO- ((NN (iNn nd- (N ccob (ONN <n in n00 0 0 0 O O 0 O O NO O-otO- NO NO in m d- C-oOt d- cdo- of d- dc-o 0 0 n O O OC 00 OO CmN in 00 CO CO (N <N T CN <N ONI X X - - • • ' U 1 X X X ' ' GO GO X GO GO GO GO GO Qh X GO 3U X GO GO GO U U R C u u D U u u u X * Pi obS Pi Pi O—N OocNoo OdN- Od>/N"3 OV'sNf3 NO-3ON- Om0n OO"3NN- OOONO Odi-Nn OdiN-n Oi^Nn1- OTmtN- OOdNO- 0O(0NN mO(NN Continued. >. o3 Pi c>N T1able. xO&N>/ tOJ4<%g0i=3U3 ec0do3 .5aa0jS3P’U£c^nd^fSl3j ^ci01§d3)) S^=b3 •'Q000§c0£bc033?n..i^c2o22SajbP* VT0cVOcidO3Xhd3) '«u130oSSd£33§—h3) P0d>i3N ‘£uIsO2>ci5—d3N '.yMcw%20d§2 %™^w2«^0 -XUcsb0lcsS3-3dlO vo CQ CQ hJ go Oh c* CQ eu O <N oo co oo oo £ 3 43 d no r^h XOa> «^ ^<of2ot. dtNfo-3is *.oCtq5§Qs 3bCs^e<d -.O#Csa2< s4•OiQ-<> d.•JtM82d8do>*dSds1r,,r^o1^^£.c5o§gQsd§2J,S^w0^-&2Pg«<bS^K323ho T4td<2>d3»u.-c£o^&saoo; cts2l0>>??H)?%u'—S<aoOgorriDdT T.0O0<333B tO.qd~Q6o9s •0>2o£dc3N~^<Xp5xoSS3 d1^R—<d1cs«33/ID33 O~~2^i3.—s5S&803u22ngi33.,O"-*^1-fdOc§OSd-f§2xdj>5e,, d-§o"J0iooo<QOS-d3DJHo0Q-Ow.S•fdg0^GO6b-*gd3j,- Q P a D U FISHES OF LA JOLLA 209 Or<u U<p CoO Zg Z2 7o5 7o1 2^ u m ^ (N on m r-H W O CO CO CO CO CO DC DC CO Ph p5 p2 Ph" Ph' Ph Ph Ph Ph' Ph' pH Ph CO Ph £ U u U P^ Pi Pi Pi 23 Pi Pi Pi 23 23 m O >n in in 7i in OVOs O71s O<!)s• TOts On O0s0 O7s) Os 1954 1965 > w O > -od aoo bij 5p G (o Lizardfish Lancetfish ^ o <d •S -S h .£ 3 «* £do •S ^ o| O 73 O o boO <Dau.JOa>-SG To3 y o ^ 2 California Longnose Q Q < £ 03 j__ o3 co Ph Pi Hh Ph ro-o^do~>"o0 5 d .*£3 ooo o—ITo) ^ 5 d £ <D 1855) aS"DodoGO3iG5^«rC; .->2Ca0§Gl3 3w-10a>O2ro-5>ojsh1 T'03—3 0=S0 2O>/23 '-.dDoi2 T^doOC53335 2T«%«-<2*H, SiiD/is2Oo4l_So-Hr *j1dd0o^2p33o "~^^S2jCK5oJ2l.n sPK<2Po3 ^c0dbi)o l(uAcyiroceesp,s lancetfi-shes fL1eo8wr3eo3,x zts0aaoo3 R^^.^2s2 PTWgb£30O-aO^t<jh3 0IDdi 0§|S'•^ tq 2ddIb£VDho ‘<j(b0-doD3Go ^-bfC<S\»S>o ’goIsl£-DH 'CSd0oD0dO33 ~6c2RPR\ jC<aJao§/-DHD 25^32dO5h ^RC2^,T5 £'3C<alOO/-UDHh dC-Cdood>Oau.Synodus AlepisauridaeAlepisaurus <i-H 13 C CO 55

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