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Home , Anarchias galapagensis, Apterichtus, Caranx lugubris, Clipperton Island, Doryrhamphus excisus, Echidna nocturna

Rev. Biol. Trop., 45(2): 813-843, 1997

An annotated che�klist of the of Clipperton ,

GeraId R. Allen 1 and D. Ross Robertson2 I Department of Aquatic Vertebrates, Westero Australian Museurn, Francis Street, Perth, WA6000, . Fax: +61.9.3288686 Email: @multiline.com.au Smithsonian Tropical Research Institute (Balboa,Panama). Mailing Address: STRI, Unit 0948, APO ÁA34002-0948, USA Fax: +507-2280516 Email: [email protected]

(Received IO-IX-1996. Corrected 24-11-1996. Accepted 28-11-1996)

Abstract: Clipperton Atoll is the most isolated, most westerly and largest reef in the tropical eastero Pacific (TEP). We collected and recorded fishes to depths of 60m during a two week visit in April 1994. Previous collections were made late last century andby expeditions from Scripps Institution of Oceanography in 1956 and 1958. One hundred and fifteen identified species from 89 genera and 43 farnilies are now known from the . The most speciose families present at the island are moray (Muraenidae, 14 species), jacks (, 11 species), wrasses (Labridae, 8 species), surgeonfishes (Acanthuridae, 8. species), squirrelfishes (Holocentridae, 5 species), (, 5 species), and (Balistidae, 5 species). These represent -14% of the TEP nearshorefish fauna and 40% of its genera. The 115 species inelude 14 offshore pelagic species, 22 inshore pelagic and midwater species, 70 demersal species that live 011 hard reef substrata, and nine demersal species tltat live in or feed on unconsolídated substrata (rubble and sand). Among the IOlnon-oceanie species, 68% ate carnivoreson mobile organisms, 9.2% feed on sessile benthie invertebrates, 12.9% are planktivorous, and 17.8% are benthie feeding herbivores. At least 70 of the non-oceanie speeies appear to have resident (i.e. self sustaining) populatións at the atoll, while 17 species probably are vagrants. Clipperton's fishes inelude 63 rranspacific species (i.e. species that also oceur on the westero side of the Bastero Pacifie Barrier) and 52 species endemie to the TEP. While most (36) of the TEP species oeeur throughout the region, four oceur o¡:¡ly at boh Clipperton and the Revillagigedos, the nearest shoal habitat,950 km to the north. Nine species or subspeeies Crom seven families are endemie to Clipperton. They represent 11.3% of Clipperton's demersal shorefishes. The sister speeies of one of them likely is a transpacifie species, while the sister species of the other eight Iikely are TEP speeies.

Key words: Bastero Pacific, , fish fauna.

Clipperton IsIand, at lOol8'N, I09°13'W, Revillagigedo IsIands (Fig. 1). It is the most is a small island at the westem edge of the isolated shoaling reef in the TEP and the most tropical Eastem Pacific (TEP) marine isolated reef in the Indo-Pacific tropics. There biogeographic region. It lies at the eastem edge has been long standing interest in relations of the 4,000-7,000 km wide Eastem Pacific bétween the eastem and westem Pacific fish Barrier (EPB) (Ekman 1953), 1,lookm SW of faunas from a biogeographicaI perspective, the American mainland and 950 km S from the because the EPB is the world's lilrgest deep nearest offshore shoal habitat, the water barrier to the dispersal ofmarine shore 814 REVISTA DE BIOLOGIA TROPICAL organisms (Ekman 1953), and the fishes of the fishes based on our 1994 collections and TEP have obvious affinities to those of the observations, scattered references in the central and westernPacific (Herre 1940, Ekman literature, and records of Clipperton specimens 1953, Briggs 1961, Springer 1962, 1967, deposited at several institutions. A companion Rosenblatt et al 1972, Leis 1984, Brothers and paper, Robertson and Allen (1996), presents a Thresher 1985, Clark 1995). Due to its zoogeographic analysis of the structure of the position Clipperton may be an important shorefish fauna of Clipperton. stepping stone allowing a connection between those faunas. The first fish recorded from Clipperton METHODS Island was Bathygobius arundelii, described by Gatman in 1899. It was collected by the The island and its marine habitats: The Hopkins-Stanford Galapagos Expedition of tiny island (Fig. 1) is roughly circular with a 1898-99. Eight other species were also maximum diameter of 4km and a combined collected and reported by Snodgrass and Heller land and area of about 10 km2• Jt (1905). The only other coBections besides our's consists of a narrow, unbroken 40-400m wide reportedhere were made by Scripps Institution ring of land arounda central lagoon. Except for of Oceanography (La Jolla, California) during a large basalt rock that rises to 21m on the two visits of several weeks in 1956 and 1958. southeastern comer, the land is less than -5m Most of those collections were made on the aboye level and consists of coralline rock shallow reef flat. Until now, they remain with a thin covering of loose rubble. The largely unreported, with the exceptions of island's terrestrial flora was described in detail descriptions of three Clipperton endemics - by Sachet (1962). Skaggs (1989) provided a Holacanthus limbaughi Baldwin (1963), comprehensive account of the history of human Myripristis gildi Greenfield (1965), and occupation of the island, which is a French Stegastes baldwini ABen and Woods (1980). territory. We encountered dozens of abandoned In addition, records of a few other species are boat anchors of a variety of sizes and ages, and scattered through the literature (Schmidt and forms of construction (sorne obviously not of Schultz 1940, Springer 1962, Limbaugh 1963, standard commercial origin) in lO-20m of water Beny and Baldwin 1966, Springer 1967, around the island, indicating a history of Rosenblatt et al 1972, Fritsche 1980, Pietsch repeated recent human usage of thereef. During and Grobecker 1987, McCosker and Randall our 1994 visit crews from a tuna boat line­ 1993, Randall 1995). fished forreef fish from work boats at lO-20m Clipperton fishes were featured in faunal depth, and several sailing yachts also anchored summaries for the TEP by ABen and at those depths. Robertson (1994) and Fischer et al (1995). Clipperton is the only coral atoll in the Descriptions of three additional Clipperton TEP and its 3.7 km2 is the largest in endemics, Thalassoma robertsoni ABen the region (Glynn et al, 1996). Steep rubble (1995a), Pseudogramma axelrodi Allen and beaches fringe the seaward edges of the ísland. Robertson (1995a), and Xyrichtys wellingtoni These beaches lead into a shallow (O.5-lm ABen and Robertson (1995b), resulted from our deep, depending on tides) reef flat, covered in 1994 expedition. Summaries of the 1994 coralline rock and rubble interspersed with Clipperton Expedition appeared in the appendix small patches of shallow sand. There are only a oíAlIen and Robertson'sFishes of the tropical few lOs of meters of beach rock. This 50-200m eastem Pacific (1994), and in a popular artiele wide habitat is bordered to seaward by a 10- by ABen (l995b), which ineluded color 15m wide semi-emergent spur and groove photographs of most reef-dwelling species. Here we present an annotated list of CIipperton Allen & Robertson: CJipperton Atoll 815

*

1 Km

Fig 1. Map of Clipperton Atoll, showing subtidal sites (stars) we sampled in 1994. Land is solid color, bordered by an irregular Une indicatingthe edge of the intertidal reeffiat. The site of the single reeffiat rotenone sample is indicated by a star on tha! fiat on theSW side of the island. system that stretches around the entire island. of about 30-45 degrees (the outer reef slope). The atoll lacks a leeward seaward exposure, The inner reef slope consists almost entirely of which is a common feature of other Pacific hard bottom, with a scattering of small (Glynn et al 1996). swells broke depressions containing a thin layer of coral continuously on this spur and groove rim rubble, but Httle sand. At the time of our visit, around the island throughout our visito Outside live coral cover was high on the reef slope; on this rim the reef slopes gently to -20m depth the order of 33-83% in many arcas (Glynn et over distances of -100-300 m (the inner reef al, 1996). There is only one significant patch slope), then plunges into deep water at an angle of fine sanu bottom in shallow water; it filIs a 816 REVISTA DE BIOLOGIA TROPICAL depression about 500m2 in area in 15m of total of 82 dives with SCUBA to depths of water offthe northwest comer of the island on SOmo The average duration of each dive was 40- fueouter partof the inner rref slope. However, 60 minutes, with au average maximum depth while diving to depths of 50m we observed of 20-25m. Those dives were scattered around extel1sÍveareas of sand and rubble bottom on a aH sides of the island (see Fig 1). In addition tenace at the base of the outer rref slope al we made half a dozen drift snorkel dives along about 60-7Om depth off the north east and the upper slope, covering dístances of -1-2Iun western sides of the island. While this "60m on each. The names and abundances of observed tenace" varies in width from about 30-100m species were recorded on plastic slates, and we around much of the island, and has a slope made coHections using mini-spears powered by ranging from 10-40°, on toe NW side of tlle smaH rubber bands, and small rotenone island ít is greatly enlarged and slopes off stations. AH but one of toe -15 rotenone gradually from SOm to 110m over a distance of collections were made on the outer reef slope at l.5km (sounding profiles in Glynn et al. 15 to 30m depths; a single collection was rnade 1996). Duriug our visir visibiiity was usually on the reef flat adjacent to shore on the in the muge of 25-40m, and surface water southwestern side oear the landing site (see Fig temperatures were 28-29°C. 1). Constant wave surge prevented liS from The ¡stand has a large (7.3 km2) central using rotenone in the groove and spur zone and lagoon. That lagoon apparently recently had a in the shaHow parts of the upper reef slope. pair of very shallow connections to toe open Fishes were fixed with 10 percent forrnalin, sea, which eventuaHy c10sed around tile end of and later transferred to 70 percent ethanol. Our the last century (Sachet 1962, Skaggs, 1989). specimens are deposited at the Members of the Scripps Expedition to Natural History Museurn, Washington, D.C. Clípperton in 1958 found evídence of a recent (USNM). Underwater photographs of many channel, thought to be the result of a violen! species are deposited at the Western Australian stonn (Wayne Baldwin, personal Museum, Perth and the Smíthsonian Tropical communication to G.R. ABen 1995). During Research Institute, Panama. If known to liS we our vísit the lagoon had no swface connection also indicate the iocation of previous to the sea and was slightly brackish. It collections from Clipperton, which a'"e supported a dense growth of freshwater weed deposited at the USNM, Scripps Institution of and sornescattered parches of reeds and sedges. Oceanography (SIQ), the Califomia Acudemy We did not find auy fishes in the lagoon duril1g of Sciences (CAS), the Univer3íty of several searches using a mask and snorkel. California at Los Angeles (UCLA), and the Hordes of midwater-swirnming isopods were Los Angeles County Museum (LACll/i). the only macroscopic Jife. Depths in Specimens coHected on the Scripps expeditions most of fue lagoon are shallow (less than about of the 19505 by W. Baldwin, C. Limbaugh, E. 10m), but Cousteau and two other divers from Reese etc. were il1itialIy deposüed at UCLA, R.V. Calypso, which visited in 1980, díved to but about half that collection was subsequently 37m in fue northern section (Skaggs, 1989). tral1�;ferred to LACM. They found no sign of lite at this depth, only The phylogenetic sequence of families inky darkness, a high level of hydrogen sulfide, appearing in the species list foHows Eschmeyer and a softmud-ooze bottom. (1990). Genera and species are arranged alphabeticaUy. Brief literature citations are The 1994 expeditioll! We visited included Ín fue text for the original description Clipperton Island aboard the 29m sport-fishing of each species (inc1uding the type locality) and boat Royal Star on a trip out of San Diego, its first record from Clipperton. Annotations California. During 13 days spent at the island for each species refer to: (1) relative abundance between14-26 April 1994 we marlea combined and depth distribuuon on the Clippelton reef; Aikn .(f.Rol)eJ:I:sOl.1!:C1ípperton Aton 817

(2) growfu stages coHected or observed (íe. populations. TIIey include sorne species that small individuals that apparently had recently were un.comrnon during our 1994 visit,but of recruited from the pelagic environment,larger which substantial numbers of specimens were juveniks, adults); (3) major activity mode coHected at Clipperton in the 1950s; Vagrants (diurnal/noctumal; benthic, pe1agic,etc.); (4) are species that were so rare during our visit, social behaviour; (5) majar feeding category; and/or for which <5 specimens from (5) reproductive morle; (6) approximate Clipperton are known from previous maximum known size; (7) general collections,that they p:obabiy recIllited from distributional range; (8) an indication of ofuer sites (although we cannot exclude the whether or not photographs were obtained at possibility that they are remnants of a local Clipperton (field is omitted if no3 photographs); population that is approaching extinction). We and (9) the number arrd size muge of collected assigned a species to one of these population specimens,or an indication that the species status only when sufficient infonnation was observed on!y. Additkmal armotations r¡;;fer appeared to be available. Uncertain - refers to to deposition of specimens at various species as rare as the vagrants, but wruch institutions, the probable population status . canoo! be assigned to either Resident or (resident or vagrant - see below), and Vagrant statusbecause they may be more systematic relationships. Lengths are given as abundant than coUectionsindicate, due either to standardlength (SL) for most species,which is their cryptic behavior and/or the possibility the distance from the tip of the SnOtlt to the that significant numbers of indi viduals could be base of the caudal fin. Tota! length (TL) is present on sand and rubble habitats of the "60m given for a few fishes which do not have a terrace". dearly defined tail (e.g eels). Fork length (FL - the distance from the snout tip to the end of the middle caudal tln rays) is used for sorne RESULTS commercial fishes suchas tunas. An annotated species lis! 01 the fishes Popullation categories: We use four known from Clipperton Atoll categories oí re1ative abunda.'1ce derivedfrom Carcharrunidae - Whaler Sharks our impre ssion of the QvetaU abw)dul){:e of each Carcharhinus albimarginatus (Rüppell, species at the is1and .in 1994: Abundan! - 1837) observed al alrnost aH SÜes in a variety Qf Ca reharías . albimarginatus RüppeH, 1837: habitats, with nundrerls to tbousands of 64 (Ras Mehomet, ). individuals routin eiy seen Oil each dive; Carcharias platyrhynchus.- Snodgrass and

Common • seen al the majority of sites, wit.h HeBer,1905: 344 (Clipperton Island). ,...50 individuals usuaUy seen on each dive; An inshare species occasional1y seen on fue

Occasional •. often localized and rwt observed oliter slope; adults only; roving predator; bu most dives, usuaHy <20 individuals seen Oll solitary or in gtoups; canllvore: fishes,rays, any dive; Rare � fewer than 20 individuals and cephalopods; viviparous;300 cmTL; Jndo­ (often iess),seen during the entire expedition. Pacific to !he ; observed only; Based on a combination of·. their abundance at Limbaugh (1963) published observations made

Clippertonduring.. 1994, and/or in collections at Clipperton in the.1950s; resident. made in the 1950s,andouroverall impressions gained fmm otherareas scattered around the Carcharhinus falciformis (Müller and Indo-Pacific we assigned the non-ocearnc Henle,1841) species to one of three population categories: Carcharias falciformisMüller and Henle, Residents species that appeared sufficientJy 1841: 47 (): abundant to have established, self-recruiting An offshore pelagic species occasionaHy 818 REVISTA DE BIOLOGlA TROPICAL seen on the outer slope, usually near the surface; adults only; roving predator; solitary or Muraenidae - Moray Eels in groups; carnivore: fishes, cephalopods; viviparous; 300 cm TL; circumtropical; galapagensis (Seale, 1940) underwater photographs. Uropterygius galapagenis Seale, 1940: 6 (Galapagos ). Carcharhinus galapagensis (Snodgrass and Abundant on reef fIat in O- 1m; juveniles Heller, 1905) and adults; cryptic; solitary; carnivore: fishes Carcharias galapagensis Snodgrass and and ; pelagic eggs; 30 cm TL; TEP, Heller, 1905: 343 (Galapagos Islands). from to the Galapagos; 23: Carcharinus galapagensis.- Limbaugh, 79-133 mm TL;re sident; specimens at USNM, 1963: 75 (Clipperton Island). LACM. A nearshore pelagic species occasionally seen on the outer slope; adults only; roving nebulosa (AhI, 1789) predator, solitaryor in groups; carnivore: fishes Muraena nebulosa Ahl, 1789: 5 and cephalopods; viviparous; 370 cm TL; (Indonesia). circumtropical; resident; specimens at SIO, Occasionally sighted on the outer slope in LACM. Limbaugh (1963) noted this species 5-15 m; juveniles and adults; cryptic; solitary; was abundant in the 1950s, with many carnivore: crustaceans; pelagic eggs; 60 cm TL; juveniles. Indo-west and central Pacific; 2: 110-169 mm TL; underwater photographs; resident; Carcharhinus limbatus (Valenciennes specimens at USNM, LACM, S10. 1839) Charcharias (Prionodon) limbatus Echidna nocturna (Cope, 1872) Valenciennes (1839), in Muller and Poecilophis nocturnus Cope 1872: 474 Henle,1839: 49 () (Rio Grande, ) Carcharhinus limbatus.- Limbaugh, 1963: Not observed or collected by us; collected in 75 (Clipperton Island) the 1950s; cryptic; solitary; carnívore: Not seen by us ; a nearshore pelagic crustaceans; pelagic eggs; 71 cm TL; species; roving predator, solitary or in groups; widespread in TEP;population status uncertain carnivore: fishes and cephalopods; viviparous; due to cryptic behavior and possible presence 247cm TL; circumtropicaI; possible vagrant; below 50m; specimens at USNM, ueLA. specimens at LACM. Limbaugh (1963) observed several individuals in the 1950s. Enchelycore octaviana (Myers and Wade, 1941) Sphymidae - Hammerhead Sharks octavianus Myers and Wade, 1941: 89 (Octavia Bay, . Sphyrna lewini (Griffith and Smith, 1834) Rare, only 3-4 individual s sighted, in 10-20 Zygaena lewini Griffith and Smith, 1834: m; adults only; cryptic; solitary; carnivore: 640 (south coast of Australia). fishes; pelagic eggs; 70 cm TL; TEP; Rare, a few seen on outer slope; a nearshore underwater photographs; population status pelagic species; adults only; roving predator, undeardue to cryptic behavior; specimens at solitary or in groups; carnivore: fishes, USNM. cephalopods; vlvlparous; 330 cm TL; circumtropical; population status uncertain. Enchelynassa canina (Quoy and Gaimard, Limbaugh (1963) observed groups of adults of 1824) an unidentifiedspecies of this during the Muraena canina Quoy and Gaimard 1824: 1950s. 247 (Vaigiou and Rllwak) Allen & Robertson: Clipperton Atoll 819

Enchelynassa canina.- Rosenblatt et al SIG, LACM. 1972: 11 (Clipperton) Not seen or collected by us; solitary, Gymnothorax flavimarginatus (Rüppell, camivore: fishes, crustaceans; pelagic eggs; 1830) 250 cm TL; westem Pacific to the Americas; Muraena flavimarginatus Rüppell, 1830: classed as a resident because numerous 119 (Red Sea). specimens were collected in the 1950s; Gymnothoraxflavim arginatus.- Rosenblatt specimens at SIG, LACM. et al 1972: 10 (Clipperton Island). Occasionally sighted on outer reef slope; Gymnomuraenazebra (Shaw, 1797) adults only; cryptic; solitary; camivore: fishes, Gymnothorax zebra Shaw, 1797: PI. 322 crustaceans; pelagic eggs; 125 cm TL; Indo­ (Pacific Gcean). Pacific to the Americas; 2: 109-125 mm TL; Echidna zebra.- Rosenblatt et al 1972: 10 underwater photographs; resident; specimens at (Clipperton Island) USNM, LACM. Rare, only two individual s sighted on upper edge of outer slope in 10m of water; adults Gymnothorax panamensis (Steindachner, only; cryptic; solitary; camivore: crustaceans; 1876) pelagic eggs; 150 cm TL; Indo-Pacific to the Gymnothorax panamensis Steindachner, Americas; underwater photographs; population 1876: 67 (Panama) status uncertaÍn due to cryptic behavior; Gymnothorax panamensis.- Randall and specimens at SIG, LACM. McCosker, 1975: 16 (Clipperton Island) Rare,only a few specimens collected by us; Gymnothorax buroensis (Bleeker, 1857) cryptic, solitary, camivore: crustaceans, fishes; Muraenaburoensis Bleeker 1857: 79 (Buru pelagic eggs; 43 cm TL; widespread in TEP; Island, Indonesia) numerous specimens collected in 19508 Gymnothorax buroensis.- Rosenblatt et al therefor probable resident; specimens at 1972: 10 (Clipperton Island) USNM, LACM. Not seen or collected by us; cryptic; solitary; carnivore: fishes; pelagic eggs; 47 cm Scuticaria tigrina (Lesson, 1829) TL; Indo-Pacific to the Americas; population Ichthyophis tigrinus Lesson, 1859: 399 status uncertain due to cryptic behavior and (Indonesia). possible occurrence below 50m; collected In Rare, none seen by us, but severa} sighted the 1950s; specimens at LACM, UCLA. by other members of our party when turning over rocks in search of shells; adults only; Gymnothorax dovii(Günther, 1870) cryptíc; solitary; camivore: fishes and Muraena dovii Günther, 1870: 103 crustaceans; pelagic eggs; 140 cm TL; Indo­ (Panama). Pacific to the Americas; underwater Gymnothorax dovii.- Allen and Robertson, photographs; population status unclear, 1994: 303 (Clipperton Island). because normally associates with rubble and Relatively common, up to 10-20 hence may occur on "60 m terrace"; not individuals seen on every dive; less cryptic than collected in the 1950s. most morays, frequently seen swimming away from cover; adults only; solitary or Síderea picta (Ahl, 1789) occasionally in pairs or small groups; Muraena picta Ahl, 1789: 6 (Indonesia). camivore: fishes; pelagic eggs; 124 cm TL; Common, numerous individuals seen and southem half of TEP, including the mainland coUected on reef flat in 0- 1 m; juveniles and and Galapagos (Fischer et al 1995); underwater adults; cryptic; solitary; camivore: crustaceans; photographs; resident; specimens at USNM, pelagic eggs; 120 cm TL; Indo-Pacific to the 820 REVISTA DE BIOLOGIA TROPICAL

Americas; 2: 43-50 mm TL; resident; benthicinvertebrates; pelagic eggs; 50 cm TL; specimens at USNM, LACM, SIO. Clipperton and only; 3: Uropterygius macrocephalus (Bleeker, 145-170 mm TL; resident; specimens at CAS, 1865) USNM, LACM, UCLA. Gymnomuraena macrocephalus Bleeker, 1865: 54 (Indonesia). Phaenomonas pinnata Myers and Wade, Common on reef flat in O-1m; juveniles 1941 and adults; cryptic; solitary; carnivore: fishes Phaenomonas pinnata Myers and Wade, and crustaceans; pelagic eggs; 47 cm TL; Indo­ 1941 Pacific to the Americas; 11: 91-284 mm TL; Not seen or collected by us, but collected resident; specimens at USNM, LACM and SIO on reef flat in the 1950s; lives in sandy (as U.necturus). habitats; nocturnal; cryptic; solitary; carnivore: benthic invertebrates;TEP, at scattered Uropterygius versutus Bussing, 1991 localities between the Gulf of California mxl Uropterygius versutus Bussing, 1991: 97 the' Galapagos; population status uncertáin; (Costa Rica). specimens at LACM. Only one subadult collected on reef flat in O-1m; cryptic; solitary; carnivore: fishes mxl Chanidae - Milkfishes crustaceans; pelagic eggs; 56 cm TL; TEP, from Gulf of California to Colombia, Chanos chanos (ForsskAl, 1775) including the Galapagos; probable resident as Mugil chanos ForsskAl, 1775: 74 (Red numerous specimens were collected in the Sea). 1950s; specimens at LACM (as U. schultzi). Rare, a single adult seen near surface over upper reef slope; forms aggregations; benthic - Snake Eels fe".... ng herbivore: algae and detritus; pelagic eggs; 160 cm FL; Indo-Pacific to the Apterichtus equatorialis (Myers and Wade, Americas; observed only; vagrant. 1941) Caecula equatorialis Myers and Wade, Antennariidae- Anglerfishes 1941: 75 (Galapagos Islands). Regularly collected with rotenone on sand Antennarius coccineus (Cuvier, 1831) patches in 10-20 m; juveniles and adults; Antennarius coccineus Cuvier, in Lesson cryptic; solitary; carnivore: fishes mxl 1831: 143 () crustaceans; pelagic eggs; 30 cm TL ; TEP, Antennarius coccineus.- Pietsch mxl from to the Galapagos Islands; 3: 19Q.; Grobecker 1987: 149 (Clipperton Island) 296 mm TL; resident; specimens at USNM. Not seen or collected by us, recorded from Clipperton,by Pietsch and Grobecker (1987); Myrichthys pantostigmius Jordan mj cryptic, solitary camivore: fishes; pelagic eggs; McGregor, 1899 9.1 cm SL; widespread in Indo-Pacific mxl Myrichthys pantostigmius Jordan mxl TEP; probably residentas numerous specimens McGregor, 1899: 2802 (; were collecterl from the reef flat in the 1950s; Revillagigedos) specímeris at LACM, UCLA. Myrichthys pantostigmius.- McCosker mxl Rosenblatt, 1993: 164 (Clipperton Island) Antennafius sanguineus Gill, 1863 Occasionally sighted in small depressions Antennarius sanguineus Gill, 1863b: 91 containing sand on outer slope, to 20 m depth, (Cabo San Lucas, Baja California). and on shallow reef flat; juveniles and adults;, Antennarius sanguineus .- Pietsch mj nocturnal benthic; solitary; carnivore: small Grobecker 1987: 158 (Clipperton Island). ABen & Robertson: Clipperton Atoll 821

Relatively common, several colIected vegetation; 25 cm FL; eastern Pacific; 1: 22 (mainly juveniles) at each rotenone station in cm FL; specimen at NMNH. 10-35 m; juveniles and adults; cryptic; solitary; camivore: fishes; pelagic eggs; 7.5 cm SL; Cheilopogon spilonotopterus (Bleeker, TEP, from Gulf of California to ; 11: 7- 1866) 52 mm SL; resident; specimens at USNM, Exocoetus spilonotopterus Bleeker 1866: LACM, UCLA. 113 (Indonesia) Not seen or collected by us, occurrence at Antennatus strigatus Gill (1863) Clipperton is based on distribution map of Antennarius strigatus Gill, 1863: 92 (Cabo Fischer et al (1995); oceanic surface dweller; San Lucas, Baja California). solitary or in groups; planktivore; eggs Antennatus strigatus. - Pietsch and attached to floating vegetation; Indo-Pacific to Grobecker 1987: 190 (Clipperton Island) the Americas; 38 cm FL. Not seen or collected by us; numerous specimens collected on reef flat in the 1950s; Fodiator acutus rostratus (Gunther, 1866) cryptic, solitary camivore: fishes; pelagic eggs; Exocoetus rostratus Gunther 1866: 280 8 cm SL; widespread in TEP; probable (Hawaiian Islands, but doubtful) resident; specimens at CAS, UCLA, SIO, Fodiator acutus rostratus. - Fischer et al LACM. (1995): 1101 (Clipperton Island) Not seen or collected by us, occurrence at Belonidae - Needlefishes Clipperton is based on distribution map of Fischer et al (1995); oceanic surface dweller; Tylosurus acus melanotus (Bleeker, 1851) solitary or in groups; planktivore; eggs Belone melanotus Bleeker, 1851a: 94 attached to floating vegetation; 19 cm FL; (Java) temperate and tropical eastern Pacific, Tylosurus acus melanotus.- Fischer et al Californiato Galapagos; another subspecies F. 1995: 925 (Clipperton Island) acutus acutus occurs in the tropical Atlantic. Common on outer slope near surface during our visit, mainly on NE side of island; adults Hemirhamphidae - Halfbeaks only; roving predator; solitary or in groups; carnivore: fishes; eggs attached to floating Euleptorhamphus viridis (van Hasselt, vegetation; 83cm SL; a circumtropical species 1823) with two subspecies, T. acus pacificus from Hemirhamphus viridis Van Hasselt .1823: TEP mainland, and T. acus melanotus from 131 (India) remainder of its range, including the Euleptorhamphus viridis.- Fischer et al Revillagigedo, Clipperton and Cocos islands in (1995): 1180 (Clipperton Island) the TEP (Fischer 1995); resident; specimens at Not seen or collected by us, collected at LACM (as Belone persimilís). Clipperton in the 1950s; Indo-Pacific to the Americas; oceanic surface dweller; solitary or Exocoetidae - FIyingfishes in groups; planktivore; eggs attached to floating vegetation; tropical and temperaté Cheilopogon heterurus hubbsi Parin, 1961 waters of Indo-Pacific; 37 cm SL; specimens at Cheilopogon heterurus hubbsi Parin, SIO, LACM. 1961: 85 (24°58'N, 117°49'W) One adult colIected from deck of Royal Oxyporhamphusmicropterus (Valencien­ Star; a warm temperate·(Fischer et al 1995), nes, 1847) oceanic surface dweller; solitary or in groups; Exocoetus micropterus Valenciennes in planktivore; eggs attached to floating Cuvier and Valenciennes 1847: 127 (King 822 REVISTA DE BIOLOGIA TROPICAL

George Sound, ) although weactively searched for this species Oxyporhamphus micropterus.- Fischer et al we may not have recorded ít because its close (1995): 1181 (Clipperton Island) resemblance to M. bemtdi (which greatly Not seen or collected by US, collected at outnumbered M.gildi in collections made in the Clipperton in the 1950s; oceanic surface 1950s); resident on the basis of its restriction dweller; solitary or in groups; planktivore; to Clipperton; specimens at UCLA, USNM, eggs attached to floating vegetation; 37 cm SL; LACM. circumtropical; specimen at LACM. There arefour members of this genus in the TEP (ABen and Robertson 1994, Fischer et al Holocentridae- Squirrelfishes 1995) M. berntdi, M. clarionensis, M. gildi, and M. leiognathus. Greenfield's (1965) non­ Myripristis berndti Jordan and Evermann, cladistic analysis indicates that, among those

1903 species, M. gildi . is most closely related to M. Myriprístis bemdti Jordan and Evermann, berntdi, and he suggested that M. gildi is 1903: 170 (). derived from an ancestor of M. berntdi that Myripristis murdjan (non Forsskiil).­ previously invaded the TEP. However, since Greenfield, 1965: 240 (Clipperton Island). M. berntdihas a number of relatives in eastem Abundant, numerous individual s seen on Oceanía the issue of the phylogenetic outer slope in caves and crevices between 15-40 relationships and origins of M. gildi remains to m; small to large adults; nocturnal benthic; be resolved through cladistic analysis of the solitary or in groups; carnivore: large Pacificand neotropical members of the genus. zooplankton ( larvae, etc.); spawns pelagic eggs; 24 cm SL; Indo-Pacific to the Plectrypops lima (Valenciennes, 1831) Americas; underwater photographs; resident; Myripristis lima Valenciennes, in Cuvier specimens at USNM (as M.murdjan), LACM. and Valenciennes, 1831: 493 (Mauritius). Rare,a single adult collected with rotenone Myripristis clarionensis Gilbert, 1897 in 30 m; nocturnal benthic; solitary; carnivore: Myripristis clarionensis Gilbert, 1897: fishesand crustaceans; pelagic eggs; 15 cm SL; 441 (CIaríon Island). Indo-Pacific to the Arnericas; 1: 63 mm SL; Myripristis clarionensis.- Greenfield, population status uncertain, as it normal1y 1965: (Clipperton Island) inhabits deep water (see Randall et al 1985); Common on outer slope in caves and specimen at USNM. crevices between 15-40 m; juveniles and adults; nocturnal benthic; solitary or in groups; Sargocentron suborbitalis (Gill, 1864) carnivore: large zooplankton (crustacean larvae, Holocentrus suborbitalis Gill, 1863a: 86 etc.); pelagic eggs; 18 cm SL; TEP, from (Cabo San Lucas, Baja California). Revillagigedo Islands and Clipperton only; 5: Sargocentron suborbitalis.- Fischer et al 70-110 mm SL; underwater photographs; 1995: 1185 (Clipperton Island) resident population; specimens at USNM, Fewer than fiveadults seen by us at -20m LACM. depth, and a dozen or so recently recruited juveniles collected by rotenone on the reef flat; Myripristis gildi Greenfield, 1965 numerous specimens collected with rotenone Myripristis gildi Greenfield, 1965: 241 on the reef flatand offshore at thesurface in the (Clipperton Island). 1950s; nocturnal benthic; solitary or in groups; Not seen or collected by us; nocturnal carnivore: crustaceans, worms, small fishes; benthic; solitary or in groups; carnivore: large pelagic eggs; 22 cm SL; TEP from Gulf of zooplankton (crustacean larvae, etc.); pelagic California to , also Galapagos Islands; eggs; 25 cm SL; endemic to Clipperton Island; underwater photographs; resident; specimens at ABen & Robertson:Clipperton Atoll 823

USNM,LACM. the eastern Pacific(Fritzsche 1980).

Aulostomidae - Trumpetfishes - Scorpionfishes Aulostomus chinensis (Linnaeus,1766) Pontinus vaughni Barnhart and Hubbs, Fistularia chinensis Linnaeus, 1766: 515 1946 (). Pontinus vaughni Barnhart and Hubbs, Aulostomus chinensis.- Rosenblatt et al 1946: 371 (Baja California) 1972: 14 (Clipperton Island). Pontinus vaughni.- Fischer et al 1995: Both yellow- and green-phase fish relatively 1552 (Clipperton Island) common,several seen on every dive in 5-60 m; Not seen or collected by us; cryptic, mainly adults and several juveniles; diurnal solitary; carnivore: fishes and crustaceans; benthic; solitary; carnivore: fishes, crustaceans; pelagic eggs; 45 cm SL; an eastern Pacific pelagic eggs; 62 cm SL; Indo-Pacific to the species that ranges from central to lower Baja Americas; underwater photographs; resident; California and has been collected at the specimens at USNM,LACM. Revillagigedos and Clipperton; probably a vagrant as only one specimen is known from Fistulariidae - Flutemouths Clipperton and it probably is a warm temperate species; however, because other members of Fistularia commersonii Rüppell, 1838 the genus live in deep water (Fischer et al Fistularia commersonii Rüppell, 1838: 1995) it is possible that it is a resident below 142 (Red Sea). 50 m; specimen at SIO. Several adults seen by us in 10-20 m on NE side of island; specimens were cóllected in Scorpaenodes xyris (Jordan and Gilbert, the 1950s; roving predator; solitary or in 1882) groups; carnivore: fishes,crustaceans; pelagic Sebastopsis xyris Jordan and Gilbert, eggs; 11 cm SL; Indo-Pacific to the Americas; 1882a: 369 (Cape San Lucas,Baja California). 1: 145 mm SL; probable resident; specimens - A secretive, rarely seen species, severat LACM, SIO. specimens collected with rotenone on rocky - Pipefishes and Seahorses slopes to depths of 35 m, numerous specimel1s collected in the 1950s; juveniles and adults; excisus excisus Kaup,1856 nocturnal benthic; solitary; carnivore: fishes Doryrhamphus excisus Kaup,1856: 54 (no and crustaceans; pelagic eggs; 7 cm SL; TEP, locality given). from Southern California to Peru, also Doryrhamphus melanopleura.- Fritzsche, Galapagos Islands; 20: 13-69; probable 1980: 189 (Clipperton Island). resident; specimens at USNM,SIO, LACM. Commonly collected in most rotenone stations in 15-35 m; juveniles and adults; Serranidae- Groupers cryptic; solitary or in pairs; carnivore: small benthic and planktonic crustaceans; male dermatolepis (Boulenger, broods eggs in pouch or on ventral surface; 4.1 1895) cm SL; Indo-Pacific to the Americas; 12: 22- dermatolepis Boulenger, 41 mm SL; resident population; specimens at 1895: 256 (Cape San Lucas,Baja California). USNM. There are two subspecies of this Epinephelus dermatolepis.- Smith 1971: species in the eastem Pacific (Dawson 1985), 162 (Clipperton Island). D. excisus paulus, which is endemic to the Abundant on outer slope in 10-40 m; Revillagigedo Islands,and D. excisus excisus, juveniles (associated with Diadema) and adults; which occurs widely throughout the Indo­ diurnal benthic; solitary; carnivore: fishes and Pacific, and from Mexico to the Galapagos in crustaceans; pelagic eggs; 80 cm SL; TEP, 824 REVISTA DE BIOLOGIA TROPICAL from Baja California to Ecuador, also Occasional, a few collected at each rotenone Galapagos Islands; underwater photographs; station in 15-30m; juveniles and adults; resident; specimens at SIO, LACM, UCLA. cryptic; solitary; carnivore: fishes and Epinephelus sp. crustaceans; pelagic eggs; 7 cm SL; endemic to Epinephelus labriformis.- Snodgrass and Clipperton Island; 16: 13-61 mm SL, HelIer, 1905: 367 (Clipperton Island). including types, which are deposited at WAM Abundant on outer slope to 60 m depth; and USNM; underwater photographs; resident; juveniles and adults; diurnal benthic; solitary; specimens at USNM, LACM (as Rhegma carnivore: fishes, crustaceans; pelagic eggs; 45 thaumasium). cm SL; a Ciipperton endemic that we will Inc1uding P. axelrodi there are six known soon describe, mainly using specimens species of Pseudogramma (ABen and collected in the 1950s, this species was Robertson 1995). P. axelrodi belongs to a previously confused with E labriformis (see complex of four species (characterised by the Smíth 1971, ABen arld Robertson 1994, presence of a dermal flap on the upper t'.dge of Fischer et al 1995, Robertson and ABen 1996); the eye) that aIso ineludes the widespread 1EP 2: 50-54 mm SL; underwater photographs; species P. thaumasium, and P. bermudensis and resident; specímens at USNM, UCLA, LACM, P. gregoryifrom the tropical western Atlantic. SIO. It seems likely that P. axelrodi and P. Smith (1971) placed E. labriformis, wmch thaumasium are sister species, since they are occurs from the Gulf of California to Peru, and most alike in general appearanceand fin counts the Revillagigedo, Cocos and Galapagos (Allen and Robertson 1995). However, further lsIands, in a group of three species that analysis is needed to test that phylogenetic includes two Atlantic species, E. guaza, from hypothesis. Brazil and the northeast Atlantic, and E drummondhayi, from Florida 3J'1d in Rypticus bicolor Valenciennes, 1846 the northwest Atlantic. Based on close Rypticus bicolor Valenciennes, 1846: 307, similarities in their morphologíes E pl.1, fig.2 (Galapagos Islands). labriformis evidently is the sister species of Occasional, a few collected at each rotenone Epinephelus sp. station and sometimes sighted in crevíces between 15-30m; juveniles and adults; cryptic; Paranthiascolonus (Valenciennes, 1855) solitary; camivore: fishes and small Serranus colonus Valenciennes, 1855: 300, crustaceans; pelagic eggs; 10 cm SL; TEP, pI. 2, fig. 1 (Galapagos Islands). from Baja California to Peru, also Galapagos furcifer.- Smith 1971: 89 Islands; 15: 32-123 mm SL; underwater (Clipperton Island). photographs; resident; specimens at USNM, Abundant on outer slope to 40 m depth; LACM. mainly small to large adults, but juveniles also present; diurnal midwater; forms aggregations; Kuhliidae - Flagtails planktivore; zooplankton; pelagic eggs; 30 cm SL; TEP, from Gulf of California to Peru, also (Schneider, 1801) Galapagos Islands; 2: 75-76 mm SL; Sciaena mugil Schneider in Boch and underwater photographs; resident; specimens at Schneider 1801: 541 () SIO, LACM, UeLA. Not seen or collected by us, however numerous specimens collected on reef flat in Pseudogramma axelrodi ABen and 1950s; diurnal, schools inshore in surf zone; Robertson, 1995 carnivore: free swimming crustaceans and small Pseudogramma axelrodi ABen and fishes; pelagic eggs; Indo-Pacific to the Robertson, 1995a: 72 (Clipperton Island). Americas; possible resident on the basis of the Allen &: Robertson: Clipperton Atoll 825 numbers of fish coUected; specimens at sro (as solitary or in groups; camivore: fishes, , K. arge), LACM (as K. taeniura). lobsters; pelagic eggs; 65 cm SL; TEP, from Southern California to Pern, al50 Galapagos Apogonidae - Cardinalfishes Islands; observed only; probable vagrant

Apagan atrir:audus lordan and McGregor, lugubris Poey, 1860 1898 Caranx lugubris Poey, 186C: 222 (Cuba). Apogon atricaudus Jordan and McGregor, Common on outer slope in 15-60m; 1898: 277 (Revillagigedo Islands). inshore-offshore; adults only; roving predator; Cornmon on outer slope in 15-40 m; adults solitary ar in groups; carnivore: fishes, crabs, only; nocturnal benthic; forms aggregations; lobsters; spawns peJagic eggs; 70 cm SL; Indo­ carnivore: crabs, shrimps, worms, fishes; male Pacific to the Americas; undelwater broods eggs in mouth; 6.5 cm SL; TEP, at photographs; resident population; specimens at Clipperton Island, Reviliagigedo Islands and, as LACM. a vagrant, at Baja California; 13: 32-77 mm SL; underwater photographs; resident; Caranxmelampygus Cuvier, 1833 specimens at LACM, US]\'¡"'M. Caranx melampygus Cuvier, in Cuvier and Valenciennes, 1833: 116 (Waigeo, Euru, Echineidae - Remoras Vanico)o, Mauritius). Caranx melampygus.- ABen and Remora remora (Linnaeus, 1758) Robertson, 1994: 303 (Clipperton Island). Echeneis remora Linnaeus (1758): 260 Abundant on outer slope to a depth of 40 (ludian ). m; inshore-offshore; adults only; roving Not seen or collected by liS; several predator; solitary or in groups; camivore: specimens coliected in 1950s; oceanic, fishes, crabs, lobsters; pelagic eggs; 80 cm SL; associated with sharks; solitary; camivore: fish Indo-Pacific to the Americas; undelwater scraps, ectoparasites, various small prey; photographs; resident population; specimens at pelagic eggs; 62 cm rL; circumtropical; LACM. specimens at LACM. Caranx sexfasciatus Quoy and Gaimard, Carangidae- Jacks 1825 Caranx sexfasciatus Quoy and Gaimard, Carangoides orthogrammus (lordan and 1825: 358 (Waigeo, Indonesia). Gilbert, 1881) Caranx sexfasciatus.- ABen and Robertson, Caranx orthogrammus lordan and Gilbert, 1994: 303 (Clipperton Island)o 1881a: 226 (Clmion Island). Common on outer slope, especi.aJly on l\¡t Rare, a few aduits seen in midwater on side of island in 15-25 m; inshol'e-offshore; outer slope in 15-25m; roving predator; adults only; roving predator; solilary or in solitary or in groups; camivore: fishes, groups; carruvore: fisnes, crabs, lobsters; crustaceans; pelagic eggs; 70 cm FL; Indo­ pelagic eggs; 85 cm SL; Indo-Pacific to the Pacific to the Americas; uuderwater Americas; underwater photographs; resident photographs; probable vagrant. population; specimens at LACM (as C. marginatus). Caranx caballus Günther, 1869 Caranx caballus Günther, 1869: 431 Decapterus macarellus (Cuvier, in Cuvier (Panama). and Valenciennes 1833) Rare, several adults seen in midwater at Caranx macarellus Cuviel' and 20m on NE side of island; roving predator; Valenciennes 1833: 40 (Martinique) 826 REVISTA DE BIOLOGIA TROPICAL

Not seen or collected by us; specimens collected in 1950s; offshore pelagic; roving Trachínotus stilbe (Jordan and McGregor, predator; fonns aggregations; carnivore: fishes, 1898) crustaceans; pelagic eggs; 32 cm TL; Zalocys stilbe Jordan and McGregor,1898 : circumtropical; specimens at LACM. 277 (CIaríon Island). Trachynotus stilbe.- Fischer et al 1995: Elegatis bipinnulata (Quoy and Gaimard, 983 (Clipperton Island) 1825) Occasional,a few schools containing about Seriola bipinnulata Quoy and Gaimard, 10-50 adults seen on NE side of island to 15 m 1825: 363 (Keeling Island,New ). depth; adults only; specimens collected in the Commonly seen in midwater on outer 1950s; roving predator; fonns aggregations; slope in 10-60 m; adults only; inshore-offshore carnivore: fishes, large zooplankton; spawns pelagic roving predator; fonns aggregations; pelagic eggs; 35 cm SL; throughout TEP, carnivore: fishes,pelagic crustaceans; spawns although mainly around offshore islands; pelagic eggs; 115 cm SL; circumtropical; underwater photographs; resident; specimens at underwater photographs; resident population; LACM. specimens at LACM. Coryphaenidae - dolphinfishes Naucrates ductor (Linnaeus,1758) Gasterosteus ductor Linnaeus, 1758: 295 Coryphaena equiselis Linnaeus, 1758 (no locality given). Coryphaena equiselis Linnaeus 1758: 261 Rare, several subadults seen with (no locality) Carcharinus fa lciformis; offshore pelagic Not seen or collected by us, specime. species; rovíng predator,swims with sharks and col1ected in 1950s; offshore pelagic; rovinl other large predators; solitary or in groups; predator; solitary or in groups; carnivore: carnivore: fish scraps; pelagic eggs; 60 cm SL; fishes,squid, zooplankton; pelagic eggs; 75 cm circumtropical; observed only; specimens at FL; circumtropical; specimens at LACM. LACM. Lutjanidae - Snappers Setar crumenophthalmus (Bloch,1793) Scomber crumenophthalmus Bloch 1793: Lutjanus viridis (Valenciennes,1855) 77 (Acara,Guinea) Diacope viridis Valenciennes, 1855: 303, Not seen or collected by us; specimens pU,fig. 2 (Galapagos Islands). collected in the 1950s; coastal pelagic; roving Lutjanus viridis.- ABen and Robertson, predator; fonns aggregations; camivore: fishes, 1994: 303 (Clipperton Island). crustaceans; pelagic eggs; 60 cm SL; Common on ro;::,: slope in 15-60 m, circumtropical;specimens at LACM. usually seen in small groups, but a huge resting aggregation containing several hundred Seriola rivoliana Valenciennes,1833 individuals sighted on one occasion; adults Seriola rivoliana Valenciennes in Cuvier only; diurnaVnocturnal, benthic; fonns and Valenciennes 1833: 207 (Greek aggregations; carnivore: fishes, crustaceans; Archipelago) pelagic eggs; 25 cm SL; TEP, from Baja Not seen or coHected by us; specimens California to Ecuador,also Galapagos Islands; collected in fue 1950s; inshore-offshore underwater photographs; resident population; pelagic-demersal; roving predator; solitary or specimens at USNM. small groups; camivore: fishes, crustaceans; pelagic eggs; 103 cm FL; circumtropical; Haemulidae - Grunts specimens at LACM (as S. colburni). AlIen & Robertson: Clipperton Atoll 827

Orthopristis cantharinus (Jenyns,18 42) Common on outer slope in 8-30 m; Pristopoma cantharinus Jenyns, 1842: 49 medium to large adults only; diurnal midwater; (Galapagos) forms aggregations; herbivore: algae; pelagic Not seen or collected by us; one specimen eggs; 32 cm SL; TEP,from Gulf of California collected in the 1950s, which was apparently to Peru, also Galapagos Islands; underwater lost, as it is not longer in UCLA collection photographs; resident population; specimens - (D. Buth personal communication 1996); sandy LACM. bottoms; diurnal; forms aggregations; carnivore: crustaceans and molluscs; 38 cm SL; Sectator ocyurus (Jordan and Gilbert, 1881) widespread in TEP; presumed vagrant; Pimelepterus ocyurus Jordan and Gilbert, specimen formerly at UCLA. 1881b: 327 (Panama). Rare,about 10 sighted on NE side to depths Mullidae - Goatfishes of 40 m; adults only; diurnal midwater; solitary or in groups; omnivore: algaeand zooplankton; Mulloidichthys dentatus (Gill, 1862) pelagic eggs; 50 cm SL; widespread in TEP; Upeneus dentatus Gill, 1862c: 256 (Cape also occurs as a vagrant in the , SanLucas, Baja California). Hawaii (repeatedly) and as far as Japan (Randall Mulloidichthys dentatus.- Allen and 1961, 1985, J.E. Randall, personal Robertson, 1994: 303 (Clipperton Island). communication 1996); underwater Common on outer slope in 15-60 m, photographs; probable vagrant. usually seen in small groups of about one dozen individuals, but several resting schools Chaetodontidae - of about 50 individuals sighted; mainly medium to large adults, but a few juveniles Forcipiger flavissimus Jordan and 'seen; carnivore: sand-dwelling invertebrates; McGregor, 1898 spawns pelagic eggs; 26 cm SL; TEP, from Forcipiger flavissimus Jordan and Gulf of California to Peru, also Galapagos McGregor, in JOrdan and Evermann, 1898: Islands; underwater photographs; resident; 1671 (ClaríonIslan d). specimens at NMNH, LACM (as Upeneus Common on outer slope in 15-40 m; small grammorhynchus). adults only; diurnal benthic; solitary or in pairs; carnivore: crustaceans and other benthlc Kyphosidae - Sea Chubs invertebrates; spawns pelagic eggs; 17.5 cm SL; Indo-Pacific to the Americas; underwater . Kyphosus analogus (Gill, 1862) photographs; resident population; specimens at Pimelepterus analogus Gill, 1862b: 245 USNM,LACM. (Cape San Lucas,Baja California). Occasionally sighted on outer slope in 15- Johnrandallia nigrirostris (Gill, 1862) 30 m; medium to large adults only; diurnal Sarothrodus nigrirostris Gill, 1862b: 243 midwater; formsaggregations; herbivore: algae; (Cape San Lucas,Baja California). pelagic eggs; 32 cm SL; TEP, from Southern Common on outer slope in 10-40 m; small California to Peru, also Galapagos Islands; adults only; diurnal benthlc; solitary or in underwater photographs; residentpopulation . groups; omnivore: algae, molluscs, and crustaceans; pelagic eggs; 16 cm SL; TEP, Kyphosus elegans (Peters, 1869) from Baja California to Panama, also Pimelepterus elegans Peters, 1869: 707 Galapagos Islands; underwater photographs; (Mazatlan, Mexico). resident population; specimens al NMNH, Kyphosus elegans.- Allen and Robertson, LACM. 1994: 303 (Clipperton Island). 828 REVISTA DE BIOLOGIA TROPICAL

adults only observed; diurnal benthic; solitary Pomacanthidae - Angelfishes or in groups; planktivore; , etc.; parental care of demersal eggs; 10 cm SL; Holacanthus clarionensis Gilbert, 1890 TEP, mainly offshore íslands; photographed; Holacanthus clarionensis Gilbert, 1890: resident; 3: 54-81 mm SL; specimens at 72 (ReviBagigedo Islands). USNM. This specíes is variable in color Rare, several large adults of this according to geographic locality. Clipperton unmistakeable species sighted on NE side of adults are entirely dark brown in life in contrast island in 20-30 m; diurnalbenthic; solitary or to the light brown individuals from the in groups; omnivore: algae, sponges, tunicates; Galapagos iHustrated by AlIen and Robertson spawns pelagic eggs; 20 cm SL; TEP, (1994). Adults from CIaríon Island have a pale abundant endemic resídent of Revillagigedo stripe at the base of tbe , similar to Islands, with vagrants not uncommon at that of the juvenile from Galapagos illustrated southerntip of BajaCalífornia; vagrant. by ABen and Robertson (1994).

Holacanthus limbaughi Baldwin, 1963 Stegastes baldwini ABen and Woods, 1980 Holacanthus limbaughi Baldwin, 1963: 3 Stegastes baldwini ABen and Woods, (Clipperton Island). 1980: 183 (Clipperton Island). Abundant on outer slope in 10-60 m; Abundant on outer slope to depths of 60 m; juveniles and adults; diurnalbenthic; solitary or diurnal benthic; juveniles (including new in groups; omnivore: algae, sponges, tunicates; recruits) and adults; solitary or in groups; spawns pelagic eggs; 25 cm SL; endemic to herbivore: algae; parental care of demersal eggs Clipperton Island; underwater photographs; (presumed males, were often observed guarding resident popuJation; specimens at USNM, benthic eggs during our visit); 9 cm SL; UCLA, LACM, SIO. endemic to Clipperton Island; 3: 19-78; The genus Holacantltus is restricted to !he underwater photographs; resident; specimens at neotropics and west Mrica (R. Pyle, Bishop USNM, SIO, UCLA, LACM. Museum, personal communication). It includes There are síx TEP species in !his genus - S. three species from the TEP, H clarionensis acapulcoensis (Mexico lo Panama), S. (endernicto !he Revillagigedos), H. limbauglti, arcifrons (Galapagos and Malpelo), S. baldwini and H passer (widespread in the region), and (Clipperton), S. leucorus beebei (Galapagos), three from the tropical west Atlantic - H S. l. leucorus (the ReviUagigedos), S. · bermudensis, H. ciliaris and H tricolor. rectifraenum (Mexíco), and S. redemptus While fin-ray counts and the juvenile color (Revillagigedos ). patterns of the three TEP species are similar ABen and Wooos (1980) proposed that S. (Baldwin 1963), the adult color pattern of H baldwini is most closely related to S. leucorus. limbaughi most c10sely resembles tllat of H An analysis of similarities in the mitochondrial passer. Although a comprehensive analysis of DNA sequences of all TEP specíes (H.A. relationships ís lacking it appears that fue Lessios, D.R. Robertson and B. Kessing, sister species of H. limbaughi líkely ís H unpublished data) supports that hypothesis and passer. indicates !hat tbe sister species of S. baldwini 1S S. leucorus beebei. Pomacentridae - Damselfishes Cirrhitidae - Hawkfishes Chromis alta Greenfield and Woods, 1980 Cltromis alta Greenfield and Woods, 1980: Cirrhitichthys oxycepltalus (Bleeker, 1855) 630 (Baja California). Cirrhites oxycephalus Bleeker, 1855: 408 Common on NE side of island in 40-6Om; (Ambon, Indonesia). AlIen & Robel"tson: Clipperton Ato!! 829

Common on outer slope in 3-30 m, usually photographs; resident; specimens at USl"TM, associated with Pocillopora coral heads; SID. LACM. juveniles and adults; diurnal benthic; solitary; carnivore: benthic cmstaceans and fishes; Novaculichthys taeniourus (Lacepede, spawns pelagic eggs; 7 cm SL; Indo-Pacific to 180l) the Americas; 4: 27-60 mm SL; underwater Labrus taeniourus Lacepede, 1801: 488 and photographs; resident; specimens at NMNH, 518 (lndian Ocean). LACM (as C. corallicola), SIO. Occasional, sighted in rubble-filled depressions on outer slope at lO-20m depth; juveniles and adults; diurnai benthic; solitary or Cirrhitus rivulatus Valenciennes, 1855 in pairs; camivore: gastropods, bivalves, Cirrhitus rivulatus Valenciennes, 1855: echinoden:ns, wonns, etc.; pelagic eggs; 27 cm 309 (Galapagos Islands). SL; Indo-Pacific to the Americas; undenvater Common on outer slope in 10-25 m; small photographs; resident; specimens at USNM, to large adults only; diurnal benthic; solitary; LACM. carnivore: crabs; sea urchins, brittle stars, fishes; pelagic eggs; 45 cm SL, the largest Stethojulis bandanensis (Bleeker, 1851) member of the family; TEP, from Gulf of Julis bandanensis Bíeeker, 1851b: 254 California to Colombia, also Galapagos (Indonesia) . Islands; underwater photographs; resident; Common on outer slope to 20 m depth, specimens at SIO, LACM. juveniles common in rotenone station on shallow reef flat; juveniles and adults; diurnal Mugilidae - MuHets benthic; solitary or in groups; carnivore: small benthic invertebrates; pelagic eggs; 13 cm SL; Chaenomugil proboscidens (Gunther, Western and Central Pacific; underwater 1861) photographs; resident; specimens at USNM. Mugil proboscidens Gunther 1861: 459 () Thalassoma grammaticum Gilbcrt, 1890 Not seen or collected by us; one specimen Thalassoma grammaticum Gilbert, 1890: collected in the 1950s; schools in inshore 68 (Socorro and CIarionIslands). shallow habitats; herbivore: algae and detritus; Thalassoma grammatícum.- ABen and pelagic eggs; 22 cm SL; TEP, from Mexico to Robertson, 1994: 303 (Clipperton Island) Panama; apparent vagrant; specimen at UCLA. Common on outer slope to 30m depth, juveniles confined to shallow (0-7 m) water; Labridae- Wrasses juveniles (inc1uding new recmits) a\1d adults; diurnal benthic; solitary; carruvore: benthic Bodianus diplotaenia (Gill, 1862) cmstaceans; pelagic eggs, spawning observed Harpe diplotaenia Gill, 1862a: 140 (Cabo several times in mid afternoon; 20 cm SL; San Lucas, Baja California). TEP, from Gulf of California to Pan ama, also Bodianus diplotaenius.- Snodgrass and Galapagos Islands; underwater photographs; HeBer, 1905: 391 (Clipperton Istand). spawning observed by us at Clipperton Island; Common on outer slope in 10-60 m; resident; see Randall (1995) for c1arification of mainly medium to large adults, but a few the taxonomic status of T. grammaticum and juveniles seen; diurnal benthic; solitary; other TEP species of Thalassoma; specimens camivore: sea urchins, cmstaceans and other at USNM. small benthic invertebrates; pelagic eggs; 70 cm SL; TEP, from Baja California to northern Chile, also GaJapagos Islands; underwater Thalassomapurpureum (Forsskal, 1775) 830 REVISTA DE BIOLOGIA TROPICAL

Searus purpureum Forsskftl, 1775: (Red Lessios, D.R Robertson, B. Kessing, Sea). unpublished data) indicate that those three Thalassoma purpureum.- Randall, 1995: species are ¡ndeed very c10sely related and that 675 (Clipperton Island). the sister species of T. robertsoni is T. Occasionally but regularly sighted in areas lueasanum. exposed to strong surge on upper part of outer Thalassoma virens (Gilbert, 1890) slope and on shallow reef flat, depth range 2-20 Thalassoma virens Gilbert, 1890: 68 m; juveniles and adults; diurnal benthic; (). solitary or in groups; camivore: benthic Thalassoma virens.- ABen and Robertson, crustaceans; pelagic eggs; 37 cm SL; Indo­ 1994: 207(Clipperton Island). Pacífic to the Americas; underwater OccasionalIy sighted in areas exposed to photographs; resident; specimens at USNM .. strongsurge on upper part of outer slope in 2-7 Randall (1995) reported sighting this species at m; small to medium adults only; diurnal Darwin Island, in the Galapagos. One of us benthic; solitary; camivore: benthic (DRR) sighted it at Isabela Island in the crustaceans; pelagic eggs; 25 cm SL; TEP, Galapagos in 1990, and at , off the Revillagigedo Islands and Clipperton Island; coast of Costa Rica, in 1992. However, underwater photographs; resident. Clipperton is the only TEP locality at which this species has yet been recorded in any Xyriehtys wellingtoni Allen and Robertson, abundance. Fish were seen spawning on the 1995 middle of the upper reef slope at 10m depth on Xyriehtys wellingtoni ABen and Robertson, several occasions during our visit. 1995b: 80 (Clipperton Island). Rare, about 10-15 individuals sighted on Thalassoma robertsoni ABen, 1995 the only large sand patch on the upper reef Thalassoma robertsoni Allen, 1995a: 75 slope on the NW side of the island, at 15m; (Clipperton Island). juveniles and adults; diurnal benthic; solitary; Thalassoma sp.. - Allen and Robertson, camivore: small benthic and planktonic 1994: 207 (Clipperton Island). invertebrates; pelagic eggs; 6 cm SL; TEP, Abundant on outer slope to 50 m depth; endemic to Clipperton; 6: 19-65 mm SL, perhaps the most abundant fish at Clipperton; including type specimens at USNM and WAM; diurnal benthic; forros aggregations while underwater photographs; areas of deep sand on feeding in midwater; juveniles (inc1uding the "60 m terrace" presumably support a abundant new recruits) and adults; planktivore; resident population; specimens at USNM. mainly crustaceanzooplankton ; pelagic eggs; 9 Five species of this genus are presently cm SL; TEP, endemic to Clipperton; 29: 15- known from the TEP -x pavo (lndo-Pacific to 77 mm SL, inc1uding type specimens at the Americas), X perlas (known only from USNM and WAM; underwater photographs; Panama - see Wellington et al 1995), X. vietori resident; specimens at USNM. We frequently (Galapagos only - Wellington 1992), and X saw this species spawning at midafternoon in wellingtoni (Clipperton only). In general lO-20m depth on the reef slope. appearance and color pattern X wellingtoni Strong resemblances in the structure of most closely resembles X. perlas and they may their color patterns (see Allen 1995) indicate be sister specíes. However, the three TEP that T. robertsoni belongs to a group of three members of this genus were discovered only Pacific species of this genus T. recently, and the fish fauna of the amblyeephalum, which is widespread in the ReviUagigedos has not been thoroughly Indo-Pacific, and T. lueasanum, which occurs documented. throughout the TEP except at Clipperton. Similarities in their mitochondrial DNA (H.A. Scaridae - Parrotfishes Allen & Roberlson: Clipperton AtolJ 831

bentrnc eggs during our visit; specimens at Scarus rubroviolaeeus Bleeker, 1847 USNM, UCLA, LACM, SIO. Searus rubroviolaceus Bleeker, 1847: 162 Ophioblennius is restricted to the TEP, and (Java). the east and west Atlantic (Springer 1962). o. Occasionally sighted on outer slope in 10- steindaehneri, the only species in the TEP, 60 m; mainly adults, but a few large juveniles has two subspecies - o. s. clippertonensis, seen; diurnal benthic; solitary or in groups; which is endemic to Clipperton, and O. s. herbivore: algae; pelagic eggs; 48 cm SL; Indo­ steindachneri, which occurs in the remainder Pacific to the Amencas; underwater of the TEP, from the Gulf of California to photographs; resident; specimens at USNM. Peru

Blenniidae - Blennies Gobiidae - Gobies

Entomacrodus ehiostictus (lordan and Bathygobius arundelii (Garman, 1899) Gilbert, 1882) Gobius arundelii Garman, 1899: 63 Salarias ehiostietus Jordan and Gilbert, (Clipperton Island). 1882c: 363 (Mazatlan, Mexico). Mapo sopo rato r.- Snodgrass and Heller Entomacrodus chiostictus.- Springer 1967: 1905: 416 (Clipperton Island). 108 (Clipperton Island). Common on reef f1at in 0-1 m; juveniles A few adults collected with rotenone on reef and adults; diurnal benthic; solitary or in flat next to beach; intertidal and shallow reefs ; groups; omnivore: algae and benthic diurnal benthi c; solitary; herbivore: algae; invertebrates; parental care of dernersa! eggs; 9 parental care of demersal eggs; 5.5 cm SL; cm SL; TEP, endemic to Clipperton; 9: 2.8-6 TEP, from Gulf of California to Panama and cm SL; resident; specimens al USNM, 510, offshore islands; 3: 37-50 mm SL; probable LACM. resident as many specimens were coUected in We provisionally recognize the Clippertol1 the 1950s; specimens at USNM, LACM, SID. populations as a distinct species (see also Springer (1967) notes that, Fischer et al 1995). However, there is need for morphologicaHy the Clipperton forro is more revison of TEP Bathygobius, which include closely allied to the Revillagigedos form than five named species - B. andrei (Costa Rica to the mainland one, which ranges from Mexico Ecuador), B. arundelii (Clipperton), B. lineatus to Peru. (Galapagos and Peru) , B. longipinnis (Revillagigedos) and B. ramosus (Baja Ophioblennius steindachneri Californialo Peru). clippertonensis Sprínger, 1962 Ophioblennius steindachneri lordan and Zanclidae - Moonsh IdoIs Evermann, 1898: 2401 (Tres Marias Islands, Mexico). Zanclus comutus Linnaeus, i 758 Ophioblennius steíndachneri Zanclus cornutus Linnaeus, 1758: 273 clippe rtonensis Springer 1962: 426 (lndies). (Clipperton Island) Cornmon on outer slope in 1-60 rn, with Common on outer slope to 40 m depth, large adults concentrated between S-12m; also present on reef f1at; mainly small adults, rnainly smaU to medium-sized adults; diurna! but a few large adults and new recruits seen; benthic; solitary or in groups; omnivore: diurnalbenth ic; solitary; herbivore: algae; maJe sponges, algae, etc.; pelagic eggs; 14 cm SL; parental care of demersa! eggs; resident; Indo-Pacífic to the A.rnencas; underwater endemic to Clipperton: 20 cm SL; 4: 48-100 photographs; resident; specimens at USNM, mm SL; males often seen guarding clutches of LACM. 832 REVISTA DE BIOLOGIA TROPICAL

photographs; resident; specimens at USNM, Acanthuridae - Surgeonfishes LACM, S10.

Acanthurus achilles Shaw, 1803 Acanthurus xanthopterus Valenciennes, Acanthurus achilles Shaw, 1803: 383 (No 1835 locality) Acanthurus xanthopterus Valenciennes, in Not seen or collected by us; known from a Cuvier and Valenciennes 1835: 215 single pelagic juvenile (58mm SL) coIlected at (). a night light on a Tuna boat near Clipperton in Rare, a few adults sighted on outer slope at 1969. The late-stage pelagic juvenile of this depth of 20 m; diurnal benthic; solitary oc in species is characterisedby its large size and the groups; herbivore: algae, diatoms, detritus, presence of dark spots on the body (Randall hydroids, etc.; pelagic eggs; 42.5 cm SL, fue 1956). Those characteristics were used by R largest member in the genus; Indo-Pacific to Feeney of LACM (personal communication fue Americas; observed only; population status 1996) to confirm the identity of that specimen; uncertaín - occurs elsewhere to 90m (Randall surge zone; diurnal, benthic; solitary or in 1985a) and may have been more eommon on groups; herbivore: algae; pelagic eggs; 20 cm the "6Om terrace". SL; central Paeifie; one specimen at LACM (# 34354). Ctenochaetus marginatus (Valencíennes, 1835) Acanthurus nigricans (Linnaeus, 1758) Acanthurus marginatus Valencíennes, in nigricans Linnaeus, J 758: 274 Cuvier and Valencíennes, 1835: 221 (Caroline (Red Sea). Islands). Hepatus aliala.- Snodgrass and Heller, Ctenochaetus marginatus.- Allen and 1905: 403 (Clipperton Island). Robertson, 1994: 303 (Clipperton Island). Abundant on outer slope in 5-40 m; Abundant on outer slope to 40 m depth, but juveniles andadults; diurnal benthic; solitary or large adults concentrated in surge zone and in groups; herbivore: algae; pelagic ' eggs; 16 upper reef slope to 10m depth; juveniles cm SL; Indo-Paeifie to the Amerieas; (including new recruits) and adults; diurnal underwaterphotograph s; resident; specimens at benthic; solitary or in groups; herbivore: algae USNM. and detritus; pelagic eggs; 18 cm SL; scattered localities in the central Pacífic including fue Acanthurus triostegus triostegus Mortlock Group (), , (Linnaeus, 1758) the Phoerux Islands, and the Marquesas, and, in Chaetodon triostegus Linnaeus, 1758: 274 the TEP, from Cocos Island (where it is (Indies). common), Gulf of Chiriqui (Panama), and Acanthu rus triostegus.- Sehmidt and Clipperton Island; underwater photographs; Schultz 1940: 8 (Clipperton Island) resident; specimens at LACM. Abundant on upper edge of outer slope to 10 m depth, new recruits common on reef flat; Naso annulatus (Quoy and Gaírnard, 1825) juveniles and adults; diurnal benthic; forms Priodon annulatus Quoy and Gaimard, aggregations; herbivore: algae; pelagic eggs; 1825: 377 (). 20.5 cm SL; Indo-Pacífic to the Americas; Rare, two large adults sighted on NE side of represented in the TEP by the subspecies A istand in 15-30m; diurna! midwater; solitary or triostegus triostegus, which occurs widely in in groups; planktivore; zooplankton; pelagic the Indo-Pacific (except for the Marquesas, and eggs; 85 cm SL; Indo-west Pacific; observed the Hawaíian region, each of which has its own only; vagrant; new record for fue TEP. subspecies - Randan 1956); underwater Alien & Robertson: Clipperton Atoll 833

Naso hexacanthus (Bleeker, 1855) circumtropical; specimens nol retained. Priodon hexacanthus Bleeker, 1855: 393 (Ambon, Indonesia). Euthynnus lineatus Kishinouye, 1920 Rare, several large adults sighted on NE Euthynnus lineatus Kishinouye, 1920: 113 side of island in 20-25 m; diurnal midwater; (Mexico). forms aggregations; planktivore; zooplankton; Offshore pelagic species; occasionally pelagic eggs; 50 cm SL; Indo-west and central caught by crew members with trolling lures; Pacific; observed only; vagrant; new record for adults only; roving predator; forms the TEP. aggregations; camivore: fishes, crustaceans, One 46.5cm SL individual was speared and squids, tunicates; pelagic eggs; 65 cm FL; its sagittal otoliths extracted. E.B. Brothers TEP, from Southern California to n01thern (personal communication) exarmned those Peru and offshore islands; specimens not otoliths and estimated that it had a larval life of retained. 150 days, and that it was between 5-11y old. Thunnus albacares (Bonnaterre, 1788) Naso lituratus (Bloch and Schneider, 1801) Scomber albacares Bonnaterre, 1788: 140 Acanthurus lituratus Bloch and Schneider, (). 1801: 216 (no locality given). Offshore pelagic species; frequent1y captured Rare; based on repeated sightings and their by crew members with trolling lures; adults distribution around half the aton there may only; roving predator; solítary or in groups; have been several dozen large adults of this carruvore: fishes, crustaceans, squíds; spawns species present in 1994, in 15-25m on upper pelagic eggs; 210 cm FL; circumtropical; reef slope; adults only; diurnal benthic; solitary specimens not retained. or in groups; herbivore: leafy brown algae; pelagic eggs; 28 cm SL; Indo-west and central Bothidae - LefteyeFlounders Pacific; observed only; probable vagrant; a new record for the TEP. Bothus mancus (Broussonet, 1782) Pleuronectes mancus Broussonet, 1782: no Sphyraenidae - Barracudas pagination (Tahiti). Sphyraena ensis Jordan and Gilbert, 1882 Bothus mancus.- Fischer et al 1995: 935 Sphyraena ensis Jordan and Gilbert, 1882b: (Clipperton Island) 106 (Mazadan, Mexico). Rare; a few medium to large adults sighted Rare, about 20 adults sighted in a group in 15-20 m over rubble bottoms; specimens near surface on outer slope; roving predator; also collected in 1950s; diurnal benthic; solitary or in groups; carruvore: fishes; pelagic solitary; carruvore: fishes, benthic eggs; 60 cm SL; TEP, from Gulf of California invertebrates; spawns pelagic eggs; 38 cm SL; to Peru; observed only; possible vagrant. Indo-Pacific to the Americas; 1: 40 mm SL; underwater photographs; probable resident due Scombridae - Tunas and Mackerels to occurrence over a 40 year period; specimens at LACM, SIG. Acanthocybium solandri (Cuvier, 1831) Cybium solandri Cuvier, in Cuvier and Balistidae - Triggerfishes Valenciennes, 1831: 192 (no locality given). Offshore pelagic species; a few individuals Balistes polylepis Steindachner, 1876 captured by crew members with trolling Jures Balistes polylepis Steindachner, 1876: 21 adjacent to the island; adults only; roving (Magdalena Bay, Mazadan, and , predator; solitary or in groups; camivore: Mexico). fishes, squids; pelagic eggs; 210 cm FL; Rare, less than 10 individuals sighted, 834 REVISTA DE BIOLOGIA TROPICAL mainly on NE side in 15-20 m; adults only; 1882) diurnal benthic; solitary; omnivore: algae, Balistes mento Jordan and Gilbert, 1882a: coral, ecmnoderms, fishes; parental care of 228 (CIarionIsland) . demersal eggs; 70 cm SL; TEP, from northern .- Berry and Baldwín, California to Chile; also recorded by Randall 1966: 457 (Clipperton Island). (1985b), as a vagrant in Hawaii; underwater Rare, several observed in 20-50 m; collected photographs; probable vagrant; specimens at in the 1950s; adults only; diurnal benthic; SIO. solitary or in groups; planktivore; zooplankton; parental careo f demersal eggs; 25 Canthidermis maculatus (Bloch, 1786) cm SL; antitropical distribution (Randall et al, Balistes maculatus Bloch, 1786: 25 1978), at scattered localities in the westem and (American seas) central Pacific including Ryukyu Islands, Not seen or collected by us; a few Marcus Island (Minami Tori Shima), Pitcairn specimens collected in the 1950s; demersal, Island, , and the Hawaiian Islands benthic on reefs or associated with floating (where it is abundant in the northwest islands, logs well offshore; forms aggregations; but rare and in deep water in the southern planktivore: zooplankton; parental care of islands; Randall 1985a, pers. comm., 1996), in demersal eggs; 50 cm TL; circumtropical; the TEP from southern California to the specimens at SIO, LACM. Galapagos, including the Revillagigedo Islands (where DRR found it in abundance in shallow Melichthys niger (Bloch, 1786) water on the south side of Socorro Island in Balistes niger Bloch, 1786: 27 (China 1991) and Clípperton Island; underwater Seas). photographs; possible vagrant; specimens at Melichthys niger.- ABen and Robertson, LACM. 1994: 303 (Clipperton Island). Abundant, aggregations containing many Monacanthidae Triggerfishes and hundreds of fish usuaUy seen at each anchorage, Leatherjackets surface to 50m; juveniles and adults; diurnal benthic; planktivore, zooplankton, occasionally Aluterus scriptus (Osbeck, 1765) live coral (see Glynn et al, 1996); parental caro Balistes scriptus Osbeck, 1765: 145 of demersal eggs; 25 cm SL; circumtropical; (China). underwater photographs; resident; specimens at Rare, a few adults seen on outer sIope in USNM, SIO, LACM. 10-20 m; diurnal benthic; solitary; omnivore: algae, seagrasses, cnidarians, tunicates; parental Sufflamen verres (Gilbert and Starks, 1904) care of demersal eggs; 70 cm SL; Balistes verres Gilbert and Starks, 1904: circumtropical; underwater photographs; 153 (Mazatlan,Mexico and Panama). probable vagrant; specimens at USNM. Balistes verres.- Snodgrass and Heller, 1905: 406 (Clipperton Istand). Cantherines dumerilii (Hollard, 1854) Common on outer slope to 30m depth; Monacanthus dumeriUi Hollard, 1854: 361 juveniles and adults; diurnal benthic; solitary; (Mauritius). omnivore: algae, crabs, molluscs, ecrunoids, Common on outer slope in 10-50 m; worms, etc.; parentalcare of demersal eggs; 32 medium to large adults; diurnal benthic; cm SL; TEP, from Gulf of California to solitary or in pairs; omnivore: , urchins, Ecuador; underwater photographs; resident molluscs, bryozoans, sponges; parental care of population; specimens at USNM, SIO, demersal eggs; 31.5 cm SL; Indo-Pacific to the LACM. Americas; underwater photographs; resident; Xanthichthys mento (Jordan and Gilbert, specimens at USNM. Allen & Robertson: Clipperton Atoll 835

Diodon holocanthus Linnaeus, 1758 Ostracüdae- Boxfishes Diodon holocanthus Linnaeus, 1758: 335 (India). Ostracion meleagris Shaw, 1796 Rare, a few adults seen on NE side of island Ostracion meleagris Shaw, in Shaw and in 15 m depth; nocturnal benthic; solitary; Nodder, 1796: PI. 253 (South Pacific). carnivore: sea urchins, gastropods, crabs; Ostracion clippertonense Snodgrass and pelagic eggs; 30 cm SL; circumtropical, Heller, 1905: 410 (Clipperton Island). widespread in the TEP, common in the Ostracion lentiginosum.- Snodgrass and Revillagigedo Islands in 1994; observed only; Heller, 1905: 410 (Clipperton Island). population status uncertain - because it is Occasionally sighted on outer slope in 10- cornmonon unconsolidated bottoms in Panama 25 m, mainly on NE side of island; juveniles (DRR personal observations) it may occur on and adults; diurnal benthic; solitary; omnivore: the "6Om terrace"; specimens USNM. algae, polychaetes, sponges, tunicates, molluscs; spawns pelagic eggs; 15 cm SL; Diodon hystrix Linnaeus, 1758 Indo-Pacific to the Americas; resident; Diodon hystrixLinnaeus, 1758: 335 (India). specimens at USNM, LACM. Rare, a few adults seen on outer slope in 15-20 m; nocturnalbenthic; solitary; carnivore: Tetraodontidae - Pufferfishes sea urchins, gastropods, crabs; pelagic eggs; 80 cm SL; circumtropical, widespread in TEP, (Bleeker, 1853) occasional in the Revillagigedo Islands in Tetrodon meleagris Bleeker, 1853: 507 1994; observed only; probable vagrant; (Asia). specimen at LACM. Tetraodon setosus.- Snodgrass and Heller, 1905: 413 (Clipperton Island). PROBABLE ERRONEOUS RECORDS Common on outer slope to 50 m depth; mainly medium to large adults, but a few large Serranidae juveniles seen; diurnal benthic; solitary; Epinephelus analogus Gill, 1864 omnivore: live corals (see Glynn et al, 1996), Epinephelus analogus Gill 1864: 163 algae, sponges, worms, echinoderms, etc.; (west coast of America) pelagic eggs; 28 cm SL; Indo-Pacific to the Epinephelus analogus.- Smith 1971: 149 Americas; resident; specimens at USNM, SIO, (Clipperton Island) LACM (as A. setosum). This is a distinctive, shallow-living TEP species (Allen and Robertson, 1994, Fischer et Canthigaster punctatissima (Günther, al 1995). We did not see or collect it at 1870) Clipperton. The distribution map for E Tetrodon punctatissimus Günther, 1870: analogus in Smith (1971) includes Clipperton, 302 (Panama). based on a "literaturereco rd". However, thetext Common on outer slope in 10-35 m; contains citations for all records shown on the juveniles (including new recruits) and adults; map, except for the one for Clipperton. The diurnal benthic; solitary or in pairs; omnivore: Clipperton record shown on the distribution algae and benthic invertebrates; spawns pelagic map for this species in Fischer et al (1995) is eggs; 6 cm SL; TEP, from Gulf of California probably based on the Smith (1971) reference to Panama, also Galapagos Islands; underwater (P. Heemstra, personal cornmunication 1996). photographs; 17 mm SL; resident; specimens There are no Clipperton specimens in at USNM, SIO, LACM. collections at the USNM, UCLA, LACM or SIO. Hence we regard Smith's (1971) Diodontidae - Porcupinefishes Clipperton record as an error. 836 REVISTA DE BIOLOGIA TROPICAL

(Labridae); Acanthurus nigricans and Aulorhynehidae Ctenochaetus marginatus (Aeanthuridae); and Melichthys niger (Balistidae). The richest area Aulorhynchus flavidus GiIl, 1862 in terms ofbot h coral growth (see Glynn et al Aulorhynchus flavidus Gill, 1862: 169 1996) and fish speeies diversity was off the (Washington) northeastern side of the island. This is a temperate speeies that oeeurs The total includes 14 oceanie forms (one between and northem Baja California, shark, three flying-fishes, two half-beaks, three (Eschmeyer et al 1983). There is a speeimen jaeks, a remora, a dolphin-fish, and three supposedly from Clipperton al LACM (W58- tunas). Three others (a needle-fish, a jaek and a 297, C68). R. Feeney of the LACM eonfmned )range between offshore and inshore the identity of that specimen (personal areas. The 98 inshore species inelude 19 eommunieation 1996). However, that record mídwater and surfaee forms (four sharks, seven appears to be an error. On the field sheet that jaeks, and single representatives of eight other included that entry, Aulostomus chinensis families), 70 demersal fishes restricted to hard­ (whieh does oecur at Clipperton) is erossed out reef habitats, and nine demersal species that and replaced with Aulorhynchus flavidus. live in or feed on uneonsolidated bottoms sueh Moreover, the length given in that entIy as rubble and sand (Apterichthys equatorialis, (156mm) differs substantially from that of the Myrichthys pantostigrnius, Phaenomonas A flavidus speeimen (125mm - measured by pinnata, probably Pontinus vaughni, R. Feeney). Orthopristis cantharinus, Mulloidichthys dentatus, Novaculichthys taeniourus, Xyrichthys wellingtoni, and Bothus mancus. DISCUSSION Species within this last group constitute about 4(': the TEP shorefish fauna but <10% of The known Clipperton fish fauna eonsists Clípperton's shorefish fauna. Because there is of 115 speeies belonging to 89 genera and 43 very little unconsolidated habitat in <60 m of families. The most speciose families, whieh water, and extensive areas of sueh between 60- eonstitute 49 pereent of the total fauna, are 100m, it is quite possible that further samplíng moray eels (Muraenidae, 14 speeies), jaeks will reveal additional species of that type at (Carangidae, 11 speeies), wrasses (Labridae, Clipperton. The non-oeeanic species consist eight speeies), surgeonfishes (Aeanthuridae, primaríly (68%) of camivores that eat various eight species), squirrelfishes (Holocentridae, rnobile . Another 9.1 % feed on benthic, five speeies), groupers (Serranídae, five sessile invertebrates while 12.9% are speeíes), and triggerfishes (Balistidae, five exclusively or largely planktivorous, and species). The most abundant reef fishes at the 17.8% are benthic feeding herbivores. ísland in 1994 were as follows (in phylogenetic Reef sharks often are abundant at offshore order): Gymnothorax dovii (Muraenidae); reefs and islands. They were surprisingly Myriprisüs berndti (and M. gildi?) uncommon at Clipperton in 1994 - we did not (Holocentridae); , see any during the first two days of diving. Dermatolepis dermatolepis and Epinephelus Their scareity appears to be a recent sp. (Serranidae); Caranx melampygus and C. development. Early reports refer to the sexfasciatus (Carangidae); Lutjanus viridis abundance of sharks (Skaggs 1989), and the (Lutjanidae); Mulloidichthys dentatus Scripps expeditíons of 1956 and 1958 found (Mullidae); Kyphosus elegans (Kyphosidae); Carcharhinus galapagensis to be common to Holocanthus limbaughi (Pomaeanthidae); the point of being a eontinuous nuisance Stegastes baldwini (Pomaeentridae); (Limbaugh 1963, W. Baldwin, personal Th alassoma grammaticum and T. robertsoni communication to G.R. ABen, 1994). The lack A1len & Robertson: Oipperton Atoll 837

of sharks in 1994 may have been due to recent thus likely are derived from species that fishing - unconfmned reports indicate that six recently invaded the 1EP from the west Pacifico Mexican boats intensively fished this S. baldwini is elosely related to the 1EP population in 1993, catching as many as species S. leucorus, and its sister species is the -2000 individuals (John Jackson, personal subspecies of S. leucorus endemic to the communication 1994). Clipperton is the only Galapagos Islands (S. leucorus beebei) rather coral reef within the range of the reef shark than the one endemic to the much nearer Triaenodon obesus where we have not found (950km vs 2,I00km) Revillagigedos (S. it. However, as it is unelear as to what extent leucorus leucorus).The probable sister species this shark is targetted by fishermen it is of Epinephelus sp. P. axelrodi, H. limbaughi uncertain whether its absence at Clipperton was and O. S. clippertonensis are widespread 1EP due to overfishing or natural causes, such as a species that belong to genera or species groups failure to recruit there. restricted to the and Atlantic. The There are approximately 825 species of phylogenetic affinities of the two other shore and near-shore marine fishes from -105 Clipperton endemics are less clear. Both families in the 1EP (Allen and Robertson Xyrichtys and Bathygobius occur in the west 1994, Fischer et aI 1995). Thus only -14% of Pacific as well as the 1EP, and the lantic, At the region's species and -40% of its families and phylogenetic relationships within both occur at Clipperton. In addition to the island's these genera remain to be elarified. extreme isolation, other faetors that probably Despite the faet that they have evolved in contribute to the small faunal size inelude its an environment dominated by live corals, none small size, reduced habitat diversity, and of Clipperton's endemic fishes can be oceanic environment (see Robertson and Allen considered to be "live-coraldepend ent", i.e. live 1996). However, sampling effort probably also only in andlorfeed exelusively on live coral at contributes, as we were not able to sample some stage of their life cycles. Ecologically the extensive areas of sand and rubble bottom on endemics are typical members of their genera, the "6Om terraee", which lie below the limits and species of all but one of those genera are of conventional SCUBA diving. common on rocky-reefs as well as coralreefs in The nine Clipperton endemics represent the 1EP: Myripristis, Epinephelus and 9.1 % of the shorefish fauna and 11.3% of the Pseudogramma species are carnivores; strictly demersal component of that fauna. Five Holacanthus species are benthic feeding of those endemics (Epinephelus sp. omnivores, and we saw no coral feeding by H. Holacanthus limbaughi, Stegastes baldwini, limbaughi at Clipperton; Stegastes species Thalassoma robertsoni, and Ophioblennius maintain territories containing algal mats; T. steindachneri clippertonensis) were among robertsoni and allied species are planktivores the most abundant fishes at the island in 1994. that also feed opportunistically on benthic The abundance of two others (Myripristis gildi animals; Ophioblennius species are territorial, and X wellingtoni) is unelear (see species benthic-feeding herbivores; while Bathygobius accounts). Bathygobius arundelii was common species are benthic feeding carJ.)ivores in our reef flat collections and similar characteristically found in rocky intertidal collections in that habitat in the 1950s. The habitats. Xyrichtys is the only genus of soft­ abundance of Pseudogramma axelrodi was bottom fishes with an endemic at Clipperton, similarto that of the related P. thaumasium in and X wellingtoni and its congeners are sand­ our collections elsewhere in the 1EP. living planktivores. Robertson and Allen Clipperton endemics appear to have a (1996) suggested that only two of Clipperton's mixture of eastem and westem Pacific origins. fishes (Cirrhitichthys oxycephalus and M.gildi and T. robertsoni both have very elose Arothron meleagris) could be regarded as being relatiyes on the westem side of the EPB, and live-coral dependent. A.meleagris that occur on 838 REVISTA DE BIOLOGIA TROPICAL

coral reefs in the TEP are almost exclusively species, Myripristis clarionensis, Apogon corallivorous, and that species reaches its atricaudus and Thalassoma virens, appear to highest densities in the TEP on coral reefs. be restricted to Clipperton and the However, that species can be quite common on Revillagigedos. In addition, populations of rocky shores, and the diet of individuals that Entomacrodus chiostictus at the live on such shores is dominated by non-coral Revillagigedos and Clipperton share ¡tems. Further, A meleagris is capable of morphologícal characterisitics that dífferentiate growing (albeit slowly) on a diet composed them from conspecific populations on fue exclusively of crustose coralline algae (Guzman mainland. Moreover, the Revillagigedos and Robertson 1989). Thus whether that endemic Holacanthus clarionensis occurs at species is truly coral-dependent probably Clipperton (and Baja California) as a vagrant. depends on whether adults can on a The Clipperton fish fauna also has affinities, coral-free diet. e oxycephalus, which is a although more limited, to the fish fauna of the micro-carnivore, is ofien found in abundance in southernTEP : Gymnothorax dovii is restricted living colonies of Pocillopora corals on reefs to the southernhalf of the TEP and Clipperton in the TEP. However, one of us (DRR) found represenls its northern-most resident this species in abundance on both coral reefs population. The sister species of one and coral-free rocky sl10res al Gorgona Island Clipperton endemic (S. baldwini) is a (Colombia), and in abundance in both Uve Galapagos endemic (S. leucorus beebei). The coral and in large areas of deadcoral rubble at remaining 36 TEP fishes found at Clipperton Christmas Island, in the central Pacific. Thus are aH widely distributed throughout the region, at most one of Clípperton's físhes might be at offshore istand andlor continental , sites. obligatoríly live-coral dependent. There is insufficient documentation of fue Clipperton's fish fauna is composed of shorefish faunas of the other offshore islands roughly equal numbers of TEP and transpacific (the Revillagigedos, Cocos, Malpelo and the species: 63 and 52, respectívely of the total, Galapagos) to allow more detailed comparison and 53 and 48, respectively of the non-oceanic of regional affínities within the TEP. component. Thís pattern reflects the island's We estimate that at least 70 of the 98 non­ great isolation and fue pattern of warm surface oceanic fishes known from Clipperton have currents that could bring fishrecruits t.o it from resident populations there, and that only 17 distant sources: Although Clipperton is such species likely are vagrants - i.e. are physically much nearer to eastern Pacific reefs represented by individuals that recmited from than to reefs on the western side of the EPB other areas, and lack self-sustaining (-1 ,000 km vs 4,000 km), it líes at the populations at Clipperton. Similar percentages northern edge of the main ea'it-bound surface 75% and 78%, respectively, of species for current from Oceanía, a curren! that flows which sufficient information seems available, eastwardsfor two thirds of the year and that is of transpacific and TEP species (other than much stronger than westward flowthat occurs Clipperton endemics) appear to be resident at from the remainder of the TEP during the the island. remainder of the year. In addition, during Unfortunately, descriptions of shorefish periodic El Nino events eastward flow that faunas often consist símply of a species list, could CaIT'j recmits from Oceanía is much and lack information on their abundance, or strongerthan normal (see Robertson and ABen habitat usage. Such information is vital for 1996, for furtherdiscussio n). zoogeographic analyses: (i) It allows The non-endemic TEP component of the estimation of the occurrence of ongoing Clipperton fauna has limited affinities with recruitment fromexternal sources, and analyses thatof the Revillagigedos (see also G1ynn et al of patterns of variation in such recmitment. (ii) 1996, for corals). Resident populations of three It enables future workers to assess change in AlIen & Robertson: Clipperton Atoll 839 populatíon status over time, why previous provided excellent logístic assistanc

AlIen, a.R. 1995a. Thalassoma robertsoni, A new species Bleeker, P. 1866. Sur les especes d'Exocet de I'Inde of wrasse (Labridae) from Clipperton Island in the archipelagique. Ned. Tijdschr. Dierk. 3: 105-129. tropical eastem Pacifico Rev. fr. Aquariol. 22: 75-79. Bloch, M.E. 1786. Naturgeschichte der Auslandíschen Allen, a.R. 1995b. Clipperton: the forgotten ¡sland. Trop. Físche. Vol. 2. Auf Kosten des Verfassers und in Fish Hobb. 44:46-7 l. Cornmission in der Buchhandlung der Realschule. Allen, a.R. & D.R. Robertson. 1992a. Serranas Berlín. xii + 260 p. socorroensis, a new species of serranid fish from the tropical eastem Pacifico Rev. fr. Aquariol. 19: 37-40. Bloch, M.E. 1793. Naturgeschíchte der Auslandíschen AlIen, a.R. & D.R. Robertson. 1992b. Two new species Fische. Vol. 7. Auf Kosten des Verfassers und in of wrasse (Labridae: Halichoeres) from the tropical Cornmission in der Buchhandlung der Realschule. eastem Pacifico Rev. fr. AquarioL 19: 47-52. Berlin. xii + 1 44 p.

Allen, a.R. & D.R. Robertson. 1994. Fishes of the tropical Bloch. M.E. & J.G. Schneider. 1801. In: M.E. BIoch. eastem Pacifico University of Hawaii Press. HonoluIu. Systema lchthyologiae. Berolini: Sumtibus Auctoris 332p. Impressumet Bibliopolio Sanderiano Commissum, Ix + 584 p. ABen, a.R. & D.R. Robertson. 1995a. Pseadogramma axelrodi, a new species of serranid fish from the Bonnaterre, 1.P. 1788. lehthyologie. TabIeau tropical eastern . Trop. Fish Hob. 44: Encyclopédique el Méthodique des Trois Régnes de la 72-75. Nature. .215 p.

AlIen, G.R. & D.R. Robertson. 1995b. Xyrichtys Boulenger, G.A. 1895. Catalogue of the perciforrn fishes weUingtoni, A new species of wrasse (Labridae) from in the BritishMuseum, second edition, Vol. 1. By order Clipperton IsIand, tropical eastcrn PacificOcean. Rev. of the Trustees. London. i-xix + 391 p. fr. Aquariol. 22: 80-82. Briggs, J.C. 1961. The Eastern Pacific Barrier and Ihe AlIen, G.R. & L.P. Woods. 1980. A review of the distribution of marine shore fishes. Evolution 15: 545- damselfish genus Slegasles from the eastern Pacific 554. with description of a new species. Rec. West. Aust. Mus. 8: 171-198. Brothers, E.B. & R. E. Thresher. 1985. Pelagic duration, dispersal, and the distributioll of lndo-Pacific coral­ Baldwin, W.J. 1963. A new chaetodontid fish, reef fishes. NOAA Symp. Ser. Undersea Res. 3: 53- Holocanthus limbaaghi, from the eastern Pacifico 68. Contrib. Scí. LA County Mus. 74: 1-8. Broussonet, P.M.A. 1782. lcthyologia, Sislens Piscium Barnhart, P.S. & c.L. Hubbs. 1946. Pontinus vaughni, a Descriptiones et leones. London. 41 p. (unnumbered). new scorpaenid fish from Baja California. Bull. Scripps Inst. Oceanogr. U. Calif. 5: 371-389. Bussing, W.A. 1991. A new species of eastern Pacific moray (Pisces: Muraenidae). Rev. Bíol. Trop. 39: Berry, F.H. & W.J. Baldwin. 1966. Triggerfishes 97-102. (Balistidae) of the eastern Pacifico Proc. Cal. Acad. Sci. 34: 429-474. Clark, T.A. 1995. Larvae of nearshore fishes in oceanic waters of Ihe central equatorial Pacifico Pacif. Sci. 49: Bleeker, P. 1849. Overzicht der te Batavia Voorkomende 134- 142. Gladschubbige Labroiden, met Beschrijving van 11 Nieuwe Soorten. Verh. Gen. 22: 1-64. Cope, E.D. 1872. Report on the recent reptiles and fishes of the Survey, collected by Campbell Carrington and Bleeker, P. 1851a. Over eenige nieuwe soorten van C.M. Dawes. p. 467-476. Jn F.V. Hayden. Preliminary Belone en Hemiramphus vanJava. Nat. Tijschr. Ned.­ report of the US Geological Survey of Montana and Indie. 1: 93-95. portions of adjacent territories; being a fifth annual report of progress, pt.4. Bleeker, P. 185lb. Bijrage tOI de Kennis del" Ichthyologische fauna van de Banda-eilanden. Na!. Cuvier, G. 1816. Le Regne Animal Distribué d'apres son Tijdchr. Ned.-Indie. 2: 225-261. Organisation pour Servir de Base a I'histoire Naturelle des Animaux .... VoLll, Deterville. Paris. xx + xviii + Bleeker, P. 1853. Bijrage tot de Kennis der 532 p. Ichthyologische fauna van Solor. Nat. Tijschr. Ned.­ Indie. 5: 67-96. Cuvier, G. 1833. In: Cuvier G. and Valencienlles 1833. Histoire Naturelle des Poissons. Vol. 9. F.G. Lean!t. Bleeker, P. 1855. Zesde Bijdrage tot de Kennis der Paris. xxix + 512 p. Icthyologische fauna van Amboina. Natuurk. Tijdschr. Ned.-Indie 8: 391-434. Cuvier, G. & A. Valenciennes. 1831. Histoire Naturelle des Poissons. Vol. 7. F.G. Levrault. Paris. xxix + 531 p. Bleeker, P. 1857. Tweede bijrage tot de Kennis der Ichthyologische fauna van Boeroe. Nat.Tidjschr. Cuvíer, G. & A. Valenciennes. 1833. Histoire Naturelle Ned.-Indie. 13: 55-82. des Poissons. VoL 9. F.G. Levrault. Paris. xxix + 512 p.

Bleeker, P. 1865. Poissons Inédits Indo-archipélagiques Cuvier. G. & A. Valenciennes. 1835. Histoire Naturelle de I'ordre des Murenes. Ned. Tijdschr. Dierk. 2: 38- des Poissons. Vol. 10. F.G. LevrauIt. Paris. xxiv + 482 54. p. A1len & RobertsoQ: Clipperton Atoll 841

Cuvier, G. & A. Valenciennes. 1847. Histoire Naturelle GiII, T.N. 1863b. Descriptions of some new species of des Poissons. Vol. 19. Paris: F.G. Levrault. xxiv + 544 Pediculati, and on the classification of the group. Proc. p. Acad. Nat. Sci. Philad. 15: 88-92.

Dawson, C.E. 1985. Indo-Pacific pipefishes (Red Sea lO Glynn, P.W.G. 1.E.N. Veron & G.M. Wellington. 1996. the Americas). Ocean Springs Mississippi. Gulf Coast Oippetton Atoll (Eastern Pacific): oceanographic Research Laboratory. 230 p. setting, geomorphology, reef-building coral cornmunities and zoogeographic Relationships. Coral Ekman, S. 1953. Zoogeography of the Sea. Sidgwick and Reefs 15: 71-99. lackson. London. 474p. Greenfield, D.W. 1965. Systematics and zoogeography of Eschmeyer, W.N. 1990. Catalog of the Genera of Recent Myripristis in the eastern tropical Pacifico Cal. FJSh Fishes. California Academy of Sciences. San Game 51: 229-247. Francisco. 697p. Greenfield, D.W. & L.P. Woods. 1980. Review of the Eschmmeyer, W.N. E.S. Herald & H. Harnman. 1983. A deep-bodied species of Chromis (Pisces: field guide to Pacific coast fishes of . Pomacentrldae) from the eastern Pacific, with Houghton Mifflin.Boston. 336 p. descriptions oC three new species. Copeja 1980: 626- 641. Fischer, W. F. Krupp, W, Schneider, C. Sommer, K.E. Carpenter & V.H. Niem. 1995. Guia FAO para la Griffith,E. 1834. The Class Pisces, arranged by the Baron identificacion de especias para los fines de la pesca; Cuvier, with supplementary additions by Edward PacificoCentro-Orien tal, Vols 1-111. FAO. Rome. Griffith, F.R.S. etc, and Lieut. Col. Charles Hamilton Smith, F.R.L.S.S. etc. etc .. London. 680 p. 62 + 3 pI. ForsskAl, P. 1775. Descriptiones Animalium Avium, Amphibiourm, Piscium, Insectorum, Vermium; quae in Günther, A. 1869. An account oC the fishes oC the states Itinere Orientali Observavit. Post mortem auctoris oC , based on collections made by edidit Carsten Niebuhr. Hauniae. 20 + xxxiv + 164 p. Capt. 1.M. Dow, F. Godman, Esq. and O. Salvin, Esq. 43pl. Trans. Zool. Soco Lond. 6: 377-494.

Fritzsche, R.A. 1980. Revision of the eastern Pacific Gunther, A. 1861. Catalogue oC the acanthopterygian Syngnathidae (Pisces: ), including fishes in the collection oC the British Museum. Vol. 3. both recent and fossil fonns. Proc. Cal. Acad. Sci. 42: British Museum (Nat.Hist .). London. 586p. 181-227. Gunther, A. 1866. Catalogue oC the acanthopterygian Garman, S. 1899. A species oC goby from the shores of fishes in the collection oC the British Museum. Vol. 6. Clipperton Island. New Engl. Zool.Club 1: 63-64. British Museum (Nat. Hist.). London. 367p.

Gilbert, C.H. 1890. XII - A Preliminary report on the Günther, A. 1870. Catalogue of the Fishes in the British fishes collected by the Steamer "Albatross· on the Museum. Vol. 8. By order oC the Trnstees. London. Pacific Coast of North America during the year 1889, xxv + 549 p. with descriptions of twelve new genera and ninety­ two new species. Proc.U.S. Nat. Mus. 13: 49-126. Guzman H. M. & D.R. Robertson. 1989. Population and Ceeding responses oC the coraIlivorous puCCerfish Gilbert, C.H. 1897. Descriptions of twenty-two new Arothron meleagris to coral morta1ities in the eastern species of fishes collected by the Steamer Pacifico Mar. Ecol. Prog. Ser. 55: 121-131. "Albatross," oC the United States Fl Sh Cornmission. Proc.U.S. Nat. Mus. 19: 437-457. Heemstra, P.C. & 1.E. Randall. 1993. An annotated and ilIustrated catalogue . oC the , rockcod, hind, Gilbert, C.H. & E.C. Starlcs. 1904. The fishes of Panama coral grouperand Iyretail species known to date. FAO Bay. Mem. Cal. Acad. ScL 4: 1-304. Fish. Synop. 125, vol 16. 382p.

GiII, T.N. 1 862a. Catalogue of the fishes of lower Hollard, H.L.G.M. 1854. Monographie de la Famille des California in the Smithsonian Institution, collected by Balistides. Ann. Sci. Nat. (Zool.) 4:39-72. Mr. l. Xantus. Proc. Acad; Nat. Sci. Philad. 14: 140- 151. lenyns, L. 1842. In: C. Darwin. The of the Voyage of H.M.S. "Beagle" under the Cornmand of GiII, T.N. 1862b. Catalogue of the fishes of lower Captain Fitzroy, R.N. during the Years 1832 lO 1836. California in the Smithsonian Institution, collected by Part4. Smith, E1der andCO. London. 172 p. Mr. l. Xantus. Pt.I1. Proc. Acad. Nat. Sci. Philad. 14: 242-246. lordan, D.S. & B.W. Evermann. 1898a. The fishes of northand middle America: a descriptive catalogue oC GiII, T.N. 1862c. Catalogue of the fishes of lower the species oC fish-lite Vertebrates Cound in the California in the Smithsonian Institution, collected by waters oC north America, north of the Isthmus of Mr. l. Xantus. Pt. 111. Proc. Acad. Nat. ScL Philad. 14: Panama. Part 11. Bull. U.S. Nat. Mus. 47 : 1241-2183. 249-262. lordan, D.S. & B.W. Evermann. 1 898b. The fishes of Gill, T.N. 1863a. Catalogue of the fishes of lower north and middle America: a descriptive catalogue of California in the Smithsonian Institution, collected by the species oC fish-lite Vertebrates found in the Mr. l. Xantus. Pt. IV. Proc. Acad. Nat. Sci. Philad. 15: waters of north America, north oC the Isthmus of 80-88. Panama. Part 111. Bul!. U.S. Nat. Mus. 47: 2183-3136. 842 REVISTA DE BIOLOGIA TROPICAL

Systema naturae síve regna tria naturae, secundum, lordan, O.S. & B.W. Evermann. 1903. Descriptions of classes, ordines, genera, species, cum characteribus, new genera and species of fishes fram the Hawaiian differentiis, synonymis, locis. Holmiae. pt.l: 1-532. Islands. Bull. U.S. Fish. COmIDo 22: 161-208. McCosker, J .E. & R.H. Rosenblatt. 1993. A revision of the Jordan, D.S. & e.H. Gilbert. 1881a. Notes on a collection snake eel genus Myrichthys (Anguilliformes: of fishes made by Lieut. Henry E. Nichols, U.S.N. on Ophichthiidae) with the description of a new eastern the west coast of Mexico, with descriptions of new Pacificspecies. Proc. Cal. Acad. Sci. 48: 153-169. species. Prac. U.S. Nat. Mus. 4: 225-233. Müller, J. & F.G.J. Henle. 1838-1841. 1838, p. 27-102; Jordan, O.S. & C.H. Gilbert. 1881b. Descriptions of 1839, p. 27-28 (reset), p. 103-200. Systematische nineteen new species of fishes from the Bay of Beschreibung der Plagiostomen. Viete & CO. Berlin. Panama. Bull. U.S. Fish COmIDo 1: 306-335. Myers, G.S. & e.B. Wade. 1941. Four new genera and Jordan, D.S. & C.H. Gilbert. 1882a. Catalogue of the ten new species of eels from the Pacific coast of fishes collected by Mr. JOM Xantus at Cape San tropical America. A. Hancock Pacif. Exped. 9: 65- Lucas, which are now in the United States National 111. Museum, with descriptions of eight new Species. Proc. U.S. Nat. Mus. 5: 353-371. Osbeck, P. 1762. Reise nach Ostindien und China. (Translated fram Swedish by J.G. Georgius). Rostock. lordan, O.S. & e.H. Gilbert. 1882b. List of fishes 552 p. collected al Mazatlan, Mexico by Charles Henry Gilbert. Bull. U.S. Fish COmIDo 2: 105-108. Parin, N.V. 1961. Contributions to the knowledge of the flyingfish fauna (Exocoetidae) of the Pacific and Jordan, D.S. & c.H. Gilbert. 1882c. Descriptions of thirty­ lndian . Trudy Inst. Okeanologii 42: 40-91 three new species of Fishes from Mazatlan, Mexico. (Translated from Russian by U.S. Bur. Comm. Fish.) Prac. U.S. Nat. Mus. 4(1881): 338-365. Peters, W.C.H. 1869. Ueber neue Oder weniger bekannte Jordall, D.S. & R.e. McGregor. 1898. List of fishes Fische des Berliner Zoologischen Museums. collected al the Revillagigedo Archipelago and Monatsber. Akad. Wiss. Berlín. 1869: 703-711. neighboring Islands. Rept. U.S. Fish Comm. 24: 271- 284. Pietsch, T.W. & D.B. Grobecker. 1987. Frogfishes of (he world: Systematics, zoogeography, and behavioral Kaup, J.J. 1856. Catalogue of Lophobranchiate Fish in the ecology. Stanford U. Press. Stanford. 420 p. Collection of the British Museum. Taylor and Francis. London. iv + 76 p. Poey, F. 1860. XLIX. Poissons de Cuba, especes nouvelles: p. 115-356. In: Memorias sobre la historia Kishinouye, K. 1920. Mexican little Tunny. Suisan Gakkai natural de la isla de Cuba, Habana. Ho 3: 113. Quoy, J.R.C. & J.P. Gaimard. 1824. Chapt. 8, Poissons. p. LacepMe, B. 1801. Histoire Naturelle des Poissons. 183-328, pI. 43-65. In: Freycinet, L.CD. (ed) Voyage Vo!.3. Chez Plassan. Paris, lxvi + 558 p. autour du Monde, entrepris par ordre du Roi, execute sur les corvettes de S.M.L. Uranie et la Physicienne, Leis, lM. 1984. Larval fishdispersal and the East Pacific pendant les anees 1817, 1818, 1819, et 1820. PilIet Barrier. Oceanogr. Trop. 19: 181-192. Allne. Paris. Vol 1 and Atlas IV, 772 p. 96 pI.

Lessoll, R.P. 1829. Description du nouveau genre Quoy, J.R.C. & P. Gaimard. 1825. Zoologie. In: Voyage Ichthyopus el de plusiers especes inedites ou peu autour du monde ... Exécuté sur les Corvettes de S.M. connues de poissons recueillis dans le voyage autour l'Uranies et la Physicienne, pendant. les Années 1817- du monde de "La Coquille". Mem. Soco Hist. Nat. Paris 1820 ...Par M. L. de Freycinet. Imprimerie Royal. 4: 397-412. Paris. Vol. 3. iv + 712 p. atlas: 96 pI.

Lesson, R.P. 1831. Poissons. p. 66-238. In L.I. Ouperrey. Randall, J.E. 1956. A revision of the surgeonfish Genus Voyage autour du Monde Sur la "Coquille" pendan! Acanthurus. Pacif.S· ::;: 159-235. les Années 1822, 1823, 1824, and 1825 ... Zoologie. Vo1.2. Arthus Bertrand. Paris. Randall, J.E. 1961. A Record of the Kyphosid Fish Sectator ocyurus (=azureus) fram the Society Islands. Lesueur, C.A. 1821. Observations on several genera and Copeia, 1961: 357-358. species of fish belonging to ¡he natural farnily of the Esoces. 1. Acad. Nat. Sci. Philad. 2: 124-138. Randall, lE. 1985. Guide to Hawaiian Reef Fishes.Treasures of Nature. Kaneohe. 79 p. Limbaugh, C. 1963. Field notes on sharks. p. 63-94. In: P.W. Gilbert, Sharks and survival. D.C. Heath and Co .. Randall, J .E. 1995. On the validity of the eastern Pacific Bostoll. ¡abrid fishes Th.a lassoma grammaticum Gilbert and T. virens Gilbert. Bull. mar. Sci. 56: 670-675. Linnaeus, C. 1758. Systema Naturae. Ed.X. Vol. 1. reformata. Systema naturae per regna tria naturae, Randall, J.E. K Matsuura, & A. Zama. 1978. A revision of secnndum, c1asses, ordines, genera, species, cum the triggerflsh genus Xanthichthys, with a description characteribus, differentiis, synonymis, locis. Holmiae. of a new species. Bull. mar. Sci. 28: 688-706. ii + 824 p. Randall,lE. & J.E. McCosker. 1975. The eels of Eastee Linnaeus, C. 1766. Systema Naturae, Ed. XII. Vol. 1. Islandwith a description of a new Moray. Contrib. Sci. Allen & Robertson: Clipperton Atoll 843

LA County Mus. 264: 1-32. Springer, V.G. 1962. A review of the bleuniid fishes of the genus Op hioblennius Gill. Copeia 1962: 426-433. Robertson, D.R. & G.R AlIen. 1996. Zoogeography of the shorefish fauna of Clipperton Atoll. Coral Reefs 15: Springer, V.G. 1%7. Revision of the circumtropical 121-130. shorefish genus Entomacrodus (Bleuniidae: Salaríinae). Proc. U.S. Nat. Mus. 122: 1-150. Rosenblatt, R.H. l.E. McCosker & 1. Rubinoff. 1972. lndo-West Pacific fishes from the Gulf of Chiriqui, Steindachner, F. 1876. Ichthyologische Beitrage (V). 1. Panama. Contrib. Sci. Nat. Hist. Mus. LA County 234: Zur Fischfauna von Panama, Acapulco und Mazatlan. 1-18. Sitzber. Akad. Wiss. Wien 74: 49-240.

Rüppell, E. 1830. Atlas zu der Reise im Nordlichen Valenciennes, A. 1833. In: Cuvier, G. and A. Afrika, Zoologie. Fische des Rothen Meers.Gedruckt Valenciennes 1833. Histoire Naturelle des Poissons. und in Cornmission bei H.L. Brornmer. Frankfurt am Vol. 9. F.G. Levraolt. París. xxix + 512 p. Main. 3: 95-141. Valenciennes, A. 1847. In: Cuvier, G. and A. Rüppell, E. 1837. Neue Wirbelthiere zu der Fauna von Valenciennes 1847. Histoire Naturelle des Poissons. Abyssinien gehOrig. In cornmission bei S. Schnerber. Vol. 19. F.G. Levrault. París.xxiv + 544 p. Frankfurt am Main: 3: 53-80. Valenciennes, A. 1845. Voyage autour du monde sur la Rüppell, E. 1838. Neue Wirbelthiere zu der Fauna von Fregate Venus. Zoologie. 351p. Abyssinien gehOrig. In cornrnission bei S. Schnerber. Frankfurt am Main. 4: 81-148. Valenciennes, A. 1846. Table and pI. I-IX. In: A. du Petit-Thouars. Atlas de Zoologie. Voyage Sachet, M.H. 1962. Geography and land ecology of autour du Monde Sur la Fregate la "Vénus," pendant Clipperton Island. Atoll Res. Boll. 86: 1-115. les Anées 1836-39. París.

Schmidt, W.L. & L.P. Schultz. 1940. List of the fishes Valenciennes, A. 1855. Ichthyologie. p. ¡j-l ll + 297-323. taken on the presidential eruise of 1938. Smith. Mise. In: A. du Petit-Thouars. Voyage autour du Monde Sur Coll. 98: 1-10. la Fregate la "Vénus," pendant les Anées 1836-39. Zoology. París. Seale, A. 1940. Report on fishes from AlIan Haneock Expeditions in the California Academy of Sciences. Van Haselt, l.C. 1823. Uittreksel uit een brief de Heer AlIan Hancock Pacif. Exped. 9: 1-46. l.C. Van Hasselt, aan den HeerCJ. Ternmick, gescbreven uit Tj ecande, Residentie Bantam, den Shaw, G. 1803. General zoology or systematic natural 29sten December, 1822. Algemeene Konsten Letter­ history. Vol. 4. G. Kearsley. London. xiü + 187-632p. Bode. 2: 130-133.

Shaw, G. & F.P. Nodder. 1790-1813. The Naturalists Wellington, G.M. 1992. Xyrichtys victon, a new speeies Miscellany ...24 Vols. London. of razorfish from the Galapagos Islands (Teleostei, Labridae). Copeia 1992: 1053-1059. Skaggs, l.M. 1989. Clipperton, a History of the Island the World forgot. Walker and Company. New York. 318 Wellington, G.M. G.R. Allen & D.R Robertson. 1994. p. Xyrichtys perlas (Labridae), a new species of Smith, l.L.B. 1971. A revision of the american groupers: razorfish from the tropical eastern Pacifico Rev. fr. Ep inephelus and allied genera. BulI. Am. Mus. Nat. Aquaríol. 21: 49-52 Hist. 146: 1-241.

Snodgrass, R.E. & E. Heller. 1905. Papers frorn the Hopkins-Stanford Galapagos Expedition, 1898-1 899. XVII. Shore fishes of the Revillagigedo, Clipperton, Cocos and Galapagos Island. Proc. Wash. Acad. Sci. 6: 333-427.