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Home , Aphareus (fish), Canthigaster solandri, Centropyge multicolor, Chaetodon falcula, Chaetodon ulietensis, Cheilodipterus macrodon

Annotated Checklist ofthe ofWake 1

Phillip S. Lobel2 and Lisa Kerr Lobel 3

Abstract: This study documents a total of 321 fishes in 64 families occurring at Wake Atoll, a atoll located at 19 0 17' N, 1660 36' E. Ten fishes are listed by only and one by family; some of these represent undescribed . The first published account of the fishes of Wake by Fowler and Ball in 192 5 listed 107 species in 31 families. This paper updates 54 synonyms and corrects 20 misidentifications listed in the earlier account. The most recent published account by Myers in 1999 listed 122 fishes in 33 families. Our field surveys add 143 additional species records and 22 new family records for the atoll. Zoogeo­ graphic analysis indicates that the greatest species overlap of Wake Atoll fishes occurs with the . Several species common at Wake Atoll are on the IUCN Red List or are otherwise of concern for conservation. Fish pop­ ulations at Wake Atoll are protected by virtue of it being a U.S. military base and off limits to commercial fishing.

WAKE ATOLL IS an isolated atoll in the cen­ and Strategic Defense Command. Conse­ tral Pacific (19 0 17' N, 1660 36' E): It is ap­ quentially, access has been limited due to the proximately 3 km wide by 6.5 km long and military mission, and as a result the aquatic consists of three islands with a land area of fauna of the atoll has not received thorough 2 approximately 6.5 km • Wake is separated investigation. Our surveys were sponsored by by 546 km of deep ocean from the nearest the Department of Defense Legacy Resource reef system on Taongi Atoll in the Marshall Management Program as part of the program Islands to the south. It is 1900 km south­ to implement conservation and west of , 1400 km southeast of protection (Kerr and Lobel 2000). Minami-tori-shima (Marcus) Island, and 2260 The first ichthyological collection at Wake km east of the Mariana Islands. Thus, it is in resulted in a checklist of 107 species (Fowler a unique position with respect to the bio­ and Ball 1925). Specimens were deposited in geography of reef fauna. This survey fills a the Bernice P. (BPBM), Ho­ major data gap in the zoogeography ofPacific nolulu, Hawai'i, with duplicates deposited in fishes (see Randall 1998). the Academy of Natural Sciences in Phila­ Wake Atoll is a U.S. Air Force installa­ delphia (ANSP). The second important collec­ tion administered by the U.S. Army Space tion was by W. Gosline and J. Randall during 1953; most of their specimens are also de­ posited in the BPBM with a few at the National 1 This project was supported by the Department of Defense Legacy Resource Management Program grants Museum of Natural History (USNM) and the DACA87-97-H-0006 and DACA87-01-H-00013 and by remaining specimens are held in the Univer­ the Army Research Office grants DAAG55-98-1-0304 sity of Hawai'i (Department of ) fish and DAADI9-02-1-0218. Manuscript accepted 9 May collection. Specimens held in the 2003. Academy of Sciences (CAS) were collected by 2 Boston University Marine Program, Marine Bio­ logical Laboratory, Woods Hole, 02543. J. Kauanui and P. Kauanui or J. Oetzel in J Department of Biology, University of Massachu­ 1956 and 1957. Specimens held in the Har­ setts, 100 Morrissey Boulevard, Boston, Massachusetts vard Museum of Comparative Zoology (MCZ) 02125. were collected by us in 1999. The collections of the and by Gosline Pacific Science (2004), vol. 58, no. 1:65 -90 and Randall were primarily in shallow water © 2004 by University of Hawai'i Press without the benefit ofscuba and based largely All rights reserved on the use of ichthyocides and conventional

65 66 PACIFIC SCIENCE· January 2004

fishing techniques. Myers' (1999) listing of numbers (i.e., BPBM 1234) are given for those fishes from Wake Atoll is based on photo­ species maintained in ichthyological collec­ graphs, or in some cases video, taken by mil­ tions. In addition, the expedition during itary personnel. which the specimens in the Bishop Museum This checklist includes all species reported were collected is designated: by (T) for the in previous publications, species not pre­ Tanager Expedition of 192 3, (K) for fishes viously verified or published from the B. P. collected by J. Kauanui in 1957, (G&R) for Bishop Museum collection, and new records fishes collected by Gosline, Randall et al. found during our comprehensive scuba sur­ during 1953, and (K&L) for fishes collected veys conducted at Wake during July 1997 and by us in 1997 and 1999. May 1999. We have updated species reported Lists of specimens from other scientific in previous publications with current nomen­ collections at Wake Atoll but that have not clature and list the synonyms. yet been published were provided by the The following checklist represents all of ANSP, BPBM, and CAS. The list of fishes from the fishes currently known to occur at Wake the USNM was taken from their Web site Atoll. A total of 143 new records is added (http://www.mnnh.si.edu/vert/fishcat/). and includes fishes from the following fami­ The identification of many of the speci­ lies: Rhincodontidae, Albulidae, Moringuidae, mens held in the BPBM was verified by J. Chlopsidae, Muraenidae (eight species), Randall and A. Suzumoto. We verified the (two species), Congridae (two identity of the specimens held in the Univer­ species), Bythitidae, Mugilidae (three spe­ sity of Hawai'i, Department of Zoology fish cies), Atherinidae, Exocoetidae, Holocentri­ collection. J. McCosker verified some of the dae (three species), Fistularidae, Scorpaenidae specimens held in the CAS; however, many (five species), Caracanthidae, of the specimens reported in the collections (eight species), , Pseudochromidae, of CAS, USNM, and ANSP were not reexamined. Priacanthidae, (eight species), Thus upon further taxonomic evaluation, Malacanthidae, Echeneidae, Carangidae (six some of these earlier identifications may be species), (two species), , revised. Mullidae (three species), Pempheridae, Chae­ Our underwater surveys were conducted todontidae (two species), , during two expeditions: 10 to 23 July 1997 Oplegnathidae, (six species), and 15 to 29 May 1999. We completed a total Labridae (18 species), Scaridae (three species), of 118 dives during that time at sites located Pinguipedidae, Creedidae, Tripterygidae, around the atoll (Figure 1). In addition, more Blenniidae (three species), Callionymidae, than a dozen shallow-water snorkel surveys (14 species), , Siga­ were also conducted at locations around the nidae, Acanthuridae (eight species), Bothidae, atoll. The majority of observations were done Balistidae (three species), Monacanthidae by visual and photographic census (using both (two species), and (four spe­ still and video photography, which are stored cies). Twenty-two of these families represent in our collection). Several collections of small new family records for Wake Atoll. cryptic fishes were made using quinaldine. These were all located at the dive site directly off the small boat harbor and ranged in depth MATERIALS AND METHODS from 10 to 50 m. Several fishes were speared Species are listed in phylogenetic sequence by or collected by nets to confirm species iden­ family with the current valid name first, fol­ tification. lowed by synonyms ifthey were cited in other Overall, our surveys provide a reliable checklists. An asterisk (*) designates those census for the majority of reef fishes but cer­ species that are our new records for Wake tainly did not adequately sample small cryptic Atoll. The author and date of description are species that remain hidden in crevices and given for each species followed by the rec­ . Future surveys should include sam­ ord(s) for Wake Atoll. Museum specimen pling the cryptic fauna. Checklist of Fishes ofWake Atoll . Lobel and Lobel 67

Reef

1 km

FIGURE 1. Fish survey locations around Wake Atoll. Black dots show survey sites; multiple dives were conducted at many of these locations.

RESULTS Family MYLIOBATIDAE (Eagle Rays) Family RHINCODONTIDAE (Whale Shark) Aetobatus narinari (Euphrasen, 1790). Myers *Rhincodon typus (Smith, 1828). Sight record 1999. reported by island divers. Family MOBuLIDAE (Manta Rays) Family CARCHARHINIDAE (Requiem Sharks) Manta birostris (Donndorff, 1798). Myers Carcharhinus amblyrhynchos (Bleeker, 1856). 1999. Myers 1999. Carcharhinus melanopterus (Quoy & Gaimard, Family ALBULIDAE (Bonefish) 1824). Myers 1999. No blacktip reef sharks *Albula glossodonta (Forsskal, 1775). were observed by us. Eulamia commersoni (Blainville). BPBM 3500 Family MORINGUIDAE (Spaghetti ) (T). Fowler 1928. Triaenodon obesus (Ruppell, 1837). Fowler and *Moringua abbreviata (Bleeker, 1863). BPBM Ball 1925, Myers 1999. 28145 (G&R). 68 PACIFIC SCIENCE· January 2004

Family CHLOPSIDAE (False Morays) maculosus (Cuvier, 1817). Mc­ *Kaupichthys sp. CAS 60094. Cosker and Rosenblatt 1993, Myers 1999. *Scolecenchelys gymnota (Bleeker, 1857). CAS Family MURAENIDAE (Moray Eels) 200128. *Anarchias sp. BPBM 12710 (G&R). Family CONGRIDAE *Anarchias cantonensis (Schultz, 1943). CAS (Conger and Garden Eels) 149959. *Anarchias seychellensis Smith, 1962. CAS *Conger cinereus Ruppell, 1830. BPBM 12668 51023. (G&R). *Echidna leucotaenia Schultz, 1943. CAS 28698. *Heteroconger hassi (Klausewitz & Eibl *Gymnomuraena zebra (Shaw in Shaw and Eibesfeldt, 1959). Photographic record. Nodder, 1797). Collected by G&R and Family CHANIDAE (Milkfish) kept in the University of Hawai'i fish col­ lection, UH 8-IV-53. Chanos chanos (Forsskal, 1775). Myers 1999. Gymnothorax buroensis (Bleeker, 1857). BPBM Family MYCTOPHIDAE (Lanternfishes) 12360 (G&R). Bohlke and Randall 2000. *Gymnothorax enig;maticus McCosker & Ran­ Myctophum spinosum (Steindachner, 1867). dall, 1982. CAS 51024. BPBM 3643 (T). Gymnothorax fiavimarginatus (Ruppell, 1828). Dasyscopelus spinosus (Schneider). Fowler CAS 37106. and Ball 1925. Lycondontis flavomarginata (Ruppell). Fowler and Ball 1925 (BPBM specimen not Family BYTHITIDAE (Livebearing Brotulas) located). *Dinematichthys ilucoeteoides Bleeker, 1855. Gymnothorax javanicus (Bleeker, 1859). BPBM BPBM 15169 (G&R). This fish is probably 12358 (G&R). Myers 1999, Bohlke and not ilucoeteoides (]. Randall, pers. comm.), Randall 2000. but it has not yet been reclassified. *Gymnothorax meleagris (Shaw in Shaw and Nodder, 1795). UH Collection. Family MUGILIDAE (Mullets) Gymnothorax pictus (Ahl, 1789). BPBM 3564, *Crenimugil crenilabis (Forsskal, 1775). BPBM 3565 (T), BPBM 12530, 12544 (G&R). 3816 (T). Omitted by Fowler and Ball Bohlke and Randall 2000. 1925. Lycodontis picta (Ahl). Fowler and Ball *Liza vaigiensis (Quoy & Gaimard, 1825). A 1925. photograph was taken of a juvenile seen Gymnothorax ruppelliae (McClelland, 1845). in the small boat harbor. This distinctive BPBM 3593 (T), BPBM 12371, 12527 (G&R). species has black pectoral fins and yellow Lycodontis petelli (Bleeker). Fowler and Ball pelvic and anal fins. 1925. *Neomyxus leuciscus (Gunther, 1872). CAS Gymnothorax undulatus (Lacepede, 1803). 150042. BPBM 12380 (G&R). Bohlke and Randall 2000. Family CLUPEIDAE (Herrings) Lycodontis undulata (Lacepede). Fowler and Ball 1925. Spratelloides sp. An unidentified clupeid that Uropterygius macrocephalus (Bleeker, 1865). may be Spratelloides was photographed in CAS 41192. McCosker et al. 1984, Bohlke the channel between Wake and Peale Is­ and Randall 2000. lands. *Uropterygius xanthopterus Bleeker, 1859. CAS Family EXOCOETIDAE (Flyingfishes) 149964. *Cypselurus poecilopterus (Valenciennes, 1847). Family OPHICHTHIDAE (Snake Eels) BPBM 10536. * (Boddaert, 1781). BPBM Exocoetus volitans Linnaeus, 1758. BPBM 3668, 12518 (G&R). 3669 (T). Fowler and Ball 1925. Checklist of Fishes of Wake Atoll . Lobel and Lobel 69

Hirundichthys rondeletii (Valenciennes, 1847). tiere (Cuvier in C and V, 1829). BPBM 3673 (T). ANSP 102854. Randall 1998a, Myers 1999. Cypselurus rondelitii (Valenciennes). Fowler Family SYGNATHIDAE and Ball 1925. (Pipefishes and Seahorses) Family HEMIRAMPHIDAE (Halfbeaks) Corythoichthys fiavofasciatus (Ruppell, 1838). Hyporhamphus acutus acutus (Gunther, 1872). BPBM 3775 (T), BPBM 37884 (K&L). MCZ 158366. Collette 1974. Myers 1999. Oxyporhampus micropterus (Valenciennes, C01ythoichthys conspicillatus (Jenyns). 1846). Myers 1999. Fowler and Ball 1925. Doryrhampus excisus Kaup, 1856. Myers 1999. Family (Soldierfishes and Squirrelfishes) Family AULosToMIDAE (Trumpetfishes) * adusta (Bleeker, 1853). Photo­ Aulostomus chinensis (Linnaeus, 1766). Fowler graphic record. and Ball 1925. Although A. chinensis is Myripristis amaena (Castelnau, 1873). BPBM listed in Fowler and Ball (1925), we did not 15173 (G&R). Randall and Greenfield observe any. Apparently the only Fowler 1996. and Ball specimen ofA. chinensis to survive Myripristis berndti Jordan & Evermann, 1903. to the 1960s was from . BPBM 3758 (T), BPBM 28241. Randall and This specimen was discarded by BPBM in Greenfield 1996, Myers 1999. Identified as 1968. Thus this record from Wake Atoll is Myripristis murdjan (Forsskal) in Fowler doubtful. and Ball 1925. Family FISTULARIIDAE (Cornetfishes) *Myripristis kuntee Cuvier in C and V, 1831. *Fistularia commersonii Ruppell, 1838. Photo­ Myripristis murdjan (Forsskal, 1775). Myers graphic record. 1999 tentatively listed this species. Family SCORPAENIDAE (Scorpionfishes) *Myripristis violacea (Bleeker, 1851). Photo­ graphic record. Pterois antennata (Bloch, 1787). BPBM 15280 opercularis (Valenciennes, 1831). (G&R). Myers 1999. BPBM 3752 (T). *Scorpaenodes guamensis (Quoy & Gaimard, Holocentrus opercularis Valenciennes. 1824). CAS 150161. Fowler and Ball 1925. Sebastapistes ballieui (Sauvage in Vaillant and (Forssk:'il, 1775). BPBM Sauvage, 1875). BPBM 4365. Identified as 3751 (T), BPBM 15110 (G&R). Sebastapistes asperella (Bennett) in Fowler Holocentrus sammara Forsskal. Fowler and and Ball 1925, a species with inadequate Ball 1925. description and no specimen (Esch­ Sargocentron melanospilos (Bleeker, 1858). meyer and Randall 1975). The BPBM lot Myers 1999. contains two specimens, one S. ballieui and Sargocentron microstoma (Gunther, 1859). one S. cyanostigma; however, the locality BPBM 3733 (T). Myers 1999. of this lot was incorrectly listed as Wake Holocentrus microstomus Gunther. Fowler (J. Randall, pers. comm.). This species is and Ball 1925. not included in the species count. Sargocentron punctatissimum (Cuvier, 1829). Sebastapistes coniorta Jenkins, 1903. Randall BPBM 3715 (T). Myers 1999. 1996, Myers 1999. Holocentrus laeteoguttatus (Cuvier). Fowler *Sebastapistes fowleri (Pietschmann, 1934). and Ball 1925. BPBM 38593 (K&L). Sargocentron spiniferum (ForsskiH, 1775). BPBM *Sepastapistes mauritiana (Cuvier, 1829). BPBM 3738 (T), BPBM 15375 (G&R). Myers 1999. 12043 (K), BPBM 13804 (G&R). Holocentrus spinifer Forsskal. Fowler and *Sebastapistes tinkhami (Fowler, 1946). MCZ Ball 1925. 158370. 70 PACIFIC SCIENCE· January 2004

Family CARACANTHIDAE *Liopropoma tonstrinum Randall & Taylor, (Orbiculate Velvetfishes) 1988. BPBM 37889 (K&L). *Plectranthias longimanus (Weber, 1913). MCZ *Caracanthus maculatus (Gray, 1831). BPBM 158369. 4422 (T), MCZ 158364. This record was *Pleetranthias nanus Randall, 1980. MCZ omitted from Fowler and Ball 1925. 158368. Caracanthus unipinna (Gray, 1831). BPBM 4425 *Plectranthias winniensis (Tyler, 1966). BPBM (T). Fowler and Ball 1925. 37890 (K&L). Family SERRANIDAE pascalus (Jordan & Tanaka, ( and Sea Basses) 1927). BPBM 37891 (K&L). Myers 1999. *Pseudanthias ventralis (Randall, 1979). BPBM *Aporops bilinearis Schultz, 1943. CAS 82471. 37892 (K&L). Cephalopholis argus (Schneider, 1801). BPBM *Pseudogramma polyacantha (Bleeker, 1856). 3951 (T), BPBM 12042 (K), BPBM 15098, BPBM 37893 (K&L). 15144 (G&R), su 500065, ANSP 88339. Variola louti (Forsskal, 1775). BPBM 3947­ Fowler and Ball 1925, Randall and Heem­ 3948 (T). Fowler and Ball 1925, Randall stra 1991, Myers 1999. and Heemstra 1991, Myers 1999. Cephalopholis spiloparaea (Valenciennes, 1828). Family CIRRHITIDAE (Hawkfishes) BPBM 3950 (T). Randall and Heemstra 1991, Myers 1999. This specimen was *Amblycirrhitus bimacula (Jenkins, 1903). CAS misidentified as Cephalopholis sonnerati (Va­ 24409. lenciennes) by Fowler and Ball 1925. Cirrhitus pinnulatus (Schneider in Bloch and *Cephalopholis urodeta (Forster, 1801). Photo­ Schneider, 1801). BPBM 4103 (T), BPBM graphic record. 15121 (G&R). Epinephelus fasciatus (Forsslcll, 1775). ANSP Cirrhitus maculatus (Lacepede, 1803). 91775,91783, BPBM 3956 (T), BPBM 15428 Fowler and Ball 1925. (G&R). Randall and Heemstra 1991, Neocirrhites armatus Castlenau, 1873. BPBM Myers 1999. 4122 (T). Myers 1999. Holocentrus erythraeus (Bloch & Schneider). Paracirrhites melanotus (Giinther). Fowler Fowler and Ball 1925. and Ball 1925. Serranus fasciatus (Forsslcll). Fowler and Paracirrhites arcatus (Cuvier in C and V, Ball 1925. 1829). Myers 1999. Epinephelus hexagonatus (Forster, 1801). ANSP Paracirrhites forsteri (Schneider in Bloch and 102856, BPBM 3976 (T) 12040. Randall Schneider, 1801). BPBM 4126 (T). Myers and Heemstra 1991, Myers 1999. Speci­ 1999. men BPBM 3976 was misidentified as Ser­ Paracirrhites hemistictus (Gunther, 1874). ranus merra by Fowler and Ball 1925. Myers 1999. Epinephelus lanceolatus (Bloch, 1790). Myers Family PSEUDOCHROMIDAE (Dottybacks) 1999. *Epinephelus merra Bloch, 1790. BPBM 12040 *Pseudochromis sp. Video taken on the eastern (G&R). side of of a pseudochromid Epinephelus polyphekadion (Bleeker, 1849). that is likely to be either P. fuscus or P. Randall and Heemstra 1991, Myers 1999. cyanotaenia. R. F. Myers (pers. comm.) Epinephelus spilotoceps Schultz, 1953. Myers noted that this is likely to be P. cyanotaenia, 1999. not P. fuscus. Epinephelus tauvina (ForsskiH, 1775). ANSP *Pseudoplesiops sp. Specimen collected in 1997. 105506, BPBM 3963 (T). Randall and This is an unidentified species. Heemstra 1991, Myers 1999. This spe­ Family PRIACANTHIDAE (Bigeyes, Glasseyes) cimen was misidentified as Serranus cor­ allicola (Valenciennes) by Fowler and Ball *Heteropriacanthus cruentatus (Lacepede, 1925. 1801). Photographic record. Checklist of Fishes ofWake Atoll· Lobel and Lobel 71

Family ApOGONIDAE (Cardinalfishes) Family CORYPHAENIDAE (Dolphinfishes) *Apogon eoeeineus Ruppell, 1838. CAS 150060. Coryphaena hippurus Linnaeus, 1758. Myers *Apogon cyanosoma Bleeker, 1853. CAS 85030. 1999. Apogon doryssa (Jordan & Seale, 1906). BPBM Family CARANGIDAE (Jacks) 3901 (T). Amia doryssa (Jordan & Seale). Fowler Carangoides ferdau (Forsskal, 177 5). Fowler and Ball 1925. Specimen could not be and Ball 1925. located. *Carangoides orthogrammus (Jordan & Gil­ Apogon exostigma (Jordan & Starks, 1906). bert, 1881). BPBM 3885 (T). Photographic Myers 1999. record. *Apogon fuscus Quoy & Gaimard, 1825. Pho­ *Caranx ignobilis (Forssk:H, 1775). Photo­ tograph was tentatively identified as A. graphic record. fuscus. Caranx lugubris Poey, 1861. Myers 1999. Apogon kallopterus Bleeker, 1856. BPBM Caranx melampygus Cuvier in C and V, 1833. 15273 (G&R), BPBM 37876 (K&L). Myers Myers 1999. 1999. Caranx sexfaseiatus Quoy & Gaimard, 1824. *Apogon savayensis (Giinther, 1871). BPBM Myers 1999. 3894 (T), BPBM 12041 (K), MCZ 158363. *Deeapturus maearellus (Cuvier, 1833). Photo­ Probably misidentified as Amia (Apogon) graphic record. bandanensis (Bleeker) by Fowler and Ball *Elagatis bipinnulatas (Quoy & Gaimard, 1925. 1824). Species identified from a photo­ Apogon susanae Greenfield, 2001. CAS 84823. graph taken by local fishermen. This specimen from Wake was misiden­ *Gnathanodon speeiosus (Forsskal, 1775). Pho­ tified as A. eoeeineus. The new species A. tographic record. susanae described by Greenfield (2001) in­ Seomberoides lysan (Forsskal, 1775). Myers cluded this specimen in the description. 1999. *Apogon taeniophorus Regan, 1905. Seriola rivoliana Valenciennes, 1833. Myers * macrodon (Lacepede, 1802). 1999. MCZ 158371. *Traehinotus baillonii (Lacepede, 1801). CAS *Cheilodipterus quinquelineata (Cuvier, 1828). 24035. BPBM 37877 (K&L). *Powleria isostigma (Jordan & Seale, 1906). Family LUT]ANIDAE (Snappers) BPBM 3916 (T). Identified as Apogoniehthys *Aphareus furea (Lacepede, 1801). Photo­ auritus (Valenciennes) (current name graphic record. Powleria aurita) by Fowler and Ball 1925. *Aprion vireseens Valenciennes, 1830. Photo­ Family MALACANTHIDAE (Sand Tilefishes) graphic record. fulvus (Schneider, 1801). BPBM 4006, *Malaeanthus brevirostris Guichenot, 1848. 4007. Myers 1999. Lutjanus marginatus (Cuvier). Fowler and Family ECHENEIDAE (Remoras) Ball 1925. Remora remora (Linnaeus, 1758). BPBM 4894 Lutjanus monostigma (Cuvier, 1828). BPBM (T). 4012 (T). Eeheneis remora (Linnaeus). Fowler and Lutjanus monostigma (Cuvier). Fowler and Ball 1925. Ball 1925. 1829). USNM 202330, *Remora osteoehir(Cuvier, Family LETHRINIDAE (Emperors) 202332. Exclusively pelagic species col­ lected from the National Marine Fisheries obsoletus (Forsskal, 1775). Myers Service R.V. Charles H. Gilbert and placed 1999. in the USNM by D. W. Strasburg (U.S. Fish *Lethrinus rubriopereulatus Sato, 1978. BPBM and Wildlife Service). 4023 (T). Originally identified as Lethri- 72 PACIFIC SCIENCE· January 2004

nus mahsena (Forsskal) by Fowler and Ball quadrimaculatus Gray, 1831. BPBM 1925. 15192 (G&R). Fowler and Ball 1925, grandoculis (ForsskiH, 1775). BPBM Myers 1999. 4032 (T). Fowler and Ball 1925, Myers Chaetodon reticulatus Cuvier in C and V, 1831. 1999. BPBM 4167 (T). This specimen was mis­ identified as Chaetodon collaris Bloch by Family MULLIDAE (Goatfishes) Fowler and Ball 192 5. Also reported by *Mulloidichthys flavolineatus (Lacepede, 1801). Myers 1999. BPBM 4089 (T). Chaetodon semeion Bleeker, 1855. BPBM 4150 *Mulloidichthys vanicolensis (Valenciennes in C (T). Fowler and Ball 1925. and V, 1831). Photographic record. Cuvier, 1831. BPBM 4195, Parupeneus barberinus (Lacepede, 1801). 4196 (T). Myers 1999. Misidentified as Myers 1999. Bloch in Fowler and Ball Parupeneus cyclostomus (Lacepede, 1801). BPBM 1925. 4077 (T). Myers 1999. *Chaetodon unimaculatus Bloch, 1787. CAS Upeneus chryserydros (Lacepede). Fowler 150086. and Ball 1925. *Forcipiger flavissimus Jordan & McGregor, Parupeneus insularis Randall & Myers, 2002. 1898. Photographic record. BPBM 4070 (T), BPBM 15310, 15442 Forcipiger longirostris (Broussonet, 1782). (G&R). Specimen BPBM 4070 was identi­ Fowler and Ball 1925, Myers 1999. fied by Fowler and Ball 1925 as Upeneus thompsoni Fowler, 192 3. indicus (Shaw). Until recendy, this species Myers 1999. was classified as P. bifasciatus in the central Heniochus acuminatus (Linnaeus, 1758). Myers Pacific (Randall and Myers 2002). 1999. Parupeneus multifasciatus (Quoy & Gaimard, Family POMACANTHIDAE (Angelfishes) 1825). Myers 1999. *Parupeneus pleurostigma (Bennett, 1830). flavissima (Cuvier, 1831). BPBM Upeneus arge Jordan & Evermann, 1903. 4221 (T), BPBM 15344 (G&R). Myers Myers 1999. 1999. Holocanthus flavissimus (Cuvier). Fowler Family PEMPHERIDAE (Sweepers) and Ball 1925. *Pempheris oualensis Cuvier, 1831. Photo­ Centropyge loricula (Giinther, 1873). Myers graphic record. 1999. *Centropyge multicolor Randall & Wass, 1974. Family CHAETODONTIDAE () Photographic record. Chaetodon auriga Forsskal, 1775. BPBM 4183. Family KYPHOSIDAE Myers 1999. (Rudderfishes, Sea Chubs) Chaetodon setifer Bloch. Fowler and Ball 1925. Kyphosus bigibbus Lacepede, 1802. BPBM 4043, Chaetodon ephippium Cuvier in C and V, 1831. 4044 (T). Myers 1999. BPBM 4149 (T), BPBM 15183 (G&R). Kyphosus fuscus (Quoy & Gaimard). BPBM Fowler and Ball 1925, Myers 1999. 4043 (T). Omitted by Fowler and Ball *Chaetodon lineolatus Cuvier in C and V, 1831. 1925. Video was taken of this species. Kyphosus cinerascens (Forsskal, 1775). Myers Chaetodon lunula (Lacepede, 1803). BPBM 1999. 4164 (T), BPBM 15193 (G&R). Fowler and Family KUHLIIDAE (Flagtails) Ball 1925, Myers 1999. Chaetodon ornatissimus Cuvier in C and V, Kuhlia sandvicensis (Steindachner, 1876). BPBM 1831. Fowler and Ball 1925, Myers 1999. 3929 (T). Fowler and Ball 1925. These Chaetodon punctatofasciatus Cuvier, 1831. specimens were determined as the true Myers 1999. sandvicensis (which ranges beyond Hawai'i), Checklist of Fishes ofWake Atoll . Lobel and Lobel 73

although some characters (large eye Dascyllus aruanus (Linnaeus, 1758). BPBM and some meristic data) do not fit well. 4443 (T), BPBM 15178, 15367 (G&R). The possibility exists that this could be Fowler and Ball 1925. some other species (Randall and Randall Plectroglyphidodon dickii (Lienard, 1839). 2001). Myers 1999. Plectroglyphidodon imparipennis (Vaillant & Family OPLEGNATHIDAE (Knifejaws) Sauvage, 1875). BPBM 4536 (T), BPBM 15341 (G&R). *Oplegnathus punctatus (Temminck & Schle­ Abudefduf imparipennis (Vaillant & Sau­ gel, 1844). BPBM 37896 (K&L). vage). Fowler and Ball 1925. Family POMACENTRIDAE (Damselfishes) Plectroglyphidodon johnstonianus Fowler & Ball, 1924. Myers 1999. Abudefdufseptemfasciatus (Cuvier, 1830). BPBM *Plectroglyphidodon lacrymatus (Quoy & Gai­ 4506 (T). Fowler and Ball 1925. mard, 1825). Photographic record. Abudefduf sordidus (Forssk:H, 1775). BPBM *Plectroglyphidodon phoenixensis (Schultz, 4490, 4491 (T). Fowler and Ball 1925. 1943). UH collection. *Abudefduf vaigiensis (Linnaeus, 1758). Pho­ (Schlegel & Muller, tographic record. 1839). No specimens found in the BPBM. Chromis acares Randall & Swerdloff, 1973. Abudefdufalbofasciatus (Schlegel & Muller). BPBM 4450 (T), MCZ 158367. Myers 1999. Fowler and Ball 1925. Identified as Chromis dimidiatus (Klun­ * (Ogilby, 1889). Photo­ zinger) by Fowler and Ball 1925. graphic record. Chromis agilis Smith, 1960. BPBM 37888 (Lacepede, 1803). BPBM (K&L). Myers 1999. 4464,4465 (T), BPBM 15177 (G&R). *Chromis vanderbilti (Fowler, 1941). BPBM Pomacentrus nigricans (Lacepede). Fowler 12300 (G&R). Photographic record. and Ball 1925. Chromis viridis (Cuvier, 1830). Specimens Family LABRIDAE () listed as Chromis caeruleus (Cuvier) in Fowler and Ball 1925 and as C. caerulea, *Ammolabrus diems Randall & Carlson, 1997. BPBM 4454 (T), BPBM 15319 (G&R). BPBM 38592 (K&L). This species is cur­ Chromis caerulea has been suppressed. rently known only from O'ahu and Wake Chromis ternatensis, which formerly had Atoll. The same species or a close relative been synonymized with C. caerulea (Allen was observed in the Ogasawara Islands, 1991), was validated as a distinct species Japan, but was not collected (Randall and and the next available name for what had Carlson 1997). been called C. caerulea was C. viridis (Ran­ Anampses caeruleopunctatus Ruppell, 1829. MCZ dall et al. 1987). 158361. Myers 1999. Chrysiptera biocellata (Quoy & Gaimard, *Bodianus anthioides (Bennett, 1831). Photo­ 1824). BPBM 4516 (T). Identified as Abu­ graphic record. defdufbrownriggii (Bennett) by Fowler and *Cheilinus chlorourus (Bloch, 1791). BPBM 4688 Ball 1925. (T), BPBM 12047 (K), BPBM 12723 (G&R) Chrysiptera brownriggii (Bennett, 1828). BPBM Originally identified as Cheilinus undulatus 4528, 4540 (T). Ruppell by Fowler and Ball 192 5. Abudefduf amabilis (De Vis). Fowler and *Cheilinus fasciatus (Bloch, 1791). Ball 1925. Cheilinus trilobatus Lacepede, 1801. Myers Abudefduf leucopomus (Cuvier). Fowler and 1999. Ball 1925. Cheilinus undulatus Ruppell, 1835. Myers *Chrysiptera glauca (Cuvier, 1830). BPBM 4542 1999. (T), BPBM 15362 (G&R). Identified as Coris aygula Lacepede, 1801. BPBM 5116, Abudefduf zonatus (Cuvier) by Fowler and 12046 (K). Fowler and Ball 1925, Randall Ball 1925. 1999. 74 PACIFIC SCIENCE· January 2004

Epibulis insidiator (Pallas, 1770). BPBM 12044 Stethojulis bandanensis (Bleeker, 1851). BPBM (K). Myers 1999. 12730, 12732, 15205 (G&R). Myers 1999. varius Lacepede, 1801. BPBM *Thalassoma hardwicke (Bennett, 1820). CAS 12045 (K). Myers 1999. 75547. Halichoeres biocellatus Schultz, 1960. Myers Thalassoma lutescens (Lay & Bennett, 1839). 1999. BPBM 4646 (T), BPBM 12680, 12687 *Halichoeres margaritaceus (Valenciennes in C (G&R). Myers 1999. and V, 1839). CAS 73712. Thalassoma purpureum (Forssk:'il, 1775). BPBM *Halichoeres ornatissimus (Garrett, 1863). BPBM 4640,4641 (T). Fowler and Ball 1925. 15347 (G&R). Thalassoma umbrostigma (Riippell). Fowler Halichoeres trimaculatus (Quoy & Gaimard, and Ball 192 5 also spelled it umbrostygma. 1824). BPBM 4577 (T), BPBM 12712, Thalassoma quinquevittatum (Lay & Ben­ 12715, 15172 (G&R). Fowler and Ball nett, 1839). BPBM 4637 (T), BPBM 15244 1925. (G&R). Myers 1999. Misidentified as Hemig;ymnus fasciatus (Bloch, 1792). Myers Thalassoma g;untheri (Bleeker) by Fowler 1999. and Ball 1925. *Iniistius sp. An unknown species with an all­ *Thalassoma trilobatum (Lacepede, 1801). white body was observed in the on BPBM 15202 (G&R). several occasions. No specimens were col­ *Xyrichtys undescribed species. BPBM 38591, lected. paratype (K&L). ]. Earle and B. Halstead Labroides bicolor Fowler & Bean, 1928. Myers collected one pair ofthis species from New 1999. Guinea in 1993 and it was photographed Labroides dimidiatus (Valenciennes, 1839). in by T. Allen. We collected speci­ Myers 1999. mens from Wake at 20 m over sand, di­ Labroides pectoralis Randall & Springer, 1975. rectly in front of the small boat harbor. BPBM 37886 (K&L). Myers 1999. For description of this species see Randall *Labroides rubrolabiatus Randall, 1958. CAS and Lobel (in press). 73544. Family SCARIDAE () Novaculichthys taeniouris (Lacepede, 1801). BPBM 4696 (T). Fowler and Ball 1925. Bolbometopon muricatum (Valenciennes, 1839). Oxycheilinus diagrammus (Lacepede, 1801). Myers 1999. Cheilinus diagrammus (Lacepede). Fowler Callyodon borborus Jordan & Evermann, 1903 and Ball 1925. reported by Fowler and Ball 1925 is a *Oxycheilinus orientalis (Giinther, 1862). of the Hawaiian endemic Chloru­ *Oxycheilinus unifasciatus (Streets, 1877). BPBM rus perspicillatus (Steindachner). No speci­ 12722 (G&R). mens were found at BPBM. This species is Pseudodcheilinus hexataenia (Bleeker, 1857). not included in the species count. BPBM 4686 (T). Fowler and Ball 1925. Chlorurus frontalis Valenciennes, 1839. Myers *Pseudocheilinus ocellatus Randall, 1999. BPBM 1999. 37887, paratype (K&L). Callyodon jonesi (Streets). Fowler and Ball *Pseudocheilinus octotaenia Jenkins, 1900. 1925. *Pseudocheilinus tetrataenia Schultz, 1960. (Bleeker, 1854). Myers *Pseudocoris aurantiofasciata Fourmanoir, 1971. 1999. BPBM 38772. These fish were observed (Forsskal, 1775). BPBM feeding in mixed schools with Acanthurus 4739, 4748 (T). thompsoni in 18 to 20 m near the wreck of Callyodon sordidus (Forsskal). Fowler and the Stoner. Ball 1925. atavai Randall & Randall, 1981. Callyodon erythrodon (Valenciennes). Myers 1999. Fowler and Ball 1925. *Pseudojuloides cerasinus (Snyder, 1904). Pho­ *Hipposcarus longiceps (Valenciennes, 1839). tographic record. Photographic record. Checklist of Fishes of Wake Atoll . Lobel and Lobel 75

Scarus forsteni (Bleeker, 1861). BPBM 4770 Cirripectes polyzona (Bleeker, 1868). Myers (T). Myers 1999. Misidentified as Callyo­ 1999. don () tricolor (Bleeker) by Fowler Cirripectes quagga (Fowler & Ball, 1924). ANSP and Ball 1925. 91191, 91013, BPBM 3420 (T), BPBM *Scarus ghobban Forssk:H, 1775. 30998. Williams 1988. *Scarus globiceps Valenciennes, 1840. Rupiscartes quagga Fowler & Ball. BPBM Scarus oviceps Valenciennes, 1840. BPBM 4766 3420,10436 (T). Fowler and Ball 1925. (T). Myers 1999. Callyodon pectoralis (Va­ Cirripectes varioloms (Valenciennes, 1836). lenciennes) reported by Fowler and Ball BPBM 4925 (T), BPBM 15369 (G&R). 1925 is a synonym of Scarus scaber Valen­ Rupiscartes variolosus (Valenciennes). ciennes, an Indian Ocean species. No Fowler and Ball 1925. specimens were found at BPBM. This was *Entomacrodus marmoratus (Bennett, 1828). probably a misidentification of S. oviceps. BPBM 4933 (T). This specimen is listed in Scarus psittacus Forssk:ll, 1775. BPBM 4749, the BPBM records as having been collected 4753 (T). by the Tanager Expedition at Wake Atoll, Callyodon brunneus (Jenkins). Fowler and but the specimen is missing (A. Suzumoto, Ball 1925. pers. comm.). This is a dubious record be­ Scarus rubroviolaceus Bleeker, 1849. BPBM 4773 cause E. marmoratus is a Hawaiian endemic (T). and needs to be confirmed by additional Callyodon rubroviolaceus (Steindachner). collections. Fowler and Ball 1925. Entomacrodus striatus (Quoy & Gaimard, 1836). BPBM 12060, 14210 (G&R). Myers Family PINGUIPEDIDAE (Sandperches) 1999. *Parapercis schauinslandii (Steindachner, 1900). Istiblennius edentulus (Bloch & Schneider, 1801). BPBM 4944 (T), BPBM 15209 Family CREEDIDAE (Sand Burrowers) (G&R), MCZ 158366. Myers 1999. *Limnichthys nitidus Smith, 1958. MCZ 158368. Salarias edentulus (Schneider). Fowler and Originally identified as L. donaldsoni Ball 1925. Schultz, 1960, which has been shown to be *Plagiotremus tapienosoma (Bleeker, 1857). CAS a synonym of nitidus (Yoshino et al. 1999). 29079. Rhabdoblennius ellipes Jordan & Starks, 1906. Family TRIPTERYGIDAE (Triplefins) Blennius rhabdotrachelus Fowler & Ball. *Enneapterygius nigricauda Fricke, 1997. BPBM BPBM 3419 (T). Fowler and Ball 1925. 4910 (T). This specimen was identified as Family CALLIONYMIDAE (Dragonets) Enneapterygius hemimelas (Kner & Stein­ dachner, 1866) by Fowler and Ball 1925. *Synchiropus laddi Schultz in Schultz et aI., *Helcogramma chica Rosenblatt in Schultz, 1960. BPBM 38302 (G&R). Specimen origi­ 1960. BPBM 37885 (K&L). nally labeled as Synchiropus kiyoae Fricke & Zaiser was reidentified as S. laddi by Family BLENNIIDAE (Blennies) J. Randall. Blenniella gibbifrons (Quoy & Gaimard, 1824). *Callionymidae unidentified species. MCZ Istiblennius gibbifrons (Quoy & Gaimard). 158373. BPBM 15364, 15378 (G&R). Springer and Family GOBIIDAE (Gobies) Williams 1990. Rupiscartes gibbifrons (Quoy & Gaimard). *Amblygobius phalaena (Valenciennes, 1837). Fowler and Ball 1925. BPBM 37878 (K&L). Blenniella paula (Bryan & Herre, 1903). BPBM *Asterropteryx semipunctatus Ruppell, 1830. 4938 (T), BPBM 15368 (G&R). Springer and BPBM 37879 (K&L). Williams 1994. A specimen listed in the Bathygobius fuscus (Ruppell, 1828). BPBM 4866 CAS (150165) as Blenniella periophthalmus (T). Fowler and Ball 1925. (Valenciennes, 1836) is probably B. paula. *Cabillus tongarevae (Fowler, 1927). USNM 76 PACIFIC SCIENCE· January 2004

166748 (G&R 1953). The identity of this Gobiodon rivulatus (Ruppell, 1830). BPBM 4823 specimen needs to be reexamined because (T). Fowler and Ball 1925. its occurrence at Wake is doubtful (J. *Paragobiodon lacunicolus (Kendall & Golds­ Randall, pers. comm.). borough, 1911). BPBM 4834 (T). Originally *Coryphopterus duospilus (Hoese & Reader, identified as Paragobiodon echinocephalus 1985). BPBM 37881 (K&L). (Ruppell) by Fowler and Ball 1925. *Coryphopterus neophytus (Gunther, 1877). *Priolepis kappa Winterbottom & Burridge, BPBM 37880 (K&L). 1993. CAS 60011. Coryphopterus sp. BPBM 4844 (T). This speci­ *Priolepis semidoliatus (Valenciennes, 1837). men collected in salt water was identified USNM 166779 (G&R). as the brackish-water Glossogobius giuris *Trimma sp. Specimen collected in 1997 may (Hamilton) by Fowler and Ball 1925. be an undescribed species. Specimens in *Ctenogobiops aurocingulus (Herre, 1935). our collection. USNM 166766 (G&R). Family MICRODESMIDAE *Ctenogobiops feroculus Lubbock & Pulonin, ( and Wormfishes) 1977. Photographic record. This species was identified from a photograph; how­ * (Jordan & Hubbs, 1925). ever, it is possible that it is another species Photographic record. (J. Randall, pers. comm.). *Ptereleotris microlepis Bleeker, 1856. Ctenogobiops pomastietus Lubbock & Pulonin, Family SIGANIDAE (Rabbitfishes) 1977. Tentatively listed in Myers 1999. Eviota alfelei Jordan & Seale, 1906. BPBM *Siganus argenteus (Quoy & Gaimard, 1825). 15343, 15166 (G&R), BPBM 37882 (K&L). Photographic record. Lachner and Karnella 1980. Family ZANCLIDAE () *Eviota epiphanes Jenkins, 1903. BPBM 4816 (T). This specimen is listed in the BPBM Zanclus cornutus (Linnaeus, 1758). Myers 1999. records as having been collected by the Family ACANTHURIDAE (Surgeonfishes) Tanager Expedition at Wake Atoll, but the specimen is missing (A. Suzumoto, pers. Acanthurus achilles Shaw, 1803. BPBM 4294 comm.). Identified as E. viridis (Waite) by (T). Myers 1999. Fowler and Ball 1925. The identification Hepatus achilles (Shaw). Fowler and Ball of this specimen as E. epiphanes is tentative. 1925. Lachner and Karnella (1980) remarked *Acanthurus blochii Valenciennes in C and V, that this species is restricted to the North 1835. Photographic record. Central Pacific and is found the entire Acanthurus guttatus Forster, 1801. BPBM 4251 length from Midway to Johnston to Kiri­ (T). Myers 1999. mati (Christmas Island); however, Kar­ Hepatus guttatus (Forster). Fowler and Ball nella and Lachner (1981) also noted a 1925. tentative identification of E. epiphanes at Acanthurus leucopareius (Jenkins, 1903). Myers Minami-tori-shima (Marcus) Island. Be­ 1999. cause other Hawaiian endemics are poten­ Acanthurus nigricans (Linnaeus, 1758). tially found at Wake, there is a possibility Hepatus nigricans (Linnaeus). Fowler and that this species is as well. Ball 1925. *Eviota saipanensis Fowler, 1945. BPBM 15187 Acanthurus nigricauda Dunker & Mohr, 1929. (G&R). BPBM 4285 (T). Myers 1999. *Favonigobius sp. Specimen collected by us in Acanthurus nigrofuscus (Forsskal, 1775). Myers 1997 is an undescribed species. 1999. Gnatholepis cauerensis cauerensis (Bleeker, Acanthurus nigroris Valenciennes, 1835. BPBM 1853). 4276 (T). Myers 1999. Originally identi­ Gnatholepis anjerensis (Bleeker). BPBM 4870 fied as Hepatus elongatus (Lacepede) by (T). Fowler and Ball 1925. Fowler and Ball 1925. Checklist of Fishes ofWake Atoll . Lobel and Lobel 77

*Acanthurus nubilus Fowler & Bean, 1929. Katsuwonus pelamis (Linnaeus, 1758). Myers BPBM 37895 (K&L). 1999. Acanthurus thompsoni (Fowler, 1923). Myers Thunnus alalunga (Bonnaterre, 1788). Myers 1999. 1999. Hepatus thompsoni Fowler. Fowler and Ball Thunnus albacares (Bonnaterre, 1788). Myers 1925. 1999. Acanthurus triostegus (Linnaeus, 1758). BPBM Thunnus obesus (Lowe, 1839). Myers 1999. 4239, 4278 (T), BPBM 15255 (G&R). Family ISTIOPHORIDAE (Billfishes) Myers 1999. Hepatus sandvicensis (Streets). Fowler and Istiophorus platypterus (Shaw & Nodder, 1792). Ball 1925. Myers 1999. Ctenochaetus cyanocheilus Randall & Clements, Makaira indica (Cuvier, 1831). Myers 1999. 2001. Makaira mazara (Jordan & Synder, 1901). *Ctenochaetus hawaiiensis Randall, 1955. Pho­ Myers 1999. The specimen mounted in tographic record. the Wake Atoll airport is labeled as Ma­ Ctenochaetus striatus (Quoy & Gaimard, kaira nigricans Lacepede, 1802, the Atlantic 1825). BPBM 4302, 4304 (T), CAS 150134. counterpart ofM. mazara. Randall and Clements 2001. Misidentified Tetrapturus angustirostris Tanaka, 1915. as C. strigosus (Bennett) by Fowler and Ball Myers 1999. 1925. Ctenochaetus strigosus is found only in Tetrapturus audax (Philippi, 1887). Myers Hawai'i and (Randall and 1999. Clements 2001). Family N OMEIDAE (Driftfishes) *Naso brevirostris (Valenciennes, 1835). Pho­ tographic record. Cubiceps pauciradiatus Gunther, 1872. Myers *Naso hexacanthus (Bleeker, 1855). Photo­ 1999. graphic record. This could also be Naso Family BOTHIDAE (Lefteye Flounders) caesius Randall & Bell. Naso hexacanthus is most common, but N caesius is antitropical Bothus mancus (Broussonet, 1782). BPBM 3687 and therefore could also occur at Wake (T). (J. Randall, pers. comm.). Platophrys mancus (Broussonet). Fowler and Naso lituratus (Bloch & Schneider, 1801). Ball 1925. BPBM 4318 (T). Myers 1999. *Bothus pantherinus (Ruppell, 1830). BPBM Acanthurus lituratus (Schneider). Fowler 15189 (G&R). and Ball 1925. Family BALISTIDAE (Triggerfishes) *Naso unicornis (Forsskal, 1775). Video and photographs were taken of this species. *Balistoides viridescens (Bloch & Schneider, *Naso vlamingii (Valenciennes, 1835). Photo­ 1801). graphic record. Melichthys niger (Bloch, 1786). BPBM 5038 Zebrasoma flavescens (Bennett, 1828). BPBM (T). Myers 1999. 4312 (T). Myers 1999. Melichthys buniva (Lacepede). Fowler and *Zebrasoma veliferum (Bloch, 1797). Photo­ Ball 1925. graphic record. Melichthys vidua (Solander in Richardson, 1844). BPBM 5004 (T). Myers 1999. Family SPHYRAENIDAE (Barracudas) Balistes vidua (Richardson). Fowler and Sphyraena barracuda (Walbaum, 1792). Myers Ball 1925. 1999. Rhinecanthus aculeatus (Linnaeus, 1758). BPBM 5022 (T). Myers 1999. Family SCOMBRIDAE (Tunas and Mackerels) Balistipus aculeatus (Linnaeus). Fowler and Acanthocybium solanderi (Cuvier in C and V, Ball 1925. 1831). Myers 1999. *Rhinecanthus rectangulus (Bloch & Schneider, Euthynnus affinis (Cantor, 1849). Myers 1999. 1801). Photographic record. 78 PACIFIC SCIENCE· January 2004

*SuJJlamen bursa (Bloch & Schneider, 1801). TABLE 1 Photographic record. Zoogeography of Wake Atoll Fishes Xanthichthys mento (Jordan & Gilbert, 1882). Myers 1999. Zoogeographic % of Wake Family MONAcANTHIDAE (Filefishes) Distribution Atoll Fishes *Aluterus scriptus (Osbeck, 1765). Photo­ Hawai'i/]apan OJ Indo-Pacific 52.2 graphic record. Indo-pan-Pacific 11.1 *Cantherhines dumerilii (Hollard, 1854). West central Pacific 11.1 Indo-West Pacific 3.2 Family OSTRACIIDAE West Pacific 1.3 (Trunkfishes, Boxfishes) Pacific Plate 5.7 Pan-Pacific 1.0 Ostracion cubicus Linnaeus, 1758. Circumtropical 4.1 Ostracion lentiginosus Bloch & Schneider. Circumglobal 1.3 Fowler and Ball 1925. Ostracion meleagris Shaw, 1796. BPBM 12067 (G&R). the atoll. The Mariana Islands are composed Ostracion sebae Bleeker. Fowler and Ball of and high islands with diverse habitat 1925. types. The contain atolls Ostracion camurum Jenkins. BPBM 5059, with habitats similar to those of Wake but 5064 (T). covering a much larger area and therefore benefiting from increased connectivity be­ Family TETRAoDoNTIDAE (Puffers) tween reef systems. In comparison, Wake is *Arothron hispidus (Linnaeus, 1758). Photo­ an isolated single atoll with relatively few graphic record. habitat types. *Arothron meleagris (Bloch & Schneider, The geomorphology of the atoll is shown 1801). BPBM 5111, 5117 (T), MCZ 158362. in Figures 2 and 3. There are four basic Identified as Tetrodon regani Gunther by aquatic habitat types within Wake Atoll. The Fowler and Ball 1925. first includes a shallow and turbid lagoon with * amboinensis (Bleeker, 1865). scattered patch reefs. About one-fourth of the BPBM 15358 (G&R). lagoon habitat consists offeatureless sand flats Canthigaster janthinoptera (Bleeker, 1855). that are fully exposed at low tide. A second BPBM 37894 (K&L). Allen and Randall habitat type is the intertidal, hard-substrate, 1977. ocean reef flat, also exposed at low tide. The *Canthigaster solandri (Richardson, 1845). third is the reef crest, ofwhich the top is also BPBM 15332 (G&R). exposed at low tide. The fourth habitat type is the outer reef, which drops in depth rapidly, Family DIODONTIDAE (Porcupinefishes) on both the exposed windward and sheltered Diodon hystrix Linnaeus, 1758. Fowler and leeward sides. We did not find sea grass, algal Ball 1925. flats, or mangroves. There are no brackish­ water habitats. The island's topography is DISCUSSION very low, with a maximum height above sea level of only a few meters. The fauna of Wake Atoll is largely composed The lagoon is very murky and shallow of wide-ranging western and Indo-Pacific (maximum depth of about 5 m). The area of fishes, with the notable occurrence of one the lagoon along Wake Island is largely ex­ Hawaiian endemic species (Table 1). The fish posed at low tide and represents about 25% fauna ofWake Atoll is much less diverse than of the total lagoon area. The lagoon has those of the Marshall and Mariana Archipel­ scattered patch reefs on a sandy bottom (Fig­ agos. This is probably due to the isolation ure 4). Water circulation in the lagoon was and low diversity of habitat types found on severely reduced when the causeway between FIGURE 2. Aerial view ofWake Atoll, looking toward the south. The open reefcrest is to the right (west). Peale Island is in the foreground separated from Wake Island by a narrow channel. The northwestern margin of the atoll (foreground) has a wide intertidal reefflat. FIGURE 3. Aerial view ofWake Atoll looking north. The eastern tip ofWake Island is on the right (Peacock Point). This view shows the leeward side of the atoll, which has a very narrow intertidal reefflat that rapidly drops off to deeper depths. FIGURE 4. Top-down aerial view of the Wake Atoll lagoon showing the western reef, which is only emergent at the lowest tides. The lagoon is mainly sand flats with few patch reefs. FIGURE 5. Underwater habitat of the outer reef of Wake Atoll showing the dense coral cover. Checklist of Fishes of Wake Atoll . Lobel and Lobel 83

Wake and Wilkes Islands was built, resulting TABLE 2 in the closure of the small boat harbor inlet Most Speciose Families of Fishes at Wake Atoll into the lagoon. There have been reports of large-scale fish die-offs in the lagoon due to No. of No. of % ofTotal high temperatures when tidal flushing is low Family Genera Species Fish Fauna resulting in low dissolved oxygen (M. Henz, pers. comm.). The outer reef is densely cov­ Labridae 17 38 11.8 Gobiidae 14 21 6.5 ered in coral and slopes rapidly to abyssal Acanthuridae 4 21 6.5 depths (Figure 5). Pomacentridae 6 20 6.2 The total number offishes identified from Serranidae 8 19 5.9 Wake Atoll (n = 321) is similar to that found Chaetodontidae 4 15 4.6 Muraenidae 4 15 4.6 on other isolated central Pacific atolls and Holocentridae 3 13 4.0 islands including Johnston Atoll (n = 301 Apogonidae 3 13 4.0 [Randall et al. 1985, Kosaki et al. 1991]), Scaridae 5 13 4.0 Rapa (n = 268 [Randall et al. 1990; but many Carangidae 8 12 3.7 new records recently have been found but not Blenniidae 7 11 3.4 Mullidae 3 8 2.5 yet reponed (J. Randall, pers. comm.)]), and Balistidae 5 7 2.2 Midway (n = 258 [Randall et al. 1993]). It has Scorpaenidae 3 7 2.2 been suggested that the relatively low number Cirrhitidae 4 6 1.9 of species occurring at these locations could Tetraodontidae 2 6 1.9 be explained by their small geographic size and limited habitat diversity, combined with their isolation or relatively large distance sep­ arating them from other population sources It is interesting that a potential ocean cur­ (Randall 1992, 1998b). This comparison is rent flow pathway seems to exist, at least very broad and the islands/atolls vary greatly periodically, between Hawai'i and Wake in their geomorphology. For example, Rapa is Atoll. A 90-day oscillation in sea level has a high island, whereas Johnston is an atoll been observed intermittently at Wake Atoll with a large lagoon. Even so the relationship based upon two decades of measurements between species diversity and island/atoll iso­ (Mitchum 1995). This oscillation tends to lation and size reveals an interesting and con­ occur about 1.5 yr after an El Nino and ap­ sistent pattern. pears to be caused by ocean eddies generated The fishes that are found most abundantly off the island of Hawai'i that then move at Wake Atoll (Table 2) are typical of those westward and impinge upon Wake Atoll also most abundant on other Pacific Islands (Mitchum 1995). These ocean eddies are (e.g., Kosaki et al. 1991, Randall et al. 1997). known to contain high concentrations of fish The fishes of Wake Atoll can be divided into larvae (Lobel and Robinson 1986). We found the following categories: coastal shore fishes further evidence of the current flow between (303 species, of which six are elasmobranchs) Hawai'i and Wake by observing a small sail­ and offshore pelagic fishes (18 species). boat washed ashore with a Hawai'i State reg­ Wake Atoll occupies a unique central lo­ istration sticker and from reports ofHawaiian cation in the northern Pacific and its fauna fish aggregation devices (FADS) recovered reflects a mixture of species with different from Wake by the U.S. Coast Guard (P. zoogeographic distributions; however, con­ Fusco and M. Timmons, pers. comm.). ventional taxonomic classification by species Based upon this evidence, one would ex­ does not always incorporate regional popula­ pect an overall close faunal affinity between tion variation (potential subspecies) that is Wake and Hawai'i; however, this does not recognized most readily by color pattern dif­ seem to be the case. For example, a common ferences (Randall 1998). Thus future research species, the convict tang, Acanthurus triosteg;us, using DNA markers may reveal more re­ is found throughout the . The gional variation in zoogeography. population of A. triosteg;us at Wake Atoll is FIGURE 6. Regional color pattern variation in Acanthurus triostegus. Note variations in the bar/spot found beneath the pectoral fin. A and B, Wake Atoll; C, Johnston Atoll; D, Kona, Hawai'i; E, Tabuaeran (Fanning) Atoll, Kiri­ bati, Line Islands; F, , ; G, Saipan, . Checklist of Fishes ofWake Atoll . Lobel and Lobel 85

FIGURE 7. Ammolabrus dicrus (Labridae), formerly considered a Hawaiian endemic found only from O'ahu, Hawai'i. It is now found at Wake Atoll. Similar fish have been observed in the Marshall Islands but have not been collected (Randall and Carlson 1997). A, Female; B, male. not the same as the distinctive Hawaiian form the populations in the Mariana and Marshall (which Randall [1956] recognized as a sub­ Islands and another that is typical of a varia­ species, A. triostegus sandvicensis). Acanthurus tion occurring in Phoenix and Line Island triostegus at Wake Atoll exhibits two color specimens (Figure 6; see also Schultz et al. pattern variations: one that is very similar to [1953:625, fig. 90] for illustrations of zoo-

TABLE 3 Zoogeographic Overlap of Wake Atoll Fishes with Those of Nearest Archipelagos

% of % of Wake Species Wake Species Location in Common Datan at Location Data' Reference

Hawaiian Islands 40 129/321 23 129/557 Randall 1996 Johnston Atoll 41 131/321 44 131/301 Randall et al. 1985, Kosaki et al. 1991 Northern Mariana Islands 87 279/321 30 279/933 Myers 1999 Southern Mariana Islands 66 213/321 50 213/427 Myers 1999 Marshall Islands 82 264/321 32 264/817 Myers 1999

n Number of Wake Atoll species found at location divided by me total number of species at Wake. DNumber of Wake Atoll species found at location divided by me total species at location. 86 PACIFIC SCIENCE· January 2004

FIGURE 8. Labroid fishes of special concern for conservation found at Wake Atoll. A, Maori , Cheilinus un­ dulatus (Labridae); B, Bumphead , Bolbometopon muricatum (Scaridae); C, Tan-faced Parrotfish, Chlorurus frontalis (Scaridae); D, Pacific Steephead Parrotfish, Chlorurus microrhinos (Scaridae).

geographic color pattern variations in this Randall & Carlson, 1997 (Figure 7). It prob­ species). ably also occurs in the Marshall Islands as One species that was previously known well (Randall and Carlson 1997). As noted in only from O'ahu, Hawai'i, is now docu­ the species listing, three other Hawaiian en­ mented from Wake Atoll: Ammolabrus dicrus demics have also been reported from Wake

TABLE 4 Pacific Fishes of Special Concern for Conservation Found at Wake Atoll

Family Species IUCN Category Common Name

Labridae Cheilinus zmdulatus VUn Ald+2cd Humphead/Maori Wrasse Scaridae Bolbometopon '/tlZwicatum Giant Bumphead Parrotfish Scaridae Chlorurus frontalis Scaridae Chlorurus microrhinus Serranidae Epinephalus lanceolatus VUA2d Giant Serranidae Epinephalus polyphekadion Shortlist for assessment Camouflage Grouper Serranidae Variola louti Shortlist for assessment Yellow-edged Lyretail

n Vulnerable based on the following meN (2001) criteria: AId, reduction in population size based on actual or potential levels of exploitation; 2cd, an observed, estimated, inferred, or suspected population size reduction of 30% over the last 10 yr or three gen­ erations, whichever is the longer, where the reduction or its causes may not have ceased OR may not be understood OR may not be reversible, based on a decline in area of occupancy, extent of occurrence, and/or quality of habitat and actual or potential levels of ex­ ploitation; Ald, an observed, estimated, inferred, or suspected population size reduction of 30% over the last 10 yr or three generations, whichever is the longer, where the reduction or its causes may not have ceased OR may not be understood OR may not be reversible, based on actual or potential levels of exploitation. FIGURE 9. This underwater scene shows a school of large parrotfish in the background and at least six large Maori Wrasses (Cheilinus undulatus). Such scenes are no longer observed on other Pacific atolls subjected to commercial fishing. 88 PACIFIC SCIENCE· January 2004

Atoll, but these are probably misidentifica­ determined. This emphasizes the need for tions (Eviota epiphanes, Entomacrodus marmo­ continued protection of Wake Atoll's coral ratus, Sebastipistes ballieui). reef ecosystem. The Wake Atoll fish fauna has the most species in common with that of the Mariana ACKNOWLEDGMENTS Islands. A zoogeographic comparison of the Wake Atoll fish fauna with those of the four We are especially grateful for the enormous nearest neighboring archipelagos was made level of assistance provided by John Randall, using our checklist and the species distri­ Helen Randall, and Arnold Suzumoto of the bution data compiled from the literature. BPBM. This paper could not have been com­ This zoogeographic analysis indicates that pleted without their help. We also thankJohn the greatest species overlap occurs with the McCosker, William Eschmeyer, and Jon northern Mariana Islands (87%) and the Fong of the Department of , CAS; Marshall Islands (82%). Although the south­ Karsten Hartell of the MCZ; the late Eugenie ern Mariana Islands have a low percentage of Bohlke and Mark Sabaj of the Department of species in common with Wake (66%), the Ichthyology, ANSP; and Rob Myers for their southern Mariana Islands had a much greater assistance. We are grateful for the logistic proportion of their total fish fauna also found support on Wake Island provided by Paul at Wake than did the other five locations Fusco, Mike Timmons, and Mark Henz. Lo­ examined (Table 3). This pattern is corrobo­ gistic support was also provided by Mark In­ rated by the Acanthurus triosteg;us example. goglia and Jeff Klein of the U.S. Air Force, Another unique aspect of Wake Atoll is Pacific. We thank Mike Cermac for help with the protection of its natural resources, in the fish distribution database. particular fisheries, by the u.s. Depart­ ment of Defense. This is achieved primarily Literature Cited through the exclusion of commercial fish­ ing ventures from Wake and the ban on reef Allen, G. R. 1991. Damselfishes of the world. fish spearfishing. Our surveys discovered four Aquarium Systems, Mentor, . labroid fishes (Figure 8) and three groupers Allen, G. R., and]. E. Randall. 1977. Review on Wake's reefs that are threatened by ex­ of the sharpnose pufferfishes (subfamily ploitation elsewhere in the Pacific (Table 4). Canthigasterinae) of the Indo-Pacific. Rec. In particular, the Maori Wrasse (Cheilinus Aust. Mus. 30 (17): 475-517. undulatus), which is listed on the IUCN Red Bohlke, E. B., and]. E. Randall. 2000. A re­ List (2001) (IUCN Grouper and Wrasse Spe­ view of the moray eels (Anguilliformes: cialist Group [http://www.hku.hk/ecology/ Muraenidae) of the , with GroupersW rasses/iucnsg/doc/species.html]) descriptions of two new species. Proc. as vulnerable to overexploitation, is abundant Acad. Nat. Sci. Phila. 150:203-278. at Wake (Figure 9). The Maori Wrasse is Collette, B. B. 1974. Geographic variation in a prized food fish and is targeted for the the central Pacific halfbeak, Hyporhamphus live reef fish trade (LRFT) in Hong Kong acutus (Gunther). Pac. Sci. 28:111-122. (Donaldson and Sadovy 2001). These fishes Donaldson, T. ]., and y. Sadovy. 2001. of concern are not fished from Wake waters Threatened fishes of the world: Cheilinus and therefore population numbers may be undulatus Ruppell 1835 (Labridae). Envi­ considered pristine; however, fishing of other ron. BioI. Fishes 62:428. species does occur by the residents and visi­ Eschmeyer, W. N., and]. E. Randall. 1975. tors of Wake. For example, very few sharks The scorpaenid fishes of the Hawaiian were observed, which is unusual for a re­ Islands including new species and new mote Pacific atoll. The local workers re­ records (Pisces: Scorpaenidae). Proc. Calif. portedly fish occasionally for sharks as well as Acad. Sci. 40 (11): 265-334. for other reef species, but whether this has Fowler, H. W. 1928. The fishes of . negatively impacted the population was not Mem. Bernice P. Bishop Mus. 10. Checklist of Fishes ofWake Atoll . Lobel and Lobel 89

Fowler, H. W., and S. C. Ball. 1925. Fishes fish genus Acanthurus. Pac. Sci. 10:159­ of , Johnston Island, and Wake 235. Island. Bernice P. Bishop Mus. Bull. 26:1­ ---. 1992. Endemism of fishes in Oce­ 31. ania. UNEP: Coastal resources and sys­ Greenfield, D. W. 2001. Revision of the Apo­ tems of the Pacific Basin: Investigation and gon erythrinus complex (Teleostei: Apogon­ steps toward protective management. idae). Copeia 2001 (2): 459-472. UNEP Regional Seas Reports and Studies WCN. 2001. WCN Red List categories and no. 147:55-67. criteria: Version 3.1. WCN Species Sur­ ---. 1996. Shore fishes of Hawaii. Natu­ vival Commission. IUCN, Gland, Swit­ ral World Press, Vida, . zerland, and Cambridge, UK. ---. 1998a. Revision of the Indo-Pacific Karnella, S. ]., and E. A. Lachner. 1981. squirrelfishes (: Holocentri­ Three new species of the Eviota epiphanes dae: ) of the genus Sargocen­ group having vertical trunk bars (Pisces: tron, with descriptions offour new species. Gobiidae). Proc. BioI. Soc. Wash. 94 (1): Indo-Pac. Fishes 27. 264-275. ---. 1998b. Zoogeography of shore fishes Kerr, L. M., and P. S. Lobel, eds. 2000. De­ of the Indo-Pacific region. Zool. Stud. 37 partment of Defense coral reef protection (4): 227-268. implementation plan. U.S. Coral Reef ---. 1999. Revision of the Indo-Pacific Task Force Report. Government Printing labrid fishes of the genus Coris, with de­ Office, , D.C. scriptions of five new species. Indo-Pac. Kosaki, R. K, R. L. Pyle, J. E. Randall, and Fishes 29. D. K Irons. 1991. New records of fishes Randall, J. E., and B. A. Carlson. 1997. Am­ from Johnston Atoll, with notes on bio­ molabrus dicrus, a new genus and species of geography. Pac. Sci. 45:186-203. labrid fish from the Hawaiian Islands. Pac. Lachner, E. A., and S. J. Karnella. 1980. Sci. 51:29-35. Fishes of the Indo-Pacific genus Eviota Randall, J. E., and K D. Clements. 2001. with descriptions of eight new species Second revision of the surgeonfish genus (Teleostei: Gobiidae). Smithson. Contrib. Ctenochaetus (Peciformes: Acanthuridae), Zool. 315. with descriptions of two new species. Lobel, P. S., and A. R. Robinson. 1986. Indo-Pac. Fishes 32. Transport and entrapment offish larvae by Randall, J. E., and D. W. Greenfield. 1996. ocean mesoscale eddies and currents in Revision of the Indo-Pacific holocentrid Hawaiian waters. Deep-Sea Res. 33 (4): fishes of the genus Myripristis, with de­ 483-500. scriptions of three new species. Indo-Pac. McCosker, J. E., and R. H. Rosenblatt. 1993. Fishes 25. A revision of the snake eel genus Myrich­ Randall, J. E., and P. C. Heemstra. 1991. thys (Anguilliformes: Ophichthidae) with Revision of Indo-Pacific groupers (Perci­ the description of a new eastern Pacific formes: Serranidae: Epinephelinae), with species. Proc. Calif. Acad. Sci. 48 (8): 153­ descriptions of five new species. Indo-Pac. 169. Fishes 20. McCosker, J. E., K Hatooka, K Sasaki, and Randall, J. E., and P. S. Lobel. Xyrichthys J. T. Moyer. 1984. Japanese morays of the halsteadi, a new labrid fish from the central genus Uropterygius. Jpn. J. Ichthol. 31 (3): and westernPacific. Bull. Mar. Sci. (in press). 261-267. Randall, J. E., and R. F. Myers. 2002. Paru­ Mitchum, G. T. 1995. The source of 90-day peneus insularis, a new central Pacific spe­ oscillations at Wake Island. J. Geophys. cies of goatfish (: Mullidae) of Res. 100 (2): 2459-2475. the P. trifasciatus complex. Zool. Stud. 41 Myers, R. F. 1999. Micronesian reef fishes. (4): 431-440. 3rd ed. Coral Graphics, Guam. Randall, J. E., and H. A. Randall. 2001. Re­ Randall, J. E. 1956. A revision of the surgeon view of the fishes of the genus Kuhlia 90 PACIFIC SCIENCE· January 2004

(Perciformes: Kuhliidae) of the central and J. L. Earle. 1997. Annotated checklist Pacific. Pac. Sci. 55:227-256. of the inshore fishes of the Ogasawara Is­ Randall, J. K, P. S. Lobel, and E. H. Chave. lands. Natl. Sci. Mus. Monogr. 11. 1985. Annotated checklist of the fishes of Schultz, L. P., K S. Herald, K A. Lachner, Johnston Island. Pac. Sci. 39:24-80. A. D. Welander, and L. P. Woods. 1953. Randall, J. E., M. L. Bauchot, and M. De­ Fishes of the Marshall and Marianas Is­ soutter. 1987. Heliases ternatensis Bleeker, lands. U.S. Natl. Mus. Bull. 202 (1). 1856 (currently Chromis ternatensis; Os­ Springer, V. G., and J. T. Williams. 1990. teichthys, Perciformes): Proposed conser­ Widely distributed Pacific Plate endemics vation, and adoption of the name Chromis and lowered sea-level. Bull. Mar. Sci. 47 viridis (Cuvier, 1830) for the fish com­ (3): 631-640. monly called C. caerulea (Cuvier, 1830). --. 1994. The Indo-West Pacific blen­ Case 2516. Bull. Zool. Nomencl. 44 (4): niid fish genus Istiblennius reappraised: A 248-250. revision of Istiblennius, Blenniella, and Pa­ Randall, J. K, C. L. Smith, and M. N. Fein­ ralticus, new genus. Smithson. Contrib. berg. 1990. Report on fish collections from Zool. 565. Rapa, French . Am. Mus. Novit. Williams, J. T 1988. Revision and phyloge­ 2966. netic relationships of the blenniid fish ge­ Randall, J. E., J. L. Earle, R. L. Pyle, J. D. nus Cirripectes. Indo-Pac. Fishes. 17. Parrish, and T Hayes. 1993. Annotated Yoshino, T, T Kon, and S. Okabe. 1999. checklist of the fishes of Midway Atoll, Review of the genus Limnichthys (Perci­ Northwestern Hawaiian Islands. Pac. Sci. formes: Creedidae) from Japan, with de­ 47:356-400. scription of a new species. Ichthyol. Res. Randall, J. E., H. Ida, K. Kato, R. L. Pyle, 46 (1): 73-83.