Review of the Fishes of the Genus Kuhlia (Perciformes: Kuhliidae) of the Central Paci®c1 John E. Randall and Helen A. Randall2 Abstract: Ten species of ®shes of the genus Kuhlia are recognized from Palau to Hawai`i in the North Paci®c and from Fiji to Easter Island in the South Paci®c: K. malo (Valenciennes) from fresh water in the Society Islands; K. marginata (Cuvier) from fresh water in the western Paci®c, east to Kosrae, Caroline Islands, and Fiji; K. mugil (Forster) (K. taeniura is a synonym) from most of the Indo-Paci®c (not the Hawaiian Islands) and the tropical eastern Paci®c; K. munda (De Vis) from fresh and brackish water in Fiji, Vanuatu, New Cale- donia, and Queensland (K. proxima Kendall & Goldsborough and K. bilunulata Herre are synonyms); K. nutabunda Kendall & Radcliffe from Easter Island; K. petiti Schultz from the Phoenix Islands, Malden Island, and the Marquesas Islands (Dules taeniurus marquesensis Fowler is a synonym); K. sandvicensis (Steindachner) from the Hawaiian Islands and other islands of the central Paci®c; K. rupestris (LaceÂpeÁde) from fresh water from East Africa to Samoa (K. caerulescens Regan from the Solomon Islands is a new synonym); K. salelea Schultz from fresh water in the Samoa Islands; and K. xenura ( Jordan & Gilbert) from the Hawaiian Islands, with a mistaken type locality of El Salvador, Central America. The name K. sandvicensis has long been used for the common endemic species in the Hawaiian Islands; however, the original description leaves little doubt that it should apply to the species widely distributed in the central Paci®c and only recently discovered in Hawai`i; it has usually been misidenti®ed as K. marginata. The endemic Hawaiian species therefore takes the only available name, K. xenura ( Jordan & Gilbert). Kuhlia sandvicensis differs from K. xenura in having a smaller eye (3.0±3.45 in head length, compared with 2.55±2.95 for K. xenura), straight dorsal pro®le of the head of adults (concave in xenura); usually 14 pectoral-®n rays (usually 15 in xenura), usually 50 lateral-line scales (usually 49 in xenura), gill rakers 38±43 (35±39 for xenura), and a dark reticular pattern dorsally on the head in life. The ®shes of the family Kuhliidae are pally on planktonic crustaceans. These ®shes moderately deep-bodied and compressed, are generally silvery, often with dark mark- with two opercular spines, a deeply notched ings on the caudal ®n, the basis for the com- dorsal ®n of 10 spines and 9±13 rays, and a mon name ¯agtails. The Hawaiian name scaly sheath at the base of the dorsal and anal aÅholehole is also sometimes used beyond ®ns. They have large eyes and are primarily Hawai`i. The family consists of a single ge- nocturnal, at least as adults, feeding princi- nus, Kuhlia, the species of which occur in tropical and subtropical waters of the Indo- Paci®c region, and one is also found in the 1 Manuscript accepted 12 November 2000. tropical eastern Paci®c. Only one species ex- 2 Bishop Museum, 1525 Bernice Street, Honolulu, ceeds 320 mm in total length. Some species Hawai`i 96817-2704. occur mainly in fresh water, whereas others are primarily marine. The latter tend to form schools by day and are typically found in- Paci®c Science (2001), vol. 55, no. 3:227±256 : 2001 by University of Hawai`i Press shore; the young may be common in tide All rights reserved pools. At least some of the marine species are 227 228 PACIFIC SCIENCE . July 2001 able to live in fresh water, and the freshwater we cannot equate this description with any species often occur in brackish environments. species of Kuhlia we know today. The strong In total 40 nominal species and subspecies black markings of the caudal ®n suggest that of Kuhlia have been described (Eschmeyer it may have been one of the freshwater spe- 1998:2343), 18 of these in the genus Dules, cies of the genus. Apogon aroubiensis is another now determined as a genus of Serranidae. species described and illustrated in Hombron Kuhlia was revised by Regan (1913), who and Jacquinot that is regarded as unidenti®- recognized 12 species, three of which he de- able (Randall and Lachner 1986). scribed as new. Six species and subspecies of We recognize 10 species of the genus the genus have been described since 1913. Kuhlia from the islands of Oceania. Of these, A second species of Kuhlia was recently only K. mugil and K. rupestris range to the discovered in the Hawaiian Islands. It is the western Indian Ocean. We provide here a key ®sh currently identi®ed as Kuhlia marginata to the species of Kuhlia from the islands of (Cuvier), described from Java and presumed Oceania, illustrations, and a diagnosis of each. to be wide-ranging throughout much of Not included in our study are the nominal K. Oceania, including Johnston Island. We ex- boninensis (Fowler) from the western Paci®c, amined the holotype of Dules marginatus in K. caudovittata (LaceÂpeÁde) from Madagascar the MuseÂum National d'Histoire Naturelle in and Mauritius, K. sauvagii Regan from Mada- Paris (mnhn 9002, 93 mm SL, not the ``syn- gascar, and K. splendens Regan from fresh types'' listed by Eschmeyer [1998:1019]; water at Rodriguez and Mauritius. Cuvier described the species from a single specimen). The holotype has 13 pectoral rays, materials and methods 41 lateral-line scales, and 7 17 gill rakers. None of these counts matches those of the Specimens of Kuhlia have been examined at ®sh of Paci®c islands now identi®ed as Kuhlia or obtained on loan from the Australian marginata. We ®rst suspected that this new Museum, Sydney (ams); Academy of Natural record of Kuhlia for Hawai`i represented an Sciences of Philadelphia (ansp); the Natural undescribed species, in spite of its broad dis- History Museum, London (bmnh); Bernice P. tribution. We referred to the original de- Bishop Museum, Honolulu (bpbm); California scription of Kuhlia sandvicensis (Steindachner, Academy of Sciences, San Francisco (cas); 1876) to make sure it applied to the common Hebrew University, Jerusalem (huj); Natural endemic Hawaiian species that has long held History Museum of Los Angeles County this name. To our dismay, we found that it (lacm); MuseÂum National d'Histoire Na- ®ts the other species. The only available turelle, Paris (mnhn); Royal Ontario Mu- name for the endemic Hawaiian species of seum, Toronto (rom); and the National Kuhlia is K. xenura ( Jordan & Gilbert, 1882), Museum of Natural History, Washington, originally named Xenichthys xenurus and mis- D.C. (usnm). takenly placed in the Haemulidae. The ques- Lengths given for specimens are standard tioned type locality of El Salvador is clearly in length (SL), the straight-line distance from error. the median anterior point of upper lip to the Hombron and Jacquinot (1853) reported base of the caudal ®n (posterior end of the on ®shes collected during Dumont d'Urville's hypural plate). Head length is measured from voyage to the South Pole by the corvettes the same anterior point to the most posterior ``Astrolabe'' and ``ZeÂleÂe,'' 1837±1840. Among end of the opercular membrane, and snout the species brie¯y described and without length from the same point to the ¯eshy edge meristic data was Dules marginata Cuvier, of the orbit. Body depth is the greatest depth illustrated as ®g. 3 on Plate 3. Boulenger measured to the base of the dorsal spines; (1895:38) redescribed this ®sh as Dules urvillii body width is the greatest width just posterior and gave the locality as ``Probably some Pa- to the gill opening. Orbit diameter is the ci®c Island.'' No type specimen is extant, and greatest ¯eshy diameter. The bony inter- Review of the Kuhlia of the Central Paci®c . Randall and Randall 229 orbital width is measured over the center of Boulengerina Fowler, 1907:512 (type species, the eye. Caudal-peduncle depth is the least Dules mato Lesson, 1831; proposed as a depth, and caudal-peduncle length is mea- subgenus; preoccupied by Boulengerina sured horizontally from the rear base of the Dollo, 1886, in Reptilia). anal ®n to the caudal-®n base. Lengths of ®n Safole Jordan, 1912:655 (replacement name spines and soft rays are taken from the tips to for Boulengerina Fowler). where they emerge from the basal scaly sheath (these lengths may be taken more ac- diagnosis: Dorsal-®n rays X,9±13, the curately than ones to the extreme base of the ®n deeply notched; anal-®n rays III,10±13; spines and rays). Caudal concavity is the hor- pectoral-®n rays 13±15; pelvic rays I,5; lateral izontal distance between verticals at the tips line complete, the pored scales 39±53; gill of the shortest and longest caudal rays. rakers long and slender, 7±13 16±30; Pectoral-ray counts include the short un- branchiostegal rays 6; gill membranes not branched upper ray (but not the still smaller joined across isthmus; vertebrae 25; body splintlike ray that is closely joined to it). moderately deep, the depth 2.55±3.1 in SL, Lateral-line scales are counted from the and compressed; head moderately pointed, upper end of the gill opening to the base of its length 3.0±3.6 in SL; mouth protractile, the caudal ®n (four to seven pored scales on oblique with lower jaw projecting, and mod- the base of the caudal ®n are not counted, but erately large, the maxilla extending to below a pored scale overlapping the end of the eye; supramaxilla not present; teeth villiform hypural plate is included). Gill-raker counts in jaws, on vomer, palatines, entopterygoids, include rudiments; the upper-limb count is and ectopterygoids; opercle with two spines; given ®rst, and the raker at the angle is con- rounded corner and lower edge of preopercle tained in the lower-limb count.