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Synopsis and Phylogenetic Analysis of the Pearlfish Subfamily Carapinae (Pisces: Carapidae)

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Synopsis and Phylogenetic Analysis of the Pearlfish Subfamily Carapinae (Pisces: Carapidae) BULLETIN OF MARINE SCIENCE. 34(3): 386-397. 1984 SYNOPSIS AND PHYLOGENETIC ANALYSIS OF THE PEARLFISH SUBFAMILY CARAPINAE (PISCES: CARAPIDAE) Jeffrey T. Williams ABSTRACT Representatives of the pearlfish subfamily Carapinae are known from every ocean of the world except the Arctic Ocean. Four carapine genera are recognized: Echiodon, Onuxodon, Carapus and Encheliophis. The last contains the subgenera Encheliophis and Jordanicus. Members of the Carapinae share the following apomorphies: ossified interarcual element, reduced ventral process ofposttemporal bone and loss of pleural ribs. A phylogenetic analysis indicates that Echiodon and Onuxodon are sister groups, which together form the sister group of Carapus and Encheliophis. The subfamily and its included genera are diagnosed, and a key to all carapid genera is provided. The family Carapidae, which includes the subfamilies Pyramodontinae and Carapinae, contains an ecologically diverse group of marine fishes. They range from tropical shallow water parasitic invertebrate symbionts to supposedly free- living fishes found on the continental shelf and slope of every major ocean except the Arctic Ocean (Arnold, 1956; Markle et aI., 1983; Trott, 1970). The last sys- tematic review of the subfamily Carapinae was that of Arnold (1956). Since his study, additional material has been collected and many taxonomic problems have surfaced. A cladistic analysis of the group was undertaken to understand and define the relationships within the Carapinae. The numerous taxonomic problems at the species level are being examined by Douglas F. Markle and John E. Olney and will not be treated here. This paper diagnoses the subfamily and its genera, and proposes a hypothesis of the phylogenetic relationships among the carapine genera based on adult osteological and morphological characters. METHODS Extensive synonymies for genera and species are in Arnold (1956). Gill rakers refer to those on the first arch (sensu Cohen and Nielsen, 1978). Pelvic bones and cartilage refer to elements located inside the body immediately behind and between the ventral junction of the cleithra. Names of bones follow Courtenay and McKittrick (1970), Gosline (1960) and Tyler (1970). Osteological character states were based on specimens cleared and stained following Dingerkus' and Uhler's method (1977) and radiographs. Transverse processes are paired bony structures extending ventrolaterally from the precaudal centra (those centra without a distinct hemal spine). The transverse processes may articulate with their respective centra (usually those of centra 1-3 to 5), or be fused with the centra (Markle et aI., 1983). Pleural ribs are the bony elements (usually small in ophidiiforms) that articulate with the ventral edge of the transverse processes. Recurved cardiform teeth are those teeth that are bent in the middle in a posterior direction at a 90° angle to form an L-shaped tooth. Each tooth is loosely hinged to the premaxillary bone, directed laterally from the premaxillary, and concealed beneath the upper lip. Small fleshy flaps above the upper lip are approximately 0.2 mm lo'ng and extend into the groove between the premaxillary and maxillary bones, and, if present, are situated immediately anterior to the ventrally directed anteriormost pore of the supraorbital series (Markle et aI., 1983). The following institutional abbreviations are used; ANSP-Academy of Natural Sciences of Phila- delphia; BM(NH)-British Museum (Natural History), London; BPBM-Bernice P. Bishop Museum, Honolulu; CAS -California Academy of Sciences, San Francisco; FAKU - Fisheries Research Station, Kyoto University, Japan; ISH-Institut fur Seefischerei, Zoological Museum, University of Hamburg, 386 WILLIAMS: SYNOPSIS OF CARAPINAE 387 West Germany; SIO-Scripps Institution of Oceanography, La Jolla; UF -Florida State Museum, University of Florida, Gainesville; USAIC- University of South Alabama Ichthyological Collection, Mobile; USNM-National Museum of Natural History, Smithsonian Institution, Washington, D.C. Material Examined.-Pyramodon ventralis: off Isla San Felix, Chile: USNM 231353 (I). Snyderidia bothrops: off French Guiana: USNM 214035 (I cleared & stained). Carapus bermudensis: Dry Tortugas, Florida: UF 11825 (I cleared & stained). Carapus mourlani: Oahu, Hawaiian Islands: BPBM 17341 (I), 17821 (I). Carapus parvipinnis: Ifaluk Atoll, Caroline Islands: CAS 48978 (12 + 2 cleared & stained). Carapus variegalUs: Israel, Gulf of Aqaba: ANSP 72147 (I para type cleared & stained). Echiodon owasianus: Owase, Japan: FAKU 34517-34519 (3). Echiodon exsi/ium: Baja California Sur: SIO 65-292 (8 + I cleared & stained). Echiodon dawsoni: eastern Gulf of Mexico: USAIC 6279 (I paratype cleared & stained), UF 30886 (I para type cleared & stained). Echiodon drummondii: off northern Scotland: BM(NH) 1967.5.4: 3-5 (2 + I cleared & stained). Echiodon cryomargarites: off Argentina: ISH 1818/68 (I + I cleared & stained). Encheliophis (E.) jordani: Buena Vista, Baja California Sur: UF 34264 (I cleared & stained). E ncheliophis (E.) vermicularis: Belau Island, Caroline Islands: BPBM 11379 (I). Encheliophis (Jordanicus) gracilis: Oahu, Hawaiian Islands: BPBM 4997 (4), 4998 (I), 4999 (I); Fanning Island, Line Islands: BPBM 25714 (2); Tutuila, Samoa Islands: BPBM 5000 (I); Guam, Marianas Islands: UF 34265 (I + I cleared & stained); Ambon Island, Moluccas: BPBM 19426 (I); Seychelles Islands: BM(NH) 1981.2.24.19 (I). Onuxodon margaritiferae: Fatu Hiva, Marquesas Islands: BPBM 11674 (I); Moorea, Society Islands: CAS 13524 (I cleared & stained); Christmas Island, Line Islands: BPBM 5001 (I); Maldive Islands: CAS 48975 (2), 48976 (I cleared & stained). Onuxodon parvibrachium: Fiji: ANSP 91016 (I paratype); Seychelles Islands: ANSP 109210 (I cleared & stained). Brotuia barbata: off Aransas, Texas: UF 34271 (I cleared & stained). Otophidium omostigmum: northeastern Gulf of Mexico: UF 34270 (2 cleared & stained). Ophidion holbrooki: Cedar Keys, Florida: UF 948 (3 + I cleared & stained). Lepophidium jeannae: northeastern Gulf of Mexico: UF 26817 (3 + I cleared & stained). Parophidion schmidti: Providencia Island Group, Colombia: UF 25086 (7 + I cleared & stained). Ophisternon aenigmaticum: Mexico: UF 16209 (I I). KEY TO CARAPID GENERA la. Pectoral-fin rays 24-30, precaudal vertebrae 15 or fewer 2 lb. Pectoral. fin rays 23 or fewer, precaudal vertebrae 17-35 3 2a. Pelvic fins present pyramodon 2b. Pelvic fins absent Snyderidia 3a. Large recurved canines isolated at tips of both jaws; no strongly recurved cardiform teeth hidden under upper lip . 4 3b. No large recurved canines isolated at tips of both jaws; row of strongly recurved cardiform teeth hidden under upper lip 5 4a. Median rocker bone present at anterior end of swim bladder (found by inserting sharp probe through tissues of posterior part of gill cavity); body width about 3 in depth; no small fleshy tabs above upper lip at snout tip Onuxodon 4b. No median rocker bone at anterior end of swim bladder; body width about 2 in depth; 2-3 small fleshy tabs above upper lip at snout tip Echiodon 5a. Posterior margin of maxillary free Carapus 5b. Maxillary adnate .....................................................................•...............................................................................6 6a. Pectoral fin not apparent externally Encheliophis (Encheliophis) 6b. Pectoral fin apparent externally Encheliophis (Jordanicus) Subfamily Carapinae Jordan and Fowler, 1902 Diagnosis. - lnterarcual element ossified; ventral process of posttemporal bone reduced or absent (Fig. 1); no pleural ribs; pectoral-fin rays 23 or fewer; precaudal vertebrae 17-35; pseudobranchial filaments 2-3. Comments. - Travers (1981) found an ossified interarcual element in the Carapi- dae (sensu Arnold, 1956) and in the Synbranchidae, but it was either cartilaginous 388 BULLETIN OF MARINE SCIENCE, VOL. 34, NO.3, 1984 A C E A G E ~ ~ U ~ ~ 0 F B 0 F B~ D ~ Figure 1. (Left) Posttemporal bone of carapine genera. Left lateral view, front of fish is to the left. A, Carapus berrnudensis(UF 11825); B, Carapus parvipinnis (CAS 48977); C, Enche/iophis (E.)jordani (UF 34264); D, Enche/iophis (Jordanicus) gracilis (UF 34265); E, Echiodon cryornargarites (ISH 1818/ 68); F, Onuxodon rnargaritiferae (CAS 13524) (note reduction or absence of ventral strut). Figure 2. (Right) Shape of supracleithrum of carapine genera. Left lateral view, front of fish is to the left. A, Carapus berrnudensis (UF 11825); B, Carapus parvipinnis (CAS 48977); C, Enche/iophis (E.) jordani (UF 34264); D, Enche/iophis (Jordanicus) graci/is (UF 34265); E, Echiodon cryornargarites (ISH 1818/68); F, Onuxodon rnargaritiferae (CAS 13524). or absent in all other groups he examined. He concluded that the ossified condition is more highly derived. This element is cartilaginous in pyramodontines. The inclusion of the Pyramodontinae in the Carapidae (Williams, 1983) is based primarily on the sharing ofa unique planktonic vexillifer larval stage (Markle and Olney, 1981; Olney and Markle, 1979) during ontogeny. As the synbranchids are most closely related to the mastacembeloids (Robert A. Travers, pers. comm.), the ossified interarcual element is considered as being independently derived in carapines and synbranchids. Thus a sister-group relationship between carapines and pyramodontines is supported by the shared vexillifer larval stage. Carapus Rafinesque, 1810 Carapus Rafinesque, 1810 (type species based on Opinion 42 of the International Commission on Zoological Nomenclature Gyrnnolus acus
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