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Osteology and Relationships of Pseudotrichonotus Altivelis (Teleostei: Aulopifornies: Pseudotrichonotidae)

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Osteology and Relationships of Pseudotrichonotus Altivelis (Teleostei: Aulopifornies: Pseudotrichonotidae) Osteology and relationships of Pseudotrichonotus altivelis (Teleostei: Aulopifornies: Pseudotrichonotidae) G. David Johnson,1 Carole C. Baldwin,1 Muneo Okiyama2 & Yoshiaki Tominaga3 1 Division of Fishes, National Museum of Natural History, MRC 159, Smithsonian Institution, WofAmgfon, DC. 20560, [ASA 2Ocean Research Institute, University of Tokyo, 1-15-1 Minamidai, Nakano-Ku, Tokyo 164, Japan ^The University Museum, University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113, Japan (deceased) (Received 17 January 1995; in revised form 8 November 1995; accepted 9 November 1995) Johnson, G. D., C. C. Baldwin, M. Okiyama and Y. Tominaga. 1996. Osteology Ichthyological and relationships of Pseudotrichonotus altivelis (Teleostei; Aulopiformes: Research Pseudotrichonotidae).Ichthyol. Res., 43: 17-45. © The Ichthyological Society of Japan 1996 Abstract The osteology of the rare Japanese fish Pseudotrichonotus altivelis is described based on several specimens collected off the Izu Peninsula. Rela- tionships of Pseudotrichonotus are discussed based on osteological comparisons with other neoteleosts. The placement of Pseudotrichonotus among iniomous fishes has been questioned because of its lower numbers of caudal-fin, pelvic-fin, and branchiostegal rays. Our investigation supports an iniomous affinity for Pseudotrichonotus, specifically as a member of the Aulopiformes. Within that group, Pseudotrichonotus belongs in a new suborder diagnosed herein, the Syn- odontoidei, which also includes the Aulopidae (Aulopus), Synodontidae (Syn- odus and Trachinocephalus), and Harpadontidae (Harpadon and Saurida). A synodontoid affinity for Aulopus has never been suggested, but numerous osteo- logical features support the monophyly of this clade. Synodontoids have a pecu- liar proximal segmentation of most principal caudal-fin rays, expanded neural and haemal spines on posterior vertebrae, cartilage extending along the ventral margin of the anterior ceratohyal, ventral displacement of the first one to three epineurals, supraneurals with large laminar expansions and six or more bran- chiostegals on the posterior ceratohyal. They lack median caudal cartilages. Among synodontoids, Pseudotrichonotus is the sister group of the Synodontidae plus Harpadontidae, with which it shares paired peritoneal pigment spots, an abrupt transition between the epipleurals in and beneath the horizontal septum, and absence of the fourth pharyngobranchial toothplate. Our study does not sup- port a previously proposed relationship between Bathysaurus and synodontids. Key words. •Pseudotrichonotus; Aulopiformes; lizardfishes; osteology; phy- logeny. Tyseudotrichonotus altivelis Yoshino & Araga bears a superficial resemblance to the perciform MT (Fig. 1) is a small bottom-dwelling fish genus Trichonotus and shares with it the habit of known from one locality off Izu Oceanic Park, diving rapidly into the sand when threatened. Izu Peninsula, Japan, where it occurs on sand Yoshino and Araga (in Masuda et al., 1975) de- bottom at 30 to 50 meters in an area of strong scribed P. altivelis in a monotypic family, tidal currents. The status of a second species, Pseudotrichonotidae, which they placed among from the Indian Ocean, P. xanthotaenia Parin, the Myctophiformes (s.l.), stating that the rea- 1992, remains uncertain (Parin, pers. comm.). sons for this alignment would be forthcoming. As the generic name implies, Pseudotrichonotus R. K. Johnson (1982) noted that several charac- 18 G. D. Johnsonet al. # Fig. 1. Pseudotrichonotusaltivelis, USNM 280366, 81.5 mm SL. ters reported in the original description (specifi- clades are (1) other aulopiforms and (2) cteno- cally, low numbers of caudal-fin rays, pelvic-fin squamates. rays and branchiostegal rays) strongly suggested The description of the osteology of Pseudotri- that the authors' alignment of this species with chonotus altivelis herein is based on examination myctophiforms was incorrect. Masuda et al. of three cleared and stained specimens: USNM (1984) retained Pseudotrichonotus in the Myc- 280366 (1, 80.0 mm SL); ZUMT 55678 (1, tophiformes without discussion. The purpose of 58.0 mm SL); and ZUMT 59882 (1, 72.0 mm this paper is to describe the osteology and cer- SL). Comparative, cleared and stained aulopi- tain other aspects of the anatomy of Pseudotri- form and outgroup material examined in this chonotus altivelis and, based on this information, study is listed below. Institutional abbreviations to formulate a hypothesis about its phylogenetic follow Leviton et al. (1985). Ahliesaurus berryi: relationships. USNM 240505 (1 specimen). Alepisaurus sp.: MCZ 60345 (1). Anotopterus pharao: USNM 140825 (1); USNM 221035 (1). Afzofopfgrws sp.: Materials and Methods SIO 62-775 (1). Aulopus jilamentosus: USNM 292105 (1). Aulopus japonicus: AMNH Terminology of the laterosensory system of 28635SW (1). Aulopus sp.: AMNH 28635 (1). the head follows Weitzman (1962). Terminology Bathymicrops regis: BMNH 1989.7.25.56.61 (2). of the bony condyles and ligaments of the upper Bathypterois pectinatus: FMNH 88982 (1). Ba- jaw follows Stiassny (1986). Other osteological thysaurus mollis: VIMS 6107 (1). Bathysaurop- terms are those in general use. Abbreviations sis gigas: AMS I. 22822001 (1). Benthalbella used in illustrations of osteological features are dentata: SIO 63-379 (I). Chlorophthalmusagas- given in Table 1. Illustrations depict left side sizi: AMNH 40829SW (I); USNM 159385 (1). of specimen unless otherwise indicated. Char- Coccorella atlanticum: USNM 235199 (1). acters were polarized following the outgroup Diplophos taenia: MCZ 55469 (1); USNM comparison algorithm of Maddison et al. (1984). 206614 (1). Evermannella indica: UH. 71-3-9 Outgroups for the Aulopiformes are (1) Cteno- (1). Gigantura vorax: AMNH 55345SW (1). squarnates (myctophiforms+acanthomorphs) and Harpadon nehereus: AMNH 17563 (1); FMNH (2) stomiiforms (Johnson, 1992). Outgroups for 179018 (2). Harpadon squamosus: FMNH polarization of characters within aulopiform 80823 (3). Ipnops agassizi: USNM 54618 (1, Osteology and Relationships cf Pseudotrichonotus 19 gill arches only). Lestidiops sp.: USNM 307290 Lestidium sp.: USNM uncat. (1). Macropara- (1). Lestidium atlanticum: USNM 201183 (1). lepis affine: USNM 302410 (1); 201184 (1). Table 1. Abbreviations for Figures Aa Anguloarticular MMP Maxillary Median Process AC Anterior Ceratohyal Mx Maxilla AHF Anterior Hyomandibular Fossa NS Neural Spine Ba Basipterygium O Opercle BBn nth Basibranchial P Palatine BH Basihyal Pa Parietal BHTP Basihyal Tooth Plate Para Parasphenoid BL Baudelot's Ligament PBn Pharyngobranchial Bo Basioccipital PC Posterior Ceratohyal Br Branchiostegal PeFR Pelvic-fin Ray CBn Ceratobranchial PeR Pelvic-fin Radial CC Cranial Condyle of Maxilla PFR Pectoral-fin Ray Cl Cleithrum Ph Parhypural Co Coracoid PIR Pleural Rib D Dentary Pmx Premaxilla DH Dorsal Hypohyal Po Preopercle DHe Dorsal Hemitrich PP Posterior Process DP Dorsal Pterygiophore PR Pectoral-fin Radial DPcl Dorsal Postcleithrum Princ Principal Caudal Ray E Epural Pro Prootic EBn Epibranchial Proc Procurrent Caudal Ray EC Ethmoid Cartilage Pt Posttemporal Ecp Ectopterygoid Pte Pterotic En Epineural Pts Pterosphenoid Enp Endopterygoid PUn Preural Vertebra Ep Epipleural Q Quadrate EP Ethmoid Process Ra Retroarticular Epo Epioccipital S Sphenotic Es Extrascapula sc Supracleithrum Ex Exoccipital SDR Supernumerary Dorsal Ray F Frontal Smx Supramaxilla H. Hypural Sn Supraneural HBn Hypobranchial So Subopercle HS Haemal Spine Sob Supraorbital Hy Hyomandibula Soc Supraoccipital Ic Intercalar Sph Sphenotic In Interhyal Sy Symplectic IE Interoperculomandibular TP Trisegmental Pterygiophore Ligament u„ Ural Vertebra Io Interopercle UNn Uroneural Ion Infraorbital UP5 5 th Upper Pharyngeal Toothplate La Lacrimal V Vomer LE Lateral Ethmoid vc Vertebral Centrum LO Lamina Orbitonasalis VH Ventral Hypohyal Me Mesethmoid VHe Ventral Hemitrich Mep Metapterygoid VPcl Ventral Postcleithrum 20 G. D. Johnsonet al. Macroparalepis sp.: FMNH 49988 (1). Mycto- cations in the ethmoid region, the lateral eth- phum obtusirostris: AMNH 29140SW (1). Neo- moids, provides the anterior structure of the scopelus macrolepidotus: USNM 188056 (2); orbit. In dorsal view, the lateral ethmoids are v- 317160 (1). Notolepis rissoi rissoi: USNM shaped bones: a medial branch flanks (and is 283485 (1). Notoscopelus resplendens: AMNH partially covered by) the anterior frontal and rep- 25928SW (1). Odontostomops normalops: resents the only lateral ossification of the neuro- USNM 235029 (1). Paralepis brevirostris: cranium anterior to the orbit; a lateral branch USNM 196109 (1). Paralepis coregonoides: projects outward and forms the anterior margin USNM 290253 (1). Parasudis truculentus: of the orbit. The supraorbital bone (see "Circum- FMNH 67150 (1); USNM 159407 (1); 159850 orbital Bones" below) lies above the posterodor- (1). Saurida brasiliensis: USNM 185852 (2); sal corner of the lateral ethmoid. 187994 (3). Saurida gracilis: USNM 256409 The area of the neurocranium between the (1). Saurida normani: USNM uncat. (1). Saurida ethmoids and the lateral ethmoids is occupied by parti: USNM 193763 (2). Saurida undosqua- the ethmoid cartilage, which continues posteri- mous: USNM 325180 (10). Scopelarchus analis: orly beyond the lateral ethmoids as the lamina USNM 234988 (1). Scopelarchus nicholsi: orbitonasalis (see de Beer, 1937), in this case a USNM 201154 (2). Scopelarchoides signifer. median wall of cartilage that extends into the USNM 274385 (1). Scopelosaurus hoedti: anterior third of the orbit.
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