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1 aqua Journal of Ichthyology and Aquatic Biology Vol. 8 (4), July 2004 Aquapress ISSN

2 aqua - Journal of Ichthyology and Aquatic Biology Managing Editor: Heiko Bleher Via G. Falcone 11, Miradolo Terme (PV), Italy Tel.: /08 - Fax: heiko@pmp.it Scientific Editor: Dr. Walter Ivantsoff Senior Research Fellow, Department of Biological Sciences, Macquarie University, N.S.W. 2109, Australia Tel Fax wivantso@rna.bio.mq.edu.au Editorial Board: Gerald R. Allen, I Dreyer Road Roleystone, W. A. Australia 6111 George W. Barlow, Department of Integrative Biology, University of California, Berkeley, CA , U.S.A. Henri J. Dumont, Rijksuniversiteit Gent, Laboratorium voor Ecologie der Dieren, Zoogeografie en Natuurbehoud, K. L. Ledeganckstraat, 9000 Gent, Belgium Jacques Géry, Chemin du Plantier, Sarlat, France Frank Kirschbaum, Institut für Gewässerökologie und Binnenfischerei, Abt. 4 Forschungsverbund Berlin e. V. Müggelseedamm 310, Berlin, Germany Friedhelm Krupp, Forschungsinstitut Senckenberg, Senckenberganlage 25, Frankfurt am Main, Germany Christian Lévêque, CNRS - Programme Environnement Vie et Sociétès, 1 Place Aristide Briand, Paris Cédex, France Volker Mahnert, Muséum d Histoire Naturelle, Route de Malagnou 1, 1211 Genève 6, Switzerland Paolo Parenti, Department of Enviromental Sciences, University of Milan-Bicocca, Piazza della Scienza 1, I Milan, Italy John E. Randall, Bishop Museum, 1525 Bernice Street, P.O. Box A, Honolulu, Hawaii, U.S.A. Wolfgang Schneider, Hessisches Landesmuseum, Darmstadt, Friedensplatz 1, Darmstadt, Germany Scope and aims aqua is an international journal which publishes original scientific articles in the fields of systematics, taxonomy, biogeography, ethology, ecology, and general biology of fishes, amphibians, aquatic invertebrates, and plants. Papers on freshwater, brackish, and marine organisms will be considered. aqua is fully refereed and aims at publishing manuscripts within 2-4 months of acceptance. With the publication of aqua we are pursuing a new concept: In view of the importance of colour patterns in species identification and animal ethology, authors are encouraged to submit colour illustrations as well as descriptions of coloration. It is our aim to provide the international scientific community with an efficiently published series meeting high scientific and technical standards. Call for papers The editors welcome the submission of original manuscripts which should be sent directly to the scientific editor. Full length research papers and short notes will be considered for publication. There are no page charges and colour illustrations will be published free of charge. Authors will receive 50 free reprints of each paper. Subscription Notice A volume (4 issues) of aqua will be published each year, each issue comprising 48 pages (including cover). The annual subscription rate (for one volumr = 4 issues) is Euro (US$ 48.00) plus postage Euro (US$ 12.00) and for priority postage Euro 17,00 (US$ 20.00). Subscription enquires should be sent to the address given below or our aquapress@pmp.it Special Volume From 2003 onwards Aquapress will publish a series of Special Volumes. The first Special Volume of aqua will be available in December From then on, Special volumes will be printed yearly, or more often. All Special volumes will contain around 100 pages or more and will only be available separately from normal issues of aqua. Enquiries about subscriptions and the prices of Special Volumes should be sent to the address given below or via aquapress@pmp.it Lothar Seegers, Grenzstraße 47b, Dinslaken, Germany Wolfgang Villwock, Universität Hamburg, Zoologisches Institut und Zoologisches Museum, Martin-Luther-King- Platz 3, Hamburg, Germany Chem Yi-yu, Institute of Hydrobiology, Academia Sinica, Wuhan Hubei, P. R. China ISSN Publisher: Aquapress, Redazione aqua, I Miradolo Terme (Pavia), Italy Printer: Grafiche Dessì s.r.l. (Torino) Italy Typesetting: Rossella Bulla 2004 aqua, Journal of Ichthyology and Aquatic Biology

3 Keywords Viviparous brotulas, Caribbean Sea, Gulf of Mexico, Atlantic Ocean, Pacific Ocean, coral reef fishes, cave fishes, new species Abstract An ongoing revision of the American dinematichthyine fishes (Ophidiiformes, Bythitidae) based on examination of about 2000 specimens will be published in two parts. This publication is Part I, which includes 209 specimens in the genera Dinematichthys (one W. Atlantic species), Gunterichthys (one W. Atlantic and two new E. Pacific species), Ogilbichthys (new genus with seven new W. Atlantic species), Pseudogilbia (new genus with one new W. Atlantic species) and Typhliasina (resurrected with one W. Atlantic cave species). The diagnostic characters are male pseudoclaspers, head pores, otoliths, gill rakers, morphometric proportions, fin ray and vertebral counts. Part II will comprise the revision of the speciose and abundant genus Ogilbia. Zusammenfassung Die noch nicht abgeschlossene Revision der amerikanischen Fischgruppe Dinematichthyini (Ophidiiformes, Bythitidae) auf der Grundlage der Untersuchung von rund 2000 Exemplaren soll in zwei Teilen veröffentlicht werden. Hiermit wird Teil I vorgelegt, der 209 Exemplare auswertet, und zwar aus den Gattungen Dinematichthys (eine westatlantische Art), Gunterichthys (eine westatlantische und zwei neue ostpazifische Arten), Ogilbichthys (neue Gattung mit sieben neuen westatlantischen Arten, Pseudogilbia (neue Gattung mit einer neuen westatlantischen Art) und Typhliasina (mit einer westatlantischen höhlenbewohnenden Art wieder errichtet). Zu den Unterscheidungsmerkmalen gehören männliche Pseudo-Haftorgane, Kopfporen, Otolithen, Kiemenrechen, morphometrische Daten, Flossenstrahlen und Wirbelzahlen. Teil II wird die Revision der artenreichen und häufig vertretenen Gattung Ogilbia enthalten. aqua, Journal of Ichthyology and Aquatic Biology Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two new genera Peter Rask Møller 1, Werner Schwarzhans 2 and Jørgen G. Nielsen 1 1) Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen Ø, Denmark. pdrmoller@zmuc.ku.dk. 2) Ahrensburgerweg 103 D, Hamburg, Germany. Accepted: Résumé Une révision en cours des Dinemaththyines américans (Ophidiiformes, Bythitidae) basée sur l examen de 2000 spécimens sera publiée en deux parties. Cette publication est la 1 e partie qui comprend 209 spécimens dans les genres Dinematichthys (espèce de l Atlantique ouest), Gunterichthys (une espèce de l Atlantique ouest et deux nouvelles espèces du Pacificique est), Ogilbichthys (nouveau genre comptant sept nouvelles espèces de l Atlantique ouest), Pseudogilbia (nouveau genre avec une espèce de l Atlantique ouest) et Typhliasina (revalidé, avec une espèce cavernicole de l Atlantique ouest). Les caractéristiques diagnostiques concernent des pseudoptérygopodes mâles, des pores céphaliques, des otolithes, des branchiospines, des proportions morphométriques, le nombre de rayons et de vertèbres. La 2 de partie comprendra la révision du genre Ogilba, riche en espèces et abondant. Sommario La revisione in corso dei pesci dinematichthyini americani (Ophidiiformes, Bythitidae) basata sull esame di circa 2000 esemplari sarà pubblicata in due parti. In questa prima parte, che racchiude 209 esemplari, sono considerati i generi Dinematichthys (una specie nell Atlantico occidentale), Gunterichthys (una specie nell Atlantico occidentale e due nuove nel Pacifico orientale), Ogilbichthys (nuovo genere con sette specie nuove dell Atlantico occidentale), Pseudogilbia (nuovo genere con una nuova specie dell Atlantico occidentale) e Typhliasina (elevato a genere con one specie cavernicola dell Atlantico occidentale). Come caratteri diagnostici sono considerati i falsi organi copulatori dei maschi, i pori cefalici, gli otoliti, i rastrelli branchiali, le proporzioni morfometriche, il numero di raggi e di vertebre. La seconda parte tratterà la revisione del genere più ricco di specie, Ogilbia. Introduction Dinematichthyine fishes, a tribe within the subfamily Bythitinae, of the viviparous family Bythitidae, are so far represented by genera and about aqua vol. 8 no

4 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera species worldwide (Nielsen et al., 1999). Dinematichthyines when newly born are mm long. Adults of some species may reach mm in length; others not more than mm. Members of this tribe live in shallow tropical to subtropical waters, hidden in holes and crevices of coral reefs, algae beds and rocky shores. Few live on muddy bottoms or in brackish to freshwater caves and sinkholes. Dinematichthyine species show little variation in their general appearance, and their live coloration tends to be uniform, the prevailing colors being yellow, orange, red and brown. Cohen and Nielsen (1978) and Nielsen et al. (1999) used ossification of the male copulatory organs (pseudoclaspers) as the main diagnostic character for the bythitid tribe Dinematichthyini. Sedor (1985) noted that sclerification (used by her in a most general sense to refer to the process of hardening) of the penis, pseudoclaspers and accessory structures is widespread but not universal within the Dinematichthyini. This statement is substantiated by our observations from radiographs that sclerification is variable within genera and species, and increases with the individual s growth. Our observations further support Sedor s (1985) proposal to define the Dinematichthyini by the apomorphic position of the copulatory organs below a covering fleshy hood in a cavity of the ventral body wall. In all other viviparous bythitids (Brosmophycini and Bythitinae), the copulatory organs are an integrated part of the fleshy genital hood, with the penis present as a small, soft papilla. Pseudoclasper-like structures, if present, are indistinct papillae with or without a slight sclerification in the form of a superficial thickening of the epidermis. The family Bythitidae was confirmed to be monophyletic by Patterson and Rosen (1989), but the status and phylogenetic relationship of the subfamilies and tribes need revision. Until the introduction of scuba-diving gear and the use of fish poison, dinematichthyine fishes were rarely caught; now these fish are collected frequently. Today, especially in the United States, museums hold more than ten thousands of specimens from American waters. Because of their abundance, dinematichthyine fishes must play an important role in the ecology of reefs. Over the years, a number of studies on the available materials had been made, but did not result in a comprehensive revision. Nielsen et al. (1999) recognized seven species from American waters: Ogilbia cayorum Evermann and Kendall, 1898; O. deroyi (Poll and van Mol, 1966); O. galapagosensis (Poll and LeLeup, 1965); O. pearsei (Hubbs, 1938); O. ventralis (Gill, 1863), Gunterichthys longipenis Dawson, 1966 and Dinematichthys minyomma Sedor and Cohen, Like many previous workers (e.g. Suarez, 1975; Thomson et al., 1979; Sedor, 1985; Robins et al., 1986; Smith-Vaniz et al., 1999) they mentioned the likely existence of numerous undescribed species. Knowing the extent of the available material, we realized that we could not examine every specimen. Consequently, we have selected series of specimens from many lots covering as many geographical areas as possible. By using a combination of characters such as pseudoclaspers, head pores, otoliths, colour of preserved specimens, gill rakers, and counts and measurements, we recognise the following taxa from the study area: Dinematichthys Bleeker, 1855 (1 species), Gunterichthys Dawson, 1966 (3 species, 2 new), Ogilbia Jordan and Evermann, 1898 (18 species, 14 new), Ogilbichthys (new genus, 7 new species), Pseudogilbia (new genus, 1 species), Typhliasina Whitley, 1951 (genus resurrected, 1 species). The five last-mentioned are endemic to American waters, while Dinematichthys is also known from 6 nominal Indo-West Pacific species (Nielsen et al., 1999). Due to the large amount of material with many new taxa we decided to deal with the American dinematichthyine fishes in two publications: Part I treats all genera except for Ogilbia and Part II will treat Ogilbia only. Dinematichthyine fishes from the Indo- West Pacific area are also being reviewed and the results will appear in subsequent publications. Material and methods Of the 2217 specimens of American Dinematichthyini examined, 209 specimens were identified as members of the genera described in this work. The remainder were identified as members of the genus Ogilbia and included many new species. The specimens examined are kept in the following collections: AMS (Australian Museum Sydney), ANSP (Academy of Natural Sciences Philadelphia), CAS (California Academy of Sciences, San Francisco), LACM (Los Angeles County Museum), NHM (The Natural History Museum, London; former BMNH), NMNZ (Museum of New Zealand Te Papa Tongarewa, Wellington), SAIAB (South African Institute for Aquatic Biodiversity, Grahamstown; former RUSI (J. L. B. Smith Institute of Ichthyology)), SIO (Scripps Institute of Oceanography, La Jolla), SAM (South African Museum, Capetown), UCR (University of Costa Rica, San Jose), UF (University of Florida, Gainesville), UMMZ (University of Michigan, Museum of Zoology, Ann Arbor), USNM (Unites States National Museum, Washington D.C.), University Libre de Bruxelles, WAM (Western Australian Museum, Perth), ZMH (Zoological Institute and Museum, Hamburg), ZMUC (Zoological Museum, University of Copenhagen). Sample sites are shown in Fig. 1. Morphometric characters are given as percent of standard length (SL) throughout. In tables the mean value are given first followed by the range. Sometimes not all of the fish were measured/counted for certain characters, due to damage, very short loan time or other logistic problems. As a rule, a minimum of 30 specimens of each species were measured/ aqua vol. 8 no

5 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen counted, if available. In the descriptions, data for the holotype are given first, followed by the range in brackets. Size of eye is measured as the horizontal diameter of the pigmented eyeball. Meristic counts were made from radiographs, except for pectoral fin rays, gill rakers, teeth and scale rows. The right otolith was removed through the gill cavity by making a small cut above the gills. The anterior right gill arch of most species is illustrated by scanning electron microscopy (SEM). Pseudoclaspers were observed by bending forward the fleshy hood covering the copulatory organs and then bending laterally or spreading them and fixing them with a thin needle. Contrary to the proposal by Sedor (1985), it is not necessary to dissect pseudoclaspers for morphological analysis. This would result Fig. 1. Sample sites of Dinematichthys minyomma, Gunterichthys bussingi, G. coheni, G. longipenis, Ogilbichthys ferocis, O. haitiensis, O. kakuki, O. longimanus, O. microphthalmus, O. puertoricoensis, O. tobagoensis, Pseudogilbia sanblasensis and Typhliasina pearsei, used in the present revision. One symbol may represent several samples. 143 aqua vol. 8 no

6 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera in loss or destruction of the more delicate characters of the pseudoclaspers. When drawn, pseudoclaspers and penis are shaded, whereas other parts, such as the fleshy hood, isthmus or outline of the copulatory cavity are shown in simple line treatment. All drawings and photographs (other than Fig. 24) are drawn and/or photographed by the authors or the museum (ZMUC) artist, Birgitte Rubaek. The biology of most of the species is poorly known, since the study was done almost exclusively on museum specimens. A number of females were examined for reproductive data e.g. number and size of embryos. Character evaluation and description Because of the somewhat inconsistent and confusing nomenclature and terminology of characters in the dinematichthyine literature, the more important characters are described: Head pore system (Fig. 2). The following pore rows are recognized: supraorbital row holds 3-4 pores. The posteriormost was previously termed the first lateral line pore (e.g. Machida 1994, Cohen and A B Fig. 2A-B. Head pore system terminology. Open pores shown as solid circles and pores hidden from lateral view as dashed circles (Ogilbichthys). A) Lateral view of head; B) Ventral view of head. aqua vol. 8 no

7 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Fig. 3A-D. Male copulatory organs terminology. A) View of left pseudoclaspers from inside (Typhliasina); B) View of left pseudoclaspers from inside (Ogilbichthys); C) Inclined lateral view of male copulatory organs (Ogilbichthys); D) Inclined lateral view of male copulatory organs (Gunterichthys). Abbreviations: isth. = isthmus, lig. = ligament, o.p. = outer pseudoclasper, i.p. = inner pseudoclasper, a.i.p. = anterior inner pseudoclasper, p.i.p. = posterior inner pseudoclasper, pock. = pocket in front of anterior inner pseudoclasper (which is sometimes sunk into it), supp. = supporters of outer pseudoclasper. McCosker 1998). It is located above the opercular spine at the upper termination of the gill opening and is part of the supraorbital row. It is tube-shaped in all species. The infraorbital row consists of 3 anterior and 2-3 posterior pores. The mandibular row consists of 3 anterior and 3 posterior pores. The preopercular row generally consists of 3 lower and 1 upper pore, but the number is reduced in some species, particularly in respect to presence or absence of the upper pore. The latter two rows form a continuous system and are often referred to in the literature as the preoperculomandibular pore row. The two lower preopercular pores are often placed in a common pore-like cavity; the upper pore can be absent in a few species. The number and relative size of the various pores is generally very stable within a specific systematic unit. Differences in head pore counts are therefore often used for generic definitions, differences in pore sizes generally for species definitions. Male copulatory organs (Fig. 3). Covered anteriorly by a thick fleshy genital hood that originates from the posterior margin of the anus. Consists of a penis flanked by 1-3 pairs of pseudoclaspers, and a sometimes indistinct, accessory organ at the base of the penis. The pseudoclaspers may contain hardened, sclerified bodies that keep them in an upright position. These are here termed supporters. The pairs of pseudoclaspers are separated by a fleshy isthmus, originating from the hood and joining the base of the penis. The outer pseudoclaspers are connected distally by a ligament that passes around the posterior base of the penis. The inner pseudoclasper is free or joined to either the outer pseudoclasper or to the isthmus between the pseudoclaspers by a ligament; in Ogilbichthys, there is an additional (anterior) inner pseudoclasper located in front of the posterior inner pseudoclasper (Fig. 3A-C). The latter is termed the inner pseudoclasper when the anterior inner pseudoclasper is missing (in all genera except for Ogilbichthys). Gunterichthys has a single pair of pseudoclaspers with two supporters, homologous to the outer pseudoclaspers in the other genera. We follow Suarez s (1975: fig. 3) terminology of paired inner and outer pseudoclaspers for practical reasons. For a detailed description of the reproductive biology and functional anatomy of the copulatory apparatus see Turner (1946) and Suarez (1975). Sedor (1985: 5) notes that the pseudoclaspers are joined at the base and therefore refers to a single pair of pseudoclaspers, either branched or not branched. In this paper one of the pseudoclaspers is usually shown from the left inside. Additionally, the entire copulatory apparatus is illustrated with the fleshy hood bent forward from an inclined lateral view, showing one pair of pseudoclaspers from the outside, the other from the inside. In previous publications, the copulatory apparatus is usually shown from a ventral view only, which does not always give an adequate view of the details. 145 aqua vol. 8 no

8 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Pseudoclasper morphology is found to be the most useful character in Dinematichthyini both for distinguishing species as well as genera. The reason for the very high degree of interspecific variation in pseudoclasper morphology is probably due to its function which may guarantee mating with a conspecific female (Suarez, 1975). Presence of one or two pairs of pseudoclaspers is, with very few exceptions, used for definition of genera, while details of the pseudoclasper morphology are used for species definition. In fact, there are only few instances where specific differences have to rely exclusively on other characters. Examination of the pseudoclaspers of more than 4000 mature male dinematichthyines has revealed that its morphological pattern is very stable, both in terms of variability and ontogenetic changes (the latter after maturity has been reached). Within the American Dinematichthyini limited allometric growth has been observed in just a few species of the genus Ogilbia. A single specimen had two pseudoclaspers on one side and only one on the other, which may have been due to injury or deformation. Male pseudoclaspers are the easiest characters to use for separating species. Juveniles and females can be difficult to identify for some species. In American waters, however, all non-ogilbia specimens can be identified if radiographs are available. Sagittal otolith (Fig. 4). Characters of the sagitta follow Schwarzhans (1993). The sagitta ( otolith in the following) of dinematichthyine fishes show little ontogenetic and intraspecific variation in contrast to certain other ophidiiform fishes. For a comprehensive review of ophidiiform otolith morphologies see Nolf (1980) and Schwarzhans (1981). The most easily recognizable character in dinematichthyine otoliths is the status of the sulcus: divided (separated colliculi) or undivided (fused colliculi) and the form of the ventral margin of the sulcus between the two. These characters are often used as additional characters for generic diagnoses. However, fusion of colliculi is a very common trend in Dinematichthyini and other ophidiiform fishes and has apparently occurred in multiple parallel lineages. We therefore have refrained from separating genera in those instances, where division or non-division of the sulcus margin and/or the colliculi is not supported adequately by other characters. Additional characters (general morphology). Diagnostically important characters used in the present revision are: Opercular spine: The tip of the spine is free in Dinematichthyini, except for Gunterichthys amongst American genera, where it is covered by skin (when visible through the skin, it is shown in drawings as a dotted line). Anterior nostril is normally placed close to tip of snout in bythitids, but in Dinematichthys s.l. and a few related, yet undescribed Indo-West Pacific genera it is positioned mid-way between snout and posterior nostril, i.e. high above the upper lip (see Fig. 8A). Head squamation usually requires removal of mucus, which often covers the head. Only a few species of Dinematichthys s.l. from the Indo-West- Pacific show complete squamation of the head. In most other Dinematichthyini head squamation is restricted to the cheeks or missing entirely from the Fig. 4. Otolith (sagitta) terminology. Median view of right otolith of Dinematichthys sp. aqua vol. 8 no

9 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen head. Such differences are used in generic as well as in species definitions. Comparative material Dinematichthyini: Beaglichthys macrophthalmus Machida, 1993: Non-type: WAM Brotulina fusca Fowler, 1946: HOLOTYPE: ANSP Dermatopsis macrodon Ogilby, 1896: Non-types: WAM (3 specimens [spms]). Dermatopsoides kasougae (Smith, 1943): HOLOTYPE: RUSI 333, only radiograph examined; Non-type: RUSI Dermatopsoides talboti Cohen, 1966: HOLOTYPE: RUSI 340, only radiograph examined; PARATYPES: RUSI 339, only radiograph examined; SAM 21800, SAM 21963). Dipulus caecus Waite, 1905: Non-types: WAM , WAM , ZMUC (3 spms.). Dipulus norfolkanus Machida, 1993: Nontypes: NMNZ 11706, NMNZ 11742, NMNZ (3 spms.), NMNZ (4 spms.). Dinematichthys dasyrhynchus Cohen and Hutchins, 1982: Non-types: WAM (2 spms.), WAM , ZMUC Dinematichthys iluocoeteoides Bleeker, 1855: BMNH Dinematichthys indicus Machida, 1994: PARATYPES: ROM (14 spms.), ROM (8 spms.). Dinematichthys megasoma Machida, 1994: Non-types: WAM (7 spms.), WAM (9 spms.). Dinematichthys riukiuensis Aoyagi, 1954: Non-types: USNM (12 spms.), ROM (5 spms.). Diancistrus longifilis (Ogilby, 1899): Non-types: AMS (3 spms.) only otolith examined. Ogilbia cayorum Evermann and Kendall, 1898: HOLOTYPE: USNM 48792, only photo and radiograph examined. Ogilbia deroyi (Poll and van Mol, 1966): PARATYPES: University Libre de Bruxelles, uncatalogued. (2 spms.), Non-types: CAS 31501, CAS (2 spm.), USNM , USNM , USNM (2 spms.). Ogilbia ventralis (Gill, 1863): SYNTYPES: USNM (3 spms.), only photos and radiographs examined. Ogilbia spp. Jordan and Evermann, 1898: more than 1000 specimens to be treated in a later publication. Brosmophycinae: Bidenichthys beeblebroxi Paulin, 1995: Non-type: ZMUC Bidenichthys capensis Barnard, 1934: Non-type: ZMUC Brosmodorsalis persicinus Paulin and Roberts, 1989: Non-type: ZMUC Brosmophyciops pautzkei Schultz, 1960: Non-type: ROM 58262, 65311, 47595, 48616, MNHN , , AMS , , , Brosmophycis marginata (Ayres, 1854): Non-type: ZMUC Fiordichthys slartibartfasti Paulin, 1995: Non-type: NMNZ 32145, 35097, Melodichthys hadrocephalus: Nielsen & Cohen, 1986: Non-type: MNHN Tribe Dinematichthyini Cohen and Nielsen, 1978 (Family Bythitidae Gill, 1861; Subfamily Brosmophycinae Gill, 1862). Diagnosis Male copulatory organs with a penis and 1-2 (rarely 3) pairs of pseudoclaspers in cavity of ventral body wall, covered by a fleshy hood. First anal fin pterygiophore slightly to strongly elongated. Head pore system generally unreduced. Posteriormost supraorbital head pore tubular. Key to genera of Dinematichthyini in American waters 147 1a Anterior nostril high on snout, midway between upper lip and posterior nostril (Fig. 8A); anterior lower gill arch with one short plate-like raker between two slightly longer rakers (Fig. 5A); first anal fin pterygiophore slightly elongated in males, not reaching parapophysis of last precaudal vertebra... Dinematichthys (D. minyomma in Caribbean Columbia, Panama, Honduras, Belize, Puerto Rico, Haiti, Barbuda) 1b Anterior nostril low on snout, closer to upper lip than to posterior nostril (e.g. Fig. 10a); anterior gill arch with 2-8 developed rakers in a uninterrupted row (Figs. 5B-D, 6); first anal fin pterygiophore very elongated in males, often reaching parapophysis of last precaudal vertebra (Fig. 19A)...2 2a Specimens longer than 20 mm SL without visible eyes, minute black dots in specimens less than 20 mm; height of posterior maxillary > 5.5 % SL, pseudobranch absent, upper preopercular pore absent, posterior infraorbital pores Typhliasina (Yucatan/ freshwater caves) 2b All specimens with visible eyes (except for Ogilbia galapagosensis longer than 40 mm SL); height of posterior maxillary < 5.0 % SL; pseudobranch present (except absent in Gunterichthys longipenis), upper preopercular pore present (except absent in Ogilbia deroyi, O. galapagosensis, Ogilbichthys microphthalmus and O. puertoricoensis); posterior infraorbital pores 3 (except for 2 in G. longipenis)...3 3a Opercular spine hidden; 5-7 prolonged rakers on anterior gill arch (Fig. 5B-C); one pair of pseudoclaspers with two equally long supporters; posterior ventral knob on maxilla well anterior to rear corner (e.g. Figs. 9, 10A)... Gunterichthys (Gulf of Mexico and Pacific Panama/ Costa Rica) 3b Opercular spine distinct; 2-4 slightly prolonged rakers on anterior gill arch; 2-3 pairs of pseudoclaspers (except for one pair in a undescribed species of Ogilbia); outer pseudoclasper with no or small anterior supporter; posterior ventral knob on maxilla situated on rear corner...4 4a Two pairs of inner pseudoclaspers, posterior inner pseudoclasper joined by ligament to isthaqua vol. 8 no

10 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera mus between the pseudoclaspers (Fig. 3B-C), pectoral fin length > 16 % SL (except for a few specimens of O. ferocis and O. haitiensis), soft interorbital width > 7 % SL (in specimens > 30 mm SL).. Ogilbichthys (Caribbean Panama, Tobago, Haiti, Bahamas, Cayman Isl.) 4b One pair of inner pseudoclaspers, free of isthmus between the pseudoclaspers, pectoral fin length < 16 % SL, soft interorbital width < 7 % SL (in specimens > 30 mm SL)...5 5a. Lower preopercular pores 1, predorsal length > 33 % SL, colliculi fused, inner pseudoclasper almost as large as outer pseudoclasper (Fig. 34C, E)...Pseudogilbia (Caribbean Panama) 5b. Lower preopercular pores 3, predorsal length < 33 % SL (except for a few juveniles < 20 mm); colliculi separate, inner pseudoclasper smaller than outer pseudoclasper...ogilbia (Caribbean, W. Atlantic and E. Pacific Oceans and Galápagos Isl. fresh water caves) Dinematichthys Bleeker, 1855 Dinematichthys Bleeker, 1855: 318 (type species D. iluocoeteoides Bleeker, 1855 by monotypy). Type locality: Batu Island, Indonesia. Dinematichthys: Cohen and Nielsen 1978: 57; Nielsen et al. 1999: 129. Diagnosis Anterior nostril placed high, midway between upper lip and posterior nostril; tip of opercular spine free; two pairs of simple pseudoclaspers, a flap-like outer pseudoclasper and a small inner pseudoclasper, half the size of the outer pseudoclasper or less; eye size variable; scales on cheeks; otoliths with separate colliculi; anterior anal fin ray pterygiophore short; 3 lower preopercular pores; maxillary knob at rear corner; 3 slightly elongated rakers on anterior gill arch. Table I. Comparison of dinematichthyine genera known in American waters. Genus Dinematichthys Gunterichthys Ogilbia Ogilbichthys Pseudogilbia Typhliasina Number of 1 3 about Species Pairs of pseudoclaspers 2 1 2* Inner free absent free connected to free free pseudoclasper isthmus Anterior nostril near posterior near upper near upper near upper near upper near upper situated nostril lip lip lip lip lip Posterior infraobital pores Lower preopercular pores Upper prepresent opercular pore present present** present/absent present absent Otolith colliculi separate yes yes/no yes yes/no no no Pseudobranchial filaments Eye visible yes yes yes*** yes*** yes no Opercular spine free hidden free free free free Scales on head yes yes/no yes/no yes/no yes no Long gill rakers Head width >14 % SL >or<14 % SL <14 % SL**** >or<14 % SL >14 % SL >14 % SL Preanal length <52 % SL >or<52 % SL <52 % SL**** <52 % SL >52 % SL >52 % SL Predorsal length >or<33 % SL >or<33 % SL < 33 % SL**** >or<33 % SL >33 % SL > 33 % SL Vertebrae Dorsal/Anal fin rays 71-79/ / / / / / * An undescribed species has only one pair of pseudoclaspers. ** Absent in Ogilbia deroyi. *** Sunk into head in some Ogilbia galapagosensis and Ogilbichthys microphthalmus. **** Few exceptions found, mainly among juveniles. aqua vol. 8 no

11 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Similarity (Table I). It is premature to discuss generic similarity as Dinematichthys is probably a polyphyletic genus (see below). Remarks Among the very extensive Indo-West Pacific material that we are currently studying, there is a number of undescribed Dinematichthys-like species. Most probably the genus will be split into 2-3 genera and D. minyomma may be assigned to one of the presently undescribed genera. Sedor and Cohen (1987) stated that minyomma is only provisionally placed in Dinematichthys. Most of the uncertainty is caused by the loss of the type material of the type species, D. iluocoeteoides (see Cohen and Nielsen, 1978, for discussion). A neotype will be designated in the forthcoming review of the Indo-West Pacific Dinematichthyini. Species Seven species have so far been described: D. minyomma, from the western Caribbean and six from the Indo-West Pacific: D. dasyrhynchus Cohen and Hutchins, 1982 from Rottnest Island, Western Australia; D. iluocoeteoides Bleeker, 1855 from eastern Indian Ocean; D. indicus Machida, 1994 from western Indian Ocean; D. megasoma Machida, 1994 from Northern Australia; D. randalli Machida, 1994 from Micronesia and D. riukiuensis Aoyagi, 1952 from Riukyu Islands. Dinematichthys minyomma Sedor and Cohen, 1987 (Figs. 5A, 7, 8; Tables II, III) Dinematichthys minyomma Sedor and Cohen, 1987: 6, Fig. 1 (type locality Bay Islands, Honduras, Caribbean Sea). A B A B C D C D Fig. 5A-D. First gill arch A) Dinematichthys minyomma, CAS , female, 52 mm SL; B) Gunterichthys coheni, ZMUC P771348, male, 32 mm SL; C) Gunterichthys longipenis, UF , female, 57 mm SL; D) Typhliasina pearsei, ZMUC P771336, female, 83 mm SL. Fig. 6A-D. First gill arch of Ogilbichthys spp. A) O. ferocis, ZMUC P771345, female, 29 mm SL; B) O. longimanus, ZMUC P771350, male, 59 mm SL; C) O. microphthalmus, ZMUC P771352, female, 37 mm SL; D) O. kakuki, ZMUC P771353, male, 47 mm SL. 149 aqua vol. 8 no

12 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 7. Dinematichthys minyomma. Paratype, USNM , female, 68 mm SL. Fig. 8A-E. Dinematichthys minyomma. A) Lateral view of head, USNM , male, 66 mm SL; B) Ventral view of head, USNM , female, 61 mm SL; C) View of left pair of pseudoclaspers from inside, USNM , male, 71 mm SL; D) Inclined lateral view of male copulatory organs of same specimen; E) Median view of right otolith, USNM , male 71 mm SL. For abbreviations see Fig. 3. aqua vol. 8 no

13 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Dinematichthys minyomma: Nielsen et al. 1999: 130; (Chen and Shao 1991: 15, Fig. 10 and Randall and Lim 2000: 596 erroneously reported D. minyomma from the South China Sea the true identity is unknown). Species 2: Sedor 1985, Tab. II, Fig. 3. Material examined (40 specimens, mm SL): Holotype: USNM , male, 66 mm SL, Bay Islands, Honduras, Caribbean Sea, N, W, collected off a coral reef, at depth 0-9 m, 21 Apr. 1967, Paratypes: USNM , 5 females and 5 males, mm SL, same data as for holotype. Non-types: ANSP , 1 female, 42 mm SL and 2 males, 49 and 54 mm SL, Haiti, Gulf of Gonave; St. Marc Channel, off Mount Rouis, 3.2 km south-east of Mount Rouis town, N, W, m, rotenone, collected by J. C. Tyler and collaborators 14 Sept. 1963; CAS , 2 females, 36 mm and 52 mm SL and 1 male, 44 mm SL, San Blas Archipelago, Panama, Caribbean Sea, N, W, collected with pronoxfish, off a reef, 13 May 1974; USNM , 3 females, mm SL, Cocoa Point, Barbuda, West Indies, collected by R. S. Cowen, 27 April 1959; USNM , 3 females and 3 males, mm SL, Isla Grande, Caribbean Columbia, N, W, 0-1 m, R/V Choco, collected by L. K. Knapp, 29 Sept. 1969; USNM , 2 males, mm SL, La Pargùera, Puerto Rico, 5 June 1965; USNM , 1 male, 59 mm SL, Carrie Bow Cay, Belize, 0-2 m, collected by G. D. Johnson and P. Keener, 2 Nov. 1984; USNM , 2 females and 1 male, mm SL, Carrie Bow Cay, Belize, collected by E. O. Wiley and party, 16 July 1991; USNM , 5 females, mm SL and 1 male, 27 mm SL, Quanaja, Bay Island, Honduras, N, W, 0-10 m, 21 April 1967; ZMUC P771344, 1 male, 52 mm SL, same data as for CAS ; ZMUC P , 1 female, 47 mm SL and 1 male, 59 mm SL, same data as for ANSP Diagnosis Body covered with imbricate scales; head with scales on cheek; eyes small and distinct % SL; 3 posterior infraorbital pores; upper preopercular pore present; tip of opercular spine free; two pseudobranchial filaments. Two pairs of small (compared to size of penis), widely-separated pseudoclaspers, outer pseudoclasper much larger than inner; otoliths with separate colliculi. Similarity This is the only species of the genus in American waters. Comparison with the many described and undescribed species in the Indo-West Pacific would be premature at this time. In American waters, D. minyomma appears to be morphometrically most similar to Ogilbichthys longimanus, in the long and broad head, wide interorbital width, long upper jaw, high body, long ventral fins and wide posterior maxillary. D. minyomma differs from it by the shorter gill rakers, predorsal length ( vs % SL), smaller eye ( vs % SL), higher pectoral fin ray count (21-23 vs 17-20), the simple structure of the pseudoclaspers (see Ogilbichthys for details) and the separated colliculi (vs. fused). Description Meristic and morphometric characters are presented in Tables II and III. For a very detailed description, see Sedor and Cohen (1987). Body compressed, head broad and snout blunt. Eyes small and distinct ( % SL). Maxillaries end far behind eyes; a distinct bony knob ventrally on posterior part of maxilla. Anterior nostril tubular, placed midway between upper lip and posterior nostril, the latter being small, about one fourth of eye diameter and situated close to eye. Opercular spine free, but hardly visible. Upper branch of anterior gill arch with one knob-like raker and 2-4 plate-shaped rakers and lower branch with rakers, all plate-formed except for the first and third being knob-formed (Fig. 5A). Two pseudobranchial filaments. Scales large, about 25 horizonthal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins naked, caudal fin partly scaled. Head naked except for a patch of large scales on cheek, 5-6 scale rows on upper cheek and 2-3 scale rows on lower cheek. Origin of dorsal fin above base of pectoral fin, which ends well in front of anal fin. Ventral fins reach almost to anal fin. Head sensory pores (Fig. 8A-B). Supraorbital pores 4: 1st about size of 2nd anterior infraorbital pore, 2nd above anterior nostril and 3rd tubular pore above and behind eye, 4th small tubular pore at upper termination of gill opening above opercular spine. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small like the three posterior infraorbital pores and 2nd and 3rd anterior pores about 3 times larger, covered by dermal flap of upper lip. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large with cirrhi at margins, 2nd pore positioned in lateral skin fold and of about size of posterior mandibular pores and 3rd pore about three times the size of posterior mandibular pores. Three posterior pores of about same size as posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint openings, 3rd pore about the size of posterior mandibular pores. Upper pore small and tubular. Lateral line configuration. A dorsal branch with about 12 neuromasts and a lateral branch of about 20 neuromasts. Dentition. Premaxillaries and dentaries with larger, 151 aqua vol. 8 no

14 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Table II. Frequency distribution for American Dinematichthyini. Holotype values in bold. aqua vol. 8 no

15 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Table III. Meristic and morphometric characters of Dinematichthys minyomma. Holotype HT+Paratypes + USNM 10 paratypes 9 non-type non-types Mean and range Mean and range Mean and range Standard length (50-73) 48.3(36-59) 55.1(36-73) Meristic characters Dorsal fin rays 75 75(71-79) 74.3(73-76) 74.7(71-79) Caudal fin rays Anal fin rays (56-61) 57.4(55-59) 57.8(55-61) Pectoral fin rays (21-22) 22.1(22-23) 21.7(21-23) Precaudal vertebrae (10-11) 10.9(10-11) 10.8(10-11) Total vertebrae (39-40) (39-40) Gill rakers on anterior arch (15-19) 17.6(15-19) 17.4(15-19) Pseudobranchial filaments Anterior dorsal fin ray above vertebra no. (D/V) 6 5.4(5-6) 5.7(5-6) 5.5(5-6) Anterior anal fin ray below dorsal fin ray no. (D/A) (21-25) 22.7(22-23) 22.4(21-25) Anterior anal fin ray below vertebra no. (V/A) (12-13) 13.2(12-14) 13.1(12-14) Morphometric characters in % of SL Head length ( ) 28.3( ) 28.0( ) Head width ( ) 15.6( ) 15.5( ) Head height ( ) 19.1( ) 20.3( ) Upper jaw length ( ) 14.9( ) 14.8( ) Maxillary height ( ) 4.9( ) 5.0( ) Diameter of pigmented eye ( ) 1.9( ) 2.0( ) Interorbital width ( ) 8.5( ) 8.1( ) Postorbital length ( ) 20.7( ) 20.3( ) Preanal length ( ) 48.4( ) 49.9( ) Predorsal length ( ) 32.3( ) 32.1( ) Body depth at origin of anal fin ( ) 20.2( ) 20.9( ) Pectoral fin length ( ) 15.5( ) 14.8( ) Ventral fin length ( ) 26.6( ) 26.5( ) Base of ventral fin to anal fin ( ) 28.3( ) 29.1( ) more pointed teeth in a medial row while the rest are smaller and conical; number of rows increase anteriorad. Vomer v-shaped with conical teeth anteriorly covering most of the bone and pointed, longer teeth in a posterior row. Palatines with pointed, larger teeth in a medial row. Otolith (Fig. 8E). Elongate with a length to height ratio of 2.2 to 2.3 (50-73 mm SL). Anterior tip broadly rounded; posterior tip variable, but usually narrower than anterior tip and more pointed. Dorsal rim shallow, with broad predorsal and no postdorsal angle; ventral rim deeply curving, deepest anterior of the middle. Inner face convex and outer face smooth and concave. Otolith length to sulcus length ratio 1.6 to 1.7. Sulcus clearly divided into a long and wide ostium and a short and narrow cauda with a distinct notch at ventral sulcus margin. Ostium about 4 times the length of cauda. Ostial colliculum flat, caudal colliculum deepened. Dorsal depression shallow, narrow, close to dorsal rim of otolith; ventral furrow distinct, close to ventral rim of otolith, branching backwardly. Axial skeleton. Anterior neural spine half the length of second spine. Neural spines 5-9 much depressed, short and with blunt tips. Tips of all other neural and haemal spines pointed. Bases of spines of vertebrae 5-9 enlarged. Parapophyses on 5-6 posterior precaudal vertebrae. Pleural ribs on all precaudal vertebrae except for the posteriormost. Epipleural ribs on vertebrae 3-10/11. Male copulatory organs (Fig. 8C-D). Two pairs of small and widely-separated pseudoclaspers. Outer pseudoclasper forming a simple flap about 2-3 times the size of inner pseudoclasper, but less than half the length of penis. Inner pseudoclasper with blunt posterior margin and an upward extended hook, and connected anteriorly to outer pseudoclasper. Isthmus between pseudoclaspers wider than base of penis. Penis short, curved, with broad base and pointed tip. Allometric growth. Positive allometric growth observed in head width and body depth; negative allometric change seen in pectoral fin length (Table III). Biology. Caught in shallows, over coral reefs. A 52 mm specimen (CAS ) contains about 200 eggs with a diameter of mm and about 75 embryos of 5-6 mm length. The eggs are found in the anteriormost part of the ovary, the embryos posteriorly. Embryos with 153 aqua vol. 8 no

16 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera large black eyes and 4-5 black, short horizontal lines medially on each side of posterior third of body. Colour in alcohol. After years of preservation the specimens are uniformly brownish with black eyes. Two black pigmented areas are located dorsally in the abdominal cavity of embryos. Distribution Known from 8 localities in the Caribbean Sea, off Honduras, Haiti and Atlantic Panama (Fig. 1). The holotype and 38 paratypes were collected together. The non-type specimens here examined are the first reported from other than the type locality. Gunterichthys Dawson, 1966 Gunterichthys Dawson, 1966: 205 (type species G. longipenis Dawson, 1966: 206 by original designation). Type locality: Gulf of Mexico, off Mississippi Sound, Davis Bayour. Gunterichthys: Cohen and Nielsen 1978: 59; Nielsen et al. 1999: 132. Diagnosis Anterior nostril placed low on snout; opercular spine weak, hidden below skin; one pair of large pseudoclaspers about two-thirds the length of penis, with two almost equally long supporters; eye size variable ( % SL); head naked or with a few scales on cheek; otoliths with fused or partly separate colliculi; 3 lower preopercular pores; maxilla with postero-ventral knob well anterior to rear corner; posterioventral maxillary knob rounded posteriorly; 5-7 prolonged rakers on anterior gill arch; anterior anal fin ray pterygiophore elongated. Similarity (Table I). Gunterichthys resembles Typhliasina in having 5-7 long gill rakers and in the short ventral fins and is similar to Ogilbichthys by sharing the large interorbital width ( and % SL) and the most often fused colliculi. Gunterichthys differs from both by having a single pair of large pseudoclaspers with two supporters, which are homologous to the outer pseudoclaspers, judged by the presence of a ligament connecting them behind the base of the penis, opercular spine hidden (partly free in the other genera). A possible close relationship of Gunterichthys to a group of South African and Australian genera (Dermatopsis and Dermatopsoides) is indicated by the morphology of the posterior maxillary, where the ventral knob is placed anteriorly to the rear corner (see Cohen 1966: Fig. 1). This group was given tribal status, Dermatopsini, by Cohen (1966), but has later been ignored (Cohen & Nielsen 1978; Nielsen et al. 1999). Further studies are needed before any conclusions can be reached in regard to the relationship between Gunterichthys and Dermatopsis / Dermatopsoides. Species Known from the type species from the Gulf of Mexico and two species, described here, from the Gulf of Panama and the Pacific coast of Costa Rica. Key to species 1a Head width > 13 % SL; dorsal fin rays 60-71, anal fin rays 41-52; posterior infraorbital pores 2; snout blunt; no pseudobranchial filaments (Atlantic)......G. longipenis 1b Head width < 13 % SL; dorsal fin rays 74-81, anal fin rays 53-59, posterior infraorbital pores 3; snout pointed; pseudobranchial filaments present (Pacific)...2 2a Head naked; interorbital width % SL; eye diameter % SL; posterior nostril small (Fig. 12a); caudal fin rays 14-15; otolith length to height ratio G. coheni 2b Head with scale-band on cheek; interorbital width 5.5 % SL; eye diamenter % SL; posterior nostril large (Fig. 10a); caudal fin rays 16; otolith length to height ratio G. bussingi Gunterichthys bussingi n. sp. (Figs. 9, 10; Tables II, IV) Material examined (2 specimens, mm SL): Holotype: LACM , female, 60 mm SL, Isla del Cano, Puntarenas, Costa Rica, 8 43 N, W, depth and method unknown to us, collected by J. and S. Perry and B. Nishimoto, 13 Mar Paratype: UCR , male (immature), 20 mm SL, same data as for holotype, Diagnosis Snout pointed; eyes distinct, large ( % SL); head with scale-band on cheek; three posterior infraorbital pores; posterior nostril large (about half of eye diameter); rays in dorsal fin 79-81, in anal fin and in caudal 16; otoliths with length to height ratio 2.5 and partly fused colliculi. No information on pseudoclaspers due to lack of an adult male. Similarity G. bussingi is most similar to another newlydescribed species, G. coheni, in that both have three posterior infraorbital pores, pointed snout and many rays in dorsal and anal fins (Table II). It differs from G. coheni by having scale-band on cheek (vs. head naked), larger eye ( vs % SL), larger posterior nostril (one half vs. one sixth diameter of eye), larger interorbital width (5.5 vs % SL) and more elongate otolith (ratio length to height 2.5 vs ). aqua vol. 8 no

17 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Fig. 9. Gunterichthys bussingi. Holotype, LACM , female, 60 mm SL. Fig. 10A-D. Gunterichthys bussingi. Holotype, LACM , female, 60 mm SL. A) Lateral view of head; B) Ventral view of head; C) Median view of right otolith; D) Ventral view of right otolith. Description Meristic and morphometric characters are presented in Tables II and IV. Head profile steep, snout pointed, mouth terminal. Postero-ventral knob on maxilla well anterior to rear corner. Anterior nostril positioned low, posterior nostril large, about half of eye diameter. Opercular spine covered by skin. Upper branch of anterior gill arch with 2 small knobs and one elongate raker; lower branch with 4 elongate rakers and 7 small knobs. Pseudobranchial filaments 2. Small imbricate scales on body, about 25 horizonthal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins naked, head with narrow band of small scales on cheek. Origin of dorsal fin just posterior to base of pectoral fin. Ventral fins short, reaching two-thirds of distance from its base to anal fin origin. Head sensory pores. Supraorbital pores 3: Anterior pore about half the size of 2nd anterior infraorbital pore, and small, tubular 2nd pore above and behind 155 aqua vol. 8 no

18 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Table IV. Meristic and morphometric characters of Gunterichthys bussingi and G. coheni. G. bussingi G. coheni Holotype Paratype Holotype HT + 11 PT s N LACM UCR BMNH Mean and range Standard length in mm (20-49) 12 Meristic characters Dorsal fin rays (71-77) 12 Caudal fin rays (14-15) 9 Anal fin rays (51-59) 12 Pectoral fin rays (20-23) 9 Precaudal vertebrae (11-12) 12 Total vertebrae (40-41) 12 Long rakers on anterior gill arch (5-6) 12 Total rakers on anterior gill arch (14-17) 11 Pseudobranchial filaments Anterior dorsal fin ray above vertebra no. (D/V) (5-6) 12 Anterior anal fin ray below dorsal fin ray no. (D/A) (21-26) 12 Anterior anal fin ray below vertebra no. (V/A) (14-15) 12 Morphometric characters in % of SL * Head length ( ) 11 Head width ( ) 11 Head height ( ) 11 Upper jaw length ( ) 10 Maxillary height ( ) 10 Diameter of pigmented eye ( ) 11 Interorbital width ( ) 11 Postorbital length ( ) 11 Preanal length (50-56) 11 Predorsal length ( ) 11 Body depth at origin of anal fin ( ) 11 Pectoral fin length ( ) 9 Ventral fin length ( ) 11 Base of ventral fin to anal fin ( ) 11 *excluding morphometric characters of a 20 mm specimen eye, posterior tubular pore at upper termination of gill opening, above opercular spine. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small and 2nd and 3rd pores large, covered by dermal flap of upper lip. Posterior infraorbital pores small, less than half the size of posterior nostril. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large, non tubular and without cirrhi, 2nd anterior pore free from lateral skin fold, about half size of posterior mandibular pores; 3rd pore large, three times the size of posterior mandibular pores. Posterior mandibular pores small, about half size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd with joint openings, 3rd pore small about the size of posterior mandibular pores. Upper pore small, non-tubular. Lateral line configuration. Comprising two rows of well separated neuromasts: a dorsolateral row from above opercular lobe to just before anal fin origin with about 16 neuromasts and a mediolateral row from behind pectoral fins to posterior end of body with 28 neuromasts. Dentition. Vomer with eight larger teeth posteriorly and granular teeth in front; posterior half of palatines edentate, anterior half with large teeth in inner row and granular teeth in outer rows, number of rows increasing much anteriorad; premaxillary with few large teeth in inner row and number of granular tooth rows increasing anteriorad; posterior third of dentary edentate, anterior part with many large teeth in inner row and granular teeth as in premaxillary. Otolith (Fig. 10C-D). Otolith very elongate, with a length to height ratio of about 2.5 (60 mm SL). Anterior tip slender, pointed inferiorly with distinct notch above pointed tip; posterior tip slender, sharply pointed centrally. Dorsal rim almost straight with widely separated distinct predorsal and obtuse postdorsal angles; ventral rim gently curving, deepest anterior of the middle. Inner face slightly convex; outer face smooth, slightly concave. Ratio otolith length to sulcus length 2.0. Colliculi separated, though rather indistinctly, flat. Ventral margin of sulcus with a weak notch. Ostium about 3 times the length of cauda. Dorsal depression moderately wide, shallow; ventral furrow distinct, anteriorly close to ventral rim, posteriorly curving upward. Axial skeleton. Tips of all neural and haemal spines pointed. First neural spine half the length of second spine. Vertebrae 3-9 with depressed neural spines. Bases of vertebrae 5-11 enlarged. Parapophyses on aqua vol. 8 no

19 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen posterior 6-11 precaudal vertebrae. Pleural ribs on vertebrae Epipleural ribs indistinct. Anterior anal fin pterygiophore moderately elongated, not reaching parapophses of last precaudal vertebrae. Male copulatory organs. No adult male known. Allometric growth. The 20 mm paratype differs just slightly from the 60 mm holotype in meristic characters and the differences in morphometric characters show the same allometric growth as found in G. longipenis. The neural spines of the precaudal vertebrae are hardly depressed as compared to the holotype. The ventral fin of the paratype is very long (27.5 % SL); the longest ventral fin observed in the remaining 27 specimens of Gunterichthys is 21 % SL. Colour in alcohol. Even after 32 years of preservation, the dorsal, caudal and anal fins are still purplebrown with whitish edges. Body light brown with many small, black dots. Eyes dark. Etymology The species is named after William A. Bussing who most kindly has provided us with many specimens from Costa Rica and Isla de Coco. Distribution Known from off Isla de Cano, Pacific coast of Costa Rica (Fig. 1). Gunterichthys coheni n. sp. (Figs. 5B, 11-12; Tables II, IV) Material examined (12 specimens, mm SL): Holotype: BMNH , male, 38 SL, Eastern Central Pacific Ocean, Panama (no precise locality available), collected by Dr. D. S. Jordan, 15 May Paratypes: BMNH , 1 female, 46 mm SL, and 1 male, 35 mm SL, same data as for holotype; USNM 50398, 1 female and 2 males, mm SL, Pacific Panama, collected by D. S. Jordan, Jan.-Feb. 1896; SIO , 1 female and 1 male, 27 and 32 mm SL, Fort Amador Causeway, Bay of Panama, 8 58 N, W, depth 5 cm, cruise JEM70, st , collected by J. E. McCosker and party, chemfish fish poison, 17 Oct. 1970; SI0 71-3, 2 females, mm SL, Fort Amador Causeway, Bay of Panama, 8 58 N, W, cruise JEM70, st. 70-7, collected by J. E. McCosker, chemfish fish poison, 27 January 1971; ZMUC P771348, 1 male, 32 mm SL, same data as for SIO ; ZMUC P771358, 1 female, 39 mm SL, same data as for USNM Diagnosis Snout pointed; eyes distinct, small ( % SL); head naked; three posterior infraorbital pores; posterior nostril small (about a quarter of eye diameter); rays in dorsal fin 74-77, in anal fin and in caudal 14-15; otoliths with length to height ratio and fused colliculi, sulcus large; one pair of pseudoclaspers with two almost equally long supporters. Similarity Gunterichthys coheni is closest to G. bussingi (see p. 154), and differs mainly by by the naked head (vs. scale-band on cheek), smaller eye ( vs % SL), shorter interorbital width ( vs. 5.5% SL) and less elongate otolith (ratio length to height vs. 2.5). Description Meristic and morphometric characters are presented in Tables II and IV. Head profile steep, snout pointed, mouth slightly inferior. Eyes small, but distinct ( % SL). Maxillaries end far behind eyes; posterior end of maxillaries rounded and with a distinct knob posteroventrally. Both nostrils with unadorned openings, posterior nostril about a quarter of eye diameter, anterior placed close to upper lip. Opercular spine covered by skin. Anterior gill arch with 3-5 small knots on upper branch, one prolonged raker in the angle and lower branch with 4-5 prolonged rakers and 7-8 knots (Fig. 5B). Length of longest gill filaments 2-3 times diameter of eye. Small, imbricate scales on body, about 20 horizontal rows above anal fin origin. Predorsal area and abdomen scaled; pectoral fin base, vertical fins and head naked. Origin of dorsal fin at vertical through origin of pectoral fin. Ventral fin reaches about two third from its base to origin of anal fin. Head sensory pores (Fig. 12A-B). Supraorbital pores 3: Anterior pore about size of 2nd anterior infraorbital pore and 2nd pore small and tubular above and behind eye, 3rd tubular pore at upper termination of gill opening above opercular spine. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small and 2nd and 3rd pores medium in size, covered by dermal flap of upper lip. Posterior infraorbital pores small, about the size of posterior nostril. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large and tubular and without cirrhi, 2nd pore free from lateral skin fold and about same size as posterior mandibular pores and 3rd anterior pore about twice the size of posterior mandibular pores. 1st and 2nd posterior mandibular pores small, about the size of posterior nostril; 3rd pore about twice the size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint openings in a common pore-like cavity, 3rd pore small about the size of posterior mandibular pores and upper pore small, non tubular. Lateral line configuration. Three rows of well separated neuromasts: a short predorsal row between neck and dorsal fin with 4 neuromasts, a dorsolateral row from above opercular lobe to just before anal fin origin with about 15 neuromasts and a medio-lateral row from behind pectoral fins to posterior end of body with about 21 neuromasts. Dentition. Vomer with 10 strong depressible teeth; 157 aqua vol. 8 no

20 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 11. Gunterichthys coheni. Holotype. BMNH , male, 38 mm SL. Fig. 12A-G. Gunterichthys coheni. A) Lateral view of head, BMNH , female 46 mm SL; B) Ventral view of head, holotype; C) Median view of right otolith, BMNH , female, 46 mm SL; D) Ventral view of right otolith; E) Ventral view of male copulatory organs; F) View of left pseudoclasper from outside; G) Inclined lateral view of male copulatory organs. For abbreviations see Fig. 3. palatines edentate on posterior third, with an inner row of large depressible teeth and outer rows granular; premaxillary with edentate posterior third, inner row with teeth increasing in length anteriorad and outer rows granular; dentary with dentition like premaxillary. Otolith (Fig. 12C-D). Otoliths elongate, with a length to height ratio of 2.1 to 2.2 (35-46 mm SL). Anterior tip broad, more or less pointed centrally; posterior tip robust, centrally pointed. Dorsal rim with rounded predorsal angle and irregularly developed, often robust postdorsal angle; ventral rim gently curving, deepest at middle. Inner face slightly convex and outer face smooth, flat to slightly concave. Otolith length to sulcus length ratio 2.1 to 2.3. Sulcus with fused, flat colliculi and ventral sulcus margin with a feeble notch. Dorsal depression narrow, shallow; ventral furrow distinct, anteriorly close the ventral rim of the otolith, posteriorly curving upward and away from it. aqua vol. 8 no

21 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Axial skeleton. Tips of neural and haemal spines pointed. First neural spine half the length of second spine. Vertebrae 3-8 with depressed neural spines. Bases of vertebrae 4-8 enlarged. Parapophyses on posterior 6-11 precaudal vertebrae. Pleural ribs on vertebrae Epipleural ribs indistinct. Anterior anal pterygiophore of equal length in females and males, strongly prolonged. Male copulatory organs (Fig. 12E-G). One pair of large (outer) pseudoclaspers forming a large, extended flap with two distinct supporters, the anterior being almost as long as posterior, otherwise very similar to those of G. longipenis. When in resting position, anterior supporters of pseudoclaspers shielding penis to about two-thirds of its length. Penis long, straight, slender, with pointed tip. Allometric growth. Paratypes differ very little from holotype (see Table IV) except for the smallest specimen (20 mm SL) the morphometric characters of which are not included in Table IV. The values expressed as percentage of SL are: head length 30.0, depth at origin of anal fin 13.0, upper jaw 14.0, horizontal diameter of pigmented eye 1.5, interorbital 4.0, postorbital 21.5, preanal 49.0, predorsal 33.5, from base of ventral fin to anal fin 28.5, length of ventral fin The differences seem to be due to allometric growth. Colour in alcohol: After about 100 years of preservation the specimens are yellowish brown with black eyes. Thirty-three year old specimens are darker uniformly brown. Etymology The species is named after our colleague Daniel M. Cohen, whose many dinematichthyine papers have been of invaluable help to us. Distribution Known from 4 localities off Pacific Panama, Eastern Central Pacific Ocean (Fig. 1). Gunterichthys longipenis Dawson, 1966 (Figs. 5C, 13-14; Tables II, V) Gunterichthys longipenis Dawson, 1966: 205, figs. 1-3 (type locality: north shore of Davis Bayou, off Mississippi Sound). Gunterichthys longipenis: Dawson, 1971: 164; Wagner 1972: 16; Suarez, 1975: 145; Moore, 1975: 156; Cohen and Nielsen, 1978: 59; Sedor, 1985; Tolley and Peebles, 1987: 43; Criscione, 1996 (comparison with Ogilbia); McEachran and Fechhelm, 1998: 745; Nielsen et al., 1999: 132. Material examined (21 specimens, mm SL): Holotype: USNM , male, 44 mm SL, Mississipi Sound, Davis Bay, Mississipi, collected by B. Henson, 27 March Non-types: UF , 3 females and 2 males, mm SL, Mississippi Sound, drainage, N, W, 14 Apr. 1969; ZMUC P77549 and P P771341, 4 spms., mm SL, same data as for UF UF 66665, female, 53 mm SL, drainage in Wakulla County, Kings Bay, off Florida, Gulf of Mexico, collected by J. Rudloe and L. Crum, 20 Sep. 1969; UF , female and 3 juveniles, mm SL, drainage in Wakulla County, Kings Bay, off Florida, Gulf of Mexico, collected by J. Rudloe and L. Crum, 21 Sep. 1969; UF , female, male and 2 juveniles, mm SL, drainage in Monroe County, off Florida, Gulf of Mexico, collected by B. Yokel, 20 July 1972; UF 77175, female, 45 mm SL, drainage in Franklin County, off Florida, Gulf of Mexico, collected by P. Sheridan, 28 Feb. 1978; USNM 50827, female, 40 mm SL, Dry Tortugas, Florida, collected by J. C. Thompson, 22 Jan Diagnosis Snout blunt; eyes distinct, small ( % SL in adults); interorbital in adults % SL, body covered with imbricate scales, head naked; two posterior infraorbital pores; dorsal fin rays 60-71, anal 41-52, caudal 12-15; otolith with fused colliculi, sulcus small; one pair of pseudoclaspers with two almost equally long supporters. Similarity Gunterichthys longipenis (from the Gulf of Mexico) is generally less similar to the two Pacific species (G. bussingi and G. coheni) than they are to each other (Tables IV and V). However, G. longipenis is more similar to G. coheni than to G. bussingi in diameter of eye and pectoral fin length, and more similar to G. bussingi than to G. coheni in interorbital width. Description Meristic and morphometric characters are presented in Tables II and V. The morphometric characters of adults and juveniles are given in two columns as some of the characters show allometric growth. The original description (Dawson, 1966) is very detailed so only the more important characters are included here. Head profile flat, snout blunt. Eyes distinct showing negative allometric growth (Table V). Maxillaries end far behind eyes; two small, rounded knobs ventrally on posterior part of slightly expanded maxilla. Both nostrils with low rim; anterior nostril placed close to upper lip; posterior nostril large, one third of eye. Opercular spine covered by skin. Anterior gill arch with 5-6 prolonged rakers and many small knobs ventrally and dorsally (Fig. 5C). Longest gill filament on anterior arch about twice diameter of eye. Pseudobranchial filaments absent. Small, imbricate scales on body, about 20 horizontal rows above anal fin origin. Predorsal area and abdomen scaled; pectoral fin base, vertical fins and head naked. Origin of dorsal fin at vertical through 159 aqua vol. 8 no

22 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 13. Gunterichthys longipenis. ZMUC 77549, male, 59 mm SL. Fig. 14A-F. Gunterichthys longipenis. A) Lateral view of head, ZMUC P77549, male, 59 mm SL; B) Ventral view of head, ZMUC P771341, female, 55 mm SL; C) View of left pseudoclasper from outside, ZMUC P77549, male, 59 mm SL; D) Inclined lateral view of male copulatory organs; E) Median view of right otolith, ZMUC P771339, female, 63 mm SL); F) Ventral view of right otolith. For abbreviations see Fig. 3. middle of pectoral fin. Origin of anal fin behind mid body. Ventral fin reaching about halfway from its base to anal fin origin. Head sensory pores (Fig. 14A-B). Supraorbital pores 3: Anterior pore small, about size of 2nd anterior infraorbital pore and posterior pore small and tubular above and behind eye, 3rd pore at upper termination of gill opening above opercular spine tubular. Infraorbital pores 5 (3 anterior and 2 posterior): 1st anterior pore small and 2nd and 3rd pores large, covered by dermal flap of upper lip. The posterior infraorbital pores small, about the size of posterior mandibular pores. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore small, non-tubular and without aqua vol. 8 no

23 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Table V. Meristic and morphometric characters of Gunterichthys longipenis. Holotype USNM Non-types Mean and range N Standard length in mm (13-63) 19 Meristic characters Dorsal fin rays (60-71)* 161* Caudal fin rays (12-15)* 160* Anal fin rays (41-52)* 162* Pectoral fin rays 19/ (17-22)* 154* Precaudal vertebrae 12 12(11-13) 14 Total vertebrae (38-40) 14 Long rakers on anterior gill arch 5 5.7(5-6) 14 Total rakers on anterior gill arch (16-20) 11 Pseudobranchial filaments Anterior dorsal fin ray above vertebra no. (D/V) 7 6.5(6-7) 14 Anterior anal fin ray below dorsal fin ray no. (D/A) (21-25) 15 Anterior anal fin ray below vertebra no. (V/A) (15-17) 14 Morphometric characters in % of SL mm SL mm SL 14 specimens 5 specimens Head length ( ) 30.2( ) Head width ( ) 14.8( ) Head height ( ) 18.4( ) Upper jaw length ( ) - Maxillary height ( ) - Diameter of pigmented eye ( ) 2.8( ) Interorbital width ( ) 5.7( ) Postorbital length ( ) 20.0( ) Preanal length (53-57) 49.1(48-50) Predorsal length ( ) 33.5( ) Body depth at origin of anal fin ( ) 14.2( ) Pectoral fin length ( ) - Ventral fin length ( ) 20.0 Base of ventral fin to anal fin ( ) 24.7( ) *including data from Dawson (1971: 164) cirrhi, 2nd pore free from lateral skin fold, about the size of posterior mandibular pores and 3rd pore small, about size of posterior mandibular pores. Three posterior pores about half size of posterior nostril. Preopercular pores 3-4 (2-3 lower and 1 upper): 1st and 2nd pores with joint openings, 3rd lower pore mostly absent, small when present and upper pore small, tubular. Lateral line configuration. Two rows of well separated neuromasts: a dorsolateral row from above opercular lobe to just before anal fin origin with about 14 neuromasts and a mediolateral row from behind pectoral fins to posterior end of body with 23 neuromasts. Dentition. Vomer with strong, depressible teeth; palatines with teeth on anterior half, inner row enlarged, outer rows granular, number of rows increases anteriorad; premaxillary with a few strong teeth near symphysis, other teeth granular; dentary with strong, depressible teeth in inner row, outer rows granular. Otolith (Fig. 14E-F). Otoliths oval in shape, thin, with a length to height ratio of 2.0 to 2.1 (52-63 mm SL). Anterior tip broad, pointed inferiorly; posterior tip narrower, pointed dorsally. Dorsal rim irregular, with rounded predorsal angle and irregularly developed or lacking postdorsal angle; ventral rim gently curving, deepest anterior of middle or at about middle. Inner face almost flat, slightly convex only close to posterior tip of otolith; outer face smooth, flat to slightly concave. Ratio of otolith length to sulcus length 2.3 to 2.5. Sulcus with fused, elevated colliculi, ventral margin of sulcus without notch. Dorsal depression very wide and deep occupying a large portion of dorsal field of inner face; ventral furrow distinct, wide, at some distance from ventral rim of otolith. Axial skeleton. Tips of all neural and haemal spines pointed. First neural spine half the length of second spine. Vertebrae 3-7 with depressed neural spines. Bases of vertebrae 4-9 enlarged. Parapophyses on posterior 7-12 precaudal vertebrae. Pleural ribs on vertebrae Epipleural ribs indistinct. Male copulatory organs (Fig. 14C-D). One pair of large (outer) pseudoclaspers forming an extended flap with two distinct almost equally long supporters. In resting position, the anterior supporters of the pseudoclaspers shield the penis to about half of its length. Penis very long, extending beyond hood, straight, conical, with narrow and pointed tip. Allometric growth. The 19 specimens here examined fall into two length-groups: mm SL (14 adults) and mm SL (five juveniles, newly born). In Table V the morphometric characters are split accordingly showing negative allometric growth in head length, eye diameter and length of ventral fin, and 161 aqua vol. 8 no

24 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 15. Eye diameter-standard length correlation of Ogilbichthys spp. positive in body depth, preanal length and distance from base of ventral fin to anal fin origin. Colour when live: Live color reported as brilliant dark red with dusky fin bases outlined by black (Wagner, 1972). Colour in alcohol: In alcohol body generally yellowish to light brown with distal part of vertical fins greyish brown. Posterior vertical fin borders said to be dark by various authors (Suarez, 1975; Criscione, 1996). Eyeball black. Newly born specimens with many, tiny brown spots on dorsal part of body. Biology G. longipenis burrows into muddy bottom at shallow depth (0-7.6 m) and gives birth to very few, about 15 mm long juveniles in each clutch (Dawson, 1971). One female (UF , 37 mm SL), was found to have four 5-8 mm long embryos. The same specimen had a 3.2 mm bivalve in the intestine. Distribution Known to occur in the Gulf of Mexico, from Texas to the west coast of Florida (Fig. 1). Ogilbichthys n. gen. Type species: Ogilbichthys longimanus n. sp. (type locality: Caribbean Panama, San Blas Archipelago, Cocos-Banderas cays, unnamed central cay, N, W). Diagnosis Anterior nostril placed low on snout; tip of opercular spines free; male copulatory organs with one outer and two inner pairs of pseudoclaspers; a ligament connecting posterior inner pseudoclasper and isthmus; eyes of varable size; scales on cheeks present or absent; otoliths with fused colliculi in 4 of 7 species; anterior anal fin ray pterygiophore long; 3 lower preopercular pores; maxillary knob at rear corner; 3-4 elongated rakers on anterior gill arch. Similarity Compared to other dinematichthyine genera Ogilbichthys differs from Brosmolus Machida, 1993; Dermatopsis Ogilby, 1896; Dermatopsoides Smith, 1948; Dipulus Waite, 1905 Gunterichthys Dawson, aqua vol. 8 no

25 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Fig. 16. Predorsal length-standard length correlation of Ogilbichthys spp and Monothrix Ogilby, 1897 by having 3 pairs of pseudoclaspers (vs. one pair). Brotulina Fowler, 1946; Diancistrus Ogilby, 1899; Dinematichthys Bleeker, 1855; Ogilbia Jordan and Evermann, 1898, Pseudogilbia n. gen. and Typhliasina Whitley, 1951, have one pair inner and one pair outer pseudoclaspers, whereas Ogilbichthys is unique in having two inner pseudoclaspers (anterior and posterior). The anterior pseudoclasper is sometimes sunk into a pocket formed by the fleshy isthmus; it is then invisible unless extracted (Fig. 2C). The posterior inner pseudoclasper is connected via a soft, anteriorly inserted ligament to the isthmus (vs. free in other genera) (Table I). Ogilbichthys most closely resembles Ogilbia but can be separated by a combination of the following characters (smallest juveniles not included): longer predorsal length ( vs % SL), longer pectoral fins ( vs % SL), wider interorbital ( vs % SL), otolith sulcus with fused colliculi (except separated in 3 species, of which, however, none has a notch at ventral sulcus margin whilst all Ogilbia spp. have a notch). The larger species of Ogilbichthys resemble D. minyomma, by having a robust head with broad interorbital, but they are easily separated by the high position of anterior nostril and unique gill raker arrangement in the latter. Pseudogilbia, Typhliasina and G. longipenis have broad heads like the large species of Ogilbichthys, but they all differ by having short ventral fins and head pore reductions. The smaller species of Ogilbichthys (O. ferocis, O. haitiensis and O. tobagoensis), with adult sizes less than 30 mm SL, represent the smallest Dinematichthyini known to date. The new genus contains seven Caribbean and West Altantic species, described below. Etymology The name refers to the general resemblance to Ogilbia, named in honour of the great Australian ichthyologist, J. Douglas Ogilby. 163 aqua vol. 8 no

26 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Key to species 1a Dentary with 2-4 very large teeth in mid section (Fig. 19); dorsal fin rays 56-65; anterior inner pseudoclasper larger or smaller than posterior inner pseudoclasper...2 1b Dentary with larger teeth posteriorad; dorsal fin rays 65-79; anterior inner pseudoclasper smaller than posterior inner pseudoclasper (often submerged in a pocket)...3 2a Few scales on head; anterior inner pseudoclasper larger than posterior inner pseudoclasper forming an inward directed disk, outer pseudoclasper lobate...o. ferocis 2b No scales on head; anterior inner pseudoclasper smaller than posterior inner pseudoclasper forming a backward oriented ridge, outer pseudoclasper triangular...o. haitiensis 3a Upper preopercular pore absent...4 3b Upper preopercular pore present...5 4a Eyes very small % SL (Fig. 15), sunk into head; head large % SL; pectoral fin rays 20-21; anterior inner pseudoclasper submerged; otolith with fused colliculi......o. microphthalmus 4b Eyes larger 2.3% SL, not sunk into head; head small 24.4% SL; pectoral fin rays 18; anterior inner pseudoclasper emerged; otolith with separate colliculi...o. puertoricoensis 5a Scales on head absent; snout with depression above eye; anal fin rays <55; anterior inner pseudoclasper submerged...o. tobagoensis 5b Scales on head present; snout without depression above eye; anal fin rays >54; anterior inner pseudoclasper submerged or emerged...6 6a Postorbital length % SL; pectoral fin length % SL; predorsal length % SL (Fig. 16); scale rows on lower cheeks 2; anterior inner pseudoclasper emerged; outer pseudoclasper wing-like with knob at tip of inner face; otolith with separate colliculi...o. kakuki 6b Postorbital length % SL; pectoral fin % SL; predorsal length % SL (Fig. 16); scale rows on lower cheeks 3; anterior inner pseudoclasper submerged; outer pseudoclasper wing-like without knob at tip of inner face; otolith with fused colliculi...o. longimanus Ogilbichthys ferocis n. sp. (Figs. 6A, 17-19; Tables II, VI) Material examined (39 specimens, mm SL): Holotype: CAS 21666, male, 25 mm SL, Caribbean Panama, San Blas Archipelago, Cocos-Banderas cays, unnamed central cay, N, W, collected with pronoxfish, over corals, 0-5 m, collected by J. E. McCosker and collaborators, 15 May Paratypes: CAS , 11 males and 12 females, mm SL, same data as for holotype; CAS , 6 males and 6 females, mm SL, Caribbean Panama, San Blas Archipelago, Morbeptopo, east end reef edge, N, W, pronoxfish, collected over corals by J. E. McCosker and D. Diener, 13 May 1974; ZMUC P771345, 1 female, 29 mm SL, same data as for CAS ; ZMUC P , 1 male, 23 mm SL and 1 female, 27 mm SL, same data as for holotype. Diagnosis Dorsal fin rays 56-65, anal fin rays 46-51; otolith with fused colliculi. Few large scales on cheek. Dentary with 2-4 large fangs in mid section. Anterior inner pseudoclasper disc-shaped and inwardly-directed, Fig. 17. Ogilbichthys ferocis. Holotype, CAS , male, 25 mm SL. aqua vol. 8 no

27 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Fig. 18A-G. Ogilbichthys ferocis. A) Lateral view of head, CAS , female, 28 mm SL; B) Ventral view of head; C) Median view of right otolith, CAS , female, 28 mm SL; D) Ventral view of right otolith; E) Ventral view of left pseudoclaspers, holotype; F) View of left pseudoclaspers from inside; G) Inclined lateral view of male copulatory organs. For abbreviations see Fig. 3. A B Fig. 19A-B. Radiographs of Ogilbichthys ferocis. A) Holotype, CAS , male, 25 mm SL, note the very elongated first anal fin ray pterygiophore. B) Paratype, CAS , female, 26 mm SL, note the moderately-elongated first anal fin ray pterygiophore and the exposed large fang-like dentary teeth. 165 aqua vol. 8 no

28 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Table VI. Meristic and morphometric characters of Ogilbichthys ferocis. Holotype Holotype + 38 Paratypes CAS Mean and range Mean and range Mean and range Male Males N Females N Sex combined N Standard length in mm (16-27) (14-29) (14-29) 39 Meristic characters Dorsal fin rays (58-65) (56-64) (56-65) 36 Caudal fin rays Anal fin rays (45-51) (46-51) (45-51) 36 Pectoral fin rays (16-19) (16-19) (16-19) 20 Precaudal vertebrae (10-11) (10-11) 39 Total vertebrae (37-39) (37-40) (37-40) 39 Long rakers on anterior gill arch (3-4) 8 3.1(3-4) 11 Pseudobranchial filaments Anterior dorsal fin ray above vertebra no. (D/V) 7 6.9(6-7) (6-7) (6-7) 36 Anterior ray below dorsal fin ray no. (D/A) (16-20) (15-19) (15-20) 35 Anterior anal fin ray below vertebra no. (V/A) (13-14) 17 12(13.1) (13.4)14 35 Morphomeric characters (% of SL) Head length ( ) ( ) ( ) 39 Head width ( ) ( ) ( ) 36 Head height ( ) ( ) ( ) 20 Upper jaw length ( ) ( ) ( ) 20 Maxillary height ( ) ( ) ( ) 33 Diameter of pigmented eye ( ) ( ) ( ) 39 Interorbital width ( ) ( ) ( ) 38 Postorbital length ( ) ( ) ( ) 19 Preanal length ( ) ( ) ( ) 19 Predorsal length ( ) ( ) ( ) 39 Body depth at origin of anal fin ( ) ( ) ( ) 39 Pectoral fin length ( ) ( ) ( ) 37 Pectoral fin base height ( ) 8 5.7( ) 9 5.6( ) 17 Ventral fin length ( ) ( ) ( ) 17 Basis of ventral fin to anal fin origin ( ) ( ) ( ) 20 larger than posterior inner pseudoclasper. Outer pseudoclasper small and lobate; otolith with fused colliculi. Similarity Most similar to O. haitiensis, by the low dorsal fin ray count (56-65 vs. 61), vertebral count (37-40 vs ) and by the long fang-like dentary teeth. O. ferocis differs from O. haitiensis by having scales on cheek (vs. no scales), outer pseudoclasper lobate (vs. triangular), large, disk-shaped anterior inner pseudoclasper (vs. smaller, ridge-shaped). Description Meristic and morphometric characters are presented in Tables II and VI. Body moderately elongate, strongly compressed laterally. Head large ( % SL), snout blunt or rounded, and mouth terminal. Eyes small, partly covered by transparent skin ( % SL). Maxillaries end far behind eyes, dorsal half covered by a prominent dermal lobe of upper lip. Posterior ventral margin of maxillaries slightly expanded and with a small, sharp knob present on the rear ventral part in adult specimens. Anterior nostril positioned low, posterior nostril small, about one third of eye diameter. Opercular spine, pointed, tip free. Anterior gill arch with 3-4 elongated rakers in the angle, and with many small knobs ventrally and dorsally (Fig. 6A). Two very short pseudobranchial filaments. Scales on body in about 20 horizontal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins naked. Head with few (4-6) large scales on upper anterior part of cheeks. Predorsal distance long ( % SL), anal fin origin at about mid body. Pectoral fins moderately long, not reaching vertical through anal opening. Ventral fins long, reaching vertical through anal opening in most specimens. Head sensory pores (Figs. 18A-b). Head pores generally large. Supraorbital pores 3-4: First supraorbital pore about twice the size of 2nd anterior infraorbital pore, 2nd pore (occasionally present) in front of eye and small tubular 3rd pore above and behind eye, posterior pore at upper termination of gill opening above opercular spine tubular. Infraorbital pores 6 (3 aqua vol. 8 no

29 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen anterior and 3 posterior): First anterior pore small and 2nd and 3rd anterior pores slightly larger, covered by dermal flap of upper lip. Third posterior pore considerably larger than the other two. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore small, about the size of 1st posterior mandibular pores, nontubular and without cirrhi, 2nd anterior pore positioned in lateral skin fold, about half the size of posterior mandibular pores and 3rd anterior pore about twice the size of posterior pores. First posterior pore about the size of posterior nostril, 2nd posterior pore about twice and 3rd posterior pore about three times the size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with separate or incompletely joint openings, and 3rd lower pore the size of 3rd posterior infraorbital pore. Upper pore the same size as 3rd lower pore, tubular. Lateral line configuration. Three rows of well separated neuromasts: a short predorsal row from above gill opening to origin of dorsal fin with 6 neuromasts, a dorsolateral row from above opercular lobe to just before anal fin origin and a mediolateral row from behind pectoral fins to posterior end of body. Dentition. Premaxilla with 2 outer rows of granular teeth and a short inner row of 1-3 fangs anteriorly at symphysis. Vomer horseshoe-shaped, with 1-2 rows of small teeth. Palatines with 8-12 teeth in 1-2 irregular rows. Dentary with 5 rows anteriorly, 3 outer rows of granular teeth and 2 inner rows of 6-7 small pointed teeth; more posteriorly a inner row with 2-4 large, curved fangs followed by 8-10 small teeth (Fig. 19). Otolith (Fig. 18C-D). Oval in shape, thin, with a length to height ratio of 2.0 to 2.1 (26-29 mm SL). Anterior tip narrow, sharply pointed; posterior tip robust, broadly rounded. Dorsal rim shallow, with broad predorsal angle and more pronounced postdorsal angle; ventral rim gently curving, deepest posterior of the middle. Inner face slightly convex and outer face smooth, flat to slightly concave. Otolith length to sulcus length ratio 2.5 to 2.7. Elevated colliculi completely fused and ventral margin without a notch. Dorsal depression shallow, narrow, close to dorsal rim of otolith, best developed above and behind the sulcus; ventral furrow distinct, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, with thin and pointed tips. Vertebrae 6-9 with depressed neural spines, shorter in length than spines of vertebrae neural spines with enlarged bases. Parapophyses present from vertebra 6 (6-7) to 11 (10-11). Pleural ribs on vertebrae Epipleural ribs indistinct. Last precaudal vertebra without pleural ribs in 90% of the specimens. First anal fin pterygiophore elongated, reaching tip of or overlapping last precaudal parapophysis (males) or to just below last precaudal parapophysis (females). Sexual dimorphism. Seen in the counts related to the anal fin. Males have one anal fin ray less than females, most clearly shown by number of dorsal fin rays anterior to first anal fin ray (D/A) mean = 17.7 (males) vs (females) and by number of vertebrae anterior to first anal fin ray (V/A) mean =13.8 (males) vs (females). The explanation for this is that the male copulatory organs (penis and pseudoclaspers) are derived from a modified anal fin ray, as suggested by Suarez (1975). The variation conceals the difference in actual anal fin ray counts (48.4 (males) vs (females)), whereas the D/A and V/A counts strongly support Suarez s hypothesis. Male copulatory organs (Fig. 18E-G). Three pairs of small pseudoclaspers, extending only above the base of penis. Outer pseudoclasper forming a simple lobate flap about twice the size of inner pseudoclasper. Anterior inner pseudoclasper disk-shaped, upward and forward oriented. Posterior inner pseudoclasper connected via strong, anteriorly inserted ligament with isthmus. Penis short, curved, with broad base and pointed tip. Allometric growth. Negative allometric growth observed in diameter of eye (Fig. 15) and positive allometric growth in depth of body at anal fin origin. Sexual dimorphism. Present in the following characters: females > males: length of head, upper jaw, postorbital length; males > females: preanal length and distance from ventral fin base to anal fin origin (Table VI). Colour in alcohol: Live colour unknown. Head and body pale to light brown, sometimes darker on dorsal parts. Abdomen light. Upper part of head with a dark brown patch behind the eyes. Anterior upper part of head translucent in smallest juveniles (<20 mm SL), showing brain lobes and otoliths. All fins translucent. Peritoneum, mouth- and branchial cavities light. Distribution Known from two localities in the San Blas Archipelago off Atlantic Panama (Fig. 1). Etymology The name ferocis, meaning fearsome; its fang-like teeth on the middle dentary give the fish a ferocious expression. Ogilbichthys haitiensis n. sp. (Figs ; Tables II, VII ) Material examined (2 specimen, mm SL): Holotype: ANSP male, 22 mm SL, Haiti, Gulf of Gonave; south of St. Marc Point and 1.6 km east of Mount Rouis on Port-au-Prince to St. Marc road, N, W, 1-5 m, collected with rotenone by J. C. Tyler and collaborators 13 Sept Paratype: ANSP , male 16.5 mm SL, Haiti, Gulf of Gonave; St. Marc Channel, off Mount Rouis, 3.2 km south-east of Mount Rouis town; N, W, m, collected with rotenone by J. C. Tyler and collaborators 15 Sept aqua vol. 8 no

30 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 20. Ogilbichthys haitiensis. Holotype, ANSP , male, 22 mm SL. Fig. 21A-E. Ogilbichthys haitiensis. Holotype, ANSP , male, 22 mm SL. A) Lateral view of head; B) Ventral view of head; C) Ventral view of male copulatory organs; D) View of left pseudoclaspers from inside; E) Inclined lateral view of male copulatory organs. For abbreviations see Fig. 3. Diagnosis Dentary with 3-4 large fangs on mid section; head without scales; dorsal fin rays 61, anal fin rays Anterior inner pseudoclasper ridge-like, backward directed, smaller than posterior inner pseudoclasper. The latter with little thorn anteriorly and connected to isthmus. Outer pseudoclasper triangular. Similarity See O. ferocis (p. 164). Description Meristic and morphometric characters are given in Tables II and VII. Body moderately elongate, laterally strongly compressed. Head variable ( % SL), snout rounded, and mouth terminal. Eyes small, partly covered by transparent skin ( % SL). Maxillaries ending far behind eyes, dorsal half of maxillaries covered by a prominent dermal lobe of upper lip. Posterior margin of maxillaries slightly expanded, and with a small thorn-like projection on the rear ventral aqua vol. 8 no

31 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Table VII. Meristic and morphometric characters of Ogilbichthys haitiensis and O. puertoricoensis. O. haitiensis O. puertoricoensis Holotype Paratype Holotype ANSP ANSP ANSP Standard length in mm Meristic characters Dorsal fin rays Caudal fin rays Anal fin rays Pectoral fin rays Precaudal vertebrae Total vertebrae Long rakers on anterior gill arch Pseudobranchial filaments Anterior dorsal fin ray above vertebra no. (D/V) Anterior anal fin ray below dorsal fin ray no. (D/A) Anterior anal fin ray below vertebra no. (V/A) Morphomeric characters (% of SL) Head length Head width Head height Upper jaw length Maxillary height Diameter of pigmented eye Interorbital width Postorbital length Preanal length Predorsal length Body depth at origin of anal fin Pectoral fin length Pectoral fin base height Ventral fin length Base of ventral fin to anal fin origin corner. Anterior nostril positioned low. Posterior nostril small, about 1/6 of eye diameter. Opercular spine pointed with tip free. Anterior gill arch with 3 elongated rakers in the angle and many small knobs ventrally and dorsally. Two pseudobranchial filaments present. Scales large, about 15 horizontal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins and head without scales. Dorsal fin origin at a vertical through middle of pectoral fin; anal fin origin at about mid-body. Pectoral fins short, not reaching a vertical through anal fin origin. Ventral fins long, almost reaching the hood. Head sensory pores (Fig. 21a-b). Supraorbital pores 3: First pore small, about half the size of 2nd anterior infraorbital pore and small 2nd pore hardly visible above and behind eye, posterior pore at upper termination of gill opening above opercular spine small and tubular. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small and 2nd and 3rd anterior pores moderate in size, covered by dermal flap of upper lip. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore small, non-tubular and without cirrhi, 2nd anterior pore positioned in lateral skin fold, small, about half the size of posterior mandibular pores and 3rd anterior pore about twice the size of posterior mandibular pores. First and 2nd posterior mandibular pores small, about the size of posterior nostril; 3rd posterior pore larger, about twice the size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint openings, and 3rd lower preopercular pore small, the size of posterior infraorbital pores. Upper pore large, about twice the size of 3rd lower pore, tubular. Lateral line configuration. Indistinct. Dentition. Premaxilla with 2 outer rows of granular teeth and a short inner row of 2 fangs at symphysis. Vomer horseshoe-shaped, with 1 row of small teeth. Palatines with 1 row of small teeth. Dentary with 4 rows anteriorly, 3 outer rows of granular teeth and one inner row of small pointed teeth; more posteriorly the inner row is provided with 3-4 large, curved fangs followed by small teeth posteriorly (same arrangement as in O. ferocis (Fig. 19)). Otolith. Dissolved by formalin. Axial skeleton. Neural and haemal spines slender, with thin and pointed tips. Vertebrae 6-9 with depressed neural spines, shorter in length than spines of vertebrae 2-5. Bases of neural spines 6-11 enlarged. Parapophyses present from vertebra 6-9. Pleural ribs on vertebrae Epipleural ribs 169 aqua vol. 8 no

32 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera indistinct. Last precaudal vertebra without pleural ribs. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis. Male copulatory organs (Fig. 21C-E). Three pairs of small pseudoclaspers, only just reaching base of penis. Outer pseudoclasper forming a simple triangular flap about twice the size of inner pseudoclaspers. Anterior inner pseudoclasper ridge-like, backward oriented. Posterior inner pseudoclasper with little thorn anteriorly and connected to isthmus with a very broad ligament. Penis short, curved, with broad base and pointed tip. Colour in alcohol: Live colour unknown. Head and body light brown, with dark brown tiny pigment spots. Upper part of head with a dark brown patch behind eyes. All fin translucent. Peritoneum, mouth and branchial cavities light. Distribution Presently only known from Haiti (Fig. 1). Etymology Named after the type locality Haiti. Remarks Ogilbichthys haitiensis is known from only two small male specimens, but the differences in pseudoclaspers are so distinct from O. ferocis that establishment of a new species is warranted. Ogilbichthys kakuki n. sp. (Figs. 6D, 22-24, Tables II, VIII) Material examined (21 specimens, mm SL): Holotype: ANSP , male, 56 mm SL, Haiti, Gulf of Gonave; St. Marc Channel, off Mount Rouis, 3.2 km south-east of Mount Rouis town, N, W, collected at 0-2 m, with rotenone, by J. C. Tyler and collaborators, 14 Sept Paratypes: ANSP , 1 male, 46 mm SL, Haiti, Gulf of Gonave; Sable Island, off south-east coast of Gonave Island, N, W, 3-5 m, collected with rotenone, by J. C. Tyler and T. Devany, 19 Sept. 1967; ANSP , 1 male, 47 mm SL and 2 females, 52 and 65 mm SL, Bahamas, Crooked Island, isolated coral head directly off Pitts Town Point, north-western tip of island, N, W, 0-4 m, collected with rotenone by J. E. Böhlke and collaborators, 1 June 1962; ANSP , 1 female, 39 mm SL, Colombia, Old Providence Island, east of Crab Cay, approximately 13 N, 81 W, 8-10 m, collected with rotenone, by J. C. Tyler and collaborators, 5 Aug. 1969; ANSP , 1 female, 66 mm SL and 1 male, 51 mm SL, Bahamas, Green Cay, north of Rose Island, coral head about 400 m north of centre of cay; N, W, 15 m, collected with rotenone, by J. E Böhlke and collaborators, 13 Nov. 1955; ANSP , 1 female, 35 mm SL, same data as for holotype; ANSP , 1 male, 46 mm SL, 3 females, 39, 56 and 60 mm SL, Bahamas, San Salvador, Cat-O-Way, approximately 24 N, W, collected by Hancock and Deacon, 8 June 1968; ANSP ; 1 female, 36 mm SL, Bahamas, Athol Island, south shore near west end, N, W, 0-6 m, collected with chemfish, by J. E. Bohlke and family, 21 Aug. 1969; ANSP , 1 male, 53 mm SL and 3 females, 42, 51 and 56 mm SL, Haiti, Port-au-Prince Bay, Pelican Cays, east edge of cay just off top of massive coral reef, N, W, 2-3 m, collected with rotenone by J. C. Tyler and collaborators, 13 Sept. 1967; ZMUC P , 2 males, 47 and 67 mm SL, same data as for ANSP ; ZMUC Fig. 22. Ogilbichthys kakuki. Holotype, ANSP , male, 56 mm SL. aqua vol. 8 no

33 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Fig. 23A-F. Ogilbichthys kakuki. Holotype, ANSP , male, 56 mm SL. A) Lateral view of head; B) Ventral view of head; C) Inclined lateral view of male copulatory organs; D) View of left pseudoclaspers from inside; E) Median view of right otolith, ZMUC P771364, male, 59 mm SL; F) From ventral side. For abbreviations see Fig. 3. Fig. 24. Ogilbichthys kakuki. Paratype, ZMUC P771364, male, 59 mm SL. Live specimen in Angelfish Blue Hole, Stocking Island, Great Exuma, Bahamas prior to capture. Photo by T. Iliffe, Texas A&M University at Galveston. 171 aqua vol. 8 no

34 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Table VIII. Meristic and morphometric characters of Ogilbichthys kakuki. Holotype Holotype + 20 Paratypes ANSP Mean and range Standard length in mm (35-67) Meristic characters Dorsal fin rays (70-79) Caudal fin rays Anal fin rays (53-61) Pectoral fin rays (18-21) Precaudal vertebrae (10-11) Total vertebrae (40-42) Long rakers on anterior gill arch 3 3.1(3-4) Pseudobranchial filaments 2 2 Anterior dorsal fin ray above vertebra no. (D/V) 6 6.1(6-7) Anterior anal fin ray below dorsal fin ray no. (D/A) (18-22) Anterior anal fin ray below vertebra no. (V/A) (13-14) Morphomeric characters in % of SL Head length ( ) Head width ( ) Head height ( ) Upper jaw length ( ) Maxillary height ( ) Diameter of pigmented eye ( ) Interorbital width ( ) Postorbital length ( ) Preanal length ( ) Predorsal length ( ) Body depth at origin of anal fin ( ) Pectoral fin length ( ) Pectoral fin base height ( ) Ventral fin length ( ) Base of ventral fin to anal fin origin ( ) P , male, 59 mm SL, Bahamas, Angelfish Blue Hole, Stocking Island, near Great Exuma, approximately 23 5 N, W, collected by hand net, about 230 m inside the cave, at depth of 26 m, by B. Kakuk and T. Iliffe, 7 Jan Diagnosis Outer pseudoclasper wing-like with distinct knob at tip of inner face, isthmus between pseudoclaspers narrower than base of penis. Broad anterior inner pseudoclasper. Scale band on cheek with 1 to 2 rows on lower cheek. Otolith with separate colliculi. Similarity Ogilbichthys kakuki resembles both O. longimanus and O. microphthalmus (see p. 173 and 176). Description Meristic and morphometric characters are presented in Tables II and VIII. Body laterally compressed, high ( % SL at anal fin origin) and robust. Head long ( % SL), wide and rounded. Lower jaw broad, spatulate in large specimens. Maxillaries ending far behind eyes, dorsal margin of maxillaries covered by upper lip dermal lobe, posterior end wide, with prominent knob on rear ventral part. Anterior nostril positioned low. Posterior nostril small, about 1/3 to 1/4 the size of eye. Opercular spine pointed, with free tip. Anterior gill arch with 3 (3-4) elongate rakers in the angle, with small knobs 9 (8-11) ventrally and 3 (2-3) dorsally. Pseudobranchial filaments 2. Scales embedded in skin, large (about 1.3 mm in diameter at midbody and with 25 horizontal rows above anal fin origin in a 56 mm SL specimen). Predorsal area, abdomen and outer pectoral fin base scaled. Vertical fins and inner pectoral fin base naked. Narrow scale band on cheeks with 3 to 4 rows dorsally and 1 to 2 rows ventrally. Predorsal length short (Fig. 16), anal fin origin just before mid-body. Pectoral fins and ventral fins long, almost reaching a vertical through anal fin origin. Head sensory pores (Fig. 23A-B). Generally small; Supraorbital pores 3: First pore about twice the size of 2nd anterior infraorbital pore and small, tubular 2nd pore above and behind eye, 3rd pore at upper termination of gill opening above opercular spine tubular. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small and 2nd and 3rd anterior pores, covered by dermal flap of upper lip. Three posterior pores small, smaller than posterior mandibular pores and about equal in size, the 3rd slightly tubular. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore small and without cirrhi, 2nd anterior pore positioned in lateral skin fold, small, about half the size of posterior mandibular pores and 3rd anterior pore small, about twice the size of posterior mandibular pores. Three posterior pores small, about half the size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with aqua vol. 8 no

35 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen joint openings, and 3rd lower pore small about the size of posterior infraorbital pores. Upper pore small, tubular. Lateral line configuration. Two rows of well separated dark neuromasts: a dorsolateral row of 14 neuromasts and a mediolateral row with almost indistinct neuromasts. The end of the dorsolateral row and the origin of the mediolateral row is situated above level of the anal opening. Dentition. Teeth straight, pointed and small. Premaxilla with 6 rows of teeth anteriorly and 3 rows posteriorly. Teeth in inner row large anteriorly, in outer rows granular, except for a few larger teeth near symphysis. Vomer horseshoe-shaped, with 4 rows of teeth, larger in inner row, smaller in outer rows. Palatine teeth in 2 rows, teeth in inner row larger than in outer. Dentary with 6 rows anteriorly and 2 rows posteriorly. Inner row with large teeth, outer 1-5 rows with small to granular teeth. Otolith (Fig. 23E-F). Otoliths elongate, with a length to height ratio of about 2.0 (59 mm SL). Anterior tip pointed, posterior tip expanded, robust. Dorsal rim with broad, rounded predorsal and very feeble postdorsal angles; ventral rim gently curving, highest anterior of its middle. Inner face markedly convex, outer face smooth and flat. Otolith length to sulcus length ratio about 2.0. Sulcus short, with separated, slightly elevated colliculi; ostium about 3 times the length of cauda; ventral sulcus margin without notch. Dorsal depression shallow; ventral furrow distinct, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, except for neural spines of vertebrae 6-11 with blunt tips. Vertebrae 5-9 with depressed neural spines, shorter in length than spines of 2nd-4th vertebrae. Bases of neural spines 5-9 enlarged. Parapophyses present from vertebra 6 (6-7) to 11 (10-11). Pleural ribs on vertebrae 2-10, absent on last precaudal vertebra in all but one specimen. Epipleural ribs indistinct. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis (males) or to just below last precaudal parapophysis (females). Male copulatory organs (Fig. 23C-D). Three pairs of pseudoclaspers of moderate size. Outer pseudoclasper twice as long as posterior inner pseudoclasper, extending across half of penis, wing-like, with distinct knob at tip of inner face. Anterior inner pseudoclasper broad. Posterior inner pseudoclasper with a large forward pointing thorn anteriorly and a broad lobe posteriorly, connected with a fleshy flaplike ligament to isthmus. Isthmus much narrower than base of penis. Penis long, curved, with broad base and pointed tip. Allometric growth. Negative allometric growth in head length and eye diameter. Sexual dimorphism not observed. Colour when live (Fig. 24): One specimen observed in Angelfish Blue Hole, Bahamas, uniformly pale yellow. Colour in alcohol: Uniformly light brown. All fins translucent, except for innermost 1/4 of pectoral fins, with brown pigmentation. Peritoneum, mouth- and branchial cavities light. Etymology Ogilbichthys kakuki is named after Mr. Brian Kakuk, Diving Safety Officer, Caribbean Marine Research Center, Lee Stocking Island, Bahamas, who kindly presented us with a newly-caught specimen. Distribution Known from 7 localities from off Colombian Old Providence Island, Haiti and Bahamas (Fig. 1). Biology This species has mostly been recorded from coral reefs, but has also been caught 230 mm inside an ocean blue hole in the Exuma Cays, Bahamas. The eye is not particularly small, indicating that it is not a very specialized cave species. A 61 mm SL female from San Salvador Isl. contains 50 immature embryons (length 4 mm) and 40 eggs between mm in diameter. Ogilbichthys longimanus n. sp. (Figs. 6B, 25-26; Tables II, IX) Material examined (23 specimens, mm SL): Holotype: ANSP male, 68 mm SL, Bahamas, Green Cay, north of Rose Island, about 400 m north of centre of cay, N, W, collected on a coral head at depth of 15 m, with rotenone, by J. E. Böhlke and collaborators, 13 Nov Paratypes: ANSP , 2 males, 42 and 64 mm SL, Bahamas, Green Cay (North of Rose Island), collected over coral head about 400 m NNW of cay, N, W, 0-15 m, collected with pronoxfish, by J. E. Böhlke and collaborators, 13 May 1955; ANSP , 1 female, 39 mm SL, same data as for holotype; ANSP , 1 female, 37 mm SL and 2 males, 46 and 68 mm SL, Haiti, Gulf of Gonave, South of St. Marc Point, 1.6 km east of Mount Rouis on Port-au-Prince to St. Marc road, N, W, collected at depth of 1-5 m, with rotenone, by J. C. Tyler and collaborators, 13 Sept. 1963; ANSP , 2 males, 48 and 70 mm SL, Haiti, Gulf of Gonave; St. Marc Channel, off Mount Rouis, 3 km south-east of Mount Rouis town, N, W, collected at depth 0-2 m, with rotenone, by J. C. Tyler and collaborators, 14 Sept. 1963; ANSP , 9 females, 33, 33, 35, 48, 53, 56, 59, 60 and 67 mm SL, 2 males, 58 and 69 mm SL, Cayman Island, Grand Cayman Island, West side of island, Paradise Rocks, offshore from North side of Georgetown, m, collected with chemfish, by C. 173 aqua vol. 8 no

36 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 25. Ogilbichthys longimanus. Holotype, ANSP , male, 68 mm SL. Fig. 26A-F. Ogilbichthys longimanus. A) Lateral view of head, holotype, ANSP ; B) Ventral view of head; C) View of left pseudoclaspers from inside, ZMUC P771350, male, 59 mm SL; D) Inclined lateral view of male copulatory organs; E) Median view of right otolith, ANSP , female, 67 mm SL; F) Ventral view of right otolith. Abbreviations see Fig. 3. aqua vol. 8 no

37 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Table IX. Meristic and morphometric characters of Ogilbichthys longimanus. Holotype Holotype +22 Paratypes ANSP Mean and range Standard length in mm (33-73) Meristic characters Dorsal fin rays (70-79) Caudal fin rays (15-16) Anal fin rays (55-64) Pectoral fin rays (17-20) Precaudal vertebrae Total vertebrae (40-43) Long rakers on anterior gill arch 3 3.0(3-4) Pseudobranchial filaments 2 2 Anterior dorsal fin ray above vertebra no. (D/V) 7 6.9(6-7) Anterior anal fin ray below dorsal ray no. (D/A) (17-21) Anterior anal fin ray below vertebra no. (V/A) (13-14) Morphomeric characters in % of SL Head length ( ) Head width ( ) Head height ( ) Upper jaw length ( ) Maxillary height ( ) Diameter of pigmented eye ( ) Interorbital width ( ) Postorbital length ( ) Preanal length ( ) Predorsal length ( ) Body depth at origin of anal fin ( ) Pectoral fin length ( ) Pectoral fin base height ( ) Ventral fin length ( ) Base of ventral fin to anal fin origin ( ) R. Gilbert and J. C. Tyler, 28 Oct. 1960; ZMUC P , 2 males, 43 and 59 mm SL, same data as for holotype; SIO 67-45, 1 female, 73 mm SL, Caribbean Panama, Toro Point, N, W, collected at depth 0-5 m, with rotenone, by I. Rubinoff and party, 23 March Diagnosis Pectoral fins long ( % SL), soft interorbital broad ( % SL), predorsal distance long ( % SL), elongate neural spines 4, rows of large scales on lower cheeks 3, outer pseudoclasper a broad, rounded lobe, fleshy isthmus between pseudoclaspers wide, anterior inner pseudoclasper submerged in pocket of isthmus; otolith with fused colliculi. Similarity O. longimanus resembles O. kakuki in most morphometric and meristic characters and in the large size (max. 73 and 67 mm SL, respectively). It differs from O. kakuki by the longer pectoral fin ( vs % SL), longer predorsal length ( vs % SL) (Fig. 16), one more elongated neural spine (4 vs. 3), more rows of scales on the lower cheeks (3 vs. 2), different shape of outer pseudoclasper (Fig. 26C-D vs. Fig. 23C-D), submerged anterior inner pseudoclasper (vs. emerged), broader fleshy isthmus between pseudoclaspers and the otolith with fused colliculi (vs. separate). Description Meristic and morphometric characters are presented in Tables II and IX. Body moderately elongate, laterally compressed, deep and robust. Head long, wide and rounded. Eyes large ( % SL) compared to most congeners. Maxillaries end far behind eyes. Posterior-ventral margin robust and expanded; a distinct knob is present at the rear ventral part. Anterior nostril positioned low, posterior nostril small, about 1/4 of eye diameter. Opercular spine, pointed, tip free. Anterior gill arch with 3 (3-4) elongated rakers in angle and 9 (7-11) small knobs on lower part and 2 (2-4) small knobs on upper part of the arch (Fig. 6B). Pseudobranchial filaments 2. Scales large (about 1 mm in diameter at mid-body), about 25 horizontal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Caudal fin and pectoral fins with a few scale rows anteriorly, vertical fins naked. Head with a band of large scales on cheeks consisting of 3 rows of scales on upper and lower cheek. Predorsal distance long ( % SL), anal fin origin just before mid-body. Pectoral and ventral fins long, almost reaching level of anal fin origin. Head sensory pores (Fig. 26A-B). Head pores generally small. Supraorbital pores 3: First pore moderate in size, about the size of 2nd anterior infraorbital pore and small 2nd, tubular pore above and behind eye, 3rd pore at upper termination of gill opening above 175 aqua vol. 8 no

38 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera opercular spine tubular. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small and 2nd and 3rd anterior pores moderate in size, covered by dermal flap of upper lip. Posterior infraorbital pores small, about the size of posterior mandibular pores. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large, tubular, with little cirrhus in front, 2nd anterior pore positioned in lateral skin fold, small, about half the size of posterior mandibular pores, and 3rd anterior pore about three times the size of posterior mandibular pores. Posterior mandibular pores small, about half the size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint openings, and 3rd lower pore large, about twice the size of posterior mandibular pores. Upper preopercular pore about as large as 3rd lower pore, tubular. Lateral line configuration. Two rows of well separated neuromasts: a dorsolateral row from above opercular lobe to just before anal fin origin with 17 neuromasts and a mediolateral row from behind pectoral fins to posterior end of body with about 27 neuromasts. Dentition. Premaxilla with 4 outer rows of granular teeth and 1-2 inner rows with small pointed teeth. Anterior teeth of all rows enlarged at symphysis. Vomer horseshoe-shaped, with 2 rows of large teeth. Palatines with 2 irregular rows of large teeth. Dentary anteriorly with 6 teeth rows: 4 outer rows of granular teeth and two inner rows of regular pointed teeth. Posteriorly, 3 rows remain, teeth of inner row being much larger than those of outer rows. The teeth of inner row gradually increasing in size. Otolith (Fig. 26E-F). Oval in shape, with length to height ratio of 1.9 to 2.0 (48-67 mm SL). Anterior tip moderately pointed (more strongly pointed in smaller specimens); posterior tip broader, directed dorsally. Dorsal rim with broad, rounded predorsal and sharper postdorsal angles; ventral rim a gentle curve, its greatest height at about its middle. Inner face strongly convex and outer face smooth and flat. Otolith length to sulcus length ratio about 2.0. Elevated colliculi fused, ventral marging without notch. Dorsal depression shallow; ventral furrow distinct, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, except for depressed neural spines of vertebrae 6-10 with blunt tips. Spines of vertebrae 2-5 elongate. Bases of neural spines 5-11 enlarged. Parapophyses on vertebrae 6 (6-7) to 11 (10-11). Pleural ribs on vertebrae Last precaudal vertebra without pleural ribs in all but one specimen. Epipleural ribs indistinct. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis (males) or just below precaudal parapophysis (females). Male copulatory organs (Fig. 26C-D). Three pairs of pseudoclaspers. Outer pseudoclasper an expanded rounded lobe with broad base and about three times the size of posterior inner pseudoclasper. The latter with a prominent forward-pointing thorn anteriorly, connected with a fleshy flap to isthmus. Isthmus as wide as base of penis. Very small anterior inner pseudoclasper sunk into a pocket of the isthmus, thus invisible unless extracted. Penis much longer than outer pseudoclasper, curved, with broad base and pointed tip. Allometric growth. Positive allometric growth was observed in width of head, height of posterior maxilla and preanal length. Negative allometric growth observed in diameter of eye and pectoral fin length. Sexual dimorphism. Preanal length longer in males ( , x = 48.4 vs , x = 46.1% SL) and pectoral fin longer in females ( , x = 19.1 vs , x = 18.1% SL). Colour in alcohol: Live colour unknown. Head and body uniformly pale to light brown, fins lighter than head and body. Etymology The name longimanus refers to the long pectoral fins, which are longer than in any other American dinematichthyine. Distribution Known from 5 localities off Cayman Islands, Bahamas and Haiti (Fig.1). Ogilbichthys microphthalmus n. sp. (Figs. 6C, 27-28) Material examined (8 specimens, mm SL): Holotype: ANSP , male, 31 mm SL, Haiti, Gulf of Gonave; St. Marc Channel, off Mount Rouis, 3.2 km south-east of Mount Rouis town, N, W, collected at depth 0-2 m, with rotenone, by J. C. Tyler and collaborators, 14 Sept Paratypes: ANSP , 2 females, 24 and 35 mm SL, Cayman Island, Grand Cayman Island, West side of island, Paradise Rocks, offshore from North side of Georgetown, at depth 6-8 m, collected with chemfish, by C. R. Gilbert and J. C. Tyler, 28 Oct. 1960; ANSP , 1 female, 32 mm SL, same data as for holotype; ZMUC P , 2 females, 37 and 38 mm SL, same data as for ANSP ; USNM , 1 female, 31 mm SL and 1 male, 32 mm SL, Honduras, Bay Islands, Guanaja (Bonaca) Island, N, W, 0-9 m, collector and method unknown, 21 April Diagnosis Eyes very small (diameter % SL) and sunk into head; no upper preopercular pore; head robust (length % SL); narrow scale band on cheek with 1 to 2 scale rows on lower part; anterior inner pseudoclasper submerged in pocket of isthmus; otolith with fused colliculi. aqua vol. 8 no

39 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Fig. 27. Ogilbichthys microphthalmus. Holotype, ANSP , male, 31 mm SL. Fig. 28A-F. Ogilbichthys microphthalmus. A) Lateral view of head, ZMUC P771351, female, 38 mm SL; B) Ventral view of head; C) View of left pseudoclaspers from inside, holotype; D) Inclined lateral view of male copulatory organs; E) Median view of right otolith, ZMUC P771352, female, 38 mm SL; F) Ventral view of right otolith. For abbreviations see Fig. 3. Similarity Ogilbichthys microphthalmus resembles O. longimanus, O. puertoricoensis and O. kakuki in the broad head although it does not seem to attain their size. It resembles O. kakuki and O. puertoricoensis in the few scale rows on lower cheek and O. longimanus in the shape of the small, simple flap-like outer pseudoclasper and the small, submerged, almost invisible anterior inner pseudoclasper. It is easily distinguished from all three by the very small, sunken eyes (Fig. 15), from O. longimanus and O. kakuki further by the lack of the upper preopercular pore and from O. puertoricoensis and O. kakuki also by the fused colliculi. Description Meristic and morphometric characters are presented in Tables II and X. Body moderately elongate. Head large ( % SL), wide and rounded. Eyes very 177 aqua vol. 8 no

40 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Table X. Meristic and morphometric characters of Ogilbichthys microphthalmus. Holotype Holotype +7 Paratypes ANSP Mean and range Standard length (24-38) Meristic characters Dorsal fin rays (66-72) Caudal fin rays Anal fin rays (53-57) Pectoral fin rays (20-21) Precaudal vertebrae (10-11) Total vertebrae (40-41) Long rakers on anterior gill arch 3 3.2(3-4) Pseudobranchial filaments 2 2 Anterior dorsal fin ray above vertebra no. (D/V) 6 6 Anterior anal fin ray below dorsal fin ray no. (D/A) (17-20) Anterior anal fin ray below vertebra no. (V/A) (13-14) Morphomeric characters in % of SL Head length ( ) Head width ( ) Head height ( ) Upper jaw length ( ) Maxillary height ( ) Diameter of pigmented eye ( ) Interorbital width ( ) Postorbital length ( ) Preanal length ( ) Predorsal length ( ) Body depth at origin of anal fin ( ) Pectoral fin length ( ) Pectoral fin base height ( ) Ventral fin length ( ) Base of ventral fin to anal fin origin ( ) small, sunk into head ( % SL). Maxillaries end far behind eyes. Posterior ventral margin of maxillary strongly expanded and with a small, distinct knob at the rear ventral part. Anterior nostril positioned low. Posterior nostril moderate in size, about 1/3 of eye diameter. Opercular spine short, pointed with tip free. Anterior gill arch with 3 (3-4) elongate rakers at the angle, with 8 (8-10) small knobs ventrally and 3 (2-4) dorsally. Pseudobranchial filaments 2. Scales large (about 0.7 mm at mid-body in holotype), with 18 horizontal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins naked. Head with a band of large scales on cheeks; 2-3 rows dorsally and 1-2 rows ventrally. Predorsal length short ( % SL); anal fin origin well before mid-body. Pectoral fins elongate, reaching behind vertical through anal fin origin. Ventral fins, almost reaching anal fin origin. Head sensory pores (Fig. 28A-B). Most head pores small. Supraorbital pores 3: 1st pore moderate in size, about the size of 2nd anterior infraorbital pore, 2nd pore small and tubular, placed above and behind eye, 3rd pore at upper termination of gill opening above opercular spine tubular. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small, 2nd and 3rd anterior pores slightly larger, covered by dermal flap of upper lip. Posterior infraorbital pores small, about the size of posterior mandibular pores. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large, tubular, anteriorly open and without cirrhi, 2nd anterior pore positioned in lateral skin fold, small, about half the size of posterior mandibular pores, 3rd anterior mandibular pore about four times the size of posterior pores. Posterior mandibular pores small, about half the size of posterior nostril. Preopercular pores 3 (3 lower, upper absent): 1st and 2nd pores with joint openings, and 3rd pore moderate in size, nearly twice the size of posterior infraorbital pores. Lateral line configuration. Indistinct. Dentition. Premaxilla with 5 rows of teeth anteriorly and 3 rows posteriorly. Teeth in inner row large anteriorly, in outer rows granular, except for a few teeth anteriorly near symphysis. Vomer horseshoe-shaped, with 2 rows of teeth, larger in inner row. Palatine teeth in 1-2 irregular rows. Dentary with 4 rows anteriorly and 2 rows posteriorly. Inner row with fang-like teeth posteriorly, larger than outer 1-3 rows with small or granular teeth. Otolith (Fig. 28E-F). Robust, oval in shape, with a length to height ratio of about 1.9 (in fish of 38 mm SL). Anterior tip moderately pointed; posterior tip directed dorsally. Dorsal rim broadly rounded anteriorly, posteriorly with a small postdorsal angle; ventral rim a gentle curve, its greatest height at about its middle. Inner face strongly convex and outer face smooth, nearly flat. Otolith length to sulcus length ratio 1.8. Sulcus with fused, elevated colliculum and ventral margin undivided. Dorsal depression indistinct; ventral furrow distinct, close to ventral rim of otolith. aqua vol. 8 no

41 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Axial skeleton. Neural and haemal spines slender, except for short depressed neural spines of vertebrae 5-9 with blunt tips and enlarged bases. Parapophyses present from vertebra 6 (6-7) to 10 (10-11). Pleural ribs on vertebrae 2-10, absent on last precaudal vertebra in 2 of 6 specimens. Epipleural ribs indistinct. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis (male) or to just below last precaudal parapophysis (females). Male copulatory organs (Fig. 28C-D). Three pairs of small pseudoclaspers, extending only to the base of penis. Outer pseudoclasper a small flap about three times the size of posterior inner pseudoclasper. The latter with a small forward-pointing thorn anteriorly, and connected posteriorly with a ligament to isthmus. Isthmus as wide as base of penis. Very small anterior inner pseudoclasper sunk into a pocket of the isthmus, thus invisible without extraction. Penis much longer than outer pseudoclasper, curved, with broad base and broad tip. Colour in alcohol: Live colour unknown. Uniformly brownish, head slightly darker than body. All fins translucent. Etymology The name micropthalmus refers to the minute eyes. Distribution Known from 3 localities off Haiti, Honduras and Cayman Islands (Fig. 1). Biology The reduced eyes indicate that O. micropthalmus could be a cave species, but so far it has been caught on coral reefs only. A 31 mm SL female had a 3.7 mm long bilobed ovary containing 50 eggs, with average diameter of 0.4 mm. Ogilbichthys puertoricoensis n. sp. (Figs , Tables II, IIV) Material examined (1 specimen, 34 mm SL): Holotype: ANSP male, Puerto Rico, Mona Island, N, W, collected over sand and rock covered with calcareous algae, in shallow caves at shore, at depth of 0-1 m, by W. F. Smith- Vaniz and P. L. Colin, 6 October 1978, method of collection unknown. Diagnosis Head and predorsal length short (24.2 and 31.1% SL, respectively), ventral fins long (29.2% SL); scale band on cheeks with 2 rows of scales on lower cheek; upper preopercular pore absent, outer pseudoclasper small, anterior inner pseudoclasper ridge-like, bent backwards, anteriorly connected to outer pseudoclasper. Posterior inner pseudoclasper much larger than anterior one; otolith with separated colliculi. Similarity Ogilbichthys puertoricoensis resembles O. microphthalmus by lacking an upper preopercular pore, but differs in several other characters such as larger eye (2.3 vs % SL), fewer pectoral fin rays (18 vs ) and well-developed anterior inner pseudoclasper (vs. submerged). O. puertoricoensis resembles O. tobagoensis by having separated colliculi, but differs by having more anal fin rays (57 vs ), no notch on snout, presence of an extruding anterior inner pseudoclasper and in several morphometric characters (e.g. shorter head 24.4 vs % SL and predorsal length 31.3 vs % SL). O. puertoricoensis resembles O. kakuki in the pseudoclasper pattern and by the separated colliculi, but differs by the morphometric characters mentioned in the diagnosis. Fig. 29. Ogilbichthys puertoricoensis. Holotype, ANSP , male, 34 mm SL. 179 aqua vol. 8 no

42 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Fig. 30A-F. Ogilbichthys puertoricoensis. Holotype, ANSP , male, 34 mm SL. A) Lateral view of head; B) Ventral view of head; C) View of left pseudoclaspers from inside; D) Inclined lateral view of male copulatory organs; E) Median view of right otolith; F) Ventral view of right otolith. For abbreviations see Fig. 3. Description Meristic and morphometric characters are presented in Tables II and VII. Body elongate, high, laterally compressed. Head short (24.4% SL), maxillaries end far behind eyes, dorsal margin covered by a prominent dermal lobe of upper lip; posterior end wide, with a small knob ventrally. Anterior nostril small, positioned low; posterior nostril small, about 1/4 the size of eye. Opercular spine pointed with free tip. Anterior gill arch with 3 short knobs, 1 developed raker on upper part and 2 developed rakers and 9 knobs on lower part. The developed rakers long. Pseudobranchial filaments 2. Scales large (about 0.4 mm in diameter at mid-body), about 25 horizontal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins naked. Head with narrow scale band on cheeks, 3 rows dorsally and 2 rows ventrally. Short predorsal length, anal fin origin before mid-body. Pectoral fins moderately long, almost reaching vertical through anus. Ventral fins very long, reaching behind anal fin origin. Head sensory pores (Fig. 30A-B). Supraorbital pore 2: 1st about twice the size of 2nd anterior infraorbital pore, 2nd at upper termination of gill opening above opercular spine small and tubular. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small, 2nd and 3rd anterior pores small, covered by dermal flap of upper lip. Posterior infraorbital pores small, about half the size of posterior mandibular pores. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore small, non-tubular, located distant from and lateral to tip of lower jaw, without cirrhi, 2nd anterior pore positioned in lateral skin fold, small, about half the size of posterior mandibular pores and 3rd anterior pore very small, about the size of posterior mandibular pores and widely separated. Posterior mandibular pores small, about the size of posterior nostril. Preopercular pores 3 (3 lower, upper absent): 1st and 2nd lower pores with joint opening, 3rd lower pore small, about the size of posterior infraorbital pores. Lateral line configuration. Indistinct. Dentition. Premaxilla with 4 outer rows of granular teeth and one inner row with small pointed teeth. Teeth at symphysis slightly enlarged. Vomer horseshoeshaped, with 2 rows of small teeth. Palatines with 1 row of small teeth. Dentary with 5 teeth rows anteriorly; 4 outer rows with granular and two inner rows with small pointed teeth. Posteriorly, 2 rows are present, the teeth of the inner row being much larger than those of outer rows. Teeth of inner row gradually increase in size. Otolith (Fig. 30E-F). The otolith is slightly affected by formalin. Oval in shape, with a length to height ratio of about 2.0 (34 mm SL). Anterior tip moderately aqua vol. 8 no

43 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen pointed, posterior tip more robust. Dorsal rim with broad, rounded predorsal and postdorsal angles; ventral rim gently curving, deepest at about its middle. Inner face of otolith strongly convex and outer face smooth and flat. Otolith length to sulcus length ratio about 2.0. Sulcus short, with separated, elevated colliculi; ostium about 3 times the length of cauda; ventral sulcus margin with an indistinct notch. Dorsal depression shallow; ventral furrow very distinct, particularly posteriorly, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, except for neural spines of vertebrae 5-10 with blunt tips. Vertebrae 6-9 with depressed neural spines, shorter in length than spines of 2nd to 4th vertebrae. Bases of neural spines 4-9 enlarged. Parapophyses from vertebra 6. Pleural ribs on vertebrae Epipleural ribs indistinct. First anal fin pterygiophore moderately elongated, not reaching tip of last precaudal parapophysis. Male copulatory organs (Fig. 30C-D). Three pairs of pseudoclaspers. Outer pseudoclasper small, extending only to base of penis, simple wing-like. Anterior inner pseudoclasper ridge-like, bent backwards and anteriorly connected to the outer pseudoclasper. Posterior inner pseudoclasper about the size of outer pseudoclasper, thick, anteriorly connected by an inserted ligament to the isthmus. Penis much longer than outer pseudoclasper, curved, with broad base and pointed tip. Colour in alcohol: Live colour unknown. Head and body uniformly pale to light brown, except for a dark patch on top of the skull; fins lighter than head and body. Etymology The name puertoricoensis is named after the type locality, Puerto Rico. Distribution Known from a single specimen collected off Puerto Rico (Fig. 1). Biology Unlike most congeners, this species was caught on sand and rock covered with calcareous algae and not on coral reefs. Remarks Ogilbichthys puertoricoensis is known from a single male specimen, but the differences in pseudoclaspers, head pores and morphometrics are so distinct from the other Ogilbichthys species that the designation of a new species is warranted. Ogilbichthys tobagoensis n. sp. (Figs ; Table XI) Material examined (7 specimens, mm SL): Fig. 31. Ogilbichthys tobagoensis. Holotype, USMN , male, 23 mm SL. 181 Holotype: USNM , male, 23 mm SL, Tobago, Pirates Bay, North of Charlotteville, Man-Of-War Bay, N, W, collected off rocky shore over corals and sand in depth 0-5 m, by J. T. Williams and party, 5 September Paratypes: USNM , 1 female, 22 mm SL and 1 juvenile, 15 mm SL, same data as for holotype; USNM , 1 male, 24 mm SL, Tobago, Bloody Bay, Mouth of Bloody Bay River and adjacent rock point, N, W, rocky wall and sandy bottom, collected at depth of 0-3 m, by J. T. Williams and party, 13 September 1990; USNM , 1 male, 18 mm SL, Tobago, Buccoo Reef, N, W, collected over coral, sand and rubble at depth of m, by J. T. Williams, and party, 10 Sepaqua vol. 8 no

44 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Table XI. Meristic and morphometric characters of Ogilbichthys tobagoensis. Holotype Holotype + USNM Paratypes Mean and range Standard length in mm (15-29) Meristic characters Dorsal fin rays (65-72) Caudal fin rays Anal fin rays (50-54) Pectoral fin rays (17-19) Precaudal vertebrae Total vertebrae (38-40) Long rakers on anterior gill arch 3 3 Pseudobranchial filaments 2 2 Anterior dorsal fin ray above vertebra no. (D/V) 6 (6.2)6-7 Anterior anal fin ray below dorsal fin ray no. (D/A) (18-20) Anterior anal fin ray below vertebra no. (V/A) (13-14) Morphomeric characters in % of SL Head length ( ) Head width ( ) Head height ( ) Upper jaw length ( ) Maxillary height ( ) Diameter of pigmented eye ( ) Interorbital width ( ) Postorbital length ( ) Preanal length ( ) Predorsal length ( ) Body depth at origin of anal fin ( ) Pectoral fin length ( ) Pectoral fin base height ( ) Ventral fin length ( ) Base of ventral fin to anal fin origin ( ) Fig. 32A-F. Ogilbichthys tobagoensis. A) Lateral view of head, holotype, USNM , male, 23 mm SL; B) Ventral view of head, USNM , male, 24 mm SL; C) View of left pseudoclaspers from inside, holotype; D) Inclined lateral view of male copulatory organs; E) Median view of right otolith, USNM , male, 24 mm SL; F) Ventral view of right otolith. For abbreviations see Fig. 3. aqua vol. 8 no

45 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen tember 1990; USNM , 1 male, 29 mm SL, Tobago, Leeward side of little Tobago Island, off Speyside, N, W, in a rock-walled channel with rubble bottom, at depth of 10 m, J. T. Williams and party, 6 September 1990; ZMUC P771357,1 female, 29 mm SL, same data as for USNM Diagnosis Snout with depression above eye; no scales on cheek; dorsal fin rays 67-72, anal fin rays 46-51; otolith with separated colliculi. Anterior part of posterior inner pseudoclasper thorn-like, directed forwards. Similarity Ogilbichthys tobagoensis resembles the other small species of the genus (O. ferocis and O. haitiensis) in most morphometric characters, but has higher meristic counts (e.g. dorsal fin rays vs and 61) (Table II) and lacks fangs at the middle of the dentary. It shares the lack of scales on cheek with O. haitiensis, but its pseudoclasper pattern is more similar to that of O. longimanus and O. microphthalmus with small and simple outer and inner pseudoclaspers. O. tobagoensis, O. kakuki and O. puertoricoensis are the only species of Ogilbichthys known to have otoliths with separated colliculi. Description Meristic and morphometric characters are presented in Tables II and XI. Body elongate, laterally strongly compressed. Head profile with a depression on snout above eye. Anterior nostril positioned low. Posterior nostril moderate in size, about 1/3rd of eye diameter. Opercular spine, pointed with tip free. Anterior gill arch with 3 elongated rakers in the the angle, 2 small knobs on upper branch, and 7-8 small knobs on lower branch. Pseudobranchial filaments 2. Squamation with 20 horizontal scale rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Head and vertical fins naked. Predorsal distance long ( % SL), anal fin origin just anterior to mid-body. Pectoral fins long, ending well behind dorsal fin origin. Ventral fins reaching a vertical through anal opening anal opening. Head sensory pores (Fig. 31A-B). Most head pores large. Supraorbital pores 3: 1st pore about size of 2nd anterior infraorbital pore, 2nd pore above and behind eye and 3rd pore above opercular spine. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small and 2nd and 3rd anterior pores large, covered by dermal flap of upper lip. Posterior infraorbital pores small, about a third in size of posterior mandibular pores. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large, tubular, with a single cirrhus anteriorly, 2nd anterior pore positioned in lateral skin fold, small, about half the size of posterior mandibular pores, 3rd anterior pore about size of posterior mandibular pores. 1st and 2nd posterior pores about size of posterior nostril, 3rd posterior pore about twice the size of posterior nostril. Preopercular pores 4 (3 lower and 1 upper): 1st and 2nd lower pores with joint opening, 3rd lower the size of posterior infraorbital pores. Upper pore tubular. Lateral line configuration. Indistinct. Dentition. Premaxilla with 2 outer rows of granular teeth and one inner row with small pointed teeth. Teeth at symphysis slightly enlarged. Vomer horseshoeshaped, with 1-2 rows of small teeth. Palatines with 1 row of small teeth. Dentary with 4 teeth rows anteriorly; 2 outer rows with granular and 2 inner rows with small pointed teeth. Posteriorly 2 rows are present; teeth of inner row gradually increase in size posteriorad. Otolith (Fig. 32E-F). Oval in shape, with length to height ratio 2.3 to 2.4 (in fishes of mm SL). Anterior tip narrow, sharply pointed; posterior tip broader. Dorsal rim shallow, with broad predorsal angle and more pronounced postdorsal angle; ventral rim a gentle curve, its greatest height at its middle. Inner face almost flat and outer face smooth, slightly convex. Otolith length to sulcus length ratio about 2.4. Ventral sulcus margin undivided, ostium and cauda nevertheless recognized by separated, flat colliculi. Ostium about 4 times the length of cauda. Dorsal depression shallow; ventral furrow distinct, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, with thin, pointed tips. Vertebrae 5-8 with depressed neural spines, shorter in length than spines of vertebrae 2-4. Bases of neural spines 6-11 enlarged. Parapophyses present on vertebrae 6 to 11. Pleural ribs on vertebrae Epipleural ribs indistinct. Male copulatory organs (Fig. 32C-D). Three pairs of pseudoclaspers. Outer pseudoclasper forming a simple wing-like flap about three times the size of posterior inner pseudoclasper and expanding over base of penis. Posterior inner pseudoclasper anteriorly with small, but distinct, forwardly directed thorn, posterior part a simple flap connected to isthmus with a ligament; a tiny anterior inner pseudoclasper sunk into a pocket of the isthmus, thus invisible without extraction. Penis not much longer than outer pseudoclasper, curved, with broad base and pointed tip. Colour in alcohol: Live colour unknown. \Light brown with dorsal parts of head and body slightly darker than flanks, abdomen light. All fins translucent. Peritoneum, mouth and branchial cavities light. Etymology The species tobagoensis is named after the type locality, Tobago. Distribution Known from 4 localities at Tobago (Fig. 1). 183 aqua vol. 8 no

46 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Pseudogilbia n. gen. Type species: Pseudogilbia sanblasensis n. sp. (type locality: Caribbean Panama, San Blas Archipelago, Morbeptopo, N, W). Diagnosis Anterior nostril placed low on snout; tip of opercular spine free; male copulatory organs with two pairs of equally sized pseudoclaspers, larger than in all other American dinematichthyine species; inner pseudoclasper formed like a forward-pointing hook; eyes large (2.9% SL); head with broad scale patch on cheek; otolith with fused colliculi; 1 lower preopercular pore; maxillary knob at rear corner, 3 elongated rakers on anterior gill arch; anterior anal fin pterygiophore long; predorsal length long (35.3% SL), ventral fins short (16.4% SL), ending well in front of anal opening. Similarity Pseudogilbia differs from all other dinematichthyine genera and species, in having only a single lower preopercular pore (vs. 2-3 in Gunterichthys, 3 in others). Pseudogilbia resembles Gunterichthys in having short ventral fins, but differs in having 3 developed gill rakers (vs. 5-7 in Gunterichthys), 2 pairs of pseudoclaspers (vs. 1 in Gunterichthys) and a free opercular spine (vs. hidden in Gunterichthys). Pseudogilbia resembles Ogilbia in most morphometric characters and in having free inner pseudoclasper, but in Ogilbia the inner and outer pseudoclaspers are different in shape and size. The two genera also differ in Pseudogilbia having fused colliculi (vs. separate in Ogilbia), long predorsal length 35.3 % SL (vs. short % SL in Ogilbia). Pseudogilbia resembles Dinematichthys minyomma and Ogilbichthys spp., in some morphometric characteristics (e.g. moderate pectoral fin length in D. minyomma ( % SL) and long predorsal length in Ogilbichthys ( % SL). However, P. sanblasensis clearly differs from D. minyomma in the low position of the anterior nostil (vs. high), the fused colliculi (vs. separate) and the larger eye 2.9% SL (vs % SL). P. sanblasensis differs from Ogilbichthys in having free inner pseudoclaspers (vs. connected by a ligament to the isthmus between the pseudoclaspers), the absence of an additional anterior inner pseudoclasper (vs. present) and by the smaller pectoral fin (14.3 vs % SL) and ventral fin (16.4 vs % SL). The genus is monotypic. Etymology The name refers to the general resemblance to Ogilbia, named in honour of the Australian ichthyologist, J. Douglas Ogilby. Pseudogilbia sanblasensis n. sp. (Figs. 33, 34; Tables II, XII) Material examined Holotype: CAS , male, 66 mm SL, Caribbean Panama, San Blas Archipelago, Morbeptopo, collected at east end of reef edge, N, W, with pronoxfish, by J. E. McCosker and D. Diener, 13 May Diagnosis See generic diagnosis. Similarity See paragraph on similarity between Pseudogilbia and other genera. Fig. 33. Pseudogilbia sanblasensis. Holotype, CAS , male, 66 mm SL. aqua vol. 8 no

47 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Table XII. Meristic and morphometric characters of Pseudogilbia sanblasensis. Holotype CAS Standard length in mm 66 Meristic characters Dorsal fin rays 69 Caudal fin rays 16 Anal fin rays 56 Pectoral fin rays 20 Precaudal vertebrae 11 Total vertebrae 41 Long rakers on anterior gill arch 3 Pseudobranchial filaments 2 Anterior dorsal fin ray above vertebra no. (D/V) 7 Anterior anal fin ray below vertebra no. (V/A) 22 Anterior anal fin ray below vertebra no. (V/A) 15 Morphomeric characters in % of SL Head length 25.7 Head width 14.1 Head height 16.6 Upper jaw length 12.9 Maxillary height 4.3 Diameter of pigmented eye 2.9 Interorbital width 5.8 Postorbital length 19.2 Preanal length 52.1 Predorsal length 35.3 Body depth at origin of anal fin 18.1 Pectoral fin length 14.3 Pectoral fin base height 5.3 Ventral fin length 16.4 Base of ventral fin to anal fin origin 33.3 Fig. 34A-E. Pseudogilbia sanblasensis. Holotype, CAS , male, 66 mm SL. A) Lateral view of head; B) Ventral view of head; C) View of left pair of pseudoclaspers from inside; D) Median view of right otolith; E) Inclined lateral view of male copulatory organs. For abbreviations see Fig aqua vol. 8 no

48 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Description Meristic and morphometric characters are presented in Tables II and XII. Body moderately elongate, laterally compressed. Head long and wide, mouth subterminal. Small, imbricate scales on body and proximal part of dorsal fin, head with broad patch of small scales on cheeks, reduced to just 2-3 scale rows on lower cheeks behind termination of maxilla. Eye diameter large (2.9% SL), snout rounded. Maxillaries ending behind eyes, dorsal margin covered by a prominent dermal lobe of the upper lip. Posterior ventral margin expanded and angular. A sharp projection present on the rear ventral part. Anterior nostril positioned low. Posterior nostril small, about 1/4 the size of the eye. Opercular spine, pointed with tip free. Anterior gill arch with 3 elongate rakers in the angle, in addition to 10 ventral and 3 dorsal small knobs. Pseudobranchial filaments 2. Scales imbedded in skin, large (0.8 mm at mid-body), 20 horizontal rows above anal fin origin. Predorsal area, abdomen and pectoral fin base scaled. Vertical fins naked. Broad scale patch on cheeks. Anal fin origin at about midbody. Ventral fins short, ending well before of anal opening. Head sensory pores (Fig. 34A-B). Supraorbital pores 3: 1st pore about size of 2nd anterior infraorbital pore, 2nd pore small, indistinct and tubular, above and behind eye, 3rd pore at upper termination of gill opening above opercular spine tubular. Infraorbital pores 6 (3 anterior and 3 posterior): 1st anterior pore small, 2nd and 3rd pores large, covered by dermal flap of upper lip. Posterior pores about the size of posterior mandibular pores and tubular. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore small and tubular, with a single cirrhus anteriorly, 2nd pore positioned in lateral skin fold, about size of posterior mandibular pores and 3rd pore large, about three times the size of posterior mandibular pores. Posterior mandibular pores very small, about 1/3rd the size of posterior nostril. Preopercular pores 2 (1 lower and 1 upper): lower pore tubular, about the size of posterior mandibular pores and probably homologous to the 3rd lower preopercular pore of other species. Upper pore small and tubular. Lateral line configuration. Two rows of well separated neuromasts: a dorsolateral row of about 14 neuromasts and a mediolateral row of about 25 neuromasts. The end of the dorsolateral row and the origin of the mediolateral row is situated above level of pseudoclasper origin. Dentition. Teeth straight, pointed and small. Premaxilla with 5 rows of teeth anteriorly and 3 rows posteriorly. Inner 1-2 rows large, outer rows granular, except for a few teeth near symphysis. Vomer horseshoe-shaped, with 2 rows of small teeth. Palatines with small teeth in 3 irregular rows, teeth in inner row larger than in outer. Dentary with 8 rows anteriorly and 2 rows posteriorly. Inner row with 15 large teeth, outer rows with small to granular teeth. Otolith (Fig. 34D). Oval in shape, with a length to height ratio of about 2.0. Anterior tip sharply pointed; posterior tip more robust. Dorsal rim with a deep curve, without prominent predorsal and postdorsal angles; ventral rim a gentle curve, its greatest height at about its middle. Inner face markedly convex; outer face smooth, flat. Ratio otolith length to sulcus length about 2.5. Sulcus with undivided margin and fused, elevated colliculi. Dorsal depression shallow, wide; ventral furrow distinct, close to ventral rim of otolith. Axial skeleton. Neural and haemal spines slender, with thin, pointed tips, except vertebrae 7-11 with blunt neural spines. Vertebrae 6-9 with depressed neural spines, number 6-10 shorter in length than spines of 2-5 vertebrae. Bases of neural spines 6-11 enlarged. Parapophyses present from vertebrae Pleural ribs on vertebrae 2-10 and epipleural ribs on vertebrae 2-9. Last precaudal vertebra without pleural ribs. First anal fin pterygiophore elongated, reaching tip of last precaudal parapophysis. Male copulatory organs (Fig. 34C, E). Two pairs of very large and elongated pseudoclaspers of about equal size; outer pseudoclasper an elongate, broad flap composed of a large supporter and a much smaller, separated supporter more anteriorly. Inner pseudoclasper about size and shape of main body of outer pseudoclasper, however, with a deep angular notch anteriorly giving it a hook-like shape and a thickened base. Penis long, curved, with broad base and slender tip with a distal groove. Colour in alcohol: Live colour unknown. Light brown, head and upper 1/3 of body darker than flanks. Abdomen light. All fins translucent, except for weakly pigmented innermost 1/4-1/2 of vertical fins and anterior 1/3 of pectoral fins. Peritoneum, mouth and branchial cavities light. Etymology The name sanblasensis refers to the type locality, San Blas Archipelago, Caribbean Panama. Distribution Known from a single specimen from the Caribbean Panama, San Blas Archipelago, Morbeptopo (Fig. 1). Typhliasina Whitley, 1951 Typhliasina Whitley, 1951: 67 for preoccupied Typhlias. Typhlias Hubbs, 1938: 287 (type species T. pearsei Hubbs, 1938: 291, type locality: pool in Balaam Canche Cave near Chichen Itza, Yucatán, Mexico), preoccupied by a rotifer Typhlias Bryce, Diagnosis Anterior nostril placed low on snout; tip of opercular spine free; two pairs of pseudoclaspers, inner pseudoclasper positioned in front of outer pseudoaqua vol. 8 no

49 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen clasper; eyes not visible (eyes minute black dots in specimens less than 20 mm SL); head naked; otolith with fused colliculi; 3 lower preopercular pores; maxillary strongly vertically expanded posteriorly, knob at rear corner; 5-7 prolonged rakers on anterior gill arch; anterior anal fin ray pterygiophore elongated; no pseudobranchial filaments. Similarity Typhliasina resembles Ogilbichthys and Gunterichthys in the broad and robust head and the fused colliculi of the otoliths. It differs from Ogilbichthys in having only 2 posterior infraorbital pores (vs. 3), no upper preopercular pore (vs. present), lack of head squamation (vs. scales on cheeks) and 5-8 long gill rakers on anterior gill arch (vs. 3); and from Gunterichthys by the absence of the upper preopercular pore (vs. present), the free opercular spine (vs. hidden below skin) and 2 pairs of pseudoclaspers (vs. 1 pair). A monotypic genus. Remarks Typhliasina was considered a junior synonym of Ogilbia by Cohen and Nielsen (1978: 60) and Nielsen et al. (1999:134). However, the evidence presented in this study suggests that Typhliasina must be resurrected as the genus can be distinguished by the following: Two posterior infraorbital pores below and behind eye (vs. 3 in Ogilbia), otoliths with fused colliculi (vs. separated colliculi in Ogilbia), upper preopercular pore absent (vs. present) and 5-8 long rakers on anterior gill arch (vs. 2-4). Typhliasina pearsei (Hubbs, 1938) (Figs. 5D, 35-36, and back cover; Table II, XIII) Typhlias pearsei Hubbs, 1938: 291, pl. 3 (type locality Balaam Canche Cave, Yucatan). Ogilbia pearsei: Cohen and Nielsen 1978: 60; Nielsen et al. 1999: 134; Proudlove et al. 2001: 214; Nielsen 2003: Ogilbia (Typhliasina) pearsei: Wilkens et al. 1989: 130. Typhliasina pearsei: Whitley 1951: 67; Solorzano 1953: 286; Thinès 1969: 172; Schemmel 1977: 191; Wilkens 1982: 262; Cumba-Segura 1983: 1; Nielsen 2002: Material examined (33 specimens, mm SL): Holotype: UMMZ Collected in pool in Balaam Canche Cave near Chichen Itza, Yucatán, Mexico, N, W, collected by Dr. A.S. Pearse, 70 m within the cave, June 8, (Holotype not examined). Paratype: UMMZ , 1 juvenile, 41 mm SL, (same locality as the holotype). Non-types (all from Yucatan, Mexico): UMMZ , female, 81 mm SL, Cenote X-ebiz, Hoctun, N, W, 26 Feb. 1971; UMMZ , 1 male, 79 mm SL, Grutas de Tzab-Nah, N, W, 26 June 1975; USNM , 1 female, 36 mm SL, Carwash Cenote, Tulum, N, W, collected on the surface by T. Iliffe, 12 Nov. 1986; ZMH 7314, 3 females, 1 male, 2 sex unknown, mm SL, Cenote Dzah Nab, 4.8 km east and 0.8 km south of Chichen Itza, collected by H. Wilkens and J. Parzefall, Sept. 1987; ZMH 7351, 1 female, 4 sex unknown, mm SL, Cenote Picarote, collected by H. Wilkens, 14 Feb. 1998; ZMH 7352 and ZMUC P771336, 2 females, mm SL, locality unknown, collected by H. Wilkens, 6 Mar. 1882; ZMH 7353, 1 female, 85 mm SL, locality Fig. 35. Typhliasina pearsei. ZMUC , male, 88 mm SL. 187 aqua vol. 8 no

50 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera unknown, collected by H. Wilkens, 8 Mar. 1982; ZMH 7354 and ZMUC P771337, 2 males, mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 1975; ZMH 7355, 1 female, 89 mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 2 May 1975; ZMH 7356, 1 female, 53 mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 28 May 1975; ZMH 7357, 1 female, 63 mm SL and 1 male, 81, mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 2 June 1975; ZMH 7358, 1 female, 78 mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 2 May 1975; ZMH 7359, 1 female, 83 mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 6 June ZMH 7360, 1 male, 91 mm SL, Cueva del Pochote, about N, W, collected by H. Wilkens, 1 May 1975; ZMH 7361, 1 female, 64 mm SL, Balaam Canche Cave, N, W, collected by H. Wilkens, 1975; ZMH 7362, 1 male, head missing, Balaam Canche Cave, N, W, collected by H. Wilkens, 15 Mar. 1975; ZMH 7363, 1 female, 42 mm SL, Balaam Canche Cave, N, W, collected by H. Wilkens, 15 Mar. 1975; ZMH 7364, 2 females, mm SL, Balaam Canche Cave, N, W, collected by H. Wilkens, 15 Mar. 1975; ZMH 7365, 1 female, 72 mm SL, Balaam Canche Cave, N, W, collected by H. Wilkens, 15 Mar. 1975; ZMH 7366, 1 female, 69 mm SL, Balaam Canche Cave, N, W, collected by H. Wilkens, 3 Aug Fig. 36A-H. Typhliasina pearsei. A) Lateral view of head, ZMH 9360, male, 91 mm SL; B) Ventral view of head, UMMZ , male, 79 mm SL; C) View of left pair of pseudoclaspers from inside, ZMUC P771337, male, 88 mm SL; D) Inclined lateral view of male copulatory organs; E) Anterior view of right otolith, ZMH 7314c, male, 97 mm SL; F) Median view of right otolith; G) ventral view of right otolith; H) Median view of right otolith, ZMH 7314a, female, 78 mm SL. For abbreviations see aqua vol. 8 no

51 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen Table XIII. Meristic and morphometric characters of Typhliasina pearsei. Holotype Paratype Paratype + non- Numbers type specimens examined*** UMMZ UMMZ Mean and range * ** Standard length in mm (28-97) 30 Meristic characters Dorsal fin rays (75-87) 26 Caudal fin rays (13-14) 24 Anal fin rays (59-68) 26 Pectoral fin rays (18-22) 18 Precaudal vertebrae Total vertebrae (40-43) 28 Long rakers on first gill arch 6 6.1(5-7) 22 Pseudobranchial filaments Anterior dorsal fin ray above vertebra no. (D/V) (7-8) 28 Anterior anal fin ray below dorsal fin ray no. (D/A) (18-25) 27 Anterior anal fin ray below vertebra no. (V/A) (14-17) 28 Morphometric characters in % of SL Head length ( ) 22 Head width ( ) 5 Head height ( ) 4 Upper jaw length ( ) 20 Maxillary height ( ) 6 Body depth at anal fin origin ( ) 22 Preanal length (50-58) 22 Predorsal length ( ) 22 Ventral fin length ( ) 10 Base of ventral fin to anal fin ( ) 20 * From original description and radiographs. ** Shrivelled condition. *** The four smallest specimens (18-19 mm SL) not included. Diagnosis See generic diagnosis. Similarity See paragraph on generic similarity above. Description Meristic and morphometric characters are presented in Tables II and XIII. Head large ( % SL) with broad snout. Eyes not externally visible in adults and larger juveniles. Maxillaries vertical expanded posteriorly, with ventral corner formed as pointed, bony process. Anterior nostril with low rim, the larger posterior nostril with unadorned aperture. Opercular spine heavy, largely covered by skin, but tip distinctly free. Upper branch of anterior gill arch with 3-4 small knobs and lower branch with 5-8 long rakers and 3-8 small knobs (Fig. 5D). Long rakers quite variable in length in adult specimens. Scales on body large (1.6 mm at mid-body), 18 horizontal rows above anal fin origin in a 83 mm SL specimen. Head, pectoral fin base and vertical fins naked; abdomen scaled. Predorsal area naked, except for a few scattered scales. Predorsal distance long ( % SL), origin of anal fin behind mid-body. Pectoral fins short, not reaching vertical through anal fin origin. Ventral fins short, very thin (often broken) reaching halfway to 2/3 from their base to vertical through origin of anal fin. Head sensory pores (Fig. 36A-B). Supraorbital pores 2, 1st anterior about size of 2nd anterior infraorbital pore, 2nd pore at upper termination of gill opening above opercular spine large and tubular. Infraorbital pores 4 (2 anterior and 2 posterior). Anterior pores about 2 times as large as small posterior pores, not covered by dermal flap of upper lip. Mandibular pores 6 (3 anterior and 3 posterior): 1st anterior pore large, non-tubular and without cirrhi, 2nd anterior pore positioned in lateral skin fold and of about same size as 1st posterior mandibular pore, 3rd pore about 1.5 times the size of posterior mandibular pores. First two posterior mandibular pores about same size as posterior nostril, 3rd about times the posterior nostril. Preopercular pores 3 (lower): 1st and 2nd pores with clearly separate openings, each about as large as 2nd posterior mandibular pore, 3rd smaller, about the size of posterior infraorbital pores. Many small sensory papillae on head. Lateral line configuration. Distinct with anterior part placed dorsally and from anal fin in a medio-lateral line. For a detailed description see Schemmel (1977, Fig. 1). Dentition. All six specimens less than 30 mm SL are without teeth, a 42 mm SL specimen with few fangs anteriorly on premaxillary and all specimens larger than 53 mm SL with many retrorse fangs. Dentary and premaxillary with fangs in inner row and granular teeth in outer rows. Vomer with fangs and granular 189 aqua vol. 8 no

52 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera teeth. Dentition of palatines varying considerably with 3 teeth in a single row in the 41 mm SL paratype and teeth in 3 rows in a 88 mm SL male, the larger teeth in inner row. Otolith (Fig. 36E-H). Elongate, robust, with a length to height ratio of 1.9 to 2.0 (in specimens mm SL). Anterior tip blunt with inferior angle; posterior tip variable, but narrower than anterior tip and more pointed. Dorsal rim with marked predorsal and postdorsal angles, in large specimens with deep central concavity in between; ventral rim deeply curving, its greatest height anterior of the middle. Inner face flat in direction of horizontal axis and slightly convex in direction of vertical axis; outer face smooth, ventrally thickened. Ratio otolith length to sulcus length Sulcus with completely fused, flat to elevated colliculi, ventral margin undivided. Axial skeleton (from radiographs). Tips of anterior eight neural spines blunt, the rest pointed. First neural spine 1/2-2/3 the length of second spine. Vertebrae 3-9 with depressed neural spines. Bases of vertebrae 5-9 enlarged. Parapophyses on posterior five precaudal vertebrae. Pleural ribs on vertebrae Epipleural ribs indistinct. Anterior anal pterygiophore strongly prolonged. Male copulatory organs (Fig. 36C-D). Two pairs of robust, large and widely-separated pseudoclaspers. Outer pseudoclasper a wing-like flap with a thick supporter about twice the size of inner pseudoclasper. Inner pseudoclasper completely separated from and placed anterior to outer pseudoclasper; a thick supporter extends distally in the flap in a broad knob-like structure. Isthmus between pseudoclaspers as wide as or wider than base of penis. Penis not much extending in length over outer pseudoclasper, curved, with broad base and broad tip. Radiographs show that five of the six known adult males have ossified pseudoclaspers. Ontogenetic changes and allometric growth. The four smallest specimens examined (18-19 mm SL) are indistinct from adults, except for the following characters: no scales and teeth developed, small (0.2 mm) but distinct, black eyes visible and few, stubby rakers on anterior gill arch (8-10). In two specimens (28-29 mm SL) the eyes cannot be seen, there are no scales or teeth and the number of gill rakers is 8-9. Two specimens (36-42 mm SL) have an adult number of gill rakers (totally knobs and 6 longer rakers), developed teeth but no scales. Specimens larger than 53 mm SL show no juvenile characters. The length of the gill filaments (1.5 mm) on the first gill arch is the same in specimens 28, 53 and 83 mm SL which is indicative of negative allometric growth. Colour when live: According to Hubbs (1938: 291) in life, the 90.5 mm SL holotype was clear white, becoming pinkish along the posterior margins and the 41 mm paratype was translucent white, turning dark on preservation. Colour in alcohol: All examined specimens (except the paratype) are light yellow and none (including six specimens smaller than 30 mm) have turned dark on preservation. Distribution Known from caves and sinkholes in the Yucatan Peninsula, Mexico (Fig. 1). Biology The following information is based on observations made in Yucatan by H. Wilkens, ZMH: Specimens were observed exclusively in caves. They are associated with the blind synbranchid fish Ophisternon infernale (Hubbs, 1938) and several blind crustacean species. In only a single cave, was T. pearsei observed together with the catfish Rhamdia guatemalensis (Günther, 1864) (Wilkens, 1982: 239). They did not react to flashlights but darted away at the slightest vibration. The newly born juveniles are light yellow and about 20 mm long. There are 2-4 specimens in a batch. T. pearsei seems to feed on troglobitic shrimps and mysids (Schemmel, 1977). Acknowledgements We wish to thank the following persons for helping with material and information: Gerald R. Allen (WAM); M. Eric Anderson (SAIAB), George Burgess (UF), William Bussing (UCR), David Catania (CAS), Daniel M. Cohen (CAS), William L. Fink (UMMZ), Butch Hulley (SAM), Tom Iliffe (Texas A&M University at Galveston), Jon Fong (CAS); Berry Hutchins (WAM); Tomio Iwamoto (CAS), Susan Jewett (USNM), Brian Kakuk (Caribbean Marine Research Center, Bahamas), Jeff Leis (AMS), James Maclaine (NHM former BMNH), Mark McGruther (AMS), Sue Morrison (WAM), V. Mthombeni (SAIAB), Kerryn Parkinson (AMS), Robert Peuchot (Bruxelles), Jack Randall (BPBM); Sandra Raredon (USNM), Sally Reader (AMS), Ross Robertson (Panama), Rob Robins (UF), Mark Sabaj (ANSP), Gudrun Schultze (ZMH), Jeff Seigel (LACM), David G. Smith (USNM), H. J. Walker (SIO), Horst Wilkens (ZMH), Jeff. T. H. Williams (USNM) and Rick Winterbottom (ROM). We thank Daniel M. Cohen (CAS), Douglas F. Markle (Oregon State University) and Nigel Merrett (fromer BMNH) for valuable comments on the earlier version of the manuscript. Also thanks to the following colleagues at ZMUC: Birgitte Rubæk for making the fish specimen drawings (head, pseudoclasper and otoliths drawn by Werner Schwarzhans), Geert Brovad for taking photos and Tammes Menne for help with x-raying and packing. The project was financed by the Carlsberg foundation and by a Visiting Collection Fellowship grant from The Australian Museum, Sydney. aqua vol. 8 no

53 Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen References Bleeker, P Bijdrage tot de kennis der ichthyologische fauna van de Batoe Eilanden. Natuurkunde Tijdschrift Nederlands Indië (NTNI), 8: Chen, L. & Shao, K-.T A review of the families Ophidiidae and Bythitidae from Taiwan. Bulletin of the Institute of the Zoology, Academia Sinica, 30: Cohen, D. M A new tribe and a new species of ophidioid fish. Proceedings of the Biological Society of Washington, 79: Cohen, D. M. & J. E. McCosker A new species of bythitid fish, genus Lucifuga, from the Galápagos Islands. Bulletin of Marine Science, 63(1): Cohen, D. M. & J. G. Nielsen Guide to the identification of genera of the fish order Ophidiiformes with a tentative classification of the order. NOAA Technical Report NMFS circular. 417: Criscione, S. F Aspects of the systematics of the viviparous bythitids, of the genera Ogilbia and Gunterichthys (Ophiidiformes, Bythitidae) in the Gulf of Mexico and Caribbean Sea. M. Sc. Thesis, Texas A&M University, Unpublished. Cumba-Segura, L Brotulidae: Typhliasina pearsei. Fauna de los Cenotes de Yucatán, No 4. Departmento de Acuacultura y Biologia Marina, Universidad de Yacatán, 9 pp. Dawson, C. E Gunterichthys longipenis, a new genus and species of ophidoid fish from the Gulf of Mexico. Proceedings of the Biological Society of Washington, 79: Dawson, C. E Supplemental observations on Gunterichthys longipenis, a northern Gulf of Mexico brotulid fish. Copeia, 1971: Gill, T. N Catalogue of the fishes of the western coast of North America. Proceedings of the Academy of Natural Sciences of Philadelphia, (1961), suppl.: Gill, T. N Notice of a collection of the fishes of California, presented to the Smithsonian Institution, by Mr. Samuel Hubbard. Proceedings of the Academy of Natural Sciences of Philadelphia, (1962): Hubbs, C. L Fishes of the caves of Yucatan. Carnegie Institute of Washington, Publication 491: Machida, Y Description of three new and one resurrected species of the bythitid genus Dinematichthys (Ophidiiformes). Japanese Journal of Ichthyology, 40(4): McEachran, J. D. & J. D. Fechhelm Fishes of the Gulf of Mexico,Volume 1: Myxiniformes to Gasterosteiformes. University of Texas Press, Austin, 1112 pp. Moore, R. H New records of three marine fish from Texas waters with notes on some additional species. Texas Journal of Science, 26: Nielsen, J. G Bythitidae (Viviparous brotulas). In: The living marine resources of the western central Atlantic, vol. 2, (Ed. K.E: Carpenter): FAO species identification guide for fishery purposes and American society of ichthyologists and herpetologists species publication No. 5. Food and agriculture organization of the united nations, Rome. Nielsen, J. G Family Bythitidae (Viviparous brotulas). In: Check list of the freshwater fishes of south and central America (CLOFFSCA), (Eds. S.O. Kullander, C.J. Ferraris, Jr., R. E. Reis): EDIPUCRS, Porto Alegre, Brazil. Nielsen, J. G., Cohen, D. M., Markle, D. F. & C. R. Robins Ophidiiform fishes of the world. FAO species catalogue, 18: I-XI Nolf, D Etude monographique des otolithes des Ophidiiformes actuels et revision des especes fossiles (Pisces, Teleostei). Mededelingen van der Werkgroep Tertiaire en Kwartaire Geologie, 17(2): Patterson, C. & D. E. Rosen The Paracanthopterygii revisited: order and disorder. In: Papers on the systematics of gadiform fishes, (Ed. D. M. Cohen): Natural History Museum of Los Angeles County, Science series 32: Proudlove, G., Medina-Gonzáles, R., Chumba- Segura, L. & T. Iliffe Threatened fishes of the world: Ogilbia pearsei (Hubbs, 1938) (Bythitidae). Environmental Biology of Fishes, 62: 214. Randall, J. E. & K. K. P. Lim A checklist of the fishes of the South China Sea. Raffles Bulletin of Zoology, Supplement No. 8: Robins, C. R., Ray, G. C., & J. Douglass A field guide to Atlantic coast fishes, North America. Houghton Mifflin Co. Boston, MA, 354 pp. Schemmel, C Zur Morphologie und Function der Sinnesorgane von Typhliasina pearsei (Hubbs) (Ophidioidea, Teleostei). Zoomorphologie, 87: Schwarzhans, W Vergleichende morphologische Untersuchungen an rezenten und fossilen Otolithen der Ordnung Ophidiiformes. Berliner Geowissenschaft. Abhandlung, 32: Schwarzhans, W A comparative morphological treatise of recent and fossil otoliths of the family Sciaenidae (Perciformes). Piscium Catalogus: Part Otolithi piscium, 1: Sedor, A. N A phylogenetic hypothesis based on the male copulatory complex in dinematichthyine fishes (order Ophidiiformes, family Bythitidae). M. Sc. Thesis, University of Southern California, unpublished. Sedor, A. N. & D. M. Cohen New bythitid fish, Dinematichthys minyomma, from the Caribbean Sea. Los Angeles County Natural History Museum, Contributions in Science, 385: Smith-Vaniz, W. F., Collette, B. B. & B. E. Luckhurst Fishes of Bermuda. The American Society of Ichthyologists and Herpetologists. Special publication 4: aqua vol. 8 no

54 Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two n. genera Solorzano, A Nota ichthylógica. Variación de Typhliasina pearsei (Hubbs). Ciencia, Mexico, 12: 286. Suarez, S. S The reproductive biology of Ogilbia cayorum, a viviparous brotulid fish. Bulletin of Marine Science, 25: Thinès, G L évoluton régressive des poisons cavernicoles et abyssaux. Masson et Cie, Paris, 394 pp. Thomson, D. A., L. T. Findleyand & A. N. Kerstitch Reef fishes of the Sea of Cortez. The rockyshore fishes of the Gulf of California. John Wiley and Sons, New York. Tolley, S. G. & E. B. Peebles Occurrence of Gunterichthys longipenis (Osteichthyes: Bythitidae) in a south-west Florida estuary. North-west Gulf Science, 9: Turner, C. L Male secondary sexual characters of Dinematichthys iluocoeteoides. Copeia, 1946: Wagner, P. R New records of the emerald sleeper, Erotelis smaragdus civitatum (Pisces: Eleotridae) and gold brotula, Gunterichthys longipenis (Pisces: Ophidiidae) in Louisiana. Procedings of the Louisiana Academy of Sciences, 34: Wilkens, H Regressive evolution and phylogenetic age: The history of colonization of freshwaters of Yucatan by fish and crustacea. Association for Mexican Cave Studies Bulletin, 8: Wilkens, H., U. Strecker & J. Yager Eye reduction and phylogenetic age in ophidiiform cave fish. Zeitschrift für Zoologie Systematic und Evolution-forschung, 27: Whitley, G. P New fish names and records. Proceedings of the Royal Zoological Society of New South Wales, : aqua vol. 8 no

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56 aqua Journal of Ichthyology and Aquatic Biology Vol. 8 (4), July 2004 Contents: Peter Rask Møller, Werner Schwarzhans and Jørgen G. Nielsen: Review of the American Dinematichthyini (Teleostei, Bythitidae). Part I. Dinematichthys, Gunterichthys, Typhliasina and two new genera Papers appearing in this journal are indexed in: Zoological Record; Biolis - Biologische Literatur Information Senckenberg; Cover photo: Ogilbichthys kakuki. Live specimen in Angelfish Blue Hole, Stocking Island, Great Exuma, Bahamas. Photo by T. Iliffe. Typhliasina pearsei photographed in Coenoses cave, Yucatan, Mexico. Photo courtesy of aqua geõgraphia. (See inside page 187).