A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placement

Helen  Larson

A phylogeny of Milyeringidae is reported and a new species, Milyeringa brooksi, is described from Cape Range National Park in the North West Cape (Cape Range Peninsula) of Australia. This species is distinguished on the basis of morphological and molecular characters from its only congener Milyeringa veritas. These diagnostic characters are related to a unique pattern of sensory papillae on the body and synapomorphies in three genes (cytochrome c oxidase I, cytochrome b, and NADH dehydrogenase 2). The new species is known only from the southern portion of the North West Cape spanning roughly 50 kilometers of subterranean habitat. This habitat is exceedingly rare and measures to preserve it and its fauna should be taken.

ISHj'; 11 73117 1292 .' ISS;\; 113 U Inx A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placelDent by Helen K. Larson Records of tile Western Australian Musell111 Supplement No. 62 Records of the Western Australian Museum The Records ofthe Western Australian Museum publishes the results of research into all branches of natural sciences, and social and cultural history, primarily based on the collections of the Western Australian Museum and on research carried out by its staff members. Collections and research at the Western Australian Museum are centred on Earth and Planetary Sciences, Zoology, Anthropology, Archaeology and History. In particular the following areas are covered: systematics, ecology, biogeography and evolution of living and fossil organisms; mineralogy; meteoritics; anthropology and archaeology; history; maritime history and maritime archaeology; and conservation. Western Australian Museum Francis Street, Perth, Western Australia 6000 Tel. (08) 9427 7000 Fax. (08) 9427 2882 E-mail ann.ousey@museum.wa.gov.au Minister for Culture And The Arts Chair of Trustees Executive Director Editors The Hon. Sheila McHale MLA Dr Ken Michael AM BE (Hons), DIC, PhD, FTS, Hon FIE Aust, FCIT, FAIM Dr Gary J. Morgan B.Sc. (Hons), Ph.D. Dr Fred E. Wells B.Sc., M.Sc., Ph.D. Dr John A. Long B.Sc. (Hons), Ph.D. The Records of the Western Australian Museum is published twice a year. A series of Supplements are also produced. The Records are available for sale and exchange, the current price being $11 plus postage per part. Each volume comprises four parts. Subscriptions can be taken out for whole volumes at a rate of $33 plus postage. Supplements can be purchased from the W.A. Museum Bookshop. Prices on request Prices include GST. Printed and published by the Western Australian Museum © Western Australian Museum, May 2001 ISBN 0 7307 1292 3 Cover: Mugilogobius abei, type species of the genus. illustration by Jill Ruse. A REVISION OF THE GOBIID FISH GENUS MUGILOGOBIUS (TELEOSTEI: GOBIOIDEI), AND ITS SYSTEMATIC PLACEMENT Records a/the Western Australian Museum Supplement No. 62 A revision. of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placement Helen K. Larson um m 2001 © Western Australian Museum, June 2001 World List Abbreviation: Rec. West. Aust. Mus. Suppl. No. 62 ISBN 0 7307 1292 3 ISSN 0 313 122X Printed and published by the Western Australian Museum, Francis Street, Perth, Western Australia 6000 v TABLE OF CONTENTS Abstract 1 Introduction 1 2 History of the subfamily Gobionellinae 5 Historical overview of the Mugilogobius-group of genera Methods 8 Measurements 8 Phylogenetic analyses 9 Abbreviations used 10 Material examined..................................................................................................................... 10 13 Phylogenetic analyses Character descriptions and polarities 13 Results of phylogenetic analyses 32 Mugilogobius and its sister-group 37 40 Genera related to Mugilogobius 42 Composition of the subfamily Gobionellinae Generic diagnoses and species descriptions 48 48 Key to Mugilogobius and related gobionelline genera Brachygobius 49 50 Preliminary list of valid species Caecogobius 55 Calamiana 56 Preliminary list of valid species 57 Chlamydogobius 62 63 List of valid species . Eugnathogobius 66 Preliminary list of valid species...................................................................................................... 66 Incertae sedis 71 Hemigobius 72 73 List of valid species Mugilogobius 76 Key to species of Mugilogobius 78 M. abei (Jordan and Snyder, 1901) 80 M. adeia Larson and Kottelat, 1992 90 M. amadi (Weber, 1913) 94 M. cagayanensis (Aurich, 1938) 97 M. cavifrons (Weber, 1909) 102 M. chulae (Smith, 1932) 109 M.fasciatus sp. novo , M.filifersp.nov M.fusCUS (Herre, 1940) M. fusculus (Nichols, 1951) M. latifrons (Boulenger, 1897) M. lepidotus sp. novo M. littoralis sp. novo M. mertoni (Weber, 1911) M. myxodermus (Herre, 1935) M. notospilus (Giinther, 1877) M. platynotus (Giinther, 1861) M. platystomus (Giinther, 1872) M. rambaiae (Smith, 1945) M. rexi sp. novo M. rivulus sp. novo M. sarasinorum (Boulenger, 1897) M. stigmaticus (De Vis, 1884) M. tigrinus sp. novo M. wilsoni sp. novo Nomina dubia Pandaka Preliminary list of valid species Pseudogobius Preliminary list of valid species. Redigobius Preliminary list of valid species Stigmatogobius Preliminary list of valid species. Tamanka T. siitensis Herre, 1927 Biogeography Acknowledgements References Appendix I. Nominal species of the Mugilogobius-group and their status 116 119 123 127 132 135 139 143 150 154 159 164 170 174 177 181 185 189 192 196 197 198 200 202 204 205 209 209 212 213 216 219 219 231 Records of the Western Australian Museum Supplment No. 62 A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placement Helen K. Larson Museum and Art Gallery of the Northern Territory, PO Box 4646, Darwin, Northern Territory 0801, Australia Abstract - The gobiid fish genus Mugilogobius and 13 closely-related genera form a monophyletic group within the subfamily Gobionellinae of the family Gobiidae. Included with Mugilogobius in this group are the genera Brachygobius, Caecogobius, Calamiana, Chlamydogobius, Eugnathogobius, Gobiopterus, Hemigobius, Mistichthys, Pandaka, Pseudogobius, Redigobius, Stigmatogobius and Tamanka, which are discussed and compared. The entire group consists of about 105 species, which have been greatly confused in the literature. The genera Mugilogobius and Tamanka are revised, and full descriptions of the species included. The genera Brachygobius, Calamiana, Caecogobius, Chlamydogobius, Eugnathogobius, Hemigobius, Pandaka, Pseudogobius, Redigobius and Stigrnatogobius are diagnosed, nominal species are listed and an indication of the probable number of valid species given. Mugilogobius includes 25 species, of which eight are described as new. The genus is defined by a combination of characters. Most species of Mugilogobius occur in estuarine to fresh waters, with some species widespread in the Indowest Pacific and others restricted to a single waterbody. There is a speciescomplex in the tectonic lakes of Sulawesi, characterised by vertebral pattern and several character reversals. Cladistic analyses of the Mugilogobius-group indicate that Chlamydogobius (restricted to Australia) is the sister-group to Mugilogobius. The monophyletic genus Tamanka is sister to the Mugilogobius-Chlamydogobius group. Hemigobius is the sister group to Pseudogobius. Brachygobius and Pandaka form a closely related group. Stigmatogobius is derived compared to Redigobius, a genus with the most plesiomorphic characters of the whole Mugilogobius-group. Eugnathogobius appears to be paraphyletic. The Gobionellinae thus includes the Mugilogobius-group and a second monophyletic group in which are placed the genera Awaous, Evorthodus, Ctenogobius, Gnatholepis, Gobionellus, Oligolepis, Oxyurichthys and Stenogobius. The relationship of Redigobius and Rhinogobius to these groups is somewhat equivocal. All these gobionellines share certain characters, particularly those of the dorsal pterygiophore formula, epural number, vertebral number, headpore arrangement and a tendency to occur in freshwater to estuarine habitats. INTRODUCTION The suborder Gobioidei comprises a very large and diverse group of percomorph fishes. The number of valid gobioid genera has been put at 270 and the number of species at about 2,000 by Hoese (1984) and Hoese and Gill (1993). Within the gobioids, the largest group is the Gobiidae, which has been estimated to include 200 genera and at least 1,500 species (Nelson, 1984). The family has a "tendency toward evolution by reduction" (Birdsong et al., 1988); that is, gobiids tend to be quite small (Nelson, 1984) and have undergone loss or fusion of a number of bones during their evolutionary history (Birdsong, 1975). Gobiids are world-wide in distribution, with the majority of species found in marine and estuarine waters of the Indo-Pacific region. The systematics and phylogeny of the Gobioidei are acknowledged as being difficult and have been variously described by workers as an "infamous and unwieldy morass" (Springer, 1983), "chaotic" (Gosline, 1971), an "insane world" (Birdsong in Winterbottom, 1984) and "troublesome" (Birdsong et al., 1988). Part of the problem has been attributed to the small size of these fishes (Miller et al., 1980; Hoese and Gill, 1993). Birdsong (1975) noted that gobioids are "remarkably similar in gross osteology", despite being a group with many osteological apomorphies when compared with other percomorph orders (Gosline, 1955; Winterbottom, 1993). In addition, gobioids, especially the smaller gobiids, are often overlooked by collectors (Miller et al., 1980; my personal H.K. Larson 2 observation), and it is only since the increased use of SCUBA (and underwater photography) and ichthyocides by scientists and other collectors that a more accurate idea of the true diversity of the group has become apparent. Resolution of gobioid phylogenetic problems has been inhibited by the difficulty in finding appropriate outgroups (Hoese, 1984) and the general lack of revisionary and osteological work on these fishes (Springer, 1983; Hoese, 1986; Hoese and Gill, 1993). The entire suborder, though well-defined by autapomorphies (Regan, 1911; Miller, 1973a; Springer, 1983; Hoese, 1984; Winterbottom, 1993), is currently unaligned with any sister group of fishes; a recent search for the gobioid sister group (Winterbottom, 1993) ended without a definite conclusion (although Winter-bottom found that hoplichthyids, gobiesocids, callionymids and trachinoids share a varying number of apomorphies with gobioids). Miller (1973a) gave a good review of the history of gobioid classification, and presented his own classification of the family Gobiidae, which comprised seven subfamilies (Eleotrinae, Pirskeninae, Xenisthminae, Gobionellinae, Tridentigerinae, Gobiinae, Kraemeriinae). This classification was rejected by Birdsong (1975), although he did not present an alternative. Miller's (1973a) classification was considered to be probably para- or polyphyletic by Springer (1983) and Birdsong (1975), and contaiped one group undefined by any apomorphy (the gobiid subfamily Eleotrinae). In an abstract, Hoese (1976) presented an alternative classification in which he recognised six families within the suborder: the Rhyacichthyidae, Pirskeniidae, Eleotridae, Gobiidae, Microdesmidae and Kraemeriidae. Hoese (1984) discussed gobioid relationships and recognised six families, among which was the family Gobiidae (and its four subfamilies the Amblyopinae, Gobiinae, Oxudercinae and Sicydiinae). This work was criticised for defining some groups by apparent symplesiomorphies (Harrison, 1989). While Birdsong et al. (1988) did not provide a classification, they presented a considerable amount of phenetic baseline data grouped as "working hypotheses", not classifications. A number of their working hypotheses support present classification, for example, the Oxudercinae as defined by Murdy (1989), and the Gobionellinae as partly defined by Pezold (1993). Hoese and Gill (1993) presented a classification of the families and subfamilies of the order Gobioidei, with apomorphies for each group apart from the gobiid subfamily Butinae. The present work follows the basic classification of gobioids given in Hoese (1984), as modified by Hoese and Gill (1993) and Pezold (1993) (Table 1). Gill (1997, in litt.) favours an alternate classification which includes familial status for the Gobiidae, Eleotrididae and the Butidae, based on Table 1 Classification of the Order Gobioidei, based on Hoese (1984), Hoese and Gill (1993) and Pezold (1993). Placement of the paedomorphic gobioid genus Schindleria is still uncertain Gohnson and Brothers 1993). Family Rhyacichthyidae Family Odontobutidae Family Gobiidae Subfamily Amblyopinae Subfamily Butinae Subfamily Eleotridinae Subfamily Gobiinae Subfamily GobioIl;ellinae Subfamily Oxudercinae Subfamily Sicydiinae Family Kraemeriidae Family Microdesmidae Subfamily Microdesminae Subfamily Ptereleotrinae Family Xenisthmidae [Family Schindleriidae] work in progress in collaboration with R. Mooi. The placement of the gobiid group under study in this paper remains the same. History of the subfamily Gobionellinae Bleeker (1874) first proposed a group called the subphalanx Gobionelli within his Phalanx Gobiini, itself a group within his subfamily Gobiiformes. Gobionellus Girard and Synechogobius Gill were the only genera included in the subphalanx Gobionelli. In other subgroups of his classification, Bleeker placed taxa now referred to the Gobionellinae. In Phalanx Latrunculini, Bleeker included Latrunculus Giinther (an unavailable name; a junior synonym of Aphia Risso), Gobiopterus Bleeker, Leptogobius Bleeker (a junior synonym of Gobiopterus) and Evorthodus Gill, and also the sicydiine Sicyopus Gill. Bleeker's subphalanx Brachygobii was composed only of (the gobionelline) Brachygobius Bleeker, 1849, and (the gobiine) Lophogobius Gill. In the Eugobii, another subphalanx of the Gobiini, Bleeker placed Awaous Valenciennes, 1837, Ctenogobius Gill, 1858, Oxyurichthys Bleeker, 1857, Rhinogobius Gill, 1859b, Stenogobius Bleeker, 1874, and Stigmatogobius Bleeker, 1874, along with those now considered true gobiines: Acentrogobius Bleeker, Amblygobius Bleeker and Callogobius Bleeker. He also provided brief diagnoses for each genus he discussed. Takagi (1963; in Japanese, quoted in Arai and Kobayasi (1973)) used the gobiid subfamily name Rhinogobiinae for a number of taxa now referred to the Gobionellinae, such as Acanthogobius, Chaenogobius, Chasmichthys, Mugilogobius, Pterogobius and Rhinogobius. Takagi's work was produced in a limited number for private use only and the names within it are not available (Article 8, International Code of Zoological Nomenclature 1999). The usage by Arai and Kobayasi (1973) is not available either, as they gave no definitions of the taxon Rhinogobiinae and only referred to Takagi's unpublished work. Miller proposed the subfamily Gobionellinae, in which he placed species with the characters: two epural bones, hypurals separate, no endopterygoid (= mesopterygoid), five branchiostegal rays, no - - _.. Revision of Mugilogobius _ _. _ - - - - - - - - - , 3 A I" .,' B j\ Figure 1 TLCP ａｏｃｐｾ POCP upper postcleithrum, no supratemp orals, scapula minute or absent, metapterygoid opposed to the quadrate and not meeting the ectopterygoid, preoperculum not the symplectic in anterior oculoscapular Trypauchen canal rarely extending across the snout, and the preopercular canal with three pores when present (Miller 1973a). He included in this subfamily a disparate group of genera and tribes: Gobionellus Girard, 1858, Gobioides Lacepede, 1800, Stenogobius Sensory pore pattern of: A, a gobiine (Yongeichthys nebulosus, NTM 5.10033-003); B, a gobionelline (Stenogobius psilosinionus, NTM 5.11107-001). Gobiines sensu Pezold (1993) have a single AlP, a pair PNP, and interorbital canal single (fused). Key to pores: ANP = anterior nasal pore, PNP = posterior nasal pore, AlP = anterior interorbital pore, PIP = posterior interorbital pore, PO = postorbital pore, IP = infraorbital pore, LC1 lateral canal pore, TLCP = terminal lateral canal pore, POP = preopercular pores. (Not drawn to scale). Bleeker, 1874, Oxyurichthys Bleeker, 1857, Paroxyuriclltllys Bleeker, 1876, the Apocrypteini and Periophthalmini (the latter two tribes are now placed in the subfamily Oxudercinae by Murdy, 1989), and the Taenionini and Trypauchenini (Koumans' Taenioididae, 1953; Hoese's Amblyopinae, 1984). Miller included Takagi's (1963) Rhinogobiinae within his own subfamily Gobiinae. Hoese (1984) recognised four subfamilies within the family Gobiidae: Amblyopinae, Gobiinae, 4 Oxudercinae and Sicydiinae. He pointed out that there were a number of previously named higher taxa within his Gobiinae. Among those that he named, which include gobionelline genera, were the Brachygobiinae, Latrunculinae and Rhinogobiinae. Hoese did not provide an exhaustive list of these available higher taxa (for example, he did not list Miller's subfamily Gobionellinae), and did not offer an opinion as to their status. Birdsong et al. (1988) placed 10 genera within their Gobionellus Group (Calamiana Herre, 1945d, Ctenogobius, Gnatholepis Bleeker, 1874, Gobionellus, Mugilogobius Smitt, 1899, Oligolepis Bleeker, 1874, Oxyurichthys, Pseudogobiopsis Koumans, 1935, Stenogobius and Tamanka Herre, 1927), united by possessing two epurals, a dorsal pterygiophore formula of 3-12210 (sensu Birdsong 1975), and 26 vertebrae (10+16). Birdsong et al. suggested that Awaous Valenciennes, Evorthodus Gill, 1859a, Rhinogobius Gill, and Schismatogobius de Beaufort, 1912, may also belong with this group. Murdy (1989) observed that Birdsong et al.'s Gobionellus Group shared several characters with the oxudercines and hypothesised Evorthodus as a possible sister group to the oxudercines (based on the form of the fifth ceratobranchial, among other characters). As pointed out by Murdy (1989), species of the Gobionellus Group and the oxudercines both occur in soft-bottom habitats and/ or estuaries. Harrison (1989) placed some of Miller's gobionelline genera within the Gobiinae, Oxudercinae and Amblyopinae, and proposed a group formed by the sicydiines and Awaous as being the sister group to the Gobionellinae. He considered this whole assemblage (Gobionellinae, Sicydiinae and Awaous) to be monophyletic, based on species sharing a long palatine in at least some taxa, a dorsal pterygiophore pattern of 3-12210, an unossified scapula and a transverse cheek papillae pattern (Harrison 1989: 344, figure 11). Harrison's gobionellines consisted of Stenogobius and two groups that he called the Ctenogobius lineage (comprising Ctenogobius, Gnatholepis, Evorthodus, Gobionellus, Gobioides and Oligolepis) and the Oxyurichthys lineage (comprising Oxyurichthys, Taenioides Lacepede, Apocryptodon Bleeker, Pseudapocryptes Bleeker, Scartelaos Swainson, Boleophthalmus Valenciennes, Periophthalmus Bloch and Schneider and Trypauchen). Pezold (1993) recognised the Gobionellinae as a subfamily, separating the group from the Gobiinae as defined by Hoese (1984). He proposed monophyly of the Gobiinae based on three synapomorphies involving the oculoscapular canals and pores (Figure 1). His Gobionellinae was not specifically defined by apomorphies. It was implied that gobionellines did not possess the combination of characters which defined Pezold's Gobiinae. The H.K. Larson x2 A os 01 , / . ., s ...:",: . B ｾ . -: .... /:b:{)\:) , .:::.\:':,:\ JP Figure 2 Transverse papillae pattern in Stigmatogobius sadanundio, CMK 6278, with papillae rows named according to Sanzo (1911): A, lateral view; B, dorsal view, pores indicated. Scale bar = 1 mm. Gobionellinae sensu Pezold may therefore be paraor polyphyletic. Pezold (1993) assigned 56 genera to the Gobionellinae, and arranged them into four subgroups: one large "main" group of 35 genera, and three putative monophyletic groups of genera: the Acanthogobius Group (seven genera), Astrabe Group (six genera) and Chasmichthys Group (eight genera). These last three groups he did not diagnose but stated that Birdsong et al. (1988) proposed them as monophyletic. Within the main group of 35 gobionelline genera, Pezold also included several genera that likely belong elsewhere, such as Deltentosteus Gill and Neogobius Hjin, but did not include several taxa considered to be gobionellines in this paper: Revision of Mugilogobius 5 quadrate and the head usually has a longitudinal papillae pattern (Figure 5). This group includes the taxa Mugilogobius, Brachygobius Bleeker, 1849, Caecogobius, Calamiana, Chlamydogobius, Eugnathogobius Smith, 1931, Hemigobius, Mugilogobius, Pandaka Herre, 1927, Pseudogobiopsis, Pseudogobius Popta, 1922, Redigobius Herre, 1927, Stigmatogobius, Tamanka Herre, 1927, and Weberogobius Koumans, 1953. In this paper, fishes in this group are referred to as the Mugilogobius group. A Historical overview of the Mugilogobius-group of genera Mugilogobius Smitt, 1899, is one of the most widespread gobionelline genera. It is known to occur in Indo-Pacific fresh waters and marine coastal habitats where there is some fresh water influence B ｾ A ｾ｀Ｎ［ -s 0 .. , • ;- , a b ·c Figure 3 Sensory pore pattern of: A, Stenogobius psilosinionus, NTM S.11107-001; B, Oxyurichthys ophthalmonema, NTM S.12731020. Scale bar = 1 mm. Hemigobius Bleeker, 1874, Caecogobius Berti and Ercolini, 1991, and Chlamydogobius Whitley, 1930. Within the "main" group of Gobionellinae sensu Pezold (1993), it was observed that a number of genera appeared to fall into two natural groups. One of these groups included genera (such as Ctenogobius, Evorthodus, Gnatholepis, Gobioides, Gobionellus, Oligolepis, Oxyurichthys and Stenogobius) which share a distinctive head pore pattern including a nasal canal with two pores, paired anterior interorbital pores and no infraorbital pore (Figure 1B). They also have a long palatine and very short pterygoid, have no metapterygoid bridge, and most species have a transverse papillae pattern (Figures 2-3). The freshwater genera Awaous and Rhinogobius appear to belong here also. In this paper, this group of genera is referred to as the Stenogobius group. The second group within the Gobionellinae sensu Pezold consists of 35 nominal genera characterised by variable loss of oculoscapular canals and pores, the anterior interorbital pore is paired, they have an infraorbital pore (Figure 4), the palatine and pterygoid are usually about equal in size, there is often a bridge from the metapterygoid to the Figure 4 Sensory papillae rows. A, exemplified by Eugnathogobius oligactis (composite of CAS 32975, CMK 4722 and CMK 5385), named according to Sanzo (1911); B, lateral canal pores indicated, named according to Hoese and Lubbock (1982) (not drawn to scale). Key to pores: AlP = anterior interorbital pore, PIP = posterior interorbital pore, PNP = posterior nasal pore, PO = postorbital pore, IP = infraorbital pore, LCl = lateral canal pore, TLCP = terminal lateral canal pore, POP preopercular pores. Scale bar 1 mm. H.K. Larson 6 .....1!.2 . r . ; ｉＧＮｾｾ＾ｹＡ ..;..---..... Figure 5 .. ' .... ,.,:::,., .. ··u ｾ｡Ｎ c ." x.1. . ...... ｾ ... .. , ,.' c ' .. .cP Longitudinal papillae pattern (Mugilogobius amadi, ZMH 7579), with papillae rows named according to Sanzo (1911). Scale bar = 1 mm. (e.g. estuaries, mangroves). This genus has been widely confused and the subject of some nomenclatural controversy ever since its creation. Species of Mugilogobius have been incorrectly referred to Ctenogobius, Ellogobius Whitley, 1933, Glossogobius Gill, Gobius Linnaeus, Tanumka, Vaimosa Jordan and Seale, 1906, Stigmatogobius, and Waiteopsis Whitley, 1930. Smitt (1899) originally created the genus Mugilogobius, as a sub-genus, in a key intended as a preliminary proposal for the classification of the genus Gobius. In this key Smitt also created the subgenera Eichwaldia and Caffrogobius. Although he designated type species for the latter two taxa, Smitt neglected to do so for Mugilogobius, indicating only that the subgenus was from India and Japan and had an interorbital width of more than 20% of the head length. The name is valid, nonetheless, as the absence of a type species designation for a new genus is acceptable for genera created before 1930 (International Code of Zoological Nomenclature, 1999). No species of the new genus Mugilogobius were designated by Smitt. Jordan et al. (1913) subsequently designated a type species for the genus: "Type Ctenogobius abei Jordan & Snyder (assigned by Jordan from Schmidt [sic] Ms.)". This was based on " ... a personal letter written by Smitt in 1903, in which he stated that his type species was the one named Ctenogobius abei in 1901" Oordan 1920; Smith 1945). Smith (1945) did not accept this "retrospective" designation and used the genus Vaimosa (type species fontinalis Jordan and Seale, 1906) for his species rambaiae Smith, 1945. Ctenogobius abei, Vaimosa fontinalis and Vaimosa rambaiae are considered to be congeneric in this paper. Smith (1945) was of the opinion that the validity of Mugilogobius (if it was to be considered as a valid name) should date from that of its appearance with a genotypic species name (abei Jordan and Snyder, 1901), so that Mugilogobius Oordan et al., 1913) would have to appear as a junior synonym of Vaimosa Oordan and Seale, 1906). However, Eschmeyer and Bailey (1990) accept Jordan and Snyder's 1901 type species designation, and it is also accepted here (based on Article 69 in the International Code of Zoological Nomenclature, permitting subsequent designation of a type species). The generic name Mugilogobius has not been used much. McCulloch and Ogilby (1919) used the name Mugilogobius devisi as an unnecessary replacement name for Gobius stigmaticus (De Vis, 1884). Other nominal new species referred to the genus are: M. luzonensis Roxas and Ablan, 1940; M. polylepis Wu and Ni, 1985; M. obliquifasciatus Wu and Ni, 1985; and M. adeia Larson and Kottelat, 1992. The genus Stigmatogobius Bleeker, 1874, has been widely applied to a variety of species within the "gobionelline" group of fishes (Hoese and Winterbottom 1979). Species of the gobionellines Mugilogobius, Redigobius, Pseudogobius and Pseudogobiopsis have been incorrectly referred to this genus (Appendix 1), and the name is often applied to misidentified gobiids in museum collections. The type species of the genus, Stigmatogobius sadanundio, was not included in their Gobionellus Group by Birdsong et al. (1988: table 33), but was placed as "unassigned". The genus does belong in their Gobionellus Group, as it has the same dorsal pterygiophore formula (3-12210; Figure 6), two epurals, 26 vertebrae, and is found in freshwaters. The genus Vaimosa was erected by Jordan and Seale (1906) (type species Vaimosa fontinalis Jordan Revision of Mugilogobius A c Figure 6 First dorsal spines and pterygiophore arrangement in: A, Mieropercops borealis, ex AMNH 10441; B, Rhinogobius brunneus, NTM 5.12121-002; C, Tamanka siitensis, ex FMNH 47512. Scale bar = 1 mm. and Seale, 1906, from Samoa). They distinguished it from Gnatholepis and Rhinogobius by its having naked cheeks and the opercles covered with large scales, and suggested that Gobius javanicus (= Pseudogobius) probably belonged to the same genus. Jordan et al. (1913) synonymised Vaimosa with Mugilogobius. Herre (1927) erected the genus Tamanka, separating Tamanka from Vaimosa Jordan and Seale, based on the " much smaller and more numerous scales (38 to 54 in a longitudinal series), and by having many small cycloid scales on the opercles instead of a few large ctenoid ones" (Herre 1927). Most of the species described by Herre as Vaimosa, however, are actually Redigobius or Pseudogobius, genera which possess headpores, unlike Mugilogobius. Herre did not use the genus name Mugilogobius. Koumans (1931: 88) suggested that Vaimosa was allied to Gnatholepis but admitted that he had not seen the types of the genera Mugilogobius or Vaimosa (Koumans 1931: 90), so was unable to decide whether to agree with Jordan et al.'s (1913) synonymy of Vaimosa with Mugilogobius. Aurich (1938) described several species of Vaimosa. He suggested that it was likely that the paedomorphic Gobiopterus-like gobies were close to "Ctenogobius, Mugilogobius, Stigmatogobius, Pseudogobius, Pandaka, Berowra", indicating that he was aware of the genus Mugilogobius, despite not using the name for the species he described. Koumans (1953) stated that Mugilogobius was known from Japan, China and Queensland. The last 7 locality record was apparently in reference to the description of Mugilogobius galwayi McCulloch and Waite, 1918, a synonym of Pseudogobius olorum (Sauvage, 1880). Koumans (1953) erected the genus Weberogobius for Weber's (1913) species Gobius amadi, and indicated that it was not related to Glossogobius (Weber's opinion as to relationship, 1913) nor Tamanka. Hoese and Winterbottom (1979) gave a brief diagnosis of Mugilogobius: " ... is characterized by an inferior mouth, lack of head pores, ＴＰｾ scale rows on the body, a depressed head, a broad interorbital, and rows of cheek papillae arranged longitudinally". The genus Chlamydogobius was reviewed by Miller (1987), who considered that it shared several apomorphies (tubular anterior nostril down-turned over or towards the upper lip; mouth subterminal, with snout protuberant in "larger forms"; sensory papillae with uniserial rows a, c, cp and b, with rows a and b not extending onto anterior part of cheek, and row c broken into two parts; and dorsal pterygiophore formula of 3-12210) with the genera Mugilogobius, Weberogobius, Pseudogobius, Redigobius, Brachygobius, Hemigobius and Pandaka. Miller also considered that, given the apparent suite of apomorphic characters present, Chlamydogobius should be maintained as a valid genus pending review of Mugilogobius and its relatives. Larson (1995) described five new species of Chlamydogobius and suggested that it may be most closely related to Mugilogobius. Birdsong et al. (1988) placed Mugilogobius in their Gobionellus Group (which also included Calamiana, Gnatholepis, Oligolepis, Oxyurichthys, Pseudogobiopsis, Stenogobius and Tamanka). These genera share the characters of two epurals, 25-26 vertebrae, dorsal pterygiophore formula of 3-12210, and having two or three anal pterygiophores before the first haemal spine. They separated Brachygobius and Pandaka out as part of a Gobiopterus Group, which comprised four genera which shared only a low vertebral number, 10+15. Berti and Ercolini (1991) described the first blind cave-dwelling gobiine (sensu Hoese 1984) species, Caecogobius cryptophthalmus, from four specimens collected from the Calbiga cave system on Samar Island, the Philippines. Berti and Ercolini commented on the similarities of their new genus to Glossogobius and Mugilogobius. The status of this species is discussed later. Hoese and Gill's (1993) discussion on eleotridine relationships provided some insight on characters which may be of value in untangling problems in gobioid systematics. In this paper, they also briefly referred to what they considered the "putatively primitive" gobiine genera Pseudogobius and Redigobius. These two genera are here considered to be gobionellines. Hoese and Gill considered H.K. Larson 8 Pseudogobius and Redigobius as plesiomorphic due to their similarity to butines and odontobutids in certain key osteological and myological characters. McKay (1993) attempted to clarify the relationships of the gobionellines Brachygobius, Pandaka, Pseudogobius and Redigobius (he referred to them as gobiines) using starch gel electrophoretic data. He investigated one or two species of each genus, and polarised the allele characters using seven gobiines as outgroups (Acentrogobius, Bathygobius, Glossogobius, Gobius, Istigobius, Pomatoschistus and Vanneaugobius). He concluded, based on the allozyme results, that Brachygobius and Pandaka were sister groups, with Redigobius as sister to this clade and Pseudogobius the most distant relative of this group, and that all four formed a " ...monophyletic lineage which is distinct from other gobiine groups". The present definitions in the literature of the genus Mugilogobius are that of a gobionelline (Pezold 1993) without headpores and usually possessing 16 caudal fin rays (Larson and Kottelat 1992; Kottelat et al., 1993) and "the appearance of widely separated interorbital papillae rows p" (Miller 1987). Larson (1995) gave the most complete definition, when comparing Mugilogobius with Chlamydogobius: Mugilogobius: s papillae on snout in at least three rows of two or more papillae, the first of which is usually the longest and runs just above upper lip fold (few species with middle row represented by only 1-2 papillae); pectoral rays 13-20; intestine simple, with one "S-bend" and no full loops; gill opening to pectoral base or further, usually with fleshy knobs or ridge along shoulder; two epurals, metapterygoid forming distinct bridge to quadrate, 26-27 vertebrae, usually 26 (10, rarely 11, precaudal and 16-17 caudal), males often with distinctly enlarged mouths. Specimens examined for this paper, agreeing with the above criteria for Mugilogobius, were initially referred to that genus if they had a subterminal mouth and rounded snout, and to Tamanka if they had a terminal mouth and a somewhat pointed snout. My preliminary research suggests that Mugilogobius is part of a natural grouping of IndoPacific species which also includes the nominal genera Brachygobius, Caecogobius, Calamiana, Chlamydogobius, Eugnathogobius, Hemigobius, Pandaka, Pseudogobiopsis, Pseudogobius, Redigobius, Stigmatogobius, Tamanka and Weberogobius. Preliminary observations indicate that all these genera share the gobionelline characters as defined by Pezold (1993), as well as the characters of a down-turned, tubular anterior nostril close to the upper lip, having knobs, lobes or a flange often present on the anterior edge of the cleithrum (pectoral girdle), usually a longitudinal sensory papillae pattern on the head, a reduction or loss of head canals and pores, often having enlarged mouths (sometimes greatly so) in mature males, and an association with water of reduced salinity or freshwaters. Initial work suggests that the group is speciose (161 nominal species in total), with nine nominal species in Brachygobius, one in Caecogobius, five in Calamiana, six in Chlamydogobius, one in Eugnathogobius, six in Hemigobius, 47 in Mugilogobius, seven in Pandaka, 12 in Pseudogobiopsis, 21 in Pseudogobius, 43 in Redigobius, nine in Stigmatogobius, three in Tamanka, one in Weberogobius and nine species for which provisional generic assignment is equivocal (can be assigned to more than one genus upon available information). Based on this preliminary information, the aim here has been to diagnose the genus Mugilogobius and revise the species within it, determine and diagnose those genera which are its closest relatives; and to determine the extent of monophyly for the Mugilogobius-group. METHODS Measurements Morphometric measurements were taken using electronic (digital) callipers and a dissecting microscope. Counts and methods generally follow Hubbs and Lagler (1958), except as indicated below. Pterygiophore formula follows Birdsong et al. (1988). Transverse scale count backwards (TRB) is made from the anal fin origin upward and backward to the second dorsal fin base. The circumpeduncular scale count is taken beginning at the first normal scale (i.e., not reduced in size or indented in centre) on the top of the caudal peduncle immediately in front of the caudal fin, and following the scale rows down and forward to the ventral edge of the peduncle, then around and back to the original scale. Head length (HL) is taken to the upper attachment of the opercular membrane. Interorbital width is taken as the least fleshy width, not least bony width (the latter is often difficult to determine in many species of this group, due to the thick fleshy interorbital skin). Proportions are given as times in HL or SL (as specified). The segmented caudal ray pattern (e.g. 9/8 or 9/7) is the number of segmented caudal rays attaching to the upper and lower hypural plates respectively (the same applies to the branched ray pattern, written as 7/7, for example). Although both left and right pectoral ray numbers were recorded, the right pectoral fin count is that used in descriptions and tables, unless otherwise indicated (in cases of damage). The skin over the anterior edge of the cleithrum (referred to as pectoral girdle) may be smooth, have a raised fleshy ridge or have several fleshy knobs or flaps. These knobs and ridges are not always identical bilaterally; if knobs are present on one side but not Revision of Mugilogobius on the other, the specimen was scored as possessing knobs. In species descriptions, an asterisk indicates counts of the holotype, syntype, lectotype or neotype (as indicated below). Numbers in parentheses after counts indicate the number of specimens with that count or range of counts. Papillae pattern terminology is based on that of Sanzo (1911), who developed a system of naming papillae rows largely based on European genera and species. Hoese (1983) considered that Sanzo's system might not always be applicable to IndoPacific gobioids, and proposed a system of his own, which numbered the longitudinal and transverse patterns. Wongrat and Miller (1991) disagreed with Hoese, and considered that there was no difficulty in reconciling Sanzo's system to the patterns found in Indo-Pacific gobioids, including the eleotridids. Akihito (1967) and Akihito and Meguro (1975c) use a third system, based on numbering the papillae rows. Takagi (1989) discussed papillae and pore patterns in 82 Japanese gobioids, presenting other methods of naming the structures of these systems. Many authors ignore papillae patterns in their descriptions (as did most of the earlier authors such as Bleeker, Herre, Smith and Weber), or else they illustrate and discuss the papillae without naming or numbering the rows (e.g. Winterbottom and Burridge, 1992; Larson, 1990; Hoese and Randall, 1982; Larson and Kottelat, 1992). Sanzo's system is used here, solely due to its use in previous literature on this group of gobies by Aurich (1938) and Miller (1987, 1989) and for ease of comparison with these works. Whether the rows are directly comparable between this Indo-Pacific group of genera and Sanzo's European genera is not of concern here. The papillae themselves are oriented along the axes of their rows either crossways or lengthways (Marshall, 1986; Wongrat and Miller, 1991); the rows a, c, cp, i and p are oriented lengthways (parallel to row axis), while the papillae of rows b, d, e, ot, oi, os and s are oriented crossways (transverse to row axis). Oculoscapular canal and pore terminology used in the literature for gobioids varies greatly (for examples, see Hoese and Gill, 1993; Sanzo, 1911; Hoese and Lubbock, 1982; Akihito et al., 1988; Miller, 1987; Watson, 1991; Springer and Randall, 1992). Rather than use a system of symbols (Sanzo 1911) or letters (Akihito et al., 1988), here the pores are named and indicated on the figures as abbreviations, basically following Hoese and Lubbock (1982). The anterior interorbital pore (AlP) (Figure 1) is paired in gobionellines (Pezold, 1993) and the posterior interorbital pore may be single or double. Many gobionelli.nes lack a postorbital pore (Pezold 1993; his pore "E"), but ｐｾ･ｵ､ｯｧ｢ｩｵｳ and Redigobius are known to possess one. The oculoscapular canal is separate from the preopercular canal in most gobiids. 9 Synonymies given are not always full synonymies, but are often only partial. These fishes have been misidentified in much of the literature, and it has not always been possible to determine from descriptions or illustrations what species or genus was referred to in various publications (for example, Stigmatogobius hoevenii could be Mugilogobius chulae or Hemigobius hoevenii). Synonymies are given for those species in which the identity can be verified by examination of specimens or the description is unequivocal. In statements referring to generic or specific characters and/ or character states in other gobioids, some of this information is from available literature (and is so indicated in the text), some is from discussion with colleagues (given as "personal communication") and some is unpublished information from my experience working with gobioid fishes (personal observation). Specific colour names given (e.g. royal blue, maize yellow) are those used in the Met/wen handbook ofcolour (Kornerup and Wanscher 1978). Phylogenetic analyses The approach taken in this paper mostly follows phylogenetic systematics methodology, based on the ideas of Hennig (1979), as this has been used by the ichthyology community and has been shown to be useful in helping to resolve some systematic problems (e.g. Parenti, 1981; Winterbottom, 1990; Springer and Williams, 1994). While it has been demonstrated that there are some difficulties with it, this methodology was considered the best approach to take. For example, Carpenter et al. (1995) point out some of the problems inherent in applying phylogenetic systematics techniques to the classification of a comparatively well-studied and conspicuous group of fishes, the scombroids, tunas and billfishes. Not a great many phylogenetic analyses have been done on gobioid taxa. Springer (1983) was first to apply phylogenetics to gobioids, in an attempt to determine the monophyly of the suborder Gobioidei. Miller et al. (1980) used a phenetic approach to 28 gobioid species based on haemoglobin analyses. Winterbottom (1990) and Winterbottom and Burridge (1993) used phylogenetics in their work on Trimmatom, Trimma and Egglestonichthys species. Murdy's (1989) paper on the oxudercines, Pezold's (1993) gobionelline paper and Hoese and Gill's (1993) work on the eleotridines and odontobutids are examples of work using phylogenetic techniques among higher taxa of gobioids. McKay (1993) used a variety of methods (phenetic and cladistic) to try to establish the relationships of Brachygobius, Pandaka, Pseudogobius and Redigobius using starch gel electrophoretic data. Gill (1994) used a cladistic analysis in an attempt to show a natural grouping 10 existed of 22 genera of gobiines with a longitudinal papilla pattern. The rhyacichthyids and odontobutids have been considered plesiomorphic "basal gobioids" for some time (Miller, 1973a; Springer, 1983; Hoese, 1984; Akihito, 1986; Hoese and Gill, 1993). Hoese and Gill (1993) suggested that the odontobutids form a separate family, and that butines and eleotridines are plesiomorphic in relation to gobiines, although they were unable to find any apomorphy defining the butines or odontobutids as monophyletic groups. Therefore, potential outgroup relationships of gobioids are well-established in part, and useful characters may be obtained from the literature and corroborated with available specimens. In phylogenetic analyses using the "simple parsimony" outgroup method of Maddison et aI. (1984), it is necessary for outgroup relationships to have been resolved before character polarities can be assigned. Simple parsimony assumes that the lowest number of character state changes have occurred, and that convergences and reversals may also occur at any stage. Outgroup comparisons may be done at any taxonomic level ("functional out-groups and in-groups") (Watrous and Wheeler, 1981). Therefore one species of the family Rhyacichthyidae (Rhyacichthys aspro) and two species of the family Odontobutidae (Micropercops borealis and Perccottus chalmersi) were used as outgroups for polarising characters in this assemblage of gobiids. Morphological and osteological details of the recently described rhyacichthyid Protogobius Watson and Pollabauer were not available, so this less specialised genus could not be used in place of Rhyacichthys. Preliminary analyses of relationships among species and genera were made using several computer programs: PAUP v.2.4 (Swofford, 1985) and Hennig86 v.1.5 (Farris, 1988) on IBM machines, with PAUP v.3.0 (Swofford, 1991) on a MacH used for final analyses. Hennig86 analyses were run using the mh*, bb* branch-swapping option, while PAUP analyses were made by heuristic TBR branch-swapping (deltrans). Uninformative characters (e.g. autapomorphies) were ignored. Characters were not weighted during these analyses, as it was not possible to find any objective criteria for doing so. Of the 42 characters polarised and used, some of these were variable in that some specimens of several species had a different condition. In these cases, the modal condition was used. While it is recognised that other methods are known which attempt to deal with this individual variation, as discussed by Wiens (1995), many of these methods involve weighting. All characters discussed here are unweighted. Analyses were usually run twice, with characters ordered and unordered, as there is controversy over the information gained or lost by exclusive use of H.K. Larson unordered or ordered characters (Wilkinson 1992; Carpenter et al., 1995). In the trees presented below (Results), characters are unordered. Abbreviations used Abbreviations for institutions generally follow Leviton et al. (1985). They are: AMNH - American Museum of Natural History, New York; AMS - The Australian Museum, Sydney; ANSP - Academy of Natural Sciences, Philadelphia; BLIH - Biological Laboratory, Imperial Household, Tokyo; BMNH - The Natural History Museum, London; BPBM - Bernice P. Bishop Museum, Honolulu; CAS - California Academy of Sciences, San Francisco; BSM - Bureau of Science, Manila; CMK - Collection Maurice Kottelat, Cornol, Switzerland; FMNH - Field Museum of Natural History, Chicago; KEW - collection of Or KE. Witte, previously of University of Constance, Konstanz; KUMF - Kasetsart University Museum of Fisheries, Bangkok; MHNG - Museum d'Histoire Naturelle, Geneva; MNHN - Museum National d'Histoire Naturelle, Paris; MZB - Museum Zoologicum Bogoriense, Bogor; NIFI - National Inland Fisheries Institute, Bangkok; NMBA Naturhistorisches Museum, Basel; NMMB - National Museum of Marine Biology, Kaohsiung; NMW National Museum, Wien; NTM - Museum and Art Gallery of the Northern Territory (formerly Northern Territory Museum), Darwin; PMBC - Phuket Marine Biological Centre; QM - Queensland Museum, Brisbane; RMNH - Nationaal Naturhistorisches Museum, Leiden; ROM - Royal Ontario Museum, Toronto; RUSI - J.L.B. Smith Institute of Ichthyology, Grahamstown; SAM - South African Museum, Capetown; SAMA - South Australian Museum, Adelaide; SMF - Senckenberg Museum, Frankfurt; SMNS - Staatliches Museum fUr Naturkunde Stuttgart; URM - University of the Ryukyus, Naha; USNM - National Museum of Natural History, Washington; UW - University of Washington, Seattle; WAM - Western Australian Museum, Perth; YCMYokosuka City Museum, Kanagawa; ZMA Zoologische Museum, Amsterdam; 2MB Zoologische Museum, Berlin; ZMH - Zoologische Museum, Hamburg; ZMUC - Kobenhavns Universitet Zoologisk Museum, Copenhagen; ZRC Zoological Reference Collection, University of Singapore; ZSI - Zoological Survey of India, Calcutta; ZSM - Zoologische Staatsammlung, Miinchen. Other abbreviations used: ANGFA - Australia New Guinea Fishes Association; HL - head length; SL; standard length. Material Examined Details of preserved specimens examined are given with each species account (descriptions), except for cleared and stained and X-rayed specimens, which are listed below (C = cleared and stained specimen, X radiograph). Revision of Mugilogobius RHYACICHTHYIDAE. Rhyacichtllys aspro: CASSU 38565, C, 1(116); NTM unregistered, C, 1(85); NTM S.1212I-ClO4, X, 1(65). ODONTOBUTIDAE. Micropercops borealis: ex AMNH 10441, C, 3(31-33.5). Odontobutis auranllllS, NTM S.13964-001, X, 2(29-35). Odontobutis obscura: NTM S.13848-001, C, 2(62-72); N1M S.13848-001, X, 3(107-143); NTM S.11697-001, X, 1(122). Perccottlls clzalmersi: holotype of Philypnus chalmersi, AMNH 8384, 1(100); ex AMNH 10456, C, 3(31.5-40.5). GOBIIDAE: BUTINAE. Bostrychus sinensis: ex NTM S.11125-007, C, 1(70); NTM S.10555-001, X, 2(77-94). Bostrychus zonatus: NTM S.11845-009, X, 1(56); NTM S.11830-001, X, 1(20). Butis butis: NTM unregistered, C, 2(39-44). Incara multisquamata: NTM S.13476-001, X, 2(43-44); NTM S.10472-028, X, 4(17-21). Oxyeleotris lineolata: ex NTM S.11876-01O, C, 1(62). "Oxyeleotris" nullipora: NTM unregistered, C, 1(24.5). Oxyeleotris selheimi: NTM unregistered, C, 1(62). Prionobutis microps: ex NTM S.10553-001, C, 1(71). GOBIIDAE: ELEOTRIDINAE. Eleotris acanthopoma: ex NTM S.12131-018, C, 2(33-56). Eleotris melanosoma: ex NTM S.11125-003, 2(37-44). Hypseleotris compressa: NTM unregistered, C, 2(4148). Mogurnda mogurnda: ex NTM S.10643-001, C, 1(46). Tateurndina ocellicauda: NTM unregistered, C, 1(31). GOBIIDAE: GOBIINAE. Acentrogobius caninus: NTM unregistered, C, 2(35-47.5). Bathygobius cocosensis: ex NTM S.12089-004, C, 1(35.5). Glossogobius aureus: ex NTM S.12632-023, C, 1(54). Gobiodon rivulatus: ex NTM S.10012-011, C, 2(20.527). Gobius buccichi: ex NTM S.12042-001, C, 1(55). GOBIIDAE: GOBIONELLINAE. Awaous acritosus: NTM unregistered, C, 2(56-86). Awaous banana: NTM S.13083-001, X, 3(46-68). Brachygobius doriae: syntypes of Gobius doriae, BMNH 1868.1.28.17-19, X, 3(24-25); NTM unregistered, C, 1(30); ex ROM 56160, C, 2(19-19.5); ZRC 669, X, 6(17-31); ex CAS 33045, X, 5(21-29). Brachygobius kabiliensis: ex ZRC 828, C, 2(15-15); ZRC 828, X, 5(13-15); ZRC 19874-8, X, 5(12.5-15.5). Brachygobius nunus: syntypes of Gobius alcockii, ZMA 114.487, X, 3(10-11.5). Brachygobius xanthomelas: paratypes, CAS 16964, X, 3(10.5-15.5); ex ZRC 13963-6, C, 1(17.5); ZRC 13963-6, X, 4(12.5-17). Brachygobius xanthozonus: RMNH 12084, C, 1(31); RMNH 12083, X, 1(25); RMNH 12082, X, 2(22-24); ex CAS 33045, X, 1(22). Caecogobius cryptophthalmus: paratype, ZSM 27189, X, 1(58.5). Calamial1a illota: holotype, ZRC 39268, X, 1(37); paratypes, NTM S.14235-002, X, 7(22.5-34); NTM ex S.14235-002, C, 1(30); ZRC 39269, X, 3(37-45). Calamiana kabilia: holotype of Vaimosa kabllia, CAS 32978, X, 1(36.5); paratypes of Vaimosa rambaiae, USNM 119647, X, 2(27-29); ex CMK 4789, C, 1(47); CMK 4789, X, 1(45); NIFI unregistered, X, 3(26-34); NTM S.14302-001, X, 3(21-23); NTM ex S.14302-001, C, 2(22.5-28). 11 Calamiana mindora, ex ROM 53371, C, 1(24.5); ROM 53371, X, 8 of 20(14-25); CMK 5366, X, 6(19.5-26). Calamiana sp. novo 2: ex AMS 1.25523-003, C, 3(1720.5); QM 1.13347, X, 3(18-21). Calamiana sp. novo 3: ex AMS 1.32051-032, C, 1(21). Calamiana variegata: holotype of Tamanka ubinensis, CAS 30964, X, 1(30); ex URM P.13341, C, 1(32); URM P.13341, X, 2(2931.5); ZRC 39270, X, 1(33); URM P.13842, X, 1(33). Chlamydogobius eremius: lectotype, SAMA F.525, X, 1(51); paralectotypes, SAMA F.7674, X, 2(35-43); , SAMA F.3509, X, 10(30.5-44); AMS 1.24493-001, X, 4(22-28); SAMA F.3999, X, 12(31-41); SAMA F.4205, X, 6(27-45.5); AMS 1.24673-002, C, 2(2527.5); ex AMS 1.24493-001, C, 1(25); 2MB 31790, X, 2(43-45). Chlamydogobius gloveri: holotype, SAMA F.3463, X, 1(30); paratypes, SAMA F.7675, X, 3(3336); SAMA F.5425, X, 4(20-27); AMS 1.27118-001, X, 1(32); SAMA F.5417, X, 4(18-24). Chlamydogobius japalpa: ex NTM S.11439-006, C, 1(34.5); NTM S.11439-006, X, 5(22-38.5); ex NTM S.11436-009, X, 15(15-44); ex NTM S.11436-009, C, 5(heads only). Chlamydogobius micropterus: holotype, QM 1.25096, X, 1(22.5); paratypes, QM 1.29552, X, 9(15-23); AMS 1.25261-001, X, 3(8.5-20.5). Chlamydogobius ranunculus: holotype, NTM S.11427-001, X, 1(28.5); paratypes, NTM S.11427-002, X, 7(20-28.5); QM 1.19003, X, 1(31); AMS 1.32051-012, X, 10(20.5-28); AMS 1.22959-001, X, 5(22-25.5); ex NTM S.11509007, C, 2(28-29); AMS 1.32051-045, C, 2(23-27). Chlamydogobius squamigenus: holotype, SAMA F.6595, X, 1(34); paratypes, SAMA F.7676, X, 8(2235); SAMA F.6738, X, 1(34); SAMA F.7184, X, 7(28.5-39). Eugnathogobius sp. nov.: ex CMK 8401, C, 1(25); CMK 8401, X, 26(11-27); ZRC 26026-7, X, 2(14.5-19.5). Eugnathogobius microps: holotype, USNM 90316, X, 1(26); paratype, USNM 11951, X, 1(26); paratypes, USNM 119593, X, 3(20-26.5); KUMF unregistered., X, 2(17.5-17.5); NTM S.13953013, X, 6(18-25.5); ex NTM S.13953-013, C, 1(24). Eugnathogobius oligactis: holotype of Vaimosa perakensis, CAS 32975, X, 1(25.5); paratypes of V. perakensis, CAS 32977, X, 6(13.5-28.5); NTM S.14239-001, X, 3(31.5-36); CMK 5385, X, 7(21-36); ex CMK 4722, C, 1(30); ex CMK 10713, C, 2(28.529). Eugnathogobius paludosus: holotype of Ctenogobius paludosus, CAS 32998, X, 1(30.5); ex CMK 7384, C, 2(18-25); CMK 7384, X, 10(12.5-29); ZRC 8411, X, 1(30); CMK 9009, X, 2(13-21); ZRC 14011, X, 1(22). Eugnathogobius siamensis: holotype of Vaimosa mawaia, CAS 29080, X, 1(24); holotype of Vaimosa jurongensis, CAS 32982, X, 1(36); paratypes of V. jurongensis, CAS 32983, X, 11(19.534); ex USNM 119637, C, 3(26.5-32.5); ANSP 63126, X, 2(27.5-28); ANSP 87453, X, 13(23-32.5); CMK 8485, X, 6(22-31). EvortllOdus lyricus: ex NTM S.12858-001, C, 2(30-37). Gnatholepis sp.: ex NTM S.10005-034, C, 2(23-24); ex NTM S.12883-015, C, 1(37). Gobiopterus brachypterus: ex CMK 7277, C, 3(19-20). Gobiopterus sp.: ex NTM S.11242-38, C, 12 7(12.5-13.5). Gobius tigrellus: holotype and paratypes, AMNH 18574/ X/ 10(15-21). Hemigobius hoevenii: ex NTM 5.11065-002/ C/ 2(18.5-21.5); AM5 1.23262-001/ X/ 5(19-32.5); ex NTM 5.14235-004/ C/ 3(20-25). Hemigobius mingi: ZRC 20263-72/ X/ 10(2846); ex URM P.6677, C/ 2(34-38). Mugilogobius abei: holotype of Ctenogobius abei, CA5 6447/ X/ 1(35); ex YCM 908/ C/ 1(35.5); ex URM P.7053, C/ 2(32-36); YCM 7055/ X/ 3(36-40); MNHN 1967-0566/ X/ 3(30.5-40.5); URM Po4099, X/ 1(41); AM5 1.20372001/ X/ 5(25.5-29). Mugilogobius adeia: holotype, MZB 5891/ X/ 1(27.5); paratypes, CMK 6513/ X/ 2(16-22.5); ex NTM 5.13068-001/ C/ 1(22). Mugilogobius amadi: leetotype of Weberogobius amadi, ZMA 112.664/ X/ 1(122); paraleetotypes, ZMA 121.293/ X/ 4(93-124); ex ZMH 7579/ C/ 2(6789); Z5M 27493/ X/ 4(69-83); CMK 5774/ X/ 4(6984). Mugilogobius cagayanensis: leetotype of Vaimosa cagayanensis, ZMH 420a, X/ 1(25); paraleetotypes, ZMH 420b, X/ 2(18.5-20); ex U5NM 122921/ C/ 2(42.5-58); U5NM 122921/ X/ 31(37-61). Mugilogobius cavifrons: paralectotypes of Gobius cavifrons, ZMA 1230468/ X/ 15(14-26.5); holotype of Vaimosa karatunensis, ZMH 421/ X/ 1(31); holotype of Glossogobius parvus, FMNH 59138/ X/ 1(33.5); holotype of Tamanka talavera, CA5 36824/ X/ 1(34); CA5 38632/ X/ 2(26-25); URM P.6686, X/ 7(33-38); ZMUC unregistered, X/ 12(11-32.5); BPBM 33930/ X/ 7(29.5-44.5); ex BPBM 33930/ C/ 2(42-45). Mugilogobius chulae: CMK 10035/ X/ 1(34); CMK 10005/ X/ 1(22.5); ZRC 22757-63/ X/ 7(28-38); CMK 8921/ X/ 11(18-30.5); ex URM 9324/ C/ 2(29.5-35); ex CMK 4830/ C/ 1(31). Mugilogobius fasciatus sp. noy.: holotype, ZRC 17099/ X/ 1(20); CMK 8316/ X/ 2(2224); NTM 5.13954-042/ X/ 1(12); AM5 1.137570-001/ X/ 1(11); NTM 5.14303-001/ X/ 1(19.5); NTM 5.13953-016/ X/ 9(8-20.5). Mugilogobius filifer sp. noy.: ex NTM 5.10694-013/ C/ 1(33); ex NTM 5.10472-002/ C/ 1(27); WAM P.25668-014, X/ 4(1732). Mugilogobius fuscus: holotype of Vaimosa fusca, CA5 32984/ X/ 1(32.5); YCM P.9167P, X/ 1(42); URM Po4840, X/ 1(42); U5NM 260525/ X/ 2(21-33); U5NM 99611/ X/ 1(35.5); YCM 9323/ X/ 1(39). Mugilogobius fusculus: ex CA5 63580/ C/ 2(15.5-17). Mugilogobius latifrons: lectotype, NMBA 1847/ X/ 1(38.5); paraleetotypes, NMBA 1848-52/ 2734/ X/ 6(18.5-29); ZMA 113.627/ X/ 2(22.5-29.5); NTM 5.12706-005/ X/ 3(23-29.5); ex NTM 5.12706-005/ C/ 1(29); ex CMK 6206/ C/ 1(26). Mugilogobius lepidotus sp. noy.: CMK 6491/ X/ 10(22-25.5); CMK 9752/ X/ 5(20.5-26); ex CMK 6491/ C/ 1(23). Mugilogobius littoralis sp. noy.: holotype, NTM 5.14293-001/ X/ 1(27.5); paratypes, ex NTM 5.10452-022/ C/ 2(19.520); NTM 5.10439-001/ X/ 4(22.5-25.5); NTM 5.14296-001/ X/ 10(30.5-39). Mugilogobius mertoni: holotype of Stigmatogobius inhacae, RU5I 207/ X/ 1(27); paratypes of Stigmatogobius inhacae, RU5I 7247/ X/ 4(22-30); CA5 69655/ X/ 3(25-30); U5NM 264948/ X/ 10(18-28); ex AM5 1.20978-012/ C/ 2(25- H.K. Larson 30); ex URM Po4387, C/ 2(32.5-33); AM5 1.23637001/ X/ 1(22.5); AN5P 73284/ X/ 1(21); AN5P 96760/ X/ 1(31); U5NM 316139/ X/ 6(13.5-15.5); U5NM 316192/ X/ 1(30); ex RG. Museum Afrieain Centrale 18877-3/ X/ 1(22). Mugilogobius myxodermus: FMNH 47058/ X/ 3(23-30); CA5 69679/ X/ 8(10.5-38.5); CA5 32579/ X/ 1(33.5); ex AMNH 37029/ C/ 2(27-31.5). Mugilogobius notospilus: holotype of Gobius notospilus, BMNH 1869.11.12.31/ X/ 1(51); holotype of Vaimosa fontinalis, U5NM 51776/ X/ 1(38.5); holotype of Stigmatogobius duospilus, AN5P 71970/ X/ 1(29); U5NM 31671/ X/ 2(27-34); BMNH 1924.12.11.35/ X/ 1(38); BMNH 1925.3.30.6/ X/ 1(39); ex AM5 1.22045-003/ C/ 2(23-24); ex ZMH 19346/ C/ 1(28). Mugilogobius platynotus: holotype of Gobius platynotus, BMNH 1859.5.7.71/ X/ 1(39); holotype of Waiteopsis paludis, AM5 IA.3917, X/ 1(37.5); paratype of W. paludis, AM5 IA.3918, X/ 1(19); holotype of Ellogobius abascantus, AM5 IA.6850, X/ 1(37); paratypes of E. abascantus, AM5 1.32053-001/ X/ 2(27.5-35.5); AM5 1.25376-001/ X/ 4(22-24); ex AM5 1.25038-001/ C/ 1(26); ex AM5 1.25376-001/ C/ 1(32.5). Mugilogobius platystomus: holotype of Gobius platystoma, BMNH 1871.9.13.179/ X/ 1(40); ex NTM 5.1867-001/ C/ 1(19); ex AM5 1.22959-002/ C/ 1(34); CA5 54690/ X/ 7 out of 18(29.5-38); NTM 5.14204-001/ X/ 3(32-38); NTM 5.14205-001/ X/ 2(3040). Mugilogobius rambaiae: holotype, U5NM 119646/ X/ 1(41.5); paratypes, U5NM 119647/ X/ 7(24-33); ZRC 26972-3/ X/ 2(29.5-33.5); CA5 36032/ X/ 10(24-30.5); BMNH 1935.5.27.27/ X/ 1(27.5); U5NM 316172/ X/ 1(23); BMNH 1937.6.14.22-23/ X/ 2(38.5-41); ZMH 7992/ X/ 1(31); ZRC 19886-95/ X/ 8(24-34.5); ex CA5 36032/ C/ 2(25-30.5). Mugilogobius rexi sp. noy.: CMK 6205/ X/ 9(14.533); ex NTM 5.12707-002/ C/ 2(24-28). Mugilogobius rivulus sp. noy.: holotype, NTM 5.14065-001/ X/ 1(27.5); NTM ex 5.14305-001/ C/ 1(28.5); NTM 5.14306-001/ X/ 1(45); ex NTM 5.13744-022/ X/ 1(21). Mugilogobius sarasinorum: NTM 5.12700-002/ X/ 4(24-46); ex NTM 5.12698-003/ C/ 2(34.5-40). Mugilogobius stigmaticus: leetotype of Gobius stigmaticus, AM5 1.358/ X/ 1(37); paralectotype of G. stigmaticus, AM5 1.361/ X/ 1(39); WAM P.28814001/ X/ 6(13-42); QM 1.25225/ X/ 11(16.5-29.5). Mugilogobius tigrinus sp. noy.: NTM 5.14288-003/ X/ 3(13-16); AM5 1.37570-002/ X/ 1(19.5); NTM 5.13953-017/ X/ 6(9.5-15.5); NTM 5.14318-001/ X/ 1(18); ex URM P.12664, C/ 1(16). Mugilogobius wilsoni sp. noy.: AM5 1.21259-004/ X/ 5(11.5-23); NTM unregistered, C/ 2(22-23.5). Oligolepis acutipennis, ex NTM 5.11125-020/ C/ 1(27); NTM 5.12122-004/ X/ 4(28-60). Oxyuricht/lYS ophthalmonema: ex NTM 5.12731-020/ C/ 2(33-45); NTM 5.12731-020/ X/ 10(28-62). Pandaka lidwilli: NTM unregistered, C/ 1(10.5). Pandaka pusilla: ex CA5 38588/ C/ 4(11-13). Pandaka pygmaea: ex CA5 47916/ C/ 2(105-11). Pandaka rouxi: CMK 8941/ X/ 6(12-16). Pseudogobius avicennia: ZRC 27007-8/ X/ Revision of Mugilogobius 2(25-26); KUMF unregistered, X, 1(30.5); ZRC 27451-3, X, 3(19.5-26); ZRC 20991-6, X, 6(23-30); ZRC 20628-34, X, 3(23-30). Pseudogobius javanicus: WAM P.30806-003, X, 10(21-35); ex NTM S.11125029, C, 2(30-31). Pseudogobius masago: holotype, N5MT 30228, X, 1(23); N5MT 34047, X, 1(19.5). Pseudogobius melanostictus: ZMH 19312, X, 3(30.536); CMK 6286, X, 3(31.5-35.5); U5NM 241842, X, 1(35.5); 5MF 18199, X, 1(40); ZRC 3521, X, 9(24-38); ex U5NM 268186, C, 2(27.5-33). Pseudogobius olorum: ex AMS 1.20158-001, C, 2(32.5-33); ex AM5 1.18478-003, C, 2(38-39); NTM unregistered, X, 7(37-53); 5AMA F.5123, X, 7(27-45). Pseudogobius poicilosomus: ex NTM 5.10426-014, C, 2(25-28). Pseudogobius sp. 8: NTM unregistered, X, 1(31.5); CMK 8310, X, 2(27-31). Pseudogobius sp. 9: ex AM5 1.20037-005, C, 1(27.5); AM5 1.21443-001, 10(2340.5); AM5 I.16960-002, X, 12(24-33). Redigobius balteatus: CMK 7184, X, 7(26.5-36); ex AM5 1.22055020, C, 3(17-27). Redigobius bikolanus: ex AM5 1.22041-014, C, 5(18-19); ex MNHN 1992.432, C, 1(29); ex NTM 5.12110-017, C, 2(17-21). Redigobius chrysosomus: NTM unregistered, C, 2(36-38); ex AM5 1.24683-003, C, 2(19-23.5). Redigobius dewaalii: 5AM 24154, X, 2(23-25.5); AM5 1.27219-001, X, 1(30). Redigobius dispar: syntypes, 2MB 6705, X, 5(31-33.5); syntypes, 2MB 6700, X, 6(40-41); syntypes, 2MB 6703, X, 2(30-36); syntypes, 2MB 6702, X, 4(38-42); ex U5NM 263330, C, 2(21.5-25.5). Redigobius macrostomus: AM5 1.16954-018, X, 9(1930); ex AM5 1.19341-002, C, 3(28-35.5). Redigobius penango: ex CMK 6143, C, 1(37); CMK 6143, X, 9(27-39.5); RMNH 12076, X, 2(42-42.5). Redigobius roemeri: syntype, 5MF 6703, X, 1(24); CA5 76087, X, 10(26.5-39). Rhinogobius brzmneus: ex NTM 5.12121002, C, 2(37-37); AMS C5G-200, C, 2(43-43). Rhinogobius sp.: ex CMK 4570, 1(29); U5NM 263425, X, 3(33-43.5). Schismatogobius sp.: ex NTM 5.12134-009, C, 1(19); AM5 CSG-276, C, 5(29-33). Stenogobius ophtlzalmoporus: ex NTM S.12134-007, C, 1(57). Stenogobius psilosinionus: NTM 5.11107001, X, 5(70-107). Stigmatogobius borneensis: syntypes, RMNH 6175, X, 8(38-46); ex NMW 4537, C, 1(48); NTM unregistered, C, 2(27-28). Stigmatogobius pleurostigma: ex AN5P 63116, C, 1(41.5). Stigmatogobius sadammdio: ex CMK 66278, C, 1(43); AN5P 77797, X, 4(34-51). Stigmatogobius sp. 4: ex U5NM 314469, C, 2(36-41); U5NM 314213, X, 8(28-37). Tamanka siitensis: AM5 C5G-383, C, 2(41-42); ex FMNH 47512, C, 2(51-73). ?Tukugobius carpenteri: RMNH 15087, X, 2(37-51); RMNH 12516, X, 2(43-45). GOBIIDAE: 5ICYDIINAE: Stiphodon elegans: ex NTM 5. 12114-002, C, 1(27). Sicydium vincente: ex NTM 5.13085-001, C, 1(37). GOBIIDAE: OXUDERCINAE. Apocryptodon madurensis: ex NTM 5.10798-037, C, 1(39). Boleophthalmus birdsongi: ex NTM S.11364-016, C, 1(51). Periophthalmus argentilineatus: NTM 13 unregistered, C, 1(50). Scartdaos lzistoplzorus: ex NTM S.10418-002, C, 2(35.5-62). GOBIIDAE: AMBLYOPINAE. Braclzyamblyopus rubristriatus: ex NTM 5.10208-003, C, 1(104). Taenioides limicola: ex NTM 5.12731-019, C, 2(67-84). MICRODE5MIDAE: Parioglossus dotui: ex NTM S.12098-014, C, 2(23.5-33.5). Ptereleotris microlepis: ex NTM 5.13237-030, C, 1(57.5). Phylogenetic analyses Character descriptions and polarities Three outgroups were used to make polarity decisions for the characters: the primitive gobioid Rlzyacich tlzys aspro (Valenciennes, 1837) (Rhyacichthyidae), and the odontobutids Micropercops borealis (Nichols, 1930) and Perccottus clzalmersi (Nichols and Pope, 1927). This last species may not actually belong in Perccottus, as it does not have vomerine teeth (personal observation). Rlzyacicllthys aspro, as the most primitive gobioid, was initially considered for use as an outgroup, but, because Rlzyacichthys also has a number of specialised characters (Miller, 1973a; 5pringer, 1983), the two odontobutid species were also used to elucidate which outgroup characters were plesiomorphic. As stated above, information on many character states of the apparently generalised rhyacichthyid Protogobius attiti Watson and Pollabauer, 1998, was not available (the description of the genus does not include any osteological information). The Odontobutidae has been established as having characters plesiomorphic to the Gobiidae (Hoese and Gill, 1993). Other gobiids initially considered for use as outgroups were the butines and eleotridines, which include a number of plesiomorphic species such as Bostryclzus sinensis and Oxyeleotris lineolata (Akihito, 1986). These two subfamilies display a wide range of character states, making polarity decisions difficult. Additionally, the relationships between butines, eleotridines, gobiines and gobionellines have not been clarified (Hoese and Gill, 1993). Therefore, no butine, eleotridine or gobiine species were used as an outgroup for polarising characters. 50me characters, differing among the outgroups, were included in the analyses to help polarise characters of the in-group (discussed below). For example, Micropercops has reduced lateral line canals and pores, while Rhyacichthys and Perccottus do not. Characters used are listed below and discussed; character state polarity (0 = primitive, 1 = derived etc.) is listed beside each character (Table 2). 5pecies are referred to by their nominal generic names in this section only, and are elsewhere referred by the generic assignments based on the outcome of the phylogenetic analyses. H.K. Larson 14 Table 2 Character matrix for phylogenetic analyses of the Mugilogobius-group of taxa. Missing data is indicated by a question mark. Rhyacichthys, Micropercops, and Perccottus are outgroups. Characters Species 000 0 0 0 0 o 0 1 1 1 1 1 1 1 1 1 122 2 2 2 2 2 2 2 2 3 333 3 3 3 3 334 1 2 3 4 5 6 7 8 9 012 34567 8 9 0 1 2 3 4 5 6 7 890 1 2 3 4 5 6 7 890 Rhyaeiehthys Micropereops Perceottus B. doriae B. kflbiliensis B. xanthomeIas B. xanthozona Ca. illota Ca. kflbilia Ca. mindora Ca. sp. noy. 2 Ca. sp. noy. 3 Ca. variegata Ch. eremius Ch. japalpa Ch. mieropterus Ch. ranunculus Ch. squamigenus E. sp. noy. E. microps E. oIigactis E. paludosus E. siamensis H. hoevenii H. mingi M. abei M. adeia M. amadi M. cagayanensis M. cavifrons M. ehulae M. fasciatus sp. noy. M. fiIifer sp. noy. M·fuseus M·fusculus M. Iatifrons M. Iepidotus sp. noy. M. Iittoralis M. mertoni M. myxodermus M. notospilus M. platynotus M. platystomus M. rambaiae M. rexi sp. noy. M. rivulus sp. noy. M. sarasinorum M. stigmatieus M. wiIsoni sp. noy. Pa. pygmaea P. avieennia P. javanicus P. melanostietus P. oIorum P. poicilosomus P. sp. 8 P. sp. 9 R. baIteatus R. bikolanus R. ehrysosomus R. dispar R. macrostomus R. penango R. roemeri S. borneensis S. pleurostigma S. sadanundio T. siitensis 0 1 0 2 2 2 2 1 2 2 2 2 1 2 2 2 2 2 1 2 0 2 1 1 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 0 0 0 0 0 0 0 1 1 1 2 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 1 o1 o1 o1 o1 o1 o1 o1 o1 o0 o0 o0 o0 o0 o0 o0 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 o1 1 0 o1 o1 o1 o1 o1 o1 o1 o0 o0 o0 o0 o0 o0 o0 o0 o0 o0 o0 0 0 0 0 0 0 0 1 1 1 1 1 1 0 0 0 0 0 ? 0 0 0 0 1 0 0 0 1 2 0 0 1 2 0 1 1 2 0 ? 1 2 0 0 2 1 0 o 1 1 101 1 001 1 001 1 001 1 001 2 111 2 111 2 ? 1 1 o2 1 1 1 o2 0 1 1 1 1 0 o 1 o 1 001 0 100 1 1 2 001 1 1 0 0 1 0 101 1 0 1 1 1 1 1 1 1 0 1 1 100 1 0 1 1 0 1 1 1 101 1 1 1 0 1 1 1 1 0 1 1 1 0 0 1 1 1 1 0 1 1 1 1 0 1 1 1 1 0 1 1 1 0 0 1 1 1 0 0 1 1 1 1 0 1 1 1 1 0 1 1 1 1 0 1 1 1 1 0 1 1 1 001 11101 1 1 1 0 1 o1 0 0 2 1 1 1 0 1 1 1 1 0 1 11001 11101 o1 0 0 1 o1 0 2 1 1 1 021 1 1 021 1 1 121 1 1 021 o 1 021 1 1 021 111 ? 1 110 0 1 111 0 2 110 0 1 110 0 1 110 0 1 010 0 1 010 0 2 010 ? 2 o1 0 0 2 o1 1 0 1 0 0 0 0 ? 0 0 1 0 1 1 1 1 1 1 1 1 1 0 0 1 0 0 o o o o o o o o o o o o o o o o o o o o o 1 o o o o o 1 o o o 1 o o o o o o o o o o o o o 1 0 0 1 1 1 1 1 1 1 1 1 1 0 0 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 2 2 2 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 2 0 1 0 2 0 1 0 2 2 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 o1 1 1 1 o1 0 1 1 1 1 1 0 0 1 1 1 0 0 1 1 ? o 0 1 1 1 o 0 1 1 ? o 0 o1 1 1 1 o1 1 1 1 o1 1 1 1 o1 1 1 1 o1 1 1 1 111 1 1 101 1 1 111 1 0 1 0 0 1 0 000 1 0 011 1 0 111 1 0 111 1 0 110 1 0 111 1 0 111 1 0 110 1 0 100 1 0 000 1 0 110 1 0 111 1 0 111 1 0 111 1 0 110 1 0 110 1 0 111 1 0 000 1 0 111 1 0 o0 0 1 0 111 1 0 110 1 0 010 2 1 1 ? 1 1 1 1 1 0 1 1 1 1 111 1 1 101 1 1 011 1 1 ? 1 1 1 1 1 1 1 0 0 1 1 1 1 0 0 1 1 0 0 ? 1 1 0 0 0 1 1 1 0 0 1 1 0 1 0 1 1 0 0 ? 1 1 0 1 1 1 2 0 1 1 1 2 0 1 1 1 2 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0 2 001 002 001 002 012 012 002 012 002 001 001 o0 ? o0 1 o0 ? 012 012 012 011 012 012 002 012 002 1 ? 2 1 1 2 0 1 2 0 1 2 0 0 2 0 1 2 0 1 2 0 1 2 002 o0 2 012 012 012 112 012 012 012 002 012 002 012 012 ? o 2 o0 ? 002 012 012 012 o0 ? o1 2 o1 2 o1 ? ? ? 1 0 1 1 0 1 2 0 1 2 0 1 ? ? 1 ? ? 1 2 ? 0 2 1 0 2 ? 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 0 0 1 1 0 1 1 0 1 1 0 1 1 o 1 001 011 001 001 o0 ? o0 1 o0 ? o0 1 1 0 1 o0 1 o0 1 1 0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 0 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 o0 1 ? 0 1 0 0 ? 0 o 1 0 o 1 0 o 1 0 o 0 0 0 ? 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 1 o2 0 0 0 0 0 0 0 0 0 o 0 0 0 0 0 0 0 0 0 0 o 0 0 0 0 0 0 0 0 0 0 o 0 1 1 0 0 0 0 1 1 0 o 0 1 1 0 0 0 0 1 1 0 o 0 1 1 0 0 0 0 1 1 0 o 0 1 1 0 0 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 0 0 1 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 000 1 1 0 o 1 1 1 0 100 1 1 0 o 1 1 1 0 100 1 1 0 o 0 1 1 ? ? ? ? 1 ? 0 o 1 1 1 0 1 0 0 1 1 0 o 0 1 1 ? ? ? ? 1 ? 0 o 0 1 1 0 1 o 0 1 1 0 o 0 1 1 0 1 o 1 1 1 0 o 0 1 0 0 1 o1 0 1 0 o 0 1 1 1 1 o 1 1 1 0 o 0 1 1 0 1 o 1 1 1 0 1 1 1 1 0 1 1 0 1 1 0 1 1 1 1 0 1 1 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 1 1 o 0 1 1 0 o 0 1 1 0 1 o 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 1 0 o 0 1 1 1 1 0 0 1 ? 0 o 0 1 1 1 1 0 0 1 1 ? ? 0 1 1 0 0 ? 0 0 1 0 1 1 1 1 ? ? ? ? 1 ? 0 1 1 1 1 o 1 1 o 1 1 0 1 1 1 1 o 1 1 o 1 1 0 0 1 1 1 o 1 1 o 1 1 0 1 1 1 1 o 0 1 o 1 ? 0 1 1 1 1 ? ? ? ? 1 ? 0 0 1 1 1 0 1 1 0 1 1 0 0 0 1 0 0 1 0 0 1 1 0 0 0 1 0 0 1 0 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 0 1 0 0 0 1 0 o 0 1 0 ? 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010 o 0 ? 010 o 0 0 010 o 1 0 000 15 Revision of MlIgilogobills Character 1 (0) Headpores present, rear portion of oculoscapular canal (over preopercle) present. (1) Headpores present, rear portion of oculoscapular canal (over preopercle) absent. (2) Headpores absent. Most eleotridines, butines and gobiines have headpores, as do Rhyacichthys and most species of odontobutids. Perccottus has a complete oculoscapular canal with six to eight pores and a preopercular canal containing five pores (Figure 7). Micropercops has a variable number of headpores, with the rear portion of the oculoscapular canal being reduced or absent (Figure 8; Hoese and Gill, 1993). Many gobioids with headpores also have an oculoscapular canal and terminal pore over the preopercle and opercle, but this is lacking in some diminutive coral reef genera such as Pleurosicya and its relatives, several species groups within Eviota, etc. (Akihito, 1986; Takagi, 1989; Pezold, 1993). Information on the recently described rhyacichthyid Protogobius was provided by Akihisa Iwata, too late to include in the phylogenetic anal yses. This fish does not share all the specialisations possessed by Rhyacichthys, and resembles Micropercops and Perccottus in papillae patterns (longitudinal) and in lateral line canals (e.g. does not have the suborbital part of the infraorbital canal), and has a complete oculoscapular canal which ends near the commencement of the lateral line along the body. Among the gobionellines, the rearmost section of the oculoscapular canal over the opercle is absent in Calamiana, Eugnathogobius, Hemigobius, Pseudogobius, Redigobius and Stigmatogobius. Redigobius and some species of Pseudogobiopsis have the canal over the preopercle present. Headpores are completely absent in species currently placed in Brachygobius, Caecogobius, Mugilogobius, Pandaka, Tamanka and Weberogobius. Calamiana and Eugnathogobius species show a range of character states. Figure 7 Perccottlls chalmersi, ex AMNH 10456, head pore and sensory papillae patterns. Scale bar 1 mm. , o '\ . . B \ \ V Figure 8 I Micropercops borealis, AMNH 10441, Shantung, China, headpores and sensory papillae, scales omitted. A, lateral view; B, dorsal view. Scale bar = 1 mm. Character 2 (0) First element in D2 and A unsegmented. (1) First element in D2 and A segmented. The first element in the second dorsal and anal fin in gobioids is nearly always an unsegmented ray, usually referred to as a spine. The derived state (first element segmented) is known among few gobioids. It is present in some members of the Amblyopinae and Oxudercinae (Hoese, 1984; Murdy, 1989). It occurs in all species of the gobionelline genera Pandaka, Brachygobius and the genera Gobiopterus, Leucopsarion and Mistichthys (Akihito et al., 1988; Kottelat et al., 1993). In the anal fin, this segmented ray is the supernumerary (anterior to first pterygiophore) element. The marine dwarf gobiine Trimmatom may have the first anal fin element (and sometimes the first element of the second dorsal fin) segmented or unsegmented, depending on species (Winterbottom, 1989). The condition in Trimmatom is not H.K. Larson 16 homologous to that in the gobionellines and Gobiopterus. In Trimmatom, the first element is formed by two hemitrichs as are the rest of the anal rays, " ...and thus technically a ray and not a spine, as in other gobiids" (Winterbottom, 1990). ,- A Character 3 Segmented caudal rays 17 or more. Segmented caudal rays 16 or fewer. (0) (1) The majority of gobioids, including the outgroup taxa, have 17 segmented caudal rays (Hoese, 1984), and this arrangement is a general character of the Percomorpha (Johnson and Patterson, 1993). There are not many gobiids with fewer than 17 segmented caudal rays apart from the eleotridines (which usually have 15); Nesogobius, a marine temperate Australian endemic gobiine of uncertain relationships (which has 13); and six of the nominal genera studied for this paper. These are Calamiana, ｾＧＺ ,, ｾ ,--, \, C 0 )'" /- /-::;. ｾ Chlamydogobius, Mugilogobius, Pseudogobius, Tamanka and Weberogobius, which all have 16 (occasionally fewer) segmented caudal rays. The type species of Tamanka, T. siitensis, has 17 segmented rays (see Phylogenetic Analyses, below; all other nominal species of this genus have 16). ｾ ., Ｎｾ E F .-- - ｾ '/ ,,/ '/ '/ J Character 4 Pectoral girdle smooth. (1) Pectoral girdle with lobes or flange. (0) Most gobioids have what is often referred to in the literature as a "smooth inner shoulder girdle" (referred to as "pectoral girdle" in this paper), actually the anterior edge of the cleithrum. In some gobionellines, there may be fleshy lobes or flaps present on the anterior edge of the pectoral girdle, or a bony or fleshy flange is visible, extending from the anterior edge of the cleithrum just below the skin (Figure 9). The flange may be thin, bony and bent laterally, fleshy or with irregular knobs. Possession of any combination of these projections was scored as (1). Awaous and Stenogobius have been characterised by having fleshy lobes or knobs on the inner edge of the pectoral girdle (e.g. Koumans, 1953; Watson, 1991), but Mugilogobius has only recently been acknowledged in the literature as possessing these structures (Larson and Kottelat, 1992; Kottelat et al., 1993). The lobes in Mugilogobius and its relatives (Figure 9C-F) differ slightly from those in Awaous and Stenogobius (Figure 9A-B) in that they are shorter and fleshier (not elongate and finger-like). In his review of Awaous, Watson (1992) did not mention these characteristic lobes in his diagnosis of the genus, referring to them only in his redescription of two species. The pectoral girdle is smooth in Caecogobius, Chlamydogobius, Hemigobius, Pandaka and Stigmatogobius and in most Brachygobius species (in ｾ Figure 9 '. !, )' ------...::..' Inner edge of gill opening (pectoral girdle) showing development of fleshy ridge, lobes and flange bearing lobes in: A, Awaolls banana, NTM 5.13083-001; B, Stenogobills psilosinionlls, NTM 5.11107-001; C, Redigobills roemeri, CMK 11334; D, Tamanka siitensis, CA5 36824; E, MlIgilogobills cavifrons, NTM 5.13482-001; F, MlIgilogobills rambaiae, NTM unregistered. Scale bar =1 mm. some specimens of at least two species of Brachygobius, the pectoral girdle has a fleshy ridge). Lobes and/or a flange are present in most Mugilogobius and in most species of Calamiana, Eugnathogobius, Pseudogobiopsis, Pseudogobius, Tamanka and Redigobius. Weberogobius amadi and Tamanka siitensis differ from other Mugilogobius and Tamanka in having a smooth-edged pectoral girdle. Aurich (1938) did describe papillae on the pectoral girdle of Vaimosa latifrons (= Mugilogobius latifrons), but this character was not referred to by any other workers until Koumans (1953), who included these structures in his definition of Tamanka (which included some nominal species of Mugilogobius and Pseudogobius. This was probably because Koumans compiled his definition mostly from the literature and used Aurich's descriptions of Vaimosa latifrons, V. cagayanensis and V. zebra. Revision of Mugilogobius 17 A ｾＬｾｾｾ B c Figure 10 Caudal skeleton of: A, Rhyacichthys aspro, CAS/SU 38565, (cartilage not shown, specimen not counterstained); B, Micropercops borealis, ex AMNH 10441; C, Acentrogobius caninlls, NTM unregistered; D, Mugilogobius notospillls, ex ZMH 19346. Cartilage shown as black. Scale bar = 1 mm. Character 5 Three epurals. (1) Two epurals. (2) One epural. Rhyacichthys has three epurals (Miller, 1973a) (Figure lOA), with all other gobioids characteristically possessing one or two; however, some odontobutids may occasionally possess three epurals (Akihito, 1986) (Figure lOB). Epural number in odontobutids may vary ontogenetically (Gilt 1997, ill litL), with two epurals splitting to form (0) three or four. The number of epurals appears to be useful for classification within the gobiids, as gobiines (sensu Pezold, 1993) have only one epural (Figure 10Cl, and gobionellines generally (but not always) have two (Figure 100). Possession of one or two epurals may be helpful in determining relationships among the gobionellines. Those possessing two epurals are Calamiana, Caecogobius, EugnatJlOgobius, Mugilogobius, Pandaka, Pseudogobiopsis, Pseudogobius, Redigobius, Stigmatogobius, Tamanka and Weberogobius. Those H.K. Larson 18 with one epural are Brachygobius and Chlamydogobius. Calamiana paludosus always has one epural (however, it shares a set of characters with C. sp. novo 6, which has two epurals). Sometimes a single epural appears to be partly split in two. This may occur in Chlamydogobius, some specimens of Mugilogobius and Pandaka (specimens of P. pygmaea often have one epural which appears to be half split in two from its ventral margin). Watson (1993) characterised and illustrated the epural of Parawaous as broad and notched ventrally; he considers this to be intermediate in form between Awaous (which has one epural) and Stenogobius (two epurals). He does not tell us if the single epural in Awaous is variable in shape or sometimes partly split (a specimen of Awaous acritosus Watson examined for this study has a narrow split or notch ventrally). Character 6 Narrow, pointed neural spines. (1) Second to fifth neural spines expanded and bifid or split. Rhyacichthys and odontobutids have relatively (0) A B c Figure 11 First four vertebrae of: A, Mugilogobius abei, ex YCM 908; B, Tamanka siitensis, ex FMNH 47512; e, Oxyurichthys ophthalmonema, ex NTM 5.12731-020; 0, Hemigobius mingi, ex URM 6677; E, Pseudogobius melanostictus, ex U5NM 268186; F, Awaous acritosus, NTM unregistered. Only first two epineurals included. Scale bar = 1 mm. Revision of Mugilogobius straight, pointed neural spines (Miller, 1973a; this study), as do most other gobioids. Murdy (1989) states that Oxuderces has a "posteriorly expanded fourth neural spine", but does not indicate whether all oxudercines have this state. MugiIogobius and several other gobionellines (Clzlamydogobius, Eugnathogobius, Hemigobius hoevenii, some Redigobius and Tamanka) modally have the first few neural spines with expanded or variably split tips (Figure 11). The character does not always appear in all specimens and species of these genera. Gilbert and Randall (1979) referred to the "morphology of neural spine lying between first and second dorsal pterygiophores" as being one of nine possibly useful taxonomic characters within the Gobionellus species-group, but did not elaborate further on this. They also discussed the close relationship between Gobionellus, Oxyurichthys and Evorthodus (all gobionellines). Pezold and Gilbert (1989) observed that a speciesgroup of Gobionellus had a "basally flared fourth neural spine". In an unpublished manuscript, Pezold also reported that in a number of Gobionellus species, the fourth neural spine has a broad flange running along most of its rear margin, with no such flange present on neighbouring spines. Pezold (in prep.) characterised Oxyurichthys as possessing a distinctly bifid, expanded third neural spine (not observed in specimen in Figure 11C). These characters mayor may not be homologous with the broad, bifid or split third to fifth neural spines in the Mugilogobius-group of gobionellines. Character 7 Gut short, S-bend shape. (1) Gut long, intestine coiled separate from stomaclz. (2) Gut long, coiled spirally around itself and stomaclz. Most gobiids are carnivorous, and have a simple "S-bend" shaped gut, with a muscular stomach separated from a short intestine that makes two bends (Geevarghese, 1983; personal observation). Geevarghese (1983) and Mok (1980) carried out surveys of gut morphology in gobioid fishes. Assignation of polarity to the different ways the gut coils other than the plesiomorphic single-loop (Sbend) form is here based on information provided by Geevarghese and Mok. The many-looped "watchspring" form (state 1) is similar to that found in tlzys and (Geevarghese, 19 A o Figure 12 E Ventral views of: A, Hemigobius mingi, stomach; B, Hemigobius mingi, intestine coiled upon stomach; C, Chlamydogobius japalpa, intestine coiled upon stomach; D, Awaous banana, intestine coiled around stomach in corkscrew manner (liver removed for clarity); E, Pseudogobius javanicus, intestine coiled around stomach in corkscrew manner (liver removed for clarity); F, Mugilogobius abei, simple looped gut. Not drawn to scale. (0) as I.F" J-lemI5?obws and all have a long, manylooped intestine, although the pattern made by the looping differs between the genera. In Clzlilfll1/,:io,I;Dbills and Hemigobills (Figure 12A-e), the intestine is tightly coiled in a series of close-packed loops (centre of coiling perpendicular to the body axis) which do not coil around the stomach ("watchspring", resembling Mok, 1980: figure 10, intestinal pattern "type SP"). In Awaous and Pseudogobius (Figure 120-E), the intestinal loops are coiled in a corkscrew manner around the body axis and around the stomach itself (which is also somewhat twisted). All others have a simple 5-bend gut form. Gut contents were not examined in detail to correlate with gut morphology. However, several studies have been made on the diet of Mugilogobius-group relatives. Gill and Potter (1993) and Humphries and Potter (1993) studied gut contents of Pseudogobius olorum from the Swan Estuary and Wilson Inlet, WA. The fish fed chiefly on diatoms, detritus, amphipods, green algae and polychaetes, taking much of their food from algal mats, and the coiled gut morphology reflects the diet. Glover (1971) found that detritus, green algae, plant fragments, diatoms and insects formed the major portions of Chlamydogobius eremius' diet. 20 H.K. Larson A " ..: : " .' .. ........... ........... . ... ':'... .":. '. ' . . :" " '0' ;' " /' . • .:: 0 .; • ,. 0 :';. ..... \ ' .. . " : . ,'. ill' :::<;./. B J: vi: : a . : 0 \.:. ./:':'<19:::>\ . : .... .. ' :' I" ,, ': , .. I: , , \ Figure 13 ｳｹｨｴ｣ｪｾｹｨｒ aspro (CAS/SU 38565) sensory paplllae arrangement (headpores omitted for clarity). A, lateral; B, dorsal views. Scale bar =lmm. Character 8 (0) Scattered papillae, no clearly distinct pattern. (1) Longitudinal papillae pattern. (2) Transverse papillae pattern. The sensory papillae of gobioid fishes have received a considerable amount of attention, but there is still little agreement on how the various patterns should be polarised for use in determining taxa and relationships (e.g. Hoese, 1983; Miller and Wongrat, 1979; Hoese and Gill, 1993). The papilla structure, their differing sizes, the innervation and orientation of the papillae within the rows has been described in varying detail (Hoese, 1983; Marshall, 1986; Wongrat and Miller, 1991). Song and Boord (1993) have shown homology and conservatism in head innervation throughout the fishes (sharks to teleosts). Rhyacichthys has a rather disorganised papillae arrangement (Figure 13) in which it is difficult to discern any definite pattern, other than that the dorsal-most papillae are approximately arranged in lines, while those on the side of the head form clusters. Specimens over 100 mm SL have the papillae covering the snout and lower side of the head proliferated into broad patches or bands. This is unlike the arrangements in other gobioids, which can be generally grouped as having a longitudinal or transverse pattern (Hoese, 1983). Protogobius, the new rhyacichthyid from New Caledonia (mentioned above; information arriving too late to include in analyses), differs considerably from Rhyacichthys in that it clearly has a distinct longitudinal papillae pattern reminiscent of Micropercops and Perccottus. Odontobutids have a longitudinal pattern, while gobiids have both, with some genera having a "mixed" pattern. Hoese and Gill (1993) discussed some of the problems of polarity and using papillae patterns in classification, but they indicated that the patterns are often correlated with other characters, such as the dorsal pterygiophore pattern. Transverse patterns can be derived from longitudinal patterns (see Hoese, 1983) and this has very probably happened on more than one occasion. Within the Gobionellinae, both transverse (Figure 2) and longitudinal (Figure 5) papillae patterns are exhibited. Calamiana, Brachygobius, Caecogobius, Chlamydogobius, Eugnathogobius, Mugilogobius, Pandaka, Hemigobius, Pseudogobiopsis, Pseudogobius, Redigobius, Tamanka and Weberogobius species have "longitudinal" patterns, with a few exceptions (Brachygobius xanthozonus, Tamanka rexi sp. nov., Redigobius chrysosomus). Some specimens of T. sarasinorum show proliferation of papillae, resembling a transverse pattern. Stigmatogobius species always have transverse patterns. Character 9 Three or more s papillae rows on snout. (1) Two s papillae rows on snout. The infraorbital s and r papillae rows (sensu Sanzo 1911) on the snout are reduced to two or three in this group of fishes. In this paper, the rows crossing the snout are referred to as s rows and any median longitudinal rows are called r rows (various designations for these rows exist in the literature). The outgroups and most species within the gobionellines have three or more short s and r rows across the snout; the anteriormost s row, usually the longest, is placed very close behind the upper lip (Figures 3B, 5). Calamiana and Chlamydogobius have only two s rows, which may be represented by only one papilla. Some specimens of Pseudogobiopsis oligactis, two nominal species of Pseudogobius, Tamanka lepidotus sp. novo and T. rexi sp. novo may also have only two s rows or a single papilla present in each row (Figure 4A). (0) Character 10 with at least one more ray than A. and A ray numbers modally equal. (2) A with one more ray than D2. Rhyacichthys has equal numbers of dorsal and (0) D2 (1) D2 Revision of Mugilogobiu5 21 anal rays (usually 1,8), or one more dorsal than anal ray (Miller, 1973a); and Akihito et al. (1988) reported counts of 1,8 second dorsal rays and 1,10 anal rays. Of the 10 Rhyacichthys specimens examined for this paper, six had character state 1, and three had state O. The odontobutids Micropercops, Odontobutis and Perccottus have state O. Many eleotridines, butines and gobiines have character states 0 or 1, with state 2 being rare among gobiines (e.g. Silhouettea has one or two more anal than second dorsal rays). Hoese (1984) indicated that the Eleotrididae (= Odontobutidae, Butinae and Eleotrinae) had a greater range of second dorsal rays than anal rays than did the Gobiinae (=Gobiinae and Gobionellinae). However, state 2 is characteristic of the gobionellines Evorthodus, Gnatholepis, Gobiopterus, Oxyurichthys and Stenogobius. Caecogobius, Chlamydogobius (c. ranunculus sometimes has state 1) and most species of Redigobius have state O. Calamiana, Brachygobius, Eugnathogobius, Hemigobius, Mugilogobius, Pandaka and Pseudogobius have state 1. Stigmatogobius species (and some specimens of Weberogobius amadi) have state 2. Character 11 Mouth terminal or lower jaw tip anterionnost when mouth closed. (1) Mouth subtenninal or upper jaw tip anterionnost when mouth closed. Odontobutids and most butines and eleotridines usually have terminal mouths, or the lower jaw tip protrudes. The specialised, fast stream-dwelling Rhyacichthys has a small subterminal mouth below a flattened snout. Gobiines and gobionellines have a variety of mouth forms, although a terminal mouth is usuaL Possession of a terminal mouth may be linked to prey capture (odontobutids, eleotridines, butines and the majority of gobiines are carnivorous). Mugilogobius, Pseudogobius, Hemigobius, Chlamydogobius and Redigobius have subterminal mouths. (0) Character 12 (0) Three or more anal pterygiophores before first haemal spine. (1) Two anal pterygiophores before first haemal spine. Higher numbers of anal pterygiophores anterior to the first caudal vertebra are generally associated with rhyacichthyids, odontobutids, eleotridines, butines and some of the gobionellines (Birdsong et al., 1988), although the character is not evenly distributed among species of these groups (Figure 14). The derived condition (fewer pterygiophores) is present in Calamial1a, two nominal species of Brachygobius, Eugl1athogobius, Hemigobius hoevel1ii, Mugilogobius, Pal1daka, Pseudogobiopsis, Pseudogobius and Stigmatogobius. The plesiomorphic condition is Figure 14 First three anal pterygiophores anterior to the first haemal spine in Redigobiu5 dispar, ex USNM 263330. Scale bar = 1 mm. present in Redigobius, two nominal species of Bracllygobius, Hemigobius mil1gi and the central Sulawesi lake species (M. adeia, W. amadi, M. latifrol1s, T. rexi sp. nov., T. sarasil10rum and T. lepidotus sp. nov.). Character 13 Metapterygoid low, narrow. (1) Metapterygoid broad, expanded dorsally. The metapterygoid is variable in shape among gobioids. It is low and narrow in Rhyacichthys, Micropercops and Perccottus, and low and relatively narrow in some gobiines and gobionellines (Figure 15; Miller 1973a; Harrison 1989). The plesiomorphic butines such as Oxyeleotris and Priol1obutis have deep metapterygoids. Chlamydogobius, Eugnathogobius, Hemigobius, Pseudogobiopsis, some species of Pseudogobius, several species of Calamial1a, Mugilogobius, Stigmatogobius, some Redigobius and some Tamal1ka have the derived (broad) condition (Figure 16). Other gobionellines such as Awaous, Evorthodus, Oligolepis, Oxyurichthys, Rhinogobius and Stenogobius generally have a low metapterygoid; in Gl1atholepis the moderately low metapterygoid has a dorsal (0) H.K. Larson 22 hyomandibular A c symplectic B o E F Figure 15 Metapterygoid shape in: A; Perccottlls chalmersi, ex AMNH 10456; B, Micropercops borealis, ex AMNH 10441; C: Gnatholepis sp., ex NTM 5.12883-015; D: Stenogobills ophthalmoporus, ex NTM 5.12134-007; E: Acentrogobills caninlls, NTM unregistered; F: Rhinogobills brunnells, ex NTM 5.12121-002. Cartilage shown as black. 5cale bar = 1 rnrn. spur. Winterbottom suggested that a possible autapomorphy for the Gobioidei was that the symplectic "is usually considerably wider than the metapterygoid" (Winterbottom, 1993: 399, figure 2). This may be the condition in Rhyacichthys, but the odontobutids Micropercops, Odontobutis and Perccottus all have the symplectic about equal in width to the metapterygoid. The depth of the metapterygoid may be sometimes sexually dimorphic. Species of the Sulawesi lake-dwelling species complex have a very low (narrow) metapterygoid in both sexes. Most other Mugilogobius (and Calamiana kabilia) have a broad, dorsally expanded metapterygoid in adult males, with adult females having a medium to broad metapterygoid which is not so broadly expanded dorsally. In several species for which only adult females were available for clearing and staining (e.g. the small species Mugilogobius tigrinus sp. nov.), the metapterygoid is low, and thus was scored as such for the data matrix. Character 14 (0) Vertebrae 28-29 (or more). (1) Vertebrae 26-27. (2) Vertebrae 24-25. The odontobutids have high vertebral counts of 29-32 (arranged in precaudalj caudal vertebral Revision of Mugilogobius 23 hyomandibular preoperculum B c E o F Figure 16 Metapterygoid shape in: A: MlIgilogobills rambaiae, ex CA5 36032; B: MlIgilogobius amadi, ex ZMH 7579; C: Mugilogobills adeia, ex NTM 5.13068-001; D: Pseudogobius melanostictus, ex U5NM 268186; E: Tamanka siitensis, ex FMNH 47512; F: MlIgilogobius sarasinorllm, ex NTM 5.12698-003. Cartilage shown as black. Scale bar = 1 mm. patterns of 15-17+14-17), higher than Rhyacichthys, which has 28 vertebrae (in 10-12+16-18 pattern) (Birdsong et al., 1988; this paper). This difference between the two groups may be due to the fact that odontobutids are temperate (gobioids in temperate regions tending to have higher vertebral numbers), while Rhyaciclzthys is tropical in distribution. Butines and eleotridines have 25-32 vertebrae, in a range of precaudal/caudal patterns. Xenisthmids have 26-45 vertebrae. The proportion of precaudal to caudal vertebrae can be useful in grouping species or genera (e.g. 11 +15 versus 10+16). An attempt was made at polarising these patterns, but proved difficult and was abandoned, with only total numbers used. Further work may clarify polarity. The first caudal vertebra was defined as that having a complete haemal arch, as the distribution of pleural ribs on the last few precaudal vertebrae is variable within gobioids (Birdsong (1975) differentiated caudal from precaudal vertebrae by absence of pleural ribs on the caudal vertebrae, in addition to their having a closed haemal arch). Gill (1994) polarises vertebral numbers in the reverse direction (26 as plesiomorphic, 27 as H.K. Larson 24 derived) in his work on the monophyly of the "longitudinal" gobies of the Bathygobius and Priolepis Groups of Birdsong et al. He was using Pseudogobius and Tridentiger as outgroups for his gobiine taxa. Pseudogobius has 26, rarely 27 vertebrae (this paper), while Tridentiger has 26 (Birdsong et al., 1988). Most gobionellines, and many gobiines and oxudercines, have 26 (10+16) total vertebrae. Chlamydogobius has 27-28 vertebrae (11+16 or more) (Larson 1995). Among the nominal species of Mugilogobius, which modally have a 10+16 pattern, the Australian temperate species M. platynotus has 27 (11 +17) vertebrae. The isolated species from the tectonic lakes of central Sulawesi (M. adeia, W. amadi, T. sarasinorum, T. rexi sp. novo and T. lepidotus sp. nov.,) have a pattern of 11+15-16, similar to that in Chlamydogobius. The "dwarfed" gobionellines Brachygobius, Gobiopterus and Pandaka all share low numbers of vertebrae: 10+14-15. Redigobius has a characteristic vertebral pattern of 11+15-17. Character 15 (0) Papilla row p continuous, with at least 10 papillae present. (1) Papillae in row p widely spaced, six or fewer papillae present. (2) Papilla row p includes short transverse rows. In odontobutids and other plesiomorphic gobiids (and Mugilogobius, Tamanka and Weberogobius), the interorbital papilla row p is composed of small, closely spaced papillae (Figure 5) wherever it is not replaced by the oculoscapular canals. In Hemigobius, Pseudogobius, Eugnathogobius, Pseudogobiopsis, Calamiana and Redigobius, the few p row papillae are widely separated (Figure 17). Stigmatogobius and Brachygobius xanthozona have several transverse p rows (Figure 3B) in the interorbital region. Character 16 Papilla row a runs dorsoposteriorly, following curve of eye. (1) Papilla row a bends rearward past eye, may be horizontal. Miller (1987) pointed out that in Weberogobius amadi, row a does not turn up to follow the curve of (0) the rear of the eye (as it does in the outgroups and other gobionellines); instead it turns sharply rearward (Figure 5). This character state, although less exaggerated, is found in Tamanka siitensis and in some specimens of T. cagayanensis, Mugilogobius notospilus and M. rivulus sp. novo The derived character state also known in some eleotridines (Miller, 1987). Although it is recognised that homology for this papillae arrangement remains to be confirmed, the character was retained for the analyses. Character 17 Mouths similar in males and females. Mouths enlarged in males, sometimes greatly. (0) (1) In the outgroups, as with the majority of gobiids, males and females have similar-sized mouths. In some gobionelline species, sexually mature males have considerably enlarged mouths (reaching to below mid-eye, or to rear of eye, or further), as in most species of Mugilogobius, Tamanka, Pseudogobiopsis, Calamiana, Eugnathogobius and Redigobius (Figure 4A). Mature males of Calamiana mindora, C. sp. nov. 3, Chlamydogobius ranunculus, Hemigobius hoevenii and several species of Pseudogobius have mouths slightly larger than females, but they do not develop the greatly elongate jaws. Species of Gobiopterus often have larger mouths and much larger teeth in males, while males of the Australian endemic eleotridine Philypnodon have considerably enlarged jaws (unlike most eleotridines). Watson and Horsthemke (1995) describe the elongate jaws of mature males of the gobionelline Awaous flavus and state that other species of Awaous do not exhibit enlarged jaws. In Rhinogobius also, males tend to have longer jaws than females (Akihito et al., 1988: pIs 248-249). Further survey of the gobionellines may indicate that this tendency is characteristic of the group. Character 18 Upper posterior extension of procurrent cartilage not over tip ofepurals. (1) Upper posterior extension of procurrent cartilage halfway over epural, but not over tip ofepural shaft. (2) Upper posterior extension of procurrent cartilage over tip ofepural(s). (0) Figure 17 Sensory papillae in Calamiana kabilia, holotype (CAS 32978); papilla row e turns onto cheek. Scale bar = 1 mm. Hoese and Gill (1993) discuss this character and its polarity in relation to eleotridine relationships, Revision of Mugilogobius finding the plesiomorphic state to be present in odontobutids, butines and some gobiines. One or the other of the derived states (Figure lOC-D) is present in all gobionellines examined for this study. In gobionellines with a single epural, cartilage may extend posteriorly over the tip of the epural shaft itself (as in Pandaka), not just its anterior flange. This condition occurs in eleotridines also (Hoese and Gill, 1993: figure 2E). Hoese and Gill (1993) discuss possible links between the derived state 2 of this character and the number of segmented caudal rays in eleotridines (15, with 17 in odontobutids and most butines). In the gobionellines examined for this study, there was no correlation found between caudal ray number and caudal cartilage extent. Character 19 (0) Papilla row efollows preopercular margin. (1) Papilla row e extends onto cheek before preopercular margin. In nearly all gobiids, the papilla row e runs along or close to the edge of the preopercle (Akihito et al., 1988; Takagi 1989). In Eugnathogobius microps, Pseudogobiopsis siamensis (Figure 17) and two Calamiana species (illata and kabilia), row e turns upward onto the cheek well in advance of the rear edge of the preopercle. 25 cartilage, and referred to "broken" and "blotched" scapula types which were actually partly ossified scapulae. He found ossified scapulae in most, but not all, gobiines and gobionellines examined, and discussed the variable ossification of the scapula in specimens of Mugilogobius abei (Akihito, 1963). The upper part of the scapula above or around the foramen is ossified in Redigobius, Stigmatogobius and some specimens of Mugilogobius abei, M. adeia, M. chulae and M. mindora. Most Mugilogobius and Tamanka have an unossified scapula. Character 22 Adductor mandibulae tendon with two attachment points. (1) Adductor mandibulae tendon with one attachment. In gobionellines, the tendon of the Alb section of the adductor mandibulae muscle inserts onto the posterior or inner face of the maxilla, at about the halfway point (Figure 18), similar to that described and illustrated by Hoese and Gill (1993: figure lA) for Oxyeleotris lineolata, in which the insertion point is closer to the head of the bone. The tendon is often broad, giving the appearance of the muscle inserting directly onto the bone. Additionally, the ligament from the Al and A2 sections of the muscle inserts near, or next to, the tendon. The ligament and tendon are often difficult to distinguish in the (0) Character 20 (0) Upper jaw teeth in two or more rows. (1) Upper jaw teeth in single row. The majority of gobiids have more than one row of teeth in each jaw, as do the outgroups. The oxudercines (except for Periophthalmodon), Evorthodus, and most sicydiines have one row of teeth in the upper jaw (Murdy, 1989). Most species of the gobionelline genus Oxyurichthys have one row of upper jaw teeth (Pezold, in lift.), as do some Gobiopterus. The form and arrangement of teeth in Calamiana variegata does not resemble that found in any of the abovementioned. The teeth are compressed, evenly sized and have the pointed tips slightly bent to one side (despite their shape, the teeth were coded as state 1). This character was ignored for analyses within the Mugilogobius Group (where it is an autapomorphy for C. variegata), but was used for analyses within the Gobionellinae. Character 21 Scapula partly ossified. (1) Scapula Imossified. The scapula is well-developed and partly ossified in odontobutids and Rhyacichthys and ossified in some gobiines and gobionellines (Akihito, 1969; Hoese and Gill, 1993). Akihito used material that was not counter-stained with alcian blue for (0) A1B - addudor mandibulae Figure 18 Adductor mandibulae insertion in: A, MlIgilogobill5 cavifr0I15, 45 mm SL, ex BPBM 33930, Hawaii; B, MlIgilogobill5 latifro115, 25 mm SL, ex NTM S.12706-005, Lake Towuti, Sulawesi; C, Redigobill5 di5par, 25.5 mm SL, ex USNM 263330, Lake Buhi, Luzon; D, MlIgilogobill5 platyl1otll5, 26 mm SL, ex AMS 1.25038-001, Brunswick Heads, NSW. Not drawn to scale. H.K. Larson 26 presence of the bony support or groove for the preopercular canal, which extends along the rear and ventral margins of the preopercle. This character is present in odontobutids (Figure 15AB), but absent or restricted to the rear margin in gobiines (Figure 15E). In gobionellines, the bony support may be present or absent, usually depending if preopercular pores are present. In some genera (Awaous, Oligolepis, Oxyuriclztlzys, Redigobius, Stenogobius) the bony support runs the length of the preopercle, while in others (Evorthodus, Gnatlzolepis, Rhinogobius, Pseudogobiopsis oligactis) it is restricted to the rear of the bone. In Calamiana, Chlamydogobius, Braclzygobius, Pseudogobiopsis siamensis, Hemigobius, Mugilo-gobius, Pandaka, Tamanka, Weberogobius and Pseudogobius the support is absent. Hemigobius hoevenii and Pseudogobius poicilosomus, which lack preopercular pores, have a low bony ridge (but no fold or groove) along the rear of the preopercle. Calamiana paludosus, which lacks preopercular pores, has a low ridge with a shallow groove posterior to it. Character 24 Haemal arch offirst caudal vertebra narrow, relatively straight or angled backward. (1) Haemal arch offirst caudal vertebra wide, curved forward toward tip ofanal pterygiophore. The derived character state of this character (Figure 19) appears to be found only in three nominal species of Stigmatogobius; in one of these, S. borneensis, the haemal arch is not always curved. Birdsong et al. (1988: figure 3C) illustrate a similar condition in the microdesmid genus Ptereleotris. Additional material needs to be examined to confirm if this is an autapomorphy for the genus Stigmatogobius. (0) Figure 19 Stigmatogobius pleurostigma, ex ANSP 63116, 41.5 mm SL, Bangkok, showing curved haemal arch. A, lateral view; B, diagrammatic anterior view of shape of first (outer line) and second (inner) haemal arches. Not drawn to scale. cleared and double-stained material that was used to examine this feature. Hoese and Gill describe a different attachment method for the ligament and tendon in the eleotridines (1993: figure 18). All gobionelline species examined for this character resembled the condition found in odontobutids and butines. The tendon inserted onto the bone in two sections, often close together, or as a continuous broad sheet; with each condition equally likely in a number of species (one specimen may have one form and another, the other form). This may be a synapomorphy shared by the gobionellines and gobiines, and the character was not included in the final analyses (character used only in initial analyses). Clzaracter 23 Bony preopercular canal support present. (1) Bony preopercular canal support absent. Hoese and Gill (1993) discuss and illustrate the (0) Character 25 Lips normal, fleshy or thick. Lips very thin, lower lip reduced. Most gobioids have fleshy to thick lips. Species of the genera Hemigobius and Pseudogobius have the lower lip fold reduced in size and thickness, so that the lip is small, thin and flattened and reduced to the mouth corner (Figure 20). The remainder of the lip is completely flat and fused to the underside of the head (resembling lips of mugilids) in Hemigobius and in some Pseudogobius, or slightly fleshy in some Pseudogobius species. The upper lip is also relatively narrow and thin in both genera. The lips in sexually mature males of Pseudogobius may be somewhat inflated and fleshy when compared with lips of females. One species, the temperate Australian P. olorum, tends to have relatively fleshy lips in both sexes. (0) (1) Revision of Mugilogobius 27 with reduction in the number of pores or complete loss of the canal having occurred many times (Takagi, 1989; Pezold, 1993), often independently of the development of the oculoscapular canal system. Most eleotridines, butines and gobiines have preopercular pores; and within the Gobiinae, the number of preopercular pores may vary within a genus, for example Callogobius and Gobiopsis. Most gobionellines with oculoscapular canals present also have preopercular pores. However, in Calamiana illota, C. paludosus, Pseudogobiopsis siamensis, Stigmatogobius borneensis and all species of Oxyurichthys, preopercular pores are absent. A Character 29 No bridge from metapterygoid to quadrate. (1) Bridge from metapterygoid to quadrate. Miller (1973a, 1987) discussed the metapterygoid bridge and its possibilities as a useful character, indicating that it occurs in "more primitive gobiines". Harrison (1989) also considered that "primitive gobiids" such as Gobius niger possessed a bridge to the quadrate. His illustration of eight gobionellines (1989: figure 7A-H) (Ctenogobius, Evorthodus, Gnatholepis, Gobionellus, Oligolepis, 'Gobius' kiensis, Stenogobius) shows none of them possessing a bridge. In Rhyacichthys, the metapterygoid is low and relatively narrow, and does not contact the quadrate. The metapterygoid bridge is absent in the odontobutids Micropercops and Perccottus (but a narrow bridge toward, but not touching, the quadrate is present in Odontobutis). It is found in the butine Bostrychus, and a number of generally unspecialised gobiines (Acentrogobius, Bathygobius, Glossogobius, Gobius). The xenisthmid Paraxenisthmus springeri Gill and Hoese has a broad metapterygoid which overlaps part of the quadrate and contacts the pterygoid. The bridge is here polarised as a derived character. Mugilogobius, Tamanka and Calamiana species always have a bridge (usually broad) overlapping the quadrate (Figure 16). Tamanka lepidotus sp. nov., a highly derived, small species from Sulawesi, has the anterior portion of the metapterygoid poorly ossified, but extending toward the quadrate. This was coded as state 0, and is considered to represent a loss, as all of its relatives in Sulawesi lakes have a bridge. A bridge is found also in Stigmatogobius pleurostigma and S. sadammdio. In genera without a bridge, other than Brachygobius and Pandaka, the metapterygoid is often expanded dorsally, and may have a flat spur of bone pointing dorsally or anterodorsally (Figure 15). Brac11ygobius and Pandaka both have very low, narrow metapterygoids which are not produced upward. (0) B Figure 20 Lower lip shape in: A, Hemigobius mingi, ZRC 20263-73; B, Pseudogobius javanicus, NTM 5.11125-016. Not drawn to scale. Character 26 Snout normal or broad, not steep or inflated. (2) (1) Snout inflated, overhanging upper lip. (3) Snout flat, shovel-like. Most eleotridines, gobiines and gobionellines have a broad to flattened, rounded or pointed snout that is not inflated or "swollen" and partly overhanging a subterminal mouth. Pseudogobius characteristically has an inflated snout, as does Hemigobius hoevenii. Some specimens of Chlamydogobius eremius, C. japalpa and C. ranunculus have a somewhat inflated snout. The specialised Rhyacichthys has a very flat laterally expanded snout (character state 3). (0) Character 27 Branchiostegal rays 6. (1) Branchiostegal rays 5. Rhyacichthys, odontobutids, xenisthmids, eleotridines and butines have six branchiostegal rays, with all other gobioids having five (Hoese and Gill, 1993). (0) Character 28 Preopercular pores present. (1) Preopercular pores absent. This character is somewhat variable in gobioids, (0) Character 30 (0) Palatine short, halfway down length of pterygoid. H.K. Larson 28 A G c Figure 21 Fifth ceratobranchial form in: A, Awaous acritosus, NTM unregistered, completely honeycombed with tiny holes (only partly indicated); B, Pseudogobius javanicus, ex NTM 5.11125-029, finely honeycombed (partly indicated) thin bones; C, Gnatholepis sp., ex NTM 5.12883-015, bones partly honeycombed; D, Hemigobius mingi, ex URM 6677, lace-like network (partly indicated) formed by two layers of interconnecting struts; E, Chlamydogobius eremius, ex AM5 1.24493-001, stout, but partly honeycombed; F, Mugilogobius chulae, ex URM 9324, only few holes present; G, Apocryptodon madurensis, ex NTM 5.10798-037, covered with lattice-work of fine straight struts overlain by honeycomb-like cells. Cartilage shown as black. Scale bar =1 mm. (1) Palatine long, reaching down nearly to, or meeting, quadrate. This is the same character as "character A" used by Harrison (1989) in his discussion of the gobioid palatopterygoquadrate complex. He found that most of the gobionellines he examined had a long palatine reaching down nearly to the quadrate, with the exceptions being one or more species of Gobiopterus, Oxyurichthys, Pandaka, Pseudogobius, Rhinogobius and Tukugobius (Harrison, 1989: table 1). In his definition of the Gobionellinae, he included only those species with a derived palatinepterygoid-quadrate arrangement and a transverse sensory papillae pattern. The derived condition is found in Calamiana (most species), Chlamydogobius, Eugnathogobius, Hemigobius, Mugilogobius, Pseudogobiopsis (most species), Tamanka, Stigmatogobius and some species of Pseudogobius and Redigobius (survey incomplete for these latter two genera). Calamiana kabilia, C. variegata, Brachygobius and Pandaka have the plesiomorphic condition. The derived state of this character may be linked to the width and shape of the pterygoid (Figure 16). Character 31 Fifth ceratobranchial stoutly built, teeth conical, moderate to stout, bone may be pierced by foramina but not excessively; ventral ridge and flange extend close to outer margin. (1) Fifth ceratobranchial finely built, offragile appearance, so perforated that it may resemble lacework; (0) Revision of Mugilogobius teeth long, very fine and slender, tips may be slightly bent; ventral ridge and flange nm close to centre line of bone or close to inner margin. The outgroups and other gobionellines such as Oligolepis, Oxyurichthys, Stenogobius and Awaous have stout, almost equilaterally triangular fifth ceratobranchials with clearly defined low ridges (character state 0; Figure 21). All the Mugilogobiusgroup genera surveyed (other than Hemigobius and Pseudogobius) have the plesiomorphic condition. The ridge and flange on the ventral surface vary in shape and height (e.g. the flange may be low and rectangular or high and triangular), and there may be species-group similarities in development of these features which could be further examined. Murdy (1989) remarked upon the broad, open lattice-like fifth ceratobranchial found in the gobionelline Evorthodus and in eight out of the ten oxudercine genera he studied (Apocryptes, Apocryptodon, Boleophthalmus, Oxuderces, Parapocryptes, Pseudapocryptes, Scartelaos and Zappa). The ceratobranchials in Hemigobius and Pseudogobius are not as broad or leaf-like in appearance as in the above genera (Figure 21B, 0, G), but are very similar in the lattice structure of the bone (possibly due to similarities in habitat and feeding behaviour of these species). The bone in Hemigobius and Pseudogobius is triangular to slightly boomerangshaped. The fifth ceratobranchials in Gnatholepis (Figure 21C) are similar in shape to that of the plesiomorphic condition, but the bones interdigitate along the median line. Stenogobius psilosinionus shows a slight interdigitation at the rear very similar to that which is highly developed in Gnatholepis. Character 32 (0) Anterior end of preopercular bone rounded or pointed, may be slender. (V Anterior end of preopercular bone blun( squared-off often broadened. Most gobioids have the anterior end of the preoperculum rounded or somewhat pointed. In Eugnathogobius microps, two species of Calamiana, two species of Pseudogobiopsis and two species of Redigobius (survey of Redigobius species incomplete), the anterior end of this bone is blunt or squared-off in males at least. This feature may be linked to the development of large cheek muscles and ritual gaping behaviour in males. The genus Calamiana includes species with some of the largest gapes in the subfamily. Large mature males have the metapterygoid, quadrate and symplectic short and broadened, forming a wide flat support for the muscular cheeks. Watson and Horsthemke (1995) observe that live Calamiana aliceae (= C. kabilia), Schismatogobius and Gillichthys mirabilis have" ... maxillaries which are 29 free from preopercular cover and movable into a subvertical position .. .ff, while in Redigobius " ... the posterior end of the maxilla is covered by the preopercle allowing only moderate opening of mouth Further survey of this character and its link to the mechanics of agonistic behaviour will be carried out during revision of Redigobius (in prep.). ff • Character 33 (0) Anterior nostril near lip (not necessarily in tube) but not overhanging lip. (]) Anterior nostril in tube overhanging lip, preorbital may be curved outward around nostril. The anterior nostril at the tip of a distinct flap overhanging the upper lip is a synapomorphy for the Oxudercinae (Murdy, 1989). A similar structure occurs in amblyopines (survey incomplete). Most other gobioids have the anterior nostril in a tube (usually short) placed on the snout behind the lip (Figure 22A). Mugilogobius and the other gobionelline genera examined have the nostril tube placed at the edge of the preorbital, oriented forward and down, partly overlapping the upper lip; the preorbital is usually curved forward around the base of the nostril tube (Figure 22B). Exceptions to this are Pseudogobiopsis oligactis and several species of Redigobius (survey for this genus incomplete), which have the plesiomorphic condition (Figure 22C). There is no fleshy flap supporting the tube as in oxudercines and amblyopines, however, the curved preorbital margin is reminiscent of this. Character 34 (0) First epineural originates on base of neural arch. (1) First epineural originates on or behind tip of parapophysis. Johnson and Patterson (1993) and Patterson and Johnson (1995) indicated that patterns observed in arrangements of intramuscular bones can be useful in determining relationships among teleosts. Johnson and Patterson (1993) stated that Gnatholepis shared a pattern (first epineural originating on the parapophysis) with synbranchoids, mastacembeloids, gasterosteiforms, atherinimorphs, mugilomorphs, Elassoma and Echeneis. Although it is likely that this character is convergent in Gnatholepis and these non-gobioid groups, within the gobioids it may help define the Gobionellinae. All of the gobionelline species examined for this paper had the derived condition (Figure 11). Rlzyacichthys, Micropercops and Perccottus have the first epineural attaching anteriorly at the base of the neural spine, as do a number of gobiines, eleotridines and butines (Patterson and Johnson 1995; this paper). Several sicydiines, an oxudercine and a microdesmid had a similar condition to that of the gobionellines. Further survey of this character within the Gobioidei is needed. r H.K. Larson 30 Within the Gobionellinae, the derived state (Figures 4, 17) is present in Calamiana, Eugnathogobius, Pseudogobiopsis, Hemigobius, Pseudogobius, and a number of species of Redigobius (survey incomplete for Redigobius species). Some Brachygobius specimens are variable; modally, species exhibit the plesiomorphic state. A / B Character 36 No fine fleshy villi present on upper surface of head. (1) Upper surface ofhead, usually interorbital, dorsal surface of snout, upper part ofopercle, preopercle and sometimes preorbital, covered with very small fleshy villi. (0) \ ) This character has not been previously used in gobiid taxonomy, as far as I can determine. Miller (1987) was very likely referring to these villi when he described the snout of the holotype of Hemigobius bleekeri as "rugose and papillose"; he thought that the feature was an autapomorphy for the species. The villi are most easily observed in wellpreserved specimens with intact mucous coats. My previous work on gobiids indicates that this character may only be present in the Mugilogobiusgroup and several specialised gobiines e.g. Paragobiodon. Larger but similar structures occur in small scorpaenoids such as Caracanthus. Dr K. Cole (University of Southwestern Louisiana, Lafayette) sectioned the head of an adult Mugilogobius cavifrons to examine the microstructure of these villi. Along the outer margin of the thick epidermis were many large spherical cells: These cells are spherical, haematoxylin-staining (Le. basophilic) and contain what appear to be fibrillar and granular secretions. I assume these cells are unicellular glands, probably producing toxin(s) (as opposed to mucus). These large cells are almost continuous along the majority of the external epidermal surface but are not present in internal integument (Le. on the oral ectoderm which lines the mouth) (K. Cole, in litt.). c Figure 22 Nostril position and form in: A, Gnatholepis sp., NTM 5.12883-015; B, MllgilogobillS filifer n. sp., NTM 5.10694-013; C, Redigobills roemeri, CMK 11334. Scale bar = 1 mm. Character 35 (0) Papillae in cheek rows a, b, c, cp and d small, closeset and evenly-sized. (1) Papillae in cheek rows band d small, close-set, evenly-sized; those of rows a, c and cp relatively larger, widely spaced and fewer in number. Rhyacichthys and most odontobutids, eleotridines, butines, gobiines and most gobionellines have closely spaced, evenly sized small papillae in rows a, b, c, cp and d (Figures 2-3, 5; Akihito et al., 1988). Within the genus Mugilogobius, the villi are generally most dense in species from estuarine or mangrove habitats (the most typical habitat for the genus), where the fish may burrow among mangrove roots or be concealed in mud. Mugilogobius littoralis sp. nov., a species inhabiting exposed habitat (e.g. beachrock pools) has very few villi present. Work is continuing, with K. Cole, on these structures (outside the scope of this paper). Character 37 Posterior (broken) portion of row c consists of two or more small papillae. (1) Posterior (broken) portion of row c is a single papilla, modally larger than those of row b. In species of the Mugilogobius-group with (0) longitudinal papillae, row c is discontinuous, with a Revision of Mugilogobius gap under the eye (e.g. Figures 4A, 5; Miller( 1987). Most species in the group have the plesiomorphic state (several papillae in the rear portion of row c). Brachygobius, Calamiana, Chlamydogobius, Eugnathogobius and Pseudogobiopsis have the derived condition. Two species of Chlamydogobius have no or one papilla in the posterior part of row c. This character is linked to the derived condition of character 35. Character 38 (0) Second dorsal and anal fin rays modally 1,8 (or more). (1) Second dorsal and anal fin rays modally 1,7. Fin ray counts are not easily polarised. As stated above (under Character 10) Rhyacichthys has equal numbers of dorsal and anal rays (usually 1,8), or one more dorsal than anal ray, with one specimen from lriomotejima, Japan, with 1,7 for both fins. Odontobutids usually have one or two more dorsal fin rays (1,7-12) than anal rays (1,5-10) (Iwata et al., 1985; this paper). Most gobionellines have equal numbers of dorsal and anal rays. The lower number of rays is considered derived. Character 39 Nape scales all similarly sized, mostly small to moderate, not entering interorbital space. (1) Anteriormost nape scale enlarged, in contrast to other scales, and often partly entering interorbital space. Most gobiids, including the outgroups, have the nape scales similar in size (all small or all large, with none particularly larger than the others). An enlarged scale directly behind the eyes is considered the derived condition. Mugilogobiusgroup species fall into two groups, one with state o and one with state 1. The "state A" group includes the nominal genera Mugilogobius, Tamanka and Vaimosa. Mugilogobius rambaiae sometimes has one to several anteriormost nape scales larger than the others, but no one single scale enlarged so as to fit close behind the eyes. These specimens were coded as possessing the derived condition. (0) Character 40 If marking present along mid-ventral line ofcaudal peduncle and anal fin base, it is a thin dusky line or broken line (not including any encircling body bars). (1). If marking present along mid-ventral line ofcaudal peduncle and anal fin base, it includes more than one distinct black spots or blotches, line often black. Many gobioids have an indistinct dusky line or series of indistinct markings along the mid-ventral edge of the caudal peduncle, as does Rhyacichthys (and Odontobutis; state not recorded for Perccottus). These markings may be variable or absent. In Pandaka, Pseudogobius, and most species of (0) 31 Redigobius and Eugnathogobius, the mid-ventral line is intensified, usually visible as a narrow black line, with a series of evenly spaced black blotches or spots (which are often also internal). These spots are most obvious in Pandaka (but are also useful characters when field-sorting very small specimens of Pseudogobius and Mugilogobius). The character was not used in the final analyses, as the initial runs showed that its inclusion further decreased the resolution between taxa. Similar internal blocks of dark pigment along the caudal peduncle and vertebral column have been noted in coral reef-dwelling gobiines such as Eviota (Lachner and Karnella, 1980) and in Bryaninops and Pleurosicya (Larson, 1985, 1990). These fishes are often transparent or translucent when live, as is Pandaka. Another possible colour character was also not used in the analyses, because its condition was not known for all taxa. In Chlamydogobius, Weberogobius and most Mugilogobius species examined, breeding males become uniformly dark in colour, often obscuring body markings, while the fins darken and any fin margin colours (e.g. blue, yellow) intensify. This darkening was especially pronounced in the land-locked freshwater taxa Chlamydogobius and Weberogobius. In Brachygobius, Calamiana, Eugnath-ogobius, Stigmatogobius, and most Pseudogobius species, the males usually intensify their colouring but do not become uniformly dark. Some species of Pseudogobius and Eugnathogobius are sexually dichromatic, i.e. the males have a different colour pattern. Further observations of fishes in breeding condition (as well as accompanying behaviour patterns) may eventually provide characters useful toward reconstructing phylogeny. The following additional characters were used in attempts to determine relationships of the Mugilogobius-group to other gobionelline genera (Table 3). Character 41 Ifheadpores present, infraorbital pore present. If headpores present, infraorbital pore absent. Perccottus and Micropercops have infraorbital pores (Perccottus may have one or two pores in this position). All of the nominal Mugilogobius-group genera possessing headpores for this study (Calamiana, Pseudogobiopsis, Hemigobius, Pseudogobius, Redigobius, Stigmatogobius) had an infraorbital pore. Murdy (1989) found that Apocryptodon, an oxudercine, possessed an infraorbital pore. He stated that the Gobionellus and Sicydium Group species of Birdsong et al. (1988) lacked pore E (infraorbital pore) but possessed pore F (supraorbital pore). Pezold (1993) found that Awaous, Gnatholepis, Gobionellus, Gobioides, (0) (1) 32 H.K. Larson Table 3 Character states for the seven gobionelline species used in analyses with members of the Mugilogobius-group. Species Characters 000000000111111111122222222223333333333444 123456789012345678901234567890123456789012 Awaous Oligolepis Oxyurichthys Gnatholepis 5tenogobius Evorthodus Rhinogobius 000121320111011002000100001001100100000010 000011310201011002000100001001100110000010 000011320201011002011100001100100100000010 000010320211111002001000011001100100000010 000111120111011002000100001001100100000010 000021320211011002010000111001100100000010 000011011111011002000000001000000100000001 Evorthodus, Oligolepis, Oxyurichthys, Stenogobius, Tukugobius, and a number of other gobionelline and sicydiine genera, lacked an infraorbital pore. He found that the gobionellines Tridentiger and Rhinogobius (in addition to those already mentioned) possessed an infraorbital pore (the absence of an infraorbital pore is correlated with the presence of anterior nasal pores, except in these two genera). Character 42 (0) Anterior nasal pore present. (1) Anterior nasal pore absent. The anterior nasal pore is absent in all genera (with headpores) in the Mugilogobius-group examined for this paper, although the posterior nasal pore is usually present. The anterior nasal pore is present in the outgroup genera and in many other gobionelline genera of the Stenogobius-group (Pezold, 1993: table 2). Characters not used The dorsal pterygiophore formula was not used in the analyses. Gobionellines characteristically have the formula 3-12210 (Birdsong et al., 1988; Pezold, 1993; this paper). The gobionelline Rhinogobius differs by often having the formula 322110 in some species (survey incomplete); as do most gobiines. Rhyacichthys has the formula of 3222100 or 3-221220 (Birdsong et al., 1988) and usually has seven first dorsal spines. Odontobutids have seven to nine first dorsal spines, with a variable dorsal pterygiophore formula, e.g. 4-221111 or 3-1212111 (Figure 6A; Birdsong et al., 1988: table 33; Hoese and Gill 1993). The butines show a correlation between a pterygiophore formula of 3-2211 and a transverse sensory papillae pattern, but no similar correlation can be found within the eleotridines (which generally have a pterygiophore formula of 3-1221) or the odontobutids (Birdsong et al., 1988; Hoese and Gill, 1993). As Hoese and Gill (1993) point out, assigning polarity to pterygiophore formulae can be difficult. Bianco and Miller (1990) determined that the "2211" pattern was plesiomorphic for gobioids and characteristic of gobiines. A number of colour-pattern characters were also considered for possible use, such as the presence of a dense black spot on the first dorsal fin, whether the first dorsal fin had a blue spot or a yellow to red spot, or if an oblique dark bar was present over the pectoral base. They were not used due to difficulties with colour variation, in assigning polarity, and the presence of some of the characters throughout the subfamily (e.g. a variably developed dense black spot on the first dorsal fin). Most species of Mugilogobius and its relatives can be distinguished by a combination of scale counts and colour patterns. These characters are not easily quantified and polarised, and were not used in the cladistic analyses. This group of fishes appears to be characterised by an exhibition of maximum homoplasy so as to prevent a clear understanding of their relationships. It is expected that taxonomic decisions made here may change after further work on species in this group reveals additional synapomorphic characters. Results of phylogenetic analyses Final analyses were made using data from the matrices in Tables 2 and 3 (and see Methods). Initial attempts were made to see if any pattern was discernible within the Mugilogobius group, with an analysis of 65 taxa (the entire matrix under study) plus Rhyacichthys, Micropercops and Perccottus as outgroups. The strict consensus tree shown here (Figure 23, consensus of 700 trees, length In, consistency' index 0.28) indicates several monophyletic groups, but very little information as to the relationships of Mugilogobius. Mugilogobius, Tamanka and Weberogobius species were unresolved, and were grouped in a large clade which included species of Brachygobius, Chlamydogobius, Pandaka and Stigmatogobius. Calamiana, Eugnathogobius, Hemigobius, Pseudogobius and Redigobius species were placed outside this large clade. The relationships of many of these taxa was considered to be obscured by homoplasy and in some cases, uncertainty due to missing data. Therefore, analyses were made using fewer species. Revision of Mugilogobius 33 rl ｾ r-- .r-e= - -----re= I r-- Figure 23 Rhyacichthys Mlcroperoops Parocottus Ch. eretTius Ch.japslps Ch. ranunculus Ch. microptarus Ch. squamlgenus B. kabiliansls B. doriae B. xanthomelas Hy. xan/hozona Pa.pygmaea T. sitensls T. saraslnorum M. cagayanensls T. rexi n. sp. T. lap/do/us n. sp. M. adeia M. fusca M.mertonl M. wUsoni n. sp. M. chulae M. fascia/us n. sp. M.filifern.sp. M. cavifrons M.latifrons M. n%spUus M.rambaiae M. abe! M. rivulus n. sp. M.fusculus M. platyno/us M. stigmaticus M. platys/oma M. ittoraHs n. sp. M. myxodermus W.amadi S. pleurostigma S. bomaensls S. sadanundio Ca. n.sp.1 Ca. variagatus Ca. n.sp. 2 Ca. n. sp.3 Ca. mindora Ca. kabllla E. microps E. slamensls E. psludosus E.n.sp. He. hoavenU Ha. min{j Ps.sp.8 Ps. avicannia Ps. poicilosoma Ps. melanostic/a Ps. javanicus Ps. sp. 9 PS.olorum E. oOgactis R.penango R. crlSpar R. chrysosoma R. ballea/us R. bikolanus R. macros/oma R.roemari Result of PAUP analysis of all taxa under study; strict consensus of 700 equally parsimonious trees. In an attempt to determine what the sister-group to Mugilogobius might be, 17 taxa comprising the type species of a range of nominal genera considered to be related to or possibly synonymous with Mugilogobius (Brachygobius, Calamiana, Chlamydogobius, Cyprinogobius Koumans, Eugnathogobius, Hemigobius, Hypogymnogobius, Mugilogobius, Ostreogobius Whitley, Pandaka, Pseudogobiopsis, Pseudogobius, Redigobius, Stigmatogobius, Tamanka, Vaimosa and Weberogobius) were analysed by PAUP. This resulted in 24 equally parsimonious trees and the consensus tree shown in Figure 24 (length 108, consistency index 0.42). The tree places Mugilogobius and Vaimosa together, but the positions of Tamanka and Weberogobius are not resolved. These latter two nominal genera resemble Mugilogobius phenotypically more than any of the other genera in this unresolved polytomy. The tree also places H.K. Larson 34 Rhyecichthys MicropercoPS Perccottus Ch. erem{us B. doriee Hy. xenlhozona PB.pygmaee He. mingi Ps. jevenicus T. siitensis V. fontinalis M.ebei the type species of each nominal genus and ensuring that a range of character states was represented among the species chosen). The first analysis was made using 34 taxa, including two Redigabius species as outgroups. The number of Mugilagabius species was reduced to thirteen, and Weberagabius amadi (endemic to Lake Poso, Sulawesi) and two Tamanka species (one from Sulawesi and one from the Philippines) were included. This analysis found 63 equally parsimonious trees, and the consensus tree shown in Figure 25 (length 102, consistency index 0.39). The resolution of relationships is not much better than that shown in Figure 24. Four Mugilagabius species form a monophyletic group (chulae, mertani, rambaiae and wilsani sp. nov.), as do two species from Sulawesi (adeia and latifrans). Brachygabius and Pandaka form a clade. Hemigabius and Pseudagabius form a clade. Chlamydagabius forms its own clade, as does Stigmatagabius. Weberagabius amadi and the two W.emedi S. sedanundio Ch. eremlus Ca. kabilia B. doriae Ch. ranunoulus Pa.pygmaea E. microps He. hoeven/i He. m/ngi P. oligactis Ps. poie/losoma Ps. javanious R. dispar T. siitensis O.macrostome T. sarasinorum M. cagayanensis Cr. chrysosome Figure 24 M. adeia M. latifrons Result of PADP analysis of the type species of a range of nominal genera considered to be possible close relatives of Mugilogobius; strict consensus of 24 equally parsimonious trees. M. meltoni M. wilsonl n. sp. M. mulae M. remba/ae M. cavifrons Hemigabius and Pseudagabius together, and within the "Mugilagabius" polytomy. These two analyses (Figures 23-24), and initial Hennig86 analyses of small groups of taxa, with Rhyacichthys aspra, Micrapercaps barealis and Perccattus chalmersi as outgroups (or Rhyacichthys alone), indicated that Redigabius species were the most plesiomorphic among the in-group. Redigabius (and Pseudagabius) were considered as "putatively primitive" by Hoese and Gill (1993). Accordingly, Redigabius dispar (type species of the genus, from the Philippines) and Redigabius macrastamus (a temperate Australian species) were used as outgroups for further analyses within the Mugilagabius group. As not all species within each nominal genus always shared the same character states, one to three species of each genus related to Mugilagabius were selected for further analyses (always including M. nolospilus M. aOOI M. tusculus M. stigmatieus M. platystoma W. amad/ S. pleurosligma S. sadanundio Ca. kabilia Ca. mindora Ca. n.sp.1 E. siamensis E. microps E.oiigeclls R. macrostoma R. dispar Figure 25 Result of PADP analysis of reduced set of taxa (type species of each genus included); strict consensus of 63 equally parsimonious trees. Revision of Mugilogobius 35 Ch. erem/us Ch. ranunculus T. saraslnorum M. BeJe/a M. latlfrons M. cagayanensls M. notospilus with some non-resolution occurring at the base of the tree. The Hemigobius-Pseudogobius clade remains separate, as does the Calamiana clade. Eugnathogobius microps, E. siamensis and E. paludosus are unresolved, and E. oligactis remains closer to Redigobius. M.mertonl M. wilsonl n. sp. M. chulae M. rambaiae M. cavlfrons M. abel M. fusculus M. stlgmat/eus M.p/alystoma T. slltensls s. pleurostigma S. sadanundlo B. doriae Pa. pygmaea H. hoevenil H. mingl Ps. poleilosoma Ps. Javanleus Ca. kabilla Discussion of systematic relationships The phylogenetic analyses described above provide !;he basis for assigning taxonomic status to the various clades. From among the 12 trees comprising the consensus tree shown in Figure 27, the preferred tree (Figure 28) was chosen as a basis for deriving a classification of the Mugilogobius-group. This tree was chosen because it is one of six trees which differ from the other six only in the relationships between the Calamiana clade (four species) and the four species of Eugnathogobius (that is, placing Eugnathogobius between Calamiana and Redigobius; the other six trees placed one or more species of Eugnathogobius closer to the Hemigobius/ Pseudogobius clade). The trees in Figures 26-27 and 28 all group Mugilogobius as a single clade, but include one nominal Tamanka (T. sarasinorum) and Ca. n. sp.1 Ca. mindora Ch. eremius Ch.japalpa .---_ _ T. saraslnorum E. slamensls E. microps E. oligactis 1- M. &dela M. latifrons M. fusculus 1..- M. plalystoma R. dlspar R. macrostoma Figure 26 Result of PADP analysis of reduced set of taxa, but Mugilogobius amadi excluded; strict consensus of five equally parsimonious trees. M. cagayanensls M. notosp/lus M. chulae r--t-- M. merloni M. wllsonl n. sp. Tamanka form part of an unresolved polytomy with seven other Mugilogobius. All present data indicates that W. amadi may be a derived species exhibiting several reversals (and distinctive physiognomy). Its closest relative would appear to be T. sarasinorum, which also inhabits Lake Poso. As Kluge has suggested " ... where the members of a target taxon show a high degree of modification ... the least derived member should be used in the analysis" (Kluge, in Gill 1994). Therefore, in the next analysis, W. amadi was deleted. This resulted in five trees and the consensus tree in Figure 26 (length 96, consistency index 0.41). Mugilogobius and Tamanka sarasinorum (but not T. siitensis) form a clade, with Chlamydogobius as sister-group. The addition of Calamiana sp. novo 2 and Eugnathogobius paludosus (done because the generic assignment of these two was somewhat uncertain) resulted in 12 equally parsimonious trees and the consensus tree in Figure 27 (length 106, consistency index 0.54). The Chlamydogobius, Mugilogobius and Tamanka species relationships remained unchanged, 1-- '--_ _ M. rambalea M. abel 1- M. cavifrons M. stigmaticus 1..-- T. sIltensls 1..-- S. pleurostlgma S. sadanundio B. dOliae P.pygmaea r--- Ca. kabYla Ca. m/ndora '---+_ Ca. n. sp. 1 CB. n. sp. 2 1-------------1- E. mlcropa E.paludosus 1- E. slamensls H. /KlBverllj H. mlng/ Ps. Javanlcus 1- Ps. poicilosoma E.oligaclis R.d/spar 1- R. macrostoma 1-- Figure 27 Result of PAUP analysis of reduced set of taxa, with Calamiana n. sp. 2 and Eugnathogobius paludosus added; strict consensus of 12 equally parsimonious trees. 36 H.K. Larson exclude one nominal Tamanka (the genotype, T. siitensis). The status of Tamanka is discussed later. It is recognised that the analyses have not resulted in very robust trees (low consistency indexes and a tendency to change configuration depending on the species included). The non-resolution shown in Figures 23 and 25 is partly caused by difficulty in placing the species from freshwater lakes in Sulawesi and the Philippines, and in particular, the scarcity of non-homoplasous characters. However, given the present information, it is still possible to present a hypothesis of Mugilogobius relationships, based on the tree in Figure 28. In the descriptions that follow, the clades comprising this tree and their synapomorphies at each node are discussed, and species assigned to genera according to their position. Node 1. Relationships at the first node are supported by the derived condition of the pre-anal pterygiophores (character 12), metapterygoid shape 20 ...-----=-1 Ch. eremius Ch.japalpa ...-_ _ T. ssrasinorum M. adeia 24 19 M. fatifrons 1- M. fusculus L..- M. platystoma 23 M. cagayanensis M. notospilus M. chutaa 21 ...--1-- M. merlonl 18 22 M. wllsonl n. sp. L..-_ _ 116 M. rambaiae M.sbei 1- M. cavffrons L..- M. siigmaticus T. siitensis L..- 14 17 s. pleurosiigma S. sadanundio 15 10 B. doriae P. pygmaea H. hoevenii H. mingi 11 Ps. javanicus 3 Ps. poie/losoma , - - - - - Ca. kabilia Ca. mindora Ca. n.sp.l 6 2 L -_ _ L- 4 L..L..- L..- ｾＭＭ L.- Figure 28 Ca. n. sp. 2 E. paludosus E. siamensis E. microps E.oligactis R. dispar R.macrostoma One of 12 equally parsimonious trees comprising the consensus shown in Figure 27; preferred tree on which to base the classification used in this paper. (character 13), scapula ossification (character 21) and palatine length (character 30). Redigobius species possess the plesiomorphic conditions for these characters, while a number of Redigobius species, such as R. macrostomus, have the derived condition for mouth position (character 11). Node 2. Species at this node share the derived condition of headpore pattern (character 1), the preopercular canal (character 73), preopercular pores (character 28) and the anterior nostril (character 33). Here also is a reversal for the pectoral girdle edge (character 4). Most Redigobius have the derived condition for this character. Eugnathogobius oligactis has the derived condition of the snout s papillae rows (character 9) and the plesiomorphic condition of the pectoral girdle edge (character 4). Node 3. This node is supported by the derived condition for papilla row e (character 19). Node 4. Taxa here share the derived state for pectoral girdle edge (character 4). Eugnathogobius microps has the derived condition for the preopercle (character 32). Node 5. Taxa at this node share the derived condition for the metapterygoid bridge (character 29). Eugnathogobius siamensis shows a partial reversal in headpore pattern (character 1). Node 6. Species above this node share the derived condition of the segmented caudal rays (character 3). Eugnathogobius paludosus is characterised by the derived state for epurals (character 5), and reversals in procurrent cartilage (partial; character 18), papilla row e (character 19), bony preopercular canal (character 23) and fin ray number (character 38). Node 7. Species above this node are considered to be Calamiana. Relationships at this node are supported by the derived condition of the snout s papillae rows (character 9) and a reversal of the preoperculum tip (character 32). Calamiana kabilia has the derived condition of the neural spines (character 6) and a reversal of the palatine length (character 30). Node 8. These species share the derived condition of nape scales (character 39). Calamiana sp. novo 2 shows reversals in mouth size (character 17) and fin ray number (character 38). Node 9. Calamiana mindora and C. illota (sp. novo 1 in Figures 26-28) show reversals in headpores (character 1), metapterygoid shape (character 13) and mouth size (character 17). Node 10. There is a reversal at this node, to the plesiomorphic condition of preoperculuffi tip (character 32). Node 11. Relationships are supported by the derived condition of the mouth position (character 11), lip morphology (character 25), snout shape (character 26), fifth ceratobranchial (character 31) and a reversal from the most derived state of headpore pattern (character 1). Revision of Mugilogobius Node 12. Hemigobius is supported by the derived condition of gut morphology (character 7) and villi (character 36). Additionally, H. mingi has the derived form of the neural spines (character 6) and shows reversals in the pre-anal pterygiophores (character 12) and in mouth size (character 17). Node 13. Pseudogobius is supported by the derived conditions for segmented caudal rays (character 3), pectoral girdle edge (character 4) and gut morphology (character 7). Species at this node show a reversal in metapterygoid shape (character 13) (and some vary in snout 5 papillae rows (character 9) and in preopercular canal (character 23». Pseudogobius javanicus shows a reversal in mouth size (character 17) and P. poicilosomus shows reversals for scapula ossification (character 21) and palatine length (character 30). Node 14. At this node are two reversals, for mouth size (character 17) and cheek papillae (character 35). Node 15. Taxa here share the derived conditions of fin ray segmentation (character 2) and vertebral number (character 14), and reversals in metapterygoid shape (character 13) and palatine length (character 30). Brachygobius has the derived conditions for epural number (character 5) and villi (character 36), and shows reversals in pre-anal pterygiophores (character 12) and papilla row p (character 15). Pandaka exhibits reversals in nostril position (character 33) and papilla row c (character 37). Node 16. This node is characterised by the derived condition of the metapterygoid bridge (character 29) and reversals in papilla row c (character 37) and fin ray number (character 38). Node 17. Stigmatogobius is supported by the derived condition for papillae pattern (character 8), dorsal and anal ray ratio (character 10), haemal spines (character 24, the derived state may be an autapomorphy for the genus); nape scales (character 39) and papilla row p (character 15), a reversal in scapula ossification (character 21) and a partial reversal in headpore pattern (character 1). Stigmatogobius pleurostigma has a reversal in mouth size (character 17). Node 18. Relationships here are supported by the derived conditions of the neural spines (character 6) and villi (character 36), and a reversal in papilla row p (character 15). Tamanka siitensis is characterised by having the derived condition for papilla row a (character 16). Node 19. The derived condition for condition for caudal ray segmentation (character 3) and mouth position (character 11) is shared by species above this node. Node 20. Species of the genus Chlamydogobius have the derived condition for epural (character 5), gut morphology (character 7), snout 5 papillae rows (character 9), snout shape (character 26) and fin ray number (character 38). It has reversals for dorsal 37 and anal rays (character 10), vertebrae (character 14), procurrent cartilage (character 18) and metapterygoid bridge (character 30).' Node 21. Species here share the derived conditions for pectoral girdle edge (character 4) and mouth size (character 17). The grouping includes all of the nominal Mugilogobius and Tamanka sarasinorum, and species above this node are considered to be Mugilogobius. The abeigroup (abei is the type species of Mugilogobius) all have modal second dorsal and anal fin ray counts of 1,8 (plesiomorphic condition of character 38) and small, evenly sized predorsal scales (plesiomorphic condition of character 39). All species, other than those above node 22, share these characters. Mugilogobius stigmaticus shows a reversal for neural spine shape (character 6). Node 22. The Mugilogobius species at this node have the derived condition for fin ray number (character 38) and nape scales (character 39). They have an enlarged scale behind the eyes, and modal second dorsal and anal fin ray counts of 1,7, and are hereafter referred to as the chulae-group; M. chulae being a typical and widely distributed species. Mugilogobius rambaiae appears at the base of this clade because the condition of its nape scales usually resembles the derived state and has been scored as such (see species description). Mugilogobius mertoni has a reversal for scapula ossification (character 21), while M. wilsoni sp. novo has a relatively low metapterygoid (character 13; reversal). Node 23. The two species here both share the derived condition for papilla row a (character 16). Mugilogobius cagayanensis also shows a reversal in mouth position (character 11). Node 24. Here is a reversal for metapterygoid shape (character 13). Mugilogobius fusculus and M. platystomus both have relatively low metapterygoids but do not possess other character states agreeing with the species at node 25. Node 25. The species here show reversals in anal pterygiophore number (character 12) and mouth size (character 17). Tamanka sarasinorum often has a terminal mouth (character 11, a reversal), while M. adeia and M. latifrons, which have subterminal mouths, both show a reversal in neural spine shape (character 6). A character which was not polarised, but which supports the clade, is vertebral pattern. Species of this clade have 11+15-16 vertebrae (other Mugilogobius species have 10+16 vertebrae). Mugilogobius and its sister-group As indicated in the Introduction, the main aims of this work were to diagnose the genus Mugilogobius, revise the species assigned to it, and, using phylogenetic systematic techniques, try to 38 determine its closest relatives, determine whether Mugilogobius and its relatives are a monophyletic group, and discover what might be their closest relatives. The clade at node 21 (Figure 28) includes 13 nominal Mugilogobius species and Tamanka sarasinorum. Tamanka sarasinorum and Mugilogobius are here considered to be congeneric; as they share the derived states for characters 1 (headpores absent), 3 (caudal ray segmentation), 4 (pectoral girdle lobes), 6 (neural spines), 17 (mouth size), 31 (metapterygoid bridge) and 38 (villi on head). Tamanka sarasinorum differs from most other Mugilogobius in having the plesiomorphic state for character 11 (mouth position). It shares with some Mugilogobius (M. fusculus, M. platystomus) the plesiomorphic state for character 13 (metapterygoid shape). Ten species of Mugilogobius were not included in the final analyses (other than that which produced Figure 23). Seven of these 10 species share the characters of node 21, but were not included in the final analyses because data was either missing or redundant. Three problematic species (W. amadi, T. lepidotus sp. novo and T. rexi sp. nov.) are related to the node 25 clade from Sulawesi, and are further discussed below. Therefore Mugilogobius (node 21) can be defined by having the derived condition for headpores (character 1, pores absent), segmented caudal ray number (character 3, 16 rays), pectoral girdle (character 4, lobes present), neural spines on first few vertebrae (character 6, spines expanded), mouth position (character 11, modally subterminal), metapterygoid shape (character 13, broad, expanded), mouth size (character 17, enlarged in males), metapterygoid bridge to quadrate (character 31, present), and villi on head (character 38, present). As defined here, the species of the genus Mugilogobius share nine synapomorphies, but no autapomorphy has been found for the genus. Most species of Mugilogobius fall into one of two groups (the abei and chulae groups) of which the abei-group is the most speciose. Additionally, there is a third group formed by the freshwater lake endemics of Sulawesi and the Sulu Archipelago (seven species). Within Mugilogobius, a grouping of taxa includes the nominal monotypic genus Weberogobius (amadi), a nominal Tamanka (sarasinorum), two new species resembling Tamanka (lepidotus sp. nov., rexi sp. nov.) and two nominal Mugilogobius (adeia, latifrons). Three taxa from this group are identified at node 25 in Figure 28 (adeia, latifrons, sarasinorum). Three other taxa (amadi, rexi sp. nov., lepidotus sp. nov.) are not shown at node 25, but they share the following characters with that clade: reversals in pre-anal pterygiophore number (character 12, three or more), metapterygoid shape (character 13, low H.K. Larson and narrow), and mouth size (character 18, not enlarged in males. In preliminary analyses (not shown), amadi grouped with sarasinorum or with rexi sp.nov. Species of this group of six taxa agree with Mugilogobius in that they have the derived condition for headpores (character 1), segmented caudal rays (character 3), metapterygoid bridge to quadrate (character 31), and modally have the derived condition for the pectoral girdle (character 4). They are here included in the genus Mugilogobius. All six species are central Sulawesi freshwater lake endemics. These species modally possess terminal mouths, have rather pointed snouts, 11+15-16 vertebral pattern, two species have a smooth pectoral girdle, while one species has slender neural spines and lacks a bridge to the metapterygoid. All six Sulawesi endemic Mugilogobius have the plesiomorphic condition of character 12 (three or more anal pterygiophores before the first haemal arch), unlike other Mugilogobius, which modally have two (in M. myxodermus and M. rambaiae, about half the specimens examined had three). Of this group, only Mugilogobius adeia and M. latifrons had previously been assigned to Mugilogobius (Larson and Kottelat, 1992; Kottelat et al., 1993). Apart from the 11+15-16 vertebral pattern and narrow metapterygoid, these two species are typical Mugilogobius, possessing subterminal mouths (character 1), fleshy lobes on the pectoral girdle (character 4), and fine villi on the head (character 38). Mugilogobius sarasinorum had previously been assigned to Tamanka (Larson and Kottelat, 1992; Kottelat et al., 1993), but is here placed in Mugilogobius based on shared characters. Mugilogobius amadi was retained by Miller (1987) as a separate genus Weberogobius; he recognised that it appeared to share common ancestry with Chlamydogobius and Mugilogobius. He considered that M. amadi differed from these two genera by having apomorphies of " ... much more numerous papillae, greatly elongate rear part of row c, and short irregular tracts of papillae between rows d and cp, probably derived from the latter" (Miller, 1987). These characters show variation among the specimens examined for this paper. Firstly, the number of papillae may be proportional to the length and size of the fish's cheek (compare Figures 90, 215, M. cagayanensis and T. siitensis, both large long-headed species, with Figures 95, 103, M. cavifrons and M. chulae, species of moderate size and head shape). Among this group of fishes Mugilogobius amadi has the most slender, long-jawed head and reaches the greatest size. Secondly, the length of the rear part of row c is variable within M. amadi, with Miller's Figure 7 showing the greatest extent (Figure 5, this paper, shows a much shorter example). Thirdly, the proliferation of papillae between rows cp and d (Figure 5; Miller,.1987: figure Revision of Mugilogobius 7) closely resembles that seen in M. sarasinorum which is syntopic with M. amadi in Lake Poso. The rearward-pointing row a in M. amadi (Figure 5; Miller, 1987: figure 7) can be seen in T. siitensis, M. cagayanensis and in several other species of Mugilogobius and its shape and orientation varies somewhat. As Miller (1987) pointed out, this feature resembles that seen in eleotridids sensu Wongrat (1977; and see Figure 7, Perccottus chalmersi) and some species of Glossogobius. Mugilogobius amadi differs from other species in having a high 'second dorsal and anal ray count, and a combination of characters: smooth pectoral girdle, terminal mouth, three or more pre-anal pterygiophores and no villi on the head. It is also quite different in appearance to the whole Mugilogobius-group, being large, with a relatively long, narrow head and "predatory" jaws. Despite these differences, it is placed in Mugilogobius, as it shares with that genus the characters indicated above. Mugilogobius rexi sp. novo is one of the smallest of the Sulawesi endemics. It has some transverse rows of papillae under the eye, villi are absent and the first few neural spines are always slender and pointed (M. amadi has the tips expanded). Mugilogobius amadi and M. rexi sp. novo both have the plesiomorphic condition of character 4 (smooth pectoral girdle). Mugilogobius lepidotus sp. nov., also a small Sulawesi species, differs from all the others in the Mugilogobius,;group by two autapomorphies: papillose pads in place of gill rakers and unique striped colouring. It has a somewhat protrusible mouth which tends to point downward, a low metapterygoid without a bridge to the quadrate (which has a low, poorly ossified upper limb), narrow, pointed neural spines, a terminal mouth, and males and females have jaws of similar length. The condition of the last three of these characters are considered to be reversals. Mugilogobius lepidotus sp. novo usually lacks, as does M. rexi sp. nov., the fine villi on the head characteristic of other Mugilogobius (these villi are variably present in M. adeia, M. latifrons and M. sarasinorum but absent in M. amadi). Its closest relative may be M. rexi sp. nov., a diminutive species which also inhabits Lake Towuti. The Sulawesi endemic species of Mugilogobius all differ in size, morphology, distribution, preferred habitat and behaviour. For example, the large, elongate M. amadi (endemic to Lake Poso) inhabits very deep water, probably hovering in groups or schools, the dwarf M. rexi sp. novo (restricted to Lakes Mahalona and Towuti) and M. lepidotus sp. novo (endemic to Lake Towuti) hover in the water column, near shoreline vegetation. Mugilogobius sarasinorum (endemic to Lake Poso) is mediumsized, benthic and hides among rocks and sandy 39 areas and is most active at night. Mugilogobius adeia (endemic to Lake Matano) and M. latifrons (from Lakes Mahalona, Matano and Towuti and their tributaries) are generally small, benthic, and behave similarly to estuarine Mugilogobius, living among stones, leaf litter and logs in the shallows. A characteristic of the freshwater lake endemic species-group is that they have developed different colour patterns to other Mugilogobius. The Sulawesi lake endemics are generally plain, not conspicuously spotted or banded (although there is an exception to prove the rule: M. adeia) and do not have a distinct black spot on the first dorsal fin. These six species show a range of generally subdued colour patterns: plain dark brown to black (amadi); plain speckled brown (latifrons); yellow to grey (rexi sp. nov.); plain grey with horizontal stripes (lepidotus sp. nov.); plain brown to yellow with narrow vertical bars (sarasinorum); and one species with distinct black and white bands and spots (adeia). The Sulu Archipelago endemic Mugilogobius cagayanensis is also relatively plain: dark brown, with any bars or blotches obscured, with an intensification of the light submarginal band on the unpaired fins (colouring similar to Tamanka siitensis). It is possible that the relatively plain and inconspicuous colour of these endemics is a response to the exposed and precarious habitats available to gobies in the lakes; the fish adopting colouring which helps to camouflage them from potential predators. Mugilogobius sarasinorum was observed to be most active at night, over exposed areas in which they were not seen during the day. This behaviour may be a response to the small amount of available cover and a way of avoiding predators, such as egrets. The Sulawesi lakes are, by their limited shelter, reminiscent of the waterbodies in which are found the five inland species of Mugilogobius's sister genus, Chlamydogobius. Species of this genus are relatively plain brown, marbled and barred in inconspicuous patterns, probably so that the fish can be camouflaged in their harsh, exposed habitats. Most estuarine and stream-dwelling Mugilogobius species have distinct bars, spots and blotches in a variety of combinations and a dense black spot on the first dorsal fin. Many of these species inhabit mangroves, a habitat which provides refuges in the form of soft mud burrows, logs, leaf litter and, periodically, turbid water. Two species of Mugilogobius (M. fasciatus sp. novo and M. tigrinus sp. nov.) are known which are relatively small, strongly banded, and restricted to peninsular Thailand, Malaysia and Singapore, localities in which the conspicuously marked Brachygobius and Pandaka also occur. In mangroves in Singapore, all four genera can be found syntopically: the two banded Mugilogobius species, Brachygobius kabiliensis 40 and Pandaka pygmaea. It is possible that in more complex and sheltered habitats, more conspicuous markings are adopted. The sister-group to Mugilogobius is Chlamydogobius, which is supported by four synapomorphies. Chlamydogobius had been suggested as a sister-group to Mugilogobius (Miller, 1987; Larson, 1995). Here this genus is shown to be separated from Mugilogobius at node 19, both genera sharing the possession of 16 segmented caudal rays and a subterminal mouth. Chlamydogobius has the derived condition of characters 5 (one epural), 7 (gut long and coiled), 9 (two s papillae rows on snout) and 39 (rear portion of papilla row c). It has the plesiomorphic condition for characters 4 (smooth pectoral girdle), 14 (28-29 or more vertebrae) and 31 (no bridge to quadrate). Chlamydogobius ranunculus is, in some ways, the least derived of the genus, having 10+16 vertebrae in five out of 30 specimens X-rayed, and by its inhabiting coastal estuarine areas (the other five species inhabit very isolated desert springs near the centre of the Australian continent). It should be noted that from here onward, in the text and figure legends, species are referred to genera according to the classification adopted in this paper (e.g. Weberogobius amadi is now called Mugilogobius amadi). Genera related to Mugilogobius In the following discussion, genera are described in order according to the proximity of their relationship to Mugilogobius and its immediate sister-group, Chlamydogobius. The closest relative to these is the apparently monotypic Tamanka. Tamanka siitensis has 17 segmented caudal fin rays (16 segmented rays in all other closely related species), a smooth oblique pectoral girdle and is the type species of the genus Tamanka. It consistently remained separate from other nominal Tamanka and Mugilogobius in the analyses (e.g. Figures 24, 26-27). In keeping with cladistic methodology, Tamanka siitensis is retained as a monotypic genus, despite its sharing many other characters with Mugilogobius. The only character which consistently separates it from Mugilogobius is its possession of 17 segmented caudal fin rays (the plesiomorphic condition for the whole Mugilogobius clade), with its single derived character (rearward slope of papilla row a) being shared with M. amadi and some specimens of other species of Mugilogobius (M. cagayanensis, M. chulae, M. fusculus, M. myxodermus, M. notospilus and M. rivulus sp. nov.). Tamanka siitensis exhibits a mixture of derived and plesiomorphic characters, differing from Mugilogobius by having the plesiomorphic condition for characters 3 (caudal ray segmentation), 4 (pectoral girdle), 11 (mouth H.K. Larson position) and 17 (mouth size). However, it agrees with Mugilogobius in having the derived condition for characters 1 (headpores), 6 (neural spines), 13 (metapterygoid shape), 31 (metapterygoid bJ;'idge) and 38 (villi). It shares possession of a terminal mouth and derived condition of papilla row a (character 16) with M. cagayanensis. Mugilogobius cagayanensis and T. siitensis are very similar in appearance. Mugilogobius cagayanensis has 16 segmented caudal fin rays (typical of Mugilogobius). Both species have second dorsal and anal ray counts and predorsal scales typical of the abei-group, have a 10+16 vertebral pattern, and are endemic to lakes in small, isolated islands in the Sulu Archipelago, Philippines. Unlike the Sulawesi lake-dwelling species, these two have broad metapterygoids (as do most Mugilogobius). Tamanka siitensis could be a plesiomorphic, old species resembling the ancestral population of the Mugilogobius clade that has not developed any autapomorphies. However, it must be borne in mind that the only other Philippine land-locked species of the "node 21" group, M. cagayanensis, shows variation in the number of segmented caudal rays, with three out of 33 specimens examined having 17 segmented caudal rays, one with 18 and two with 15. Thus the similarity of M. cagayanensis and T. siitensis may be due to convergence. Stigmatogobius is characterised by the derived characters 8 (transverse sensory papillae), 16 (papilla row p in short transverse rows), 25 (haemal arch shape) and 39 (single enlarged scale behind the eyes). The anal fin also has at least one more ray than the second dorsal fin (character 10). Three species are also all very strongly marked with conspicuous dark spots or bands on the body. In initial analyses with characters ordered, Stigmatogobius was usually placed further away from the Mugilogobius clade, closer to the Hemigobius/pseudogobius clade. Brachygobius and Pandaka (Figures 24, 26) both share the derived condition for characters 2 (all second dorsal and anal rays segmented) and 14 (2425 vertebrae). Pandaka sometimes has 16 segmented caudal rays, and has a smooth pectoral girdle; Brachygobius may have the derived condition for the pectoral girdle. Brachygobius species always have one epural while Pandaka has one or two. Both genera have 10+15 vertebrae and slender, reduced bones in the palatopterygoid complex, and a reduced metapterygoid. The two genera are here retained as separate, pending further work. It should be noted that the paedomorphic genus Gobiopterus, not dealt with in this paper, shares at least three characters (unsegmented spines in second dorsal and anal rays, 10+15 vertebrae, one or two epurals) with these two genera. The relationship of Mugilogobius to the Brachygobius-Pandaka clade has been discussed to Revision of Mugilogobius some extent by Miller (1987, 1989) and McKay (1993). Miller observed that Brachygobius, Pandaka, Mugilogobius, Chlamydogobius, Weberogobius, Pseudogobius, Redigobius and Hemigobius shared"... a suite of apomorphies which indicate phyletic affinities ..." (Miller, 1987). He illustrated sensory papillae and osteological features, and stated that: All the gobiids with uniserial ('longitudinal') suborbital rows and the 12210 dorsal pterygiophore formula, i.e. ChlamydogobiuslMugilogobius-Pandaka, and the North Pacific genera, are believed by the author to share common ancestry with 22110 longitudinal forms, such as Glossogobius, Favonigobius Whitley and many other genera, all of which have a single anterior interorbital pore y when head canals are present (Akihito et al., 1984). (Miller, 1987). The results of this paper are in general agreement with Miller's (1987) observations. Miller (1989) later indicated that he considered Mugilogobius and Pseudogobius to be more"generalised" in relation to Brachygobius because Brachygobius had a narrow metapterygoid, a single epural and had lost one vertebra. As has been shown, the narrow metapterygoid is considered here to be a plesiomorphic character, and a single epural also occurs in Chlamydogobius, Pandaka and Eugnathogobius paludosus, while both Brachygobius and Pandaka have 24-25 vertebrae. Akihito and Meguro (1975b) and McKay (1993) noted the dark spots along the mid-ventral margin of the caudal peduncle and anal fin base, which are so conspicuous in Pandaka. These spots are also found in species of Eugnathogobius, Pseudogobius and Redigobius, usually in conjunction with a thin black mid-ventral line (personal observation). Brachygobius is well-known for its striking black banded colour pattern; however, in some species (B. xanthomelas and B. nunus) some of the bands are partly broken into bars and blotches. The midventral black line is absent. Pandaka species have variably developed dark spots on the body and a conspicuous red to yellow and black first dorsal fin. Both Brachygobius and Pandaka form schools or hover near the substrate in groups; their conspicuous markings may help the group stay together in the often turbid waters of their estuarine habitats. Hemigobius and Pseudogobius group together due to their derived lip and gut morphology, and abbreviate headpore patterns (no preopercular pores, and absence of rear part of oculoscapular canal). Each genus has an autapomorphic gut coiling pattern, differing from the rest of the Mugilogobius group of genera. The gut morphology of Pseudogobius resembles a short version of that in Awaous (see Mok, 1980). Hemigobius has 17 segmented caudal rays, while Pseudogobius has the derived condition. Pseudogobius has the derived condition for the s papillae rows on the snout (two 41 rows). Hemigobius has the derived condition for the fine villi on the head (present). Using electrophoretic techniques, McKay (1993) suggested an alternate relationship for some genera in the Mugilogobius-group: Brachygobius aggregatus, B. doriae, Pandaka lidwilli, Pseudogobius olorum, P. poicilosomus (as Pseudogobius sp.) and Redigobius cf bikolanus (as Redigobius sp.). He found Brachygobius, Pandaka and Redigobius formed a single clade of successive sister groups, with two species of Pseudogobius a.s sister-group to this clade. His outgroups were gobiines: three European species (Gobius, Pomatoschistus and Vanneaugobius) and four from the Indo-Pacific (Acentrogobius, Bathygobius, Glossogobius and Istigobius). McKay (1993) considered that Pseudogobius did not possess any autapomorphies, with Redigobius having an autapomorphy in the enlarged jaw, especially in males. This paper has shown that the enlarged jaw character is shared by a number of genera in the Mugilogobius group, and that Pseudogobius has an autapomorphy for character 7 (gut looped corkscrew-fashion about itself and the stomach). McKay is correct, however, in his conclusion that the Brachygobius lineage is quite separate from the gobiines he included in his electrophoretic study. Eugnathogobius does not appear to be monophyletic (e.g. Figures 25, 26-27). It is retained here as a genus for taxonomic convenience, with further work on the problem in progress (Larson, in prep.). The genus is presently characterised by (usually) possessing headpores, the mouth greatly enlarged in males, 17 segmented caudal fin rays, sensory papillae rows a, c, cp and p large and widely spaced, no fine villi on the head, neural spines on the first few vertebrae slender and pointed, palatine not reaching the quadrate, scapula tip sometimes ossified, metapterygoid short or broad but never reaching the quadrate, and a ridge or groove present along the rear edge of the preopercle. Calamiana is defined by the derived condition of characters 3 (16 segmented caudal rays), 16 (papilla row p), 20 (papilla row e, derived condition not present in all species), 37 (papillae rows a, c and cp) and 39 (rear portion of papilla row c). Calamiana kabilia, with its huge mouth, is very similar in appearance to Eugnathogobius, and is placed as sister to the rest of the Calamiana group due to its having the derived condition of characters 6 (expanded neural spines; present in four out of nine specimens examined), 18 (greatly enlarged mouth) and technically having the plesiomorphic condition of character 9 (three s papillae rows on the snout, but each row consists of one papilla only). The relationship of Calamiana to Eugnathogobius needs further examination, as both genera are problematic. Calamiana kabilia shares two characters with Mugilogobius: the first few neural spines are expanded in some specimens, and males have a H.K. Larson 42 greatly enlarged mouth. It also has three rows of s papillae on the snout, but each row consists of one papilla only; in Mugilogobius there are two to six papillae in each row. There are several undescribed species from Malaysia and Papua New Guinea belonging to either Calamiana or Eugnathogobius. These include species which appear to be similar to some taxa currently placed in the Stenogobius-group genus Rhinogobius. The relationship of Calamiana and Eugnathogobius to Rhinogobius needs clarification. Redigobius is the least-derived genus of this whole group. It has the derived state for characters 5 (epural number), 15 (papilla row p), 17 (mouth ｳｩｾ･Ｉ and in some species, characters 11 (mouth position) and 33 (anterior preopercular tip). In contrast to McKay's (1993) finding, the present study indicates that Redigobius is the least derived genus among the Mugilogobius-group of genera, due to its possession of a complete complement of oculoscapular canals and pores, ossified top of scapula, 11-12 precaudal and 14-16 caudal vertebrae, three or four (rarely two) anal fin pterygiophores before first caudal haemal spine, short palatine, short metapterygoid without bridge to quadrate, no fine villi on the head and the anterior nostril close to, but not overhanging the upper lip. During the course of this work, several characters, initially considered as possibly being useful in determining genera, were found to be shared by all or most of the species within the Mugilogobius group. These included the extent of procurrent cartilage (character 18), the adductor mandibulae insertion (character 22) and epineural attachment (character 34). The procurrent cartilage extended over the tip of at least the anteriormost epural in all gobionellines examined for this study. This condition was also found in eleotridines, oxudercines and Schismatogobius. In gobionellines with a single epural, the cartilage may extend posteriorly over the tip of the epural shaft itself (as in Pandaka), not just its anterior flange. Several gobiines (such as Table 4 Acentrogobius and Bathygobius) had the procurrent cartilage extending over the anterior flange of the single epural, but not reaching the tip (Figure 10C). The mandibulae adductor muscle insertion was in two sections, often close together, or as a continuous broad sheet, and specimens could have either condition. This condition may be shared by the gobionellines, gobiines, odontobutids and butines. The insertions were either widely separate or a single broad sheet on the lower half of the maxilla in the two sicydiines examined. Closer study of this character in more genera may provide useful information toward clarifying the higher relationships of gobioids. All the gobionellines examined had the first epineural attaching to or behind the tip of the parapophysis, as did two sicydiines, an oxudercine and a microdesmid. Rhyacichthys and some odontobutids, gobiines, eleotridines and butines had the first epineural attaching anteriorly at the base of the neural spine. This character bears further examination. Composition of the subfamily Gobionellinae A preliminary listing of the generic composition of the subfamily Gobionellinae was given by Pezold (1993), although he provided no defining characters for the group. This paper provides further information toward a diagnosis of the subfamily and generic groupings within it. The Gobionellinae is here recognised as a subfamily (Table 4), but it is acknowledged that, as not all genera contained within it have been revised, its status may change. The subfamily includes genera having paired anterior interorbital pores, a dorsal pterygiophore formula of 3-12210, modally two epurals (one as derived condition), 25-28 vertebrae (10-11+15-18) and two to three (rarely four) anal pterygiophores before the first haemal spine. Species tend to occur in freshwater or estuarine habitats. It was first observed by Robins and Lachner (1966) that gobionelline genera which enter brackish or freshwater include Evorthodus, Gobionellus and Awaous. Proposed classification of the Gobionellinae. Suborder Gobioidei Family Gobiidae Subfamily Gobionellinae Acanthogobius, Amblychaeturichthys, Astrabe, Chaenogobius, Chaeturichthys, Chasmichthys, Clariger, Clevelandia, Eucyclogobius, Gillichthys, Eutaeniichthys, Ilypnus, Lepidogobius, Lethops, Leucopsarion, Lophiogobius, Luciogobius, Pterogobius, Quietula, Sagamia, Suruga, Synechogobius, Typhlogobius. Mugilogobius clade Brachygobius, Caecogobius, Calamiana, Chlamydogobius, Eugnathogobius, Gobiopterus, Hemigobius, Mistichthys, Mugilogobius, Pandaka, Pseudogobius, Redigobius, Stigmatogobius, Tamanka. Stenogobius clade Awaous, Ctenogobius, Evorthodus, Gnatholepis, Gobioides, Gobionellus, Oligolepis, Oxyurichthys, Parawaous, Rhinogobius, Schismatogobius, Stenogobius, Tridentiger, Tukugobius. - - - - - ----------------------------------, Revision of Mugilogobius r---------------- 43 Rhyacichthys Perccotlus M/cropercops Ch/amydogob/us Mugilogob/us Brachygob/us Eugnathogobfus Ca/am/ana Hem/gob/us Pseudogob/us Sfigmafogob/us Awaous stenogob/us Ollgolep/s Oxyuric.hthys Gnatholep/s ancestry with the 3-22110 taxa with a single anterior interorbital pore - the Gobiinae sensu Pezold (1993). Other gobionelline genera comprising the possibly monophyletic North Pacific gobiids, Birdsong et al.'s (1988) and Pezold's (1993) Acanthogobius, Astrabe and Chasmichthys Groups, are not further dealt with in this paper. Gobionelline genera such as Gnatholepis, Gobionellus, Oligolepis, Oxyurichthys, 5tenogobius and Ctenogobius share several characters with Mugilogobius. Awaous, Rhinogobius and 5chismatogobius may be related to these fishes also (Birdsong et al., 1988; Murdy, 1989; Pezold, 1993). A PAUP analysis, based on data in Tables 2 and 3, compared seven 5tenogobius-group gobionelline species to nine representative species of the Mugilogobius-group using Rhyacichthys, Micropercops and Perccottus as outgroups. The analysis found four trees and the consensus tree shown in Figure 29 (length 108, consistency index = 0.44). Three of these trees place Rhinogobius within the 5tenogobius clade, and one is selected for illustration (Figure 30) as it is also the Evorlhodus Rh/nogobius Rhyacichthys Red/gobius Figure 29 Perccottus Result of PADP analysis of 16 representative gobionelline genera; strict consensus of four equally parsimonious trees. Micropercops Chlamydogoblus Pezold (1993) included three species-groups within the Gobionellinae (his Acanthogobius, Astrabe and Chasmichthys groups) which he did not diagnose further other than referring to Birdsong et al.'s (1988) assignment of them to three monophyletic groups. These three Northern Pacific groups may be more closely related to each other than any other group, all possessing wide variation in first dorsal spine numbers (six to eight in Acanthogobius and Chasmichthys Groups; absent in Astrabe Group sensu Birdsong et al.) and high vertebral numbers (Birdsong et al., 1988; Pezold 1993). Miller (1987) first noted that Chlamydogobius and its relatives (Mugilogobius, Brachygobius, Pandaka, Hemigobius and Weberogobius) shared a dorsal pterygiophore formula of 3-12210 with the northwestern Pacific genera Acanthogobius and Pterogobius. These two genera are part of a radiation of gobionellines on both sides of the temperate north Pacific. Miller considered that these north Pacific forms possessed an apomorphy in that papilla row cp was reduced to one or a few papillae below the rear end of a continuous row c (in the Mugilogobius group, row cp is continuous below row c which is broken into two portions, Figure 5) (e.g. Akihito et al., 1988: figures 173-178, 181-184). Miller stated that the Chlamydogobius-Mugilogobius group and the north Pacific genera shared a common Mugilogobius Brachygobius Eugnathogoblus ca/am/ana Hemigobius Pseudogobius Sligmafogobius Awaous stenogobius Oligo/epfs Oxyurichlhys Gnatholepls Evorlhodus Rhinogoblus Red/gob/us Figure 30 One of the four trees comprising the consensus in Figure 29; this single tree results if the analysis is run with all characters ordered. 44 H.K. Larson Rhyacichthys Perccottus Mlcropercops Chfamydogob/us Mug/logoblus Brachygob/us Eugnelhogoblus Ca/amia na Hemlgoblus Pseudogoblus Sligmelogoblus Awaous Slenogoblus Gnatholepis Evorthodus Oxyurichthys 5tenogobius clade, but the placement of Redigobius and other Mugilogobius-group taxa is based on Figures 26-28 and 33 (latter analysis run with Rhyacichthys alone as outgroup). Here also, it must be borne in mind that the trees in Figures 29-33 are based on fewer taxa. The Mugilogobius and 5tenogobius clades (Figure 32) are differentiated by four characters. The Mugilogobius-group is characterised by having the anterior nasal pore absent (derived state, character 42), villi present on head (derived state, character 38), infraorbital pores always present (plesiomorphic state, character 41), and a (modally) longitudinal papillae pattern (one of two derived states, character 8). The 5tenogobius-group has the anterior nasal pore present (plesiomorphic state, character 42), villi absent from the head (plesiomorphic state, character 38), (modally) absence of infraorbital pores (derived state, character 41), and (modally) a transverse papillae pattern (one of two derived states, character 8). Within the latter clade, Rhinogobius has the derived state for character 42 (anterior nasal pore absent) and the plesiomorphic state for character 41 ...- Rhyacichlhys Oiigolepis Mlcropercops Rhlnogoblus Perccottus Redlgob/us Chlamydogoblus Figure 31 One of the four trees comprising the consensus in Figure 29, and resembling it in placing Redigobius and Rhinogobius away from the two resolved clades. Mugiiogoblus L-.. L..- single tree which results if the analysis is run with characters ordered. The fourth tree (Figure 31) places Redigobius and Rhinogobius away from the two clades, while the consensus tree (Figure 29) places Rhinogobius in a trichotomy with the Mugilogobius and 5tenogobius clades. This unresolved relationship highlights the need for further work on characters and species of this subfamily. It should be noted that the relationships within the Mugilogobius clade in Figures 29-31 differ from those indicated in Figure 28, as fewer taxa (eight versus 34) are involved. The relationships shown in Figure 28 are preferable. Also, it should be borne in mind that not all of the 5tenogobiusgroup genera were included in this analysis and it is possible that the relationships shown in the cladogram would change if more taxa were added. The tree illustrated in Figure 30, which places Rhinogobius with the 5tenogobius clade, is used as the starting point for a hand-drawn tree. This handdrawn tree (Figure 32) is used as a basis for the classification below (Table 4). The tree agrees with the consensus (Figure 29) with regard to the Silgmalogoblus Brachygoblus Hemlgoblus Pseudogoblus Caiamiana Eugnalhogoblus Red/goblus L..- Awaous Slenogoblus O/igolepls Oxyurichlhys Evorlhodus Gnalholepis L..- Figure 32 Rhlnogoblus Hand-drawn tree based on Figs 26-28, 30, 33; and used as the basis for deriving the classification used in this paper., Revision of Mugilogobius 45 Rhyacichthys Chlamydogoblus Mugllogoblus Brachygobius Eugnalhogobius C81amiana Hemigobius Pseudogobius Sf;gmafogobius Redigobius Awaous Slenogobius Oligolepis Oxyurichlhys Gnatholepis Evorthodus Rhinogobius Figure 33 Result of PAUP analysis of 16 representative gobionelline genera, with Rhyacichthys aspro alone as outgroup; strict consensus of three equally parsimonious trees. (infraorbital pore present), thus resembling taxa such as Redigobius and Eugnathogobius (Figure 34). It is tempting to place the Mugilogobius and Stenogobius clades within tribes for taxonomic convenience. There are two available names for the Mugilogobius and Stenogobius clades, the Brachygobii and the Platygobii (Bleeker, 1874). In Bleeker's (1874) classification, the Brachygobii comprised only Brachygobius (a gobionelline) and Lophogobius (a gobiine). The Platygobii comprised Gillichthys, Gobiopsis, Glossogobius (all gobiines) and Platygobius (a junior synonym of the gobionelline Awaous). However, there is no single feature (synapomorphy) possessed by either clade and the temptation to place these taxa in tribes is resisted. Additionally, work remains to be done on the relationships of the "basal" taxa Redigobius and Rhinogobius. Birdsong et al. (1988) placed Evorthodus and Awaous with their Sicydium Group genera, although Miller (1973b) had earlier pointed out that EvortJlOdus was " ... obviously related to the American Gnatl101epis Bleeker and the Gobionellus Girard complex... by osteological and lateral-line characters. Murdy (1989) discussed similarities between Evorthodus and the oxudercines (characters of lower jaw teeth, fifth ceratobranchial /I morphology, retractor dorsalis muscle). He used Evorthodus as first outgroup and the sicydiines Stiphodon and Sicydium, the gobionellines GnatllOlepis, Gobioides, Ctenogobius, Mugilogobius and Oxyurichthys, and the arnblyopines Trypauchen and Brachyamblyopus as additional outgroups for polarising oxudercine characters. Murdy (1989) also pointed out the possibility of convergence between EvortllOdus and eight of the oxudercine genera in the enlarged lattice-like fifth ceratobranchials, as these fish live in similar softmud habitats and may share feeding or burrowing behaviour. Similar fifth ceratobranchials are found in Hemigobius and Pseudogobius. These two genera do not have the very widely expanded, almost leafshaped bones as exhibited by Evorthodus and the oxudercines. The bones are narrower, and are lattice- or lace-like with two layers of struts in Hemigobius and Pseudogobius, and more closely resemble those of Gnatholepis and Awaous. Harrison (1989) proposed a possible relationship between Ctenogobius, Evorthodus, Gnatholepis, Gobioides, Gobionellus, Oligolepis (his 'Ctenogobius lineage') and Stenogobius based upon palatine morphology. He placed these in the subfamily Gobionellinae sensu Miller (1987), which included the Amblyopinae and Oxudercinae. Harrison considered Stenogobius to be the most primitive gobionelline, despite having a derived palatine. Oxyurichthys was placed with eight genera of oxudercines, forming an 'Oxyurichthys lineage', based on their shared possession of a very short palatine (Harrison, 1989: figures 4, 8). As Murdy (1989) has now shown, the oxudercines form a distinct monophyletic group. Harrison (1989) presented a cladogram which united the genera mentioned above, Awaous and the Sicydiinae by possession of a long palatine which reaches or nearly reaches the quadrate and a 3-12210 dorsal Figure 34 Rhinogobius brunneus, papillae pattern. NTM 5.12121-012, Omijya River, Iriomotejima, Japan. Fine stipple indicating low groove on anterior part of cheek. Note the single cp row papilla on cheek below papilla row c. Scale bar = 1 mm. H.K. Larson 46 I ＢＭｾＭ B ) ...... J '\ " ')):' I I ..... I I ... ...... ..... .\. <::.\;.... ...ｾ ::i \ ..... ) ｾ I / Figure 35 A, Rhinogobius giurinus, NTM 5.12134-012; 8, Evorthodus lyricus, NTM 5.12858-001; showing relationship of papilla row d to row cp. Also note that the IP pore (infraorbital) is present in R. giurinus but absent in E. lyricus. Scale bar = 1 mm. pterygiophore formula (Figures 6, 36-37). The 'Ctenogobius lineage' and 'Oxyurichthys lineage' were sister groups sharing the character of papillae row d extending posteriorly past row cp (Figures 3B, 35B). Harrison characterised Oxyurichthys by an autapomorphy, a short stubby palatine which just reaches the pterygoid tip, but he did not give an autapomorphy for the 'Ctenogobius' lineage. He treated Awaous as separate from these two lineages and Stenogobius, and placed it as sister-group to the sicydiines, because of shared multiple a rows, a single epural, and apparently, shared similarities which are not explicit in his cladogram. Harrison (1989) discussed the character of papilla row d reaching past row cp in gobionellines with transverse papillae patterns, noting that Ctenogobius, Evorthodus, Gnatholepis, Gobionellus, Gobioides and Oligolepis all have this state. There are apparent homologies among the gobiines for this character, as Egglestonichthys patriciae Miller, E. bombylios Larson and Hoese, Mahidolia and Priolepis anthioides (Smith) have this arrangement. Many other gobiines (e.g. Amblygobius, Amblyeleotris, Drombus, Egglestonichthys melanoptera (Rao), Hetereleotris, Lotilia, Priolepis fallacincta Winterbottom and Burridge) and the gobionellines Awaous, Gobiopterus, Parawaous and Stenogobius have the papilla row c/ep extending down past row d (Figures 3A, 35). The pattern in Rhinogobius giurinus is similar, but the cp row barely extends downward, and could easily be assigned to either group, depending on the specimen examined (Figure 35A). The character also varies in Gnatholepis species, with row d not always extending back to meet row cfcp. Harrison (1989) indicated that Awaous may be closer to the Sicydiinae than the Gobionellinae. In this paper, Awaous is considered a gobionelline. The sicydiine osteological characters noted by Harrison (1989) shared by Awaous and the Sicydiinae were the long palatine, 3-12210 dorsal pterygiophore pattern, unossified scapula, single epural and spatulate posterior pelvic process. As has been shown here, the first three characters are found throughout the Gobionellinae and the single epural occurs in three other gobionelline genera as well as in the Sicydiinae. Harrison (1989: 344) considered the relationship of Awaous to Chlamydogobius eremius, Mugilogobius chulae, Stigmatogobius sadanundio and Tamanka ubinensis (= Calamiana variegata), based on these three characters but did not conclude that these shared characters implied kinship because this would then indicate "... that all other shared specializations (anatomical and ecological) of Awaous and the sicydiines are homoplastic for these taxa". He considered that these four Mugilogobius clade genera were "unrelated gobiines" which also possessed long palatines, and did not discuss Mugilogobius further. One character which he used, the spatulate pelvic process found in Awaous and the sicydiines, also occurs in at least one Rhinogobius, R. brunneus (personal observation; other species of Rhinogobius not yet surveyed), and in Tukugobius (Birdsong et al., 1988; Harrison, 1989). The latter is a genus which, based on available information, is a specialised relative of Rhinogobius living in swift freshwater streams in the Philippines. The validity Figure 36 Jaws and suspensorium of Awaous acritosus, female, NTM unregistered, 5tewart River, Queensland. Scale bar = 1 mm. Revision of Mugilogobius Figure 37 Jaws and suspensorium of Oligolepis acutipennis, female, ex NTM 5.11125-020, 5anur Beach mangroves, Bali. Scale bar = 1 mm. of Tukugobius awaits a review of Rl1inogobius. The spatulate pelvic process may possibly be linked to adaptation for living in fast-moving freshwater habitats. Watson provided diagnoses of the gobionellines Awaous (Watson, 1992) and Stenogobius (Watson, 1991) and considered them to be closely related and sometimes confused (Watson, 1991). He did not discuss their relationships further, nor place the genera in systematic context (not his intention in these papers). Parenti and Maciolek (1993) showed that Awaous was not a sicydiine due to the plesiomorphic conditions of the relationship between the pelvic fin spine and the first pelvic ray (not separated by gap from other rays), in the presence of the premaxillary process and the non-adnate tongue. Parenti and Maciolek (1993) referred to, and illustrated, the unossified portion of the pelvic fin spine of Awaous and Sicyopus, which has been regarded as a characteristic sicydiine feature (Birdsong et al., 1988; Harrison 1993). Parenti and Maciolek stated that this character is present among some gobionellines, oxudercines, sicydiines and in Rl1yacic11tl1ys, but absent in Hypseleotris (an eleotridine). Examination of cleared and doublestained taxa for this paper indicated that the pelvic fin spine is fully ossified in odontobutids, butines and eleotridines. The slender tip of the spine is partly non-alizarin staining, taking up relatively little alcian blue in comparison with the caudal cartilages, for example, in the butines Eleotris, Oxyeleotris and Bostrycl1us. Amblyopines, gobiines, gobionellines, oxudercines and sicydiines examined for this paper had up to half the pelvic spine apparently cartilaginous, that is, often heavily alcian blue-stained but sometimes transparent, with quite an abrupt break between the two areas. Rhyacichthys was similar, in that the alizarin-stained bone ended abruptly halfway but the distal half of the spine was translucent, unstained by alcian blue. The two genera of microdesmids examined were more similar to the butines in that the distal half of 47 the pelvic spine was not alizarin stained, but a sliver of alizarin-stained bone extended into the alcian blue stained half. Therefore, the character appears to occur in most gobioids (n.b. xenisthmids and kraemeriids not examined) and may be diagnostic of the whole group (Parenti and Maciolek, 1993) rather than the sicydiines alone. As stated in the Introduction, Pezold (1993) agreed with Miller in recognising the Gobionellinae as a subfamily, although they each included different genera within it. Pezold's Gobionellinae is paraphyletic in that it did not include all gobionellines (he left out Hemigobius, Caecogobius and Cl1lamydogobius) and included several genera which are here considered to be gobiines. Examination of all possible members of the subfamily Gobionellinae is outside the scope of this paper, but comments on 12 of the genera (Apl1ia, Deltentosteus, Hyrcanogobius, Knipowitscl1ia, Leucopsarion, Mesogobius, Neogobius, Nesogobius, Scl1ismatogobius, Synecl1ogobius, Vitraria, Vomerogobius) included in the Gobionellinae, sensu Pezold, follow. Nine of these genera were placed as "unassigned" by Birdsong et al. (1988) because they were either problematical or unique in their assemblage of characters. One genus (Scl1ismatogobius) was not examined by Birdsong et al., and the others (Leucopsarion and Synecl1ogobius) were respectively placed in their Astrabe and Synechogobius groups. Five genera placed in the Gobionellinae by Pezold (Deltentosteus, Hyrcanogobius, Knipowitscl1ia, Mesogobius and Neogobius) are here considered likely to be specialised gobiines more properly related to the European Gobius Group of Birdsong et al. (1988). Most of these have morphology and papillae patterns common to many Mediterranean to western European gobiines (Miller 1986). Vomerogobius, which Pezold assigned to the Gobionellinae, may be a gobiine. The holotype of V. flavus (type species of the genus) resembles Palatogobius, which Birdsong et al. (1988) placed in their Microgobius Group, and Pezold (1993) placed in the gobiine tribe Gobiosomini. Vitraria is a juvenile Sicyopterus (holotype of type species Vitraria clarescens Jordan and Evermann examined). Apl1ia may be a European gobiine or a relative of Gobiopterus (see discussion of Gobiopterus below). Nesogobius, an Australian endemic temperate genus, may be an unusual gobiine. It has only 13 segmented caudal rays, and shows variation in dorsal and anal spine numbers and presence or absence of headpores, as does its probable closest relative, the temperate Australian Tasmanogobius (Hoese, 1991). However, it is also possible that these two genera could be very derived gobionellines, as specimens may have a 3-12210 dorsal H.K. Larson 48 pterygiophore formula (formulae of 3-22110, 3-3210, 3-231010, 3-1222010 and 3-1221100 have also been observed), but usually have a single epural (Hoese 1991; personal observation). The European genera Aphia (placed in the Gobionellinae by Pezold), Crystallogobius and Pseudaphya (latter two not mentioned by Pezold) are similar in appearance to Gobiopterus but have an unsegmented spine in the second dorsal and anal fins (Miller, 1973b, 1986). Gobiopterus and Mistichthys, despite their odd dorsal pterygiophore formulae (variably 3-12210, 3-12200 or 3-120000 etc., due to the loss of one or more posterior dorsal fin spines), are here considered to be gobionellines, closely related to Brachygobius and Pandaka, as suspected by Birdsong et al. (1988). These four genera are all small, mostly schooling fishes which occupy similar estuarine to freshwater habitats, have low vertebral numbers (10+15), all second dorsal and anal fin elements segmented, one or two epurals, two pre-anal pterygiophores and may have transverse papillae patterns (greatly developed in Gobiopterus). The myology and possible relationships of these paedomorphic genera are under study by Tony Gill and Randy Mooi. The three genera Aphia, Crystallogobius and Pseudaphya are possibly derived from a European species and not related to the Indo-Pacific genera (Brachygobius, Gobiopterus, Mistichthys, Pandaka) at all. Leucopsarion is a derived monotypic genus from Japan, Korea and southern China, and is probably a gobionelline. Birdsong et al. (1988) placed it in their Astrabe Group, as it has high vertebral numbers (14+20), two epurals and one pre-anal pterygiophore. It has lost its first dorsal fin, so the dorsal pterygiophore formula cannot be applied. The genus also has well developed transverse papillae patterns, all segmented dorsal and anal fin rays and a deep fraenum forming a tube-like pelvic fin (Akihito et al., 1988; personal observation). These are characters also found in Gobiopterus; a genus in which reduction in the first dorsal fin also occurs. A review of Leucopsarion, as well as Gobiopterus and Mistichthys, is required. Schismatogobius appears to be a derived gobionelline with six nominal species, all from freshwaters of the Indo-west Pacific (from Sri Lanka to northern Australia). It has one epural, a dorsal pterygiophore pattern of 3-12210 or 322110, the first five neural spines sometimes expanded at the tips, a longitudinal papillae pattern, no scales, has posterior nasal pores but no anterior interorbital pores, has postorbital but no infraorbital pores, and only the opercular portion of the rear oculoscapular canal is present. Males have very large mouths with a brightly coloured lining (red, orange or yellow; personal observation; Watson and Horsthemke 1995). Species inhabit swift shallow freshwater streams, hiding among gravel. Its relationships remain to be studied, as Kottelat and Pethiyagoda (1989) pointed out. As a result of this work, I include 50 genera within the Gobionellinae. A provisional classification is presented here (Table 4), although it is recognised that the relationship between the Gobionellinae and the rest of the Gobiidae is still not fully resolved. Generic diagnoses and species descriptions One hundred and fifty-seven described species among 35 nominal genera were found to form a group of gobionellines which appeared to be related to (and included) Mugilogobius, with an additional 21 undescribed species discovered. Of the 35 nominal genera, 21 were found to be synonyms of other genera. Phylogenetic placement of Caecogobius, Gobiopterus, Mistichthys and problematic species such as Gobius tigrellus was not attempted. The status of Caecogobius will not be known until additional material becomes available. Two genera, Mugilogobius and Tamanka, are described and discussed in full. The other genera, which include species often confused with Mugilogobius in the literature, are diagnosed, and nominal species of each genus listed but not described (Brachygobius, Calamiana, Chlamydogobius, Eugnathogobius, Hemigobius, Pandaka, Pseudogobius, Redigobius, Stigmatogobius). These genera have been or are being revised as separate projects outside the scope of this paper (Larson, 1995; Larson, 1999a-b; Larson, in preparation). All the 157 nominal species within this group of fishes and their present status are listed in Appendix 1. 1. lA. 2. 2A. 3. Key to Mugilogobius and related gobionelline genera All elements in second dorsal and anal fins segmented 2 First element in second dorsal and anal fins unsegmented 4 Sensory papillae with reduced longitudinal pattern; if transverse rows of papillae present, opercle with scales; body and/or first dorsal fin distinctly spotted or banded. .......................................................................... 3 Sensory papillae on head and nape always in distinct transverse pattern; head and nape naked; body compressed and slender; body and fins transparent when live, usually with few melanophores Gobiopterus Adults very small, at most reaching 15 mm SL; two anal pterygiophores before first caudal vertebra; one or two epurals; opercle naked; first dorsal fin usually distinctly Revision of Mugilogobius 49 marked, rest of body spotted or marbled, not banded; freshwater to estuarine habitats; Indo-Australian Archipelago . .............................................................. Pandaka 3A. jaws subterminal, mouth small, lower lip reduced, thin and folded forward . ......................................................... Hemigobius 10. Adults larger, up to 25 mm SL; three anal pterygiophores before first caudal vertebra; one epural; opercle with scales; robustbodied with distinctive banded body colour; restricted to freshwater or mangroves of South-east Asian Archipelago . ...................................................... Brachygobius 4. Eyes present, normal; body variably patterned .......................................................................... 5 4A. Eyes reduced to pigment spots below skin; body pigment absent Caecogobius 5. Headpores present at least in interorbital region; if absent, then distinct rows of transverse papillae also present or papillae in cheek rows a and c few, rows composed of widely spaced papillae which are larger than those in rows b and d 6 5A. Headpores always absent; papillae in cheek rows a and c numerous, and same size as those in rows b and d 11 6. 17 segmented caudal fin rays; preopercular pores and rear portion of oculoscapular canal present or absent 8 6A. 16 segmented caudal fin rays; preopercular pores and rear portion of oculoscapular canal always absent 7 7. Mouth terminal, lower jaw tip anteriormost; gut short, "S-bend" shape; headpores absent in three species Calamiana 7A. Mouth subterminal, upper jaw tip anteriormost, with round snout overhanging mouth; gut long, spirally coiled about its longitudinal axis; headpores always present Pseudogobius 8. 8A. 9. 9A. Head papillae longitudinal; if transverse rows present, then rear portion of oculoscapular canal present also; colour pattern variable .. 9 Head papillae with many transverse rows; headpores usually present but rear portion of oculoscapular canal always absent; distinctively spotted or banded colour pattern Stigmatogobius Preopercular pores and rear portion of oculoscapular canal usually present (absent in three species of Eugnathogobius); gut short CS-bend"), or with two or three loops; jaws terminal to subterminal, often greatly enlarged in males, both lips fleshy 10 Preopercular pores and rear portion of oculoscapular canal always absent; gut long and coiled into many loops (12 or more); Second dorsal fin modally with one more ray than in anal fin; modally, scapula ossified at tip or around foramen; three to four anal pterygiophores before first caudal vertebra; vertebral pattern 11 + 15 to 12+14; most species stocky, head and body somewhat compressed; estuarine to freshwater, widespread across the Indo-Pacific . ......................................... Redigobius lOA. Second dorsal and anal fins with equal numbers of rays; scapula modally cartilaginous, tip above foramen sometimes ossified; two to three anal pterygiophores before first caudal vertebra; vertebral pattern 10+16; head usually depressed; restricted to freshwaters of South-east Asian region Eugnathogobius 11. S papillae on snout usually in two rows, rarely three (each row consists of only one or two papillae), first row just above upper lip absent; pectoral rays 11-14; intestine long and coiled into three loops; pectoral girdle smooth Chlamydogobius 11A. S papillae on snout in at least three rows of two or more papillae, the first of which is usually the longest and runs just above upper lip fold (few species with each row represented by only one or two papillae); pectoral rays 13-20; intestine simple, with one "S-bend"; most species with knobs or fleshy ridge along pectoral girdle .. ...................................................... Mugilogobius Brachygobius Bleeker, 1874 Brachygobius Bleeker, 1874 (type species: Gobius doriae Giinther, 1868: 265, plate 12, figure A, Sarawak, by monotypy). original designation and Hypogynmogobius Bleeker, 1874 (type species: Gobius xanthozona Bleeker, 1849: 34, Surabaya, Indonesia, monotypy). by original designation and Thaigobiella Smith, 1931 (type species: Thaigobiella sua Smith, 1931: 35, figure 17, Thailand, by original designation and monotypy). Diagnosis Distinguished by following combination of characters. Second dorsal and anal fin rays 7-9, all segmented; 15-19 pectoral rays; segmented caudal rays 17-22, modally 17, in 9/8 pattern; 21-44 scales in lateral series; TRB 7-17; 12-24 circumpeduncular H.K. Larson 50 A --...........---. . ."rO @ ".; it···. / '\ " • c:? #'," • . .... . . -: .----\ .I , • . • (/I ,.,a b • I • '"0 ... '0. ••• . / .. I :: ",' ••••. ·cp. : ':. ..... (j .. ｩｾ '. • ｾ ..:--: .'.. 1 v!'" . ·....·.. . . .- r . . . ·;···· ..· . 1'// '; Ｍｾ / ［Ｚｾ B Figure 38 Papillae patterns in Brachygobius. A, B. doriae r NTM 5.12812-001, Brunei; B, B. xanthozonus, ex CA5 33045, Borneo, Scale bar = 1 mm, scales; predorsal scales absent or up to 14 present; fine villi sometimes present on dorsal surface of head; gill opening short, reaching to lower pectoral base or to just under opercle; distinct rounded to pointed lobe on rear edge of branchiostegal membrane; pectoral girdle (anterior edge of cleithrurn) usually smooth, sometimes with fleshy flange; gut short, with one simple loop; sensory papillae generally small, evenly sized, rows c and cp sometimes including relatively large papillae (Figure 38); reduced longitudinal or transverse papillae pattern; vertically oriented r rows may be present on snout; no headpores; jaws short, mouth terminal with lower jaw anteriorrnost, jaw length similar in males and females; teeth in several rows, small, stout and pointed; unpaired fins low, rounded; genital papilla in male fleshy and pointed, usually short, and rounded and bulbous in female; body stout and robust to slender and rather compressed; colour pattern of two to five dark bands crossing white to yellow head and body. Pterygiophore formula 3-12210; one or two epurals, modally one; vertebrae 10+14-16, modally 10+15; two to three pre-anal pterygiophores; neural spines of first few vertebrae slender, pointed; palatine and pterygoid stout, almost equal in size, palatine not reaching quadrate; metapterygoid very low, narrow, not reaching toward quadrate (Figure 39); fifth ceratobranchial short, triangular, with stout curved teeth, tall flange ventrally; scapula unossified; sometimes one gill-raker ossified; first epineural inserts on or just behind parapophysis. Remarks Inger (1958) and Miller (1989) commented on the species of Brachygobius, and gave brief accounts of the taxonomic history of the genus, the nominal species and their synonymies. Neither author undertook a thorough revision (not the stated aim of either author). Miller (1989) resurrected the genus Hypogymnogobius, which is here considered to be a synonym of Brachygobius (see Remarks under B. xanthozonus). Horsthemke (1990) provided a useful summary of species as observed in the aquarium industry. After examination of type material of all described species and of recently collected specimens held in other museums, I found that several species were not identifiable. A revision of the genus is in progress (Larson, unpublished data; in part with Chavalit Vidthayanon, NIFI). Nine nominal species are known and there are probably about nine valid species, which may not all correspond with available names. The lobe and/or notch on the branchiostegal membrane is present to some degree in most of the Mugilogobius-clade, but is especially pronounced in Brachygobius (Figure 38), notwithstanding SmithVaniz and Johnson's (1990: 212) comment as to its being unique to the Plesiopidae. Gill and Mooi (1993) describe the notch in the branchiostegal membrane of gobioids, with gobiids having the notch between the second and third branchiostegal rays. PRELIMINARY LIST OF VALID SPECIES Brachygobius aggregatus Herre, 1940 Figure 43 Brachygobius aggregatus Herre, 1940a: 361, plate 4 (Kabili River, Dumaguete, Philippines). Material Examined Lectotype CAS 32990, 1(12), Durnaguete, Negros Oriental, Philippines, A.W. Herre, 26 December 1936. 51 Revision of Mugilogobius Other Material CAS 38461, 33(10-14), Dumaguete, Negros Oriental, A. Herre, 12 August 1940. Brachygobius doriae (Giinther, 1868) Figures 38-41, 43 Gobius doriae Giinther, 1868: 265, plate 12, figure A (Sarawak); Tortonese 1963: 343. Hypogymnogobius doriae: Bleeker 1983: plate 432, ･ｲｾｩｦ 10. Material Examined Syntypes BMNH 1868.1.28.17-19, 3(24-25), Sarawak, Malaysia, Doria coll. Other Material Indonesia: CMK 6599, 5(22-24), Medan, Sumatra, Vivaria Indonesia, May 1990; ZMH 21228, 11(2026), Belawan-deli, Sumatra, Ladiges, July 1935. Malaysia: ZRC 669, 8(16.5-31), Bukit Stigang, 5 of Kuching, Sarawak, B.L. Urn, 20 January 1969. ZRC 1409, 8(15.5-24.5), Bukit Kuda Road, Klang, Selangor, B.L. Urn, 13 August 1960. Brunei: NTM 5.12812-001,2(21-27), Nypa forest, Kedalayan River, R. Hanley and S. Choy, 7 April 1989. Singapore: CAS 76136, 4(22-26.5), M. Rakowicz, January 1947. Brachygobius kabiliensis Inger, 1958 Figure 43 B Brachygobius kabiliensis Inger, 1958: 110, figure 19 (Kabili River, Sandakan District, East Coast Residency, North Borneo). Figure 39 Figure 40 Jaws and suspensoria of: A, Brachygobius xanthozonus, female, RMNH 12084, Java; B, Brachygobius doriae, female, NTM unregistered, aquarium specimen. Scale bar =lmm. Material Examined Holotype FMNH 47991, 1(14), Kabili River, East Coast Residency, Sandakan, Sabah, Malaysia, A.W. Herre, 1937. Brachygobius doriae. Syntype of Gobius doriae Giinther, 24 mm SL, BMNH 1868.1.28.17-19, Sarawak. 52 H.K. Larson Figure 41 Brachygobius doriae, 29.5 mm SL, ZRC 669, Sarawak. Paratypes Malaysia: Sabah: USNM 171753, 1(13), same data as holotype; FMNH 44989, 2(11.5-12), mile 2, East Coast Residency, Sandakan, J.A. Tubb, 20 May 1948. Other Material Singapore: CAS/SU 33046, 1(10), AW. Herre, 10 March 1937; CMK 8339, 10(9-16.5), Kranji mangrove, P. Ng et al., 30 January 1992; NTM S.14235-OO1, 15(7.5-16.5), Sungei Buloh mangroves, K. Lim, 30 January 1992. Damrey, Phnom Penh, Mekong Delta, 12 March, 1962, F. D'Aubenton; CMK 4786, 3, 9-12 mm SL, same data as preceding. Other Material KUMF uncatalogued, 30(12-14.5), Nakorn Phanom market (possibly from Huay Nong Yat, municipal reservoir), 26-31 May 1990, coll. T.R. Roberts. Brachygobius nunus (Hamilton, 1822) Figure 43 Gobius nunus Hamilton, 1822: 54, 366 (Ganges River, Brachygobius mekongensis Larson and Vidthayanon, 2000 ?Brachygobius aggregatus (?): Horsthemke, 1990: 543, figure 9. Brachygobius mekongensis Larson and Vidthayanon, 2000: 2-6 (Laos, Savanakhet Province, Xe Champhon). Halo type ZRC 43853, male, 14.5 mm SL, Xe Champhon, between bridge at Muang Kengkok and about 4 km upstream, Xe Biang Hiang Basin, Savannakhet, Laos, 23 April 1997, M. Kottelat. Paratypes Laos: CMK 13558, 67, 10-16.5 mm SL, same data as holotype; NTM 5.14871-001, 15(12.5-16), same data as holotype; ZRC 43854, 15, 12.5-16 mm SL, same data as holotype; LARRI uncatalogued, 15, 1014.5 mm SL, same data as holotype. Thailand: NIFI 2959, 49, 10.5-15 mm SL, Kud Thing, Bueng Kan, Nongkhai Province, 25 March 1996; NIFI 3071,1, 10 mm SL, Nong Han Sakhon Nakhon Prince, December 1991, D. Santana; NIFI 3073, 7, 14.8-17.5 mm SL, Mukdahan Market, Mekong River, 5 April 1990, C. Vidthayanon; NTM 5.14868-001, 7, 14-15.5 mm SL, same data as preceding; NIFI 3072, 6, 8-13.5 mm SL, grassy marsh area where Udon-Nong Khai road crosses railway, about 6 km N of Udon, November 1995, J. Kittipong; NTM 5.14867-001, 5, 11-12.5 mm SL, same data as preceding. Cambodia: MNHN 1985-797, 8, 9-13 mm SL, Beng Kebal below Calcutta, India) (no types exist). Gobius alcockii Annandale, 1906: 201 (India). Material Examined Syntypes of Gobius alcockii ZMA 114.487, 3(10-11.5), Lower Bengal at Port Canning, India, N. Annandale, January 1906. Other Material India: SMF 24052, 4(12-14), Calcutta, A Werner, 1955; CAS 34762, 138(8-16), Uttarbhag, AW. Herre, 14 April 1937; ZMH 9569, 13(10.5-12), Yanali River, Raimona, Assam, V. Maydell, 29 March 1957. Sri Lanka: USNM 246753, 4(7.5-12), N of Kallu, Batticalom district, Eastern Province, T. Iwamoto, 11 June 1970. Brachygobius sabanus Inger, 1958 Figure 43 Brachygobius sabanus Inger, 1958: 113, figure 20 (Lamag, Kinabatangan District, East Coast Residency, North Borneo). Material Examined Holotype FMNH 47990, 1(15.5), East Coast Residency, Sabah, Malaysia, J.A Tubb, 1949. Paratypes FMNH 44988, 4(16-26), Kinabatangan River, Abai, Revision of Mugilogobius Figure 42 53 Brachygobius xanthozonus, 32 mm SL, RMNH 12082, Surabaya, Java. East Coast Residency, Sandakan, Sabah, Malaysia, 24 June 1949; USNM 171752, 1(18.5), same data as preceding. Brachygobius xanthozona: Fowler 1937: 248-250. Brachygobius sua: Smith 1945: 549-550. Hypogymnogobius xanthozona: Bleeker 1983: plate 432, figure 7. Other Material Malaysia: USNM 316181, 16(12-21), Muar River, West Malaysia, T. Roberts, 11-12 May 1973. Thailand: USNM 119566, 8(12-16), tidal ditch, Bangkok, H.M. Smith, 10 May 1934; CAS 27453, 694(13-18), freshwater pond, Gulf of Thailand, R. Rofen, 24 November 1957. Hypogymnogobius xanthozonus: Miller 1989: 376-382. Material Examined Holotype of Gobius xanthozona RMNH 4541, 1(25.7), "Java, Sumatra, Borneo". Other Material Brachygobius xanthomelas Herre, 1937 Figure 43 Brachygobius xanthomelas Herre in Herre and Myers, 1937: 43, plate 4 (Mawai district, Johore, Malay Peninsula). Material Examined Holotype CAS-SU 30953, 1(18), Mawai district, Johore, Malaysia, A.W. Herre, 20 March 1934. Paratypes CAS 16964, 8(8.5-15.5), same data as holotype (24 paratypes exist with this number, but remainder on loan elsewhere and unavailable for study); USNM 101226, 7(11-16), same locality as holotype, 22 March 1934; FMNH 47982, 1(16.5), same data as holotype. Other Material Malaysia: ZRC 13963-13966, 4(12.5-17), Sungai Mupor, Kota Tinggi, Johore, M. Kottelat and K. Lim, 22 January 1991. Indonesia: Borneo: CMK 6745, 14(12-16), Sungei Tekam, Kalimantan Barat, M. Kottelat, 23 April 1990. Brachygobius xanthozonus (Bleeker, 1849) Figures 38-39, 42-44 Gobius xanthozona Bleeker, 1849: 34 (Surabaya). Thaigobiella sua Smith, 1931: 35, figure 17 (Bangkok, Thailand). Malaysia: Borneo: ex CAS 33045, 1(22), Kabili River, A.W. Herre, 31 January 1937. Indonesia: ZMH 19009, 1(32), "East Indies"; RMNH 12084, 1(31), Osthoek, Java, P. Buitendijk, 1921; RMNH 12083, 1(25), Tandjang, Sumatra, P. Buitendijk, November 1927; RMNH 12082, 2(19-30.5), Surabaya, Java, P. Buitendijk, December 1929. Remarks Koumans (1931) and Smith (1945) placed Hypogymnogobius as a junior synonym of Brachygobius. Miller (1989) resurrected Hypogymnogobius as a separate genus, based on its transverse papillae pattern (Figure 38B). However, as Miller himself stated and illustrated (1989: 379, 381, figures 2-3), there is a "general correspondence between number of transverse rows in Hypogymnogobius and the number of individual papillae in rows a and c of the Brachygobius species". The transverse pattern in Brachygobius xanthozonus is an extreme form of "proliferation" as can also be seen in Mugilogobius sarasinorum and M. rexi sp. nov., for example. All other characters fall within the definition of Brachygobius. The holotype of Gobius xanthozona is included in RMNH 4541. The bottle also contained two other specimens, 33-35 mm SL, too large to be types; Bleeker gave the length of Gobius xanthozona as 33 mm TL in his original description. The illustrated specimen in Bleeker (1983) is 40 mm TL and about 33 mm SL, and so may not be the holotype. The unique holotype of Thaigobiella sua Smith (USNM 90315) is presumed to be lost, and its ｾ 30' 30' , , 10' ｬｾＭＬｾ, ;0 , .' : -" 10' ,t CAROUNE IS. 4t,;t,r, • , Kosrae ':j <.t"'o" •••• KIRllAn O· .. '; ｾＮ ". 10' • • o Mauritius 20' ... o • 30' ao' aggregatus doriae kabiliensis nunus sabanus xanthomelas xantholOna 0' ... c::::;::::; ｾＢ 60' " ':: .. .':. ".ARSHALLIS, ' :.: V . MARlANAIS, )I r:,.q LACCAOIVE IS.' ... Maldtv.$ " 20' ＮｾＺＧ '.' ,e.\ SOLOMON IS. '. " 10' " ,0:.. SA"';" ｾ '. ｆＱｊｉｾＮＬ .. <C:)", : ＮＧｾ ¥ AUSTRAUA N:). ｾｏｙａｬｔ ' ", 20' IS. 30' 100' 120' 160' :c ｾ r-' Il> Figure 43 Distribution of Brachygobius, based on type localities and material examined, Ul o ::l Revision of Mugilogobius Figure 44 55 Brachygobius xanthozonus, 22 mm SL, ex CAS/SU 33045, Kabili River, Sabah, Malaysia. identity has been in question. It was not found in the USNM during a search I made in 1991. A note on a USNM file card says that the specimen was not received from Smith as of 27 January 1934. It was discovered that several types from Smith's collection were never sent to the USNM (Wongrat personal communication). A search I made at KUMF (Bangkok) in 1993 provided no clues as to the whereabouts of the type of Thaigobiella sua. Smith's original notebooks and card index gave no further information as to the specimen's fate, and Smith apparently obtained no further specimens of the species. The KUMF collection (including Smith's collection and types) has been moved several times (Wongrat, personal communication), and it is thought that the specimen, never sent to the USNM, was lost during one of these moves. About 100 fish specimens sent by Smith to USNM never actually arrived (1930 correspondence files, USNM acc. no. 112170). Material which Inger (1958) referred to as "topotypes" of B. sua (USNM 119566) are indistinguishable from type specimens of his species B. sabanus. The specimens which Inger says he examined do not agree with the characters he gave for B. sua in his 1958 key to species, nor with Smith's (1931) drawing of the holotype of Thaigobiella sua. Horsthemke's photograph of B. sua appears to be of B. sabanus (Horsthemke, 1990: 545). No specimens in museum collections, identified as B. sua from Thailand, were found to resemble Smith's description and figure; indeed, such specimens invariably turned out to be B. sabanus. A chance discovery of a specimen of B. :mnthozonus among seven B. doriae from the Kabili River, Borneo (CAS 33045, Figure 44) solved the mystery of the identity of B. sua, as it agrees almost exactly with Smith's figure (Smith 1931: 36, figure 27). The CAS B. xanthozonus specimen had the black body band below the second dorsal fin split into two, with a dusky area between, as is shown in Smith's figure. The position of other markings agree with Smith's figure, as do the rather high-set small eyes. Smith's specimen had a low lateral scale count (26), as does the Kabili River specimen (27) (Braehygobius :mnthozonus has a lateral scale count range of 36-46, according to Miller (1989». Smith's specimen had nine dorsal rays (given as 1,8, as Smith did not observe that the first ray was segmented, an error which Miller also made in his diagnoses of Braehygobius and Hypogymnogobius), and so does the holotype of Bleeker's Gobius :mnthozona. The Kabili River specimen has only seven soft dorsal and anal fin rays and lower counts of pectoral rays and all scale counts. A ZMH specimen has eight anal rays. The species appears to be genuinely rare, and insufficient specimens (seven in all) are available to determine the variability of counts. [Note in press: a 30 mm SL female has recently been collected in Brunei]. Caeeogobius Berti and Ercolini, 1991 Caeeogobius Berti and Ercolini, 1991 (type species: Caeeogobius eryptophthalmus Berti and Ercolini, 1991: 130, Philippines, by original designation and monotypy). Diagnosis (Partly compiled from original description). Moderately sized gobiid with second dorsal rays 1,6-7; anal rays 1,5-7; pectoral rays 15-17; 17 segmented caudal rays, in 9/8 pattern; lateral scales 28-29; predorsal scales 16-17; TRB 7-9; circumpeduncular scales 13 (one specimen examined); headpores absent; head papillae pattern longitudinal, rows composed of small, evenly sized papillae, papillae in rows a, e, cp and p small and closely spaced, row e broken under eye; three s rows present, of three or more papillae; no fine villi on dorsal surface of head (in available specimen); eye greatly reduced, not visible externally (embedded, reduced to pigment patches below skin, visible only after clearing); gill opening wide, extending well under preopercle; pectoral girdle smooth; mouth large, terminal; lips smooth; teeth small, pointed, evenly sized, arranged in five to six rows; no 56 H.K. Larson enlarged teeth in female paratype; tongue tip broad, concave to rather bilobed; snout with pronounced knob in centre formed by ascending premaxillary process; anterior nostril in tube, oriented forward over upper lip and posterior nostril placed close behind anterior; body scales thin and cycloid anteriorly, apparently ctenoid posteriorly; opercle naked or with few scales; breast, belly and pectoral base with cycloid scales; pigmentation absent; female genital papilla short, rounded and bulbous. Pterygiophore formula 3-12210; two epurals; 6/6 procurrent fin rays; vertebrae 26, 10 precaudal and 16 caudal. Remarks The monotypic Caecogobius cryptophthalmus was described from four specimens (holotype MSNVR 1262, 76.5 mm SL; paratypes MSNVR 1262a, 57.5 mm; MSNVR 1262b, 42.5 mm; ZSM 27189, 58.5 mm) collected from the deep Calbiga karst cave system on Samar Island, the Philippines. It is the only known blind cave-dwelling gobionelline. It is one of the few troglodytic gobioids known from the IndoPacific region (e.g. the butines Milyeringa veritas Whitley in north-western Australia, Oxyeleotris caeca AlIen from Papua New Guinea, an undescribed Bostrychus from Indonesia, and an undescribed eleotridine, Eleotris, from Guam). Most gobiids have relatively large eyes, with eye loss or extreme eye reduction being rare among gobiines (for example, Austrolethops wardi Whitley), although common in the mud-burrowing amblyopines. Berti and Ercolini suggested possible affinities with Glossogobius and Mugilogobius, based on sensory papillae and other characters. X-ray examination of a paratype (ZSM 27189) showed that the fish had the dorsal pterygiophore formula of 3-12210 and two epurals, while external examination showed that the species had other characters similar to Mugilogobius species. These include similarities in head papillae pattern (Figure 45), in lacking headpores, and having similar scalation and general physiognomy, although the eyes are almost absent and pigment is totally lacking (Figure 46). Glossogobius differs in possessing headpores, a different dorsal pterygiophore pattern (3-22110), a different sensory papillae pattern (e.g. cheek rows a to d continuous, row os modally in contact with row at dorsally; see Akihito et al., 1988) and in having only one epural. From the characters given above, Caecogobius appears to be close to Tamanka siitensis and the two may share a common ancestor. Caecogobius was not included in the phylogenetic analyses or further discussion in this paper, due to insufficient data. Caecogobius cryptophthalmus Berti and Ercolini, 1991 Figures 45-46 .":., . .. , ........ Figure 45 Caecogobius cryptophthalmus papillae. Paratype, CMK 7249, 5amar, Philippines. Scale bar = 1 mm. Caecogobius cryptophthalmus Berti and Ercolini, 1991: 129-138 (Cabilga Cave System, Samar Island, Philippines). Material Examined Paratype ZSM 27189, 1(58.5), Luzon Cave, Calbiga Karst, near Catbalogan, Samar Island, Philippines, Verona Expedition Samar, January-February 1987. Calamiana Herre, 1945 Calamiana Herre, 1945d (type species: Calamiana magnoris Herre, 1945d: 80, Calamianes, Philippines, by original designation and monotypy). Gnathogobius Smith, 1945 (type species: Gnat/lOgobius aliceae Smith, 1945: 523, figure 104, Bangkok, Thailand, by monotypy). original designation and Diagnosis Distinguished by combination of characters. Second dorsal rays 1,6-9; anal rays 1,5-8, rays modally equal in number; pectoral rays 14-18; 16 segmented caudal rays in 9/7 pattern; lateral scales 27-63; TRB 8-22; predorsal scales variable, absent or 2-25; 12-25 circumpeduncular scales; pectoral girdle (anterior edge of cleithrum) smooth or with bony or fleshy smooth flange, no individual fleshy lobes present; gut short, S-bend shape; genital papilla in male usually slender and flattened, and short, rounded and bulbous in female; jaws terminal with lower jaw tip usually anteriormost, jaws usually not enlarged in males (greatly enlarged in one species); inner edges of lips smooth, without fimbriae; profile slightly pointed, snout short, flat; fine villi absent from naked areas of interorbital and snout; sensory papillae in longitudinal pattern; modally two but up to three s papillae rows on snout; papillae rows p, a and c composed of few large widely spaced papillae, other papillae small and close-set; papilla Revision of Mugilogobius Figure 46 57 Caecogobius cryptophtlUllmus, holotype, 76.5 mm SL, MSNVR 1262, Samar, Philippines. From colour slide by Maurice Kottelat. row c broken under eye, rear portion consisting of one papilla; headpores, if present, reduced (no rear part of oculoscapular canal, no preopercular pores, no nasal pores), two species with, and three species without, headpores. Pterygiophore formula 3-12210, modally 26 vertebrae (10+16), two epurals; two anal pterygiophores before first haemal spine, anterior tip of preoperculum rounded (blunt in adult males of one species), metapterygoid forming distinct low to moderate dorsally expanded bridge to quadrate, pterygoid shorter than palatine, palatine either reaching or falling short of quadrate, fifth ceratobranchial narrow and triangular, with distinct flange ventrally, neural spines on first few vertebrae slender, tips not bifid or broadened except in one species, five to eight rakers on first gill arch ossified, scapula unossified. Remarks Calamiana is most similar in appearance to Mugilogobius and Pseudogobius, and is distinguished by a combination of characters. Calamiana differs from Mugilogobius in that the mouth is terminal (versus subterminal), not enlarged in most species (versus enlarged, often greatly so, in males); the cheek rows a and b are composed of few widely spaced, large papillae (versus all cheek papillae small and evenly sized); the neural spines on the first few vertebrae are usually slender and pointed (versus bifid or broadened); modally two (versus three) s rows of papillae present on snout; and two species have headpores (versus headpores always absent). Additionally, fine fleshy villi are absent from the unsealed surface of the head in Calamiana, but are present in Mugilogobius. Calamiana differs from Pseudogobius in that it has a short, 5-bend shaped gut (versus corkscrew-spiral coiled gut); has a terminal mouth with short flat snout (versus subterminal mouth and inflated snout); an absence of headpores in one species (versus headpores always present); has a low metapterygoid, with bridge overlapping quadrate (versus no bridge, metapterygoid short and often expanded dorsally); and has fleshy upper and lower lips (versus thin lips, especially the lower, which is often a reduced fold of skin). Calamiana is similar to Pseudogobius, however, in that at least one species in each genus has compressed, specialised teeth in the upper jaw (a single row present only in Calamiana vanegata). Calamiana shares some similarity with Eugnathogobius microps Smith, 1931. The two share the characters of headpores absent; sensory papillae in cheek rows c and cp being large and widely spaced, with papillae in rows b and d small and close-set; papilla row p composed of widely spaced papillae, and fine villi absent from the dorsal surface of the head. Calamiana differs from Eugnathogobius in that it has 16 segmented caudal rays (versus 17), has jaws enlarged in males in one species only (versus jaws are greatly enlarged in all males and relatively large in females), has a low to moderate metapterygoid (versus broad (deep) metapterygoid) with a process or bridge overlapping the quadrate (versus no process reaching the quadrate), and has two s rows of papillae on the snout (versus three rows). Further work is required on the species and relationships of these two genera. Mugilogobius polylepis Wu and Ni, 1985, probably belongs in Calamiana. The species was described from three specimens: the 34 mm SL holotype (Shanghai Fisheries College S-OOOl), and two paratypes 22-23 mm SL (Shanghai Fisheries College S-0002, 5-0003). The paratypes now appear to be lost in the postal system (Wu Han-ling, personal communication). The holotype was not available for study. My initial impression from Wu and Ni's illustration and the English abstract given in the original description was that Mugilogobius polylepis greatly resembled an undescribed Calamiana from northern Australia in colour pattern but had VI first dorsal spines (versus IV-VI, modally V) and more predorsal scales (25-34 versus 0-16). There are at least three undescribed species (Larson 1999a). PRELIMINARY LIST OF VALID SPECIES Calamiana illata Larson, 1999 Figure 47 Calamiana illota Larson, 1999a: 260-265 (Sungei Buloh mangroves, Singapore). I III OD 80' 60' ｬＩｾ f '. v(;) [I 140' 160' :" 30' ;; RYUKYU IS. • 160' 180' 30' ｾ '. , . " ｾ LACCADlV£ IS•• : : la' Maldlve. ｾＢｾＭＬ 20' • :l MARlANA IS. ,' .. ' .. ' ". '::. .' • .': MARSHAU 1$. CAROUN£ IS. ;/ 4,.' ｾ＼ｲＧＮ KaOfOe '::, ｾ /. o' la' "': ..... ＮＺｾＧ c:::::::fJ ' ｾＬ 'I '." ,'to la' '. " KIRlllAn ,0 ,-. '. , ,.: n. sp 1 o kabilia .. mindora n. sp 2 o n, sp 3 variegatus • ' ｆｉｊｾ .. la' .. ', SAMOA L:\.', . '. :a. ,', '. l\:\. •• ｾ LOYALlY IS• * "* 30' '. . o· Mauritius '20' 0' ., SOLOMON IS. "1· '''I) 'Q, ' '''''o,f . 20' 30' TASMANSEA 60' 80' lOO' 120' 160' 180' 160' :I: ?:: Figure 47 Distribution of species of the genus Calamiana (n.b. n. sp. 1 = illata). rU1 o ::I Revision of Mugilogobius Figure 48 59 Calamiana kabilia. Holotype of Vaimosa kabilia Herre, 36.5 mm SL, CAS 32978, Kabili River, North Borneo. Material Examined Holotype ZRC 39268, 37 mm SL female, Sungei Buloh mangroves, Singapore, K. Lim, 30 January 1992. Calamiana kabilia (Herre, 1940) Figures 17,47-52 ?Glossogobius mas Hora, 1923: 742-743, figure 23 (Chilka Lake: off Samal Island, Rambha Bay, off Barkul). Vaimosa kabilia Herre, 1940a: 19, plate 14 (Kabili Paratypes Singapore: NTM S.14235-002, 7(22.5-34), same data as holotype; NTM S.13959-011, 4(19-32.5), Sungei Buloh mangroves, H. Larson and K. Lim, 4 January 1994; AMS 1.37147-001, 1(38), Sungei Pandan, D. Hoese, P. Ng and K. Lim, 22 December 1993; ZRC 39269, 3(37-45), tidal creek near Woodlands, P. Ng and K. Lim, 30 January 1992. Thailand: CAS 76093, 3(22-31.5), SE bay on Goh Mak Island, SW of Trat Bay, Gulf of Thailand, H.A Fehlmann, 30 October 1957; CAS 206724, 3(28-35), same data as preceding; NTM S.13953-012, 1(40), mangrove creek, Klong Bang Sai, Phuket, H. Larson, D. Hoese and party, 8 October 1993. Brunei: NTM S.13053-OO2, 1(24), Nypa forest, Pulau Berambang, S. Choy, 26 September 1990. Philippines: USNM 316045, 1(27), fish-market at Sorsogon, Sorsogon Bay, Bikol region, Luzon, T. Roberts, 29-30 April 1976; CAS 38650, 1(36.5), Capiz, Panay, AW. Herre, 3 August 1940. River, north Borneo); Koumans 1953: 388. Calamiana magnoris Herre, 1945d: 80 (Calamianes, Philippines); Herre 1953b: 732; Roberts 1989: 168. Gnathogobius aliceae Smith, 1945: 523-524, figure 104 (Bangkok); Suvatti 1981: 203; Kottelat 1989a: 19. Vaimosa rambaiae (in part) Smith, 1945: 538-540 (Bangkok). Calamiana kabilia: Roberts 1989: 168; Kottelat et al., 1993: 146. Mugilogobius kabilia: Kottelat et al., 1993: 146. Calamiana aliceae: Watson and Horsthemke 1995: 9192. Material Examined Holotype ofVaimosa kabilia CAS 32978, 1(36.5), Kabili River, British North Borneo, Sabah, A.W. Herre, 1936-37 Oriental Expedition, January 1937. Holotype ofGnathogobius aliceae 1(38), central canal, Bangkok, Thailand, H.M. Smith, 2 May 1931. Paratype ofGnathogobius aliceae 1(31), same data as preceding. Paratypes ofVaimosa rambaiae ex USNM 119647, 2 (27-29), from shallow slough behind Department of Fisheries, Bangkok, N. Pongse on 28 May 1931, preserved 2 December 1931. Holotype of Calamiana magnoris Figure 49 Jaws and suspensorium of Calamiana kabilia, male, ex NTM S.14302-001, Sarawak. CAS 39881, 1(31), Busuanga, Palawan Province, Philippines, AW. Herre, 1 July 1940. H.K. Larson 60 Other Material Malaysia: FMNH 51667, 1(15), East Coast Residency, Sungai Gana, tributary of Little Kretam River, just above Nypa belt, Kinabatangan, North Borneo, Sabah, R.F. Inger, 12 May 1950; NTM 5.14302-001,5(20.5-28), Kampong Pangkang Kuap, Sarawak, B.L. Lirn, 1 January 1969. Thailand: CMK 4789,2(45-47), aquarium specimens exported by K. Derwanz, 1985; NIFI uncatalogued, 5(26-34), Bangpakong River, Nongnuch. Sri Lanka: SMF uncatalogued, freshwater, A. Heyrner, December 1987. No Definite Locality: AMS 1.35823-001, 1(36), Singapore: probable aquarium specimen, I. Benoit, Sorbonne. Calamiana mindora (Herre, 1945) Figures 47, 53-54 B Vaimosa mindora Herre, 1945a: 13 (Hacienda Waterous, Mangarin, Mindoro, Philippines). Vaimosa zebrinus Herre, 1950: 74 (Layia, Batangas Province, Philippines). Vaimosa zebrina: Herre 1953b: 769. Calamiana mindora: Larson 1999a: 266. Figure 50 Calamiana knbilia papillae pattern. Holotype of Vaimosa knbilia Herre (CAS 32978). A, lateral view; B, ventral view of chin area (not to scale). Scale bar = 1 mm. Material Examined Holotype ofVaimosa mindora CAS 36826, 1(21.5), Hacienda Waterous, Mangarin, Mindoro, Philippines, A.W. Herre, 21 July 1940. Figure 51 Calamiana knbilia male, 45 mm SL, CMK 4789, Thailand. Figure 52 Calamiana knbilia female, 26 mm SL, NIFI unregistered, Bangpakong River, Thailand. Revision of Mugilogobius Figure 53 Jaws and suspensorium of Calamiana mindora, male, ex ROM 53371, Negros Oriental, Philippines. Scale bar = 1 mm. Holotype ofVaimosa zebrinus USNM 202515, 1(24.5), brackish estuary, Layia, Batangas Province, Philippines, A.W. Herre, 30 June 1948. Paratypes ofVaimosa zebrinus USNM 202572, 3(22.5-23), same data as holotype. 61 survey, 7 November 1972; AMS 1.19580-002, 4(1826), Serpentine Creek, Moreton Bay, CSIRO prawn survey, 7 November 1972; WAM P.28816-o01, 3 (1822), Moreton Bay, V. Wadley and P. Young, 7 November 1972; AM5 1.23262-004, 1(20), The Esplanade, Cairns, D. Hoese and D. Rennis, 2 October 1982. Northern Territory: NTM S.14236001, 19.5 mm SL male, among mud and mangrove roots on bank of tidal creek, Reichardt Creek, Darwin Harbour, R. Hanley, 9 February 1993; NTM 5.11933-001, 1(19.5), mud surface, mouth of Adelaide River, R. Hanley, 23 May 1985; NTM 5.10419-007, 2(14-19), tiny puddle on mud, East Arm of Darwin Harbour, near Elizabeth River mouth, H. Larson, 29 March 1982; NTM S.14075001, 1(17), on mud surface, Nayarnpi Creek, Roper River, R. Hanley and L. Banks, 7 September 1994; A /-" . . , \, ' " t'(O,\'. (95. -0 . '.' " ｾＡＺＬＧＮＭｾ • • . , •• b t. '. ＺＬｾ｜ＮＭＧｾｾＮ ..'...1/) \ .. / ;. I ｾ 0...../ I . ｾ (.-J .... Other Material Philippines: USNM 268602, 5(18.5-23.5), tidal lagoon, NE side of Siquijor Island, Negros Oriental, L. Knapp and party, 14 May 1979; USNM 244050, 2(18-18.5), South Bais Bay, Negros Oriental, K. Carpenter and party, 15 May 1978; ROM 53371, 20(14-25), mangrove at Banlas Point, Daco Island, Negros Oriental, R. Winterbottom and party, 21 May 1987; ROM 53370, 1(22), same locality as preceding, 20 May 1987. Thailand: CMK 5366, 6(19.5-26), estuary at Ao Kammala, Phuket, M. Kottelat, 23 April 1985. Indonesia: U5NM 296896, 1(17), near 5amei Island, Irian Jaya, B. Collette, 4 July 1979; USNM 297054, 14(8.5-18), Marchesa Bay, Batanta Island, Irian Jaya, B. Collette, 2 July 1979. Australia: Queensland: AM5 1.21259-008, 14(10.526.5), S of Cape Tribulation, D. Hoese, 11 August 1979; QM 1.21855, 1(26.5), brackish creek, Holloway's Beach, R. Johnson, 23 November 1985; AM5 1.21273-013, 2(12.5-18.5), mangrove in Esk River, 5 of Cooktown, D. Hoese, 20 September 1979. Fiji: ROM 45998, 1(26), mangrove swamp S of 5uva, Viti Levu, A. Emery and party, 5 March 1983; ROM 45847, 1(23), mangrove creek W of Lami, Viti Levu, R. Winterbottom and J. McKinnon, 21 April 1984. Calamiana sp. novo 2 Figure 47 Material Examined Australia: Queensland: QM 1.23883, 2(22.5-24), in subsurface mangrove mud, Serpentine Creek, Moreton Bay, 5 August 1972; QM I.13374, 3(17.521), Serpentine Creek, Moreton Bay, CSIRO prawn ｾ Figure 54 \ Calamiana mindora, sensory papillae. A, lateral view; B, dorsal view. ROM 53370, Negros Oriental, Philippines. Scale bar mm. =1 H.K. Larson 62 NTM 5.14074-001, 1(21.5), on mud surface, Nayarnpi Creek, Roper River, R. Hanley and L. Banks, 8 September 1994; NTM 5.14080-001, 1(20), on mud surface, Roper River, R. Hanley and L. Banks, 10 September 1994; NTM 5.11936-001, 2(1821), in mud, Rhizophora-lined channel, Uttle Lucky Creek, McArthur River, R. Hanley, 5 August 1985. Western Australia: AM5 1.25521-007, 1(22.5), Crab Creek, Broome, 0-2 m, D. Hoese and D. Rennis, 20 September 1985; AM5 1.25523-003, 25(13.5-23), King Sound, NE of Derby, 0-1 m, D. Hoese and D. Rennis, 22 September 1985. and party, 8 December 1993; KUMF unregistered, 1(29.5), Narathinat Province, P. 5irimontaporn, 15 January 1984. Indonesia: CMK 7263, 2(20.5-24.5), Takjong Mayong, Padang Island, Riau, M. Kottelat, 12 February 1991. Chlamydogobius Whitley, 1930 Chlamydogobius Whitley 1930: 122, central Australia (type species Gobius eremius Zietz, 1896, by original designation and monotypy). Chalamydogobius: Lake 1971: 44 (lapsus). Calamiana sp. novo 3 Figure 47 Material Examined Australia: Northern Territory: NTM 5.14287-001, 22 mm 5L male, mouth of East Alligator River, Australia, T. Davis, April 1979; AM5 1.32051-032, 12(12.5-22), same data as previous; NTM 5.14287002, 2(20-20.5), same data as previous. Calamiana variegata (Peters, 1868) Figure 47 Apocryptes variegatus Peters, 1868: 267 (Singapore). Tamanka ubinensis Herre in Herre and Myers, 1937: 41, plate 3 (Pulau Ubin, between Singapore and Johore); Koumans 1940: 153; Koumans 1953: 157. Gobiopterus variegatus: Koumans 1953: 244-245. Calamiana variegata: Larson 1999a: 273. Material Examined Lectotype of Apocryptes variegatus 2MB 6749, 1(33.5), Singapore, F. Jagor. Paralectotypes of Apocryptes variegatus 2MB 32775, 2(24-32.5), same data as lectotype. Holotype ofTamanka ubinensis CAS 30964, 1(30), Pulau Ubin, Singapore, A.W. Herre, March 1934. Other Material Singapore: NTM 5.13959-010, 6(19.5-28), Sungei Buloh Bird Reserve, H. Larson and K. Urn, 4 October 1993; ZRC 39270, 1(33), Sungei Buloh mangroves, K. Urn, 30 January 1992; CMK 8311, 29(23-37), Kranji mangrove, M. Kottelat and D. Murphy, 8 April 1992. Thailand: URM P.13341, 2(29-31.5), Kung Kraben Bay, H. Senou, C. Vidthayanon, 13 December 1983; URM P.13842, 1(33), Bangpoo, near Paknam, H. Senou, C. Vidthayanon, 27 January 1984; NTM 5.13953-011, 4(25-30.5), mangrove at Klong Bang Sai, H. Larson Diagnosis Genus distinguished by combination of characters: second dorsal fin rays 1,6-8; anal rays 1,5-8; usually one more soft dorsal than anal ray; 11-15 pectoral rays; segmented caudal rays 15-18, modally 16; 32-57 lateral scales; 11-19 TRB; predorsal scales 0-22; 15-25 circumpeduncular scales; fine villi present on naked dorsal surface of head; gill-opening restricted to pectoral base; pectoral girdle smooth, without bony flange, or fleshy bumps or flaps; jaws not much longer in males compared with females; headpores absent; sensory papillae all small and evenly sized (Figure 55), broken row c on cheek below eye much reduced to absent, s rows on snout usually consisting of one (rarely two) papilla each, only two s rows present in most species, usually only one f row mental papilla on each side behind symphysis; genital papilla in male moderate to long, flattened, with blunt to rounded tip, female papilla rounded, bulbous to conical; long gut with at least two full loops in the intestine; very dark-pigmented peritoneum. Pterygiophore formula 3-12210; epural single, sometimes epural partly split; vertebrae 27-29, 1011 + 16-18; neural spines of first three vertebrae often stout, broadened or bifid at tip, usually second and third spines bifid or broad-tipped; modally two (occasionally one) anal fin pterygiophores before haemal spine of first caudal vertebra; palatine broad anteriorly, becoming quite slender ventrally, not quite reaching to quadrate and sometimes falling well short; pterygoid short, broad-based; metapterygoid broad, not extending forward toward quadrate, with pointed, broad-based flange or spur extending anterodorsally (Figure 56); quadrate rather forked, with lower limb slender; symplectic without spur or process extending toward preopercle; preopercle narrow, without groove posteriorly; scapula unossified. Remarks The taxonomic status of the genus and its possible relationships were first discussed by Miller (1987), who considered it to be a monotypic valid genus most closely related to Mugilogobius. Larson (1995) Revision of Mugilogobius reviewed the genus and described five new species. Chlamydogobius is shown to be most closely related to Mugilogobius in the cladistic analyses. The two genera can be distinguished by the following characters: Mugilogobius: s papillae on snout in at least three rows of two or more papillae, the first of which is usually the longest and runs just above upper lip fold (few species with middle row represented by only one to two papillae); pectoral rays 13-20; intestine simple, with one "S-bend" and no full loops; gill opening to pectoral base or further, usually with fleshy knobs or ridge along pectoral girdle; two epurals; metapterygoid forming distinct bridge to quadrate; 26-27 vertebrae, usually 26 (10, rarely 11, precaudal and 16--17 caudal); males often with distinctly enlarged mouths. Chlamydogobius: s papillae on snout usually in two rows, rarely three (each row consists of only one or two papillae), first row just above upper lip absent (Figure 56); pectoral rays 11-14; intestine long and coiled into three loops separate from stomach; gill opening restricted to pectoral base, pectoral girdle smooth; one epural; metapterygoid not forming bridge to quadrate; 27-29 vertebrae, usually 28 (IQ-11 precaudal and 16--18 caudal); males with mouths not much larger than those of females. 63 B Figure 56 LIST OF VALID SPECIES Cltlamydogobius eremius (Zietz, 1896) Figures 55-58 Gobius eremius Zietz, 1896: 180, plate 16, figure 5 (Central Australia); McCulloch 1917: 183-184, plate 31, fig.l; McCulloch and Ogilby 1919: 257; McCulloch 1929: 372. Chlamydogobius eremius: Whitley 1930: 122; Koumans 1931: 159-160; Whitley 1964: 123; Jaws and suspensoria of: A, Chlamydogobius eremi us, male, AMS 1.24493-001, Finniss Springs, South Australia; B, Chlamydogobius ranunculus, female, ex NTM 5.11509-007, Leanyer Swamp, Northern Territory. Scale bar = 1 mm. Glover 1971: 1-147; Glover and Inglis 1971: 5, figure 6; Glover 1973: 8--10; Scott et al., 1974: 271272; Glover and Sim 1978: 38; Lake 1978: 73, 153; Glover 1982: 242-244; Thompson 1983: 17-20; Merrick and Schmida 1984: 309-310; Miller 1987: 687-705; Glover 1989: 90-91; AlIen 1989: 203204; Horsthemke 1989: 288; Larson and Martin 1990: 62-63 (in part); Glover 1990a: 189, 191, figure Id; Glover 1990b: 75; Larson 1995: 22-28. Chalamydogobius eremius: Lake 1971: 44 (lapsus). Material Examined . cp d'" Lectotype of Gobius eremius SAMA F.525, 1(51), Coward Springs, South Australia, Australia, bore by railway, 4 May 1894, Horn Expedition. Paralectotypes of Gobius eremius SAMA F.7674, 5(13-43), same data as lectotype. Figure 55 Chlamydogobills eremius papillae. SAMA F.3509, Coward Springs Bore, SA. Scale bar = 1 mm. Other Material Australia: South Australia: AMS 1.24673-001, 24(10-34.5), including two cleared and stained, 6 km ESE of Coward Springs, D.F. Hoese and S. Reader, 64 Figure 57 H.K. Larson Chlamydogobius eremius. Lectotype of Gobius eremius Zietz, 51 mm SL, SAMA F.525, Coward Springs Bore, SA. Reproduced with permission from Larson (1995). 23 August 1984; SAMA F.3509, 10(30-44), Coward Springs Bore, J. Glover, 2 July 1968; SAMA F.4204, 6(27-45.5), dam near Coward Springs, J. Barry, April 1976; SAMA F.3999, 13(32.5-41), Margaret River, T. Sim, 3 March 1975; AMS 1.26285-001, 4(14.5-18.5), spring near Well Spring, Freeling Springs, W. Ponder, 2 June 1988; AMS 1.24493-001, 4(28-41), including one cleared and stained, Finniss Springs, W. Ponder, R. Hershler, 28 January 1984; AMS 1.27116-001, 10(13-29), Ockenden Spring and Bore, W. Ponder, 1 June 1983. 120' Chlamydogobius gloveri Larson, 1995 Figure 58 Chlamydogobius eremius: Glover 1971: 77,99, table I, Appendix A; Ivantsoff and Glover 1974: 95. Chlamydogobius n. sp.: Harris 1987: 9; Jackson 1993: 24. Chlamydogobius sp. novo (Dalhousie goby): Glover 1989: 90, figure 31.1g; Glover 1990a: 191. 130' lOO' ISO' 10' ｓＮａｾ • *it r-----r-, ｾ Jo 13C' Figure 58 ｬｾｑＧ Distribution of the genus Chlamydogobius. Stars = C. eremius, dark squares = C. gloveri, downward triangles = C. japalpa, upward triangles = C. micropterus, circles = C. ranunculus, open squares = C. squamigenus. Reproduced with permission from Larson (1995). Revision of Mugilogobius ChlamydogobillS sp.: AlIen 1989: 224; Kodric-Brown and Brown 1993: 1849; Morton et al., 1995: 30,95. Clllamydogobius sp. 2 (undescribed species): Glover 1990b: 75. Chlal1lydogobius gloveri Larson, 1995: 28-32 (Dalhousie Springs, South Australia). Material Examined Holotype SAMA F.3463, 1(30), Dalhousie Springs, South Australia, Australia, J. Glover, August 1968. Pa ra types Australia: South Australia: SAMA F.5425, 12(1327.5), Spring "G"ab, J. Glover and T. Sim, 4 June 1985; SAMA F.7675, 3(33-36), same data as holotype; SAMA F.5417, 4(18-24), Spring B4, J. Glover, T. Sim, 3 June 1985; AMS 1.27118-001, 1(32), pool on top of mound, low mound springs, W. Ponder, 29 May 1983; AMS 1.25881-001, 4(7-16), Cold Spring Cc1B, outflow of medium active spring, W. Ponder, D. Winn, 6 June 1985; AMS 1.25879-001, 1(20), warm pool in swamp, W. Ponder, D. Winn, 13 June 1985; AMS 1.25880-001, 1(18), warm pool 20 m upstream from main pool, W. Ponder, D. Winn, 14 June 1985. Chlamydogobius japalpa Larson, 1995 Figure 58 Chlal1lydogobius erel1lius: Larson and Martin 1990: 62-63 (in part). Chlamydogobius japalpa Larson, 1995: 32-35 (Ormiston Creek, Finke River, Northern Territory). Material Examined Holotype NTM S.11436-007, 1(44), Ormiston Creek, at junction of Pioneer Creek, Finke River, Northern Territory, Australia, 23°40'S 132°42'E, H. Larson and P. Homer, 15 September 1984. Pa ra types Australia: Northern Territory: Finke River: NTM S.11436-009, 41(15-44), same data as holotype; NTM S.11439-006, 10(15-38.5), including one cleared and stained, off Palm Valley Road just N of Park border, H. Larson, 17 September 1984; AMS 1.35467-001, 8(15-28), same data as preceding; NTM S.11437-005, 25(12-45), just above Glen Helen N of main road crossing, H. Larson and P. Homer, 16 September 1984; NTM S.11639-001, 1(40), Hermannsburg Rockhole, R. Moses, 21 June 1983; 5AMA F.7677, 3(25-34), same data as preceding; NTM 5.11628-001, 1(29.5), Boggy Hole, R. Moses, 23 JWle 1983; NTM 65 S.11650-002, 2(25.5-27), Running Waters, R. Moses, 14 July 1983; NTM S.11632-001, 1(37), Palm Valley, 25 June 1983. Chlamydogobius micropterus Larson, 1995 Figure 58 Chlal1lydogobills n. sp.: Harris 1987: 8; Jackson 1993: 23. Chlamydogobius sp. nov: Glover 1989: 98, 110. Chlal1lydogobius sp. A: Wager and Jackson 1993: 85. Chlamydogobius sp.: Morton et al., 1995: 53,119. Chlamydogobills micropterlls Larson, 1995: 35-38 (Elizabeth Springs, Springvale Station, Queensland). Material Examined Holotype QM 1.25096, 1(22.5), Elizabeth Springs, Springvale Station, Queensland, Australia, J. Covacevich, P. Couper, 27 April 1988. Para types Australia: Queensland: AMS 1.25261-001, 3(8.520.5), Springvale, W. Ponder, P. Colman, 10 September 1984; AMS 1.25256-001, 6(10.5-21.5), same data as preceding; QM 1.29552, 27(9.5-23), same locality data as holotype. Chlamydogobius ranunculus Larson, 1995 Figures 56, 58 Mugilogobius sp. 9: Gee and Gee 1991: 19,21-26. Clzlamydogobius ranunculus Larson, 1995: 38-42 (Beatrice Lagoon, Adelaide River, Northern Territory). Material Examined Holotype NTM S.11427-001, 1(27.5), partly dried-up buffalo wallow, Beatrice Lagoon, Adelaide River drainage, Northern Territory, Australia, 12°37'S 131°21'E, H. Larson, 20 May 1984. Para types Australia: Northern Territory: NTM S.11427-002, 7(19-27.5), same locality data as holotype; NTM S.11509-007, 118(4-31), bombholes at Leanyer Swamp, NT. Fisheries, 9 October 1984; AMS 1.32051-022, 5(20.5-31), Alligator River mouth, T. Davis, April 1979; ex AMS 1.32051-020, 2(23-27), cleared and stained, Alligator River, T. Davis, April 1979. Queensland: AMS 1.22959-001, 5(22-25.5), lagoon behind wharf, Townsville, J. Gee, 16 June 1981; QM 1.19003, 1(31), Norman River near L_ _ H.K. Larson 66 Karumba, D.J. Russell, 11 June 1981; QM 1.19005, 1(27), same data as preceding; AMS 1.20928-001, 27(9.5-17.5), Smith Point, Prince of Wales Island, Torres Strait, D.E Hoese, 1979. Chlamydogobius squamigenus Larson, 1995 Figure 58 Chlamydogobius n. sp.: Jackson 1993: 23. Chlamydogobius sp.: Tappin 1995: 19. Chlamydogobius squamigenus Larson, 1995: 42-46 ("Western" spring, NE of Edgbaston Homestead, Queensland). Material Examined Holotype SAMA F.6595, 1(34), "Western" spring, approximately 2.4 km NE of Edgbaston Station Homestead, Queensland, Australia, J. Glover, T. Sim and T. Scott, 7 May 1989. Paratypes Australia: Queensland: SAMA F.7676, 8(21-34.5), same locality data as holotype; SAMA F.6738, 1(34), small mound artesian spring approximately 3.3 km SSE of Edgbaston Homestead, W. Zeidler, 4 May 1988; SAMA F.7184, 35 (14-39), "Crossmoor Flowing Bore", Crossmoor Station, on LongreachMuttaburra Road, T. Sim and P. Num, 3 June 1993; SAMA F.7206, 1(35), Crossmoor Station, Bore Number 1652, south arm of drain, T. Sim and P. Num, 5 June 1993. Eugnathogobius Smith, 1931 Eugnathogobius Smith 1931 (type species: Eugnathogobius microps Smith, 1931: 37, figure 18, lower Bangpakong River, central Siam, by original designation and monotypy). Eugathogobius Koumans, 1931: 69 (error for Eugnathogobius). Pseudogobiopsis Koumans, 1935 (type species: Gobiopsis oligactis Bleeker, 1875: 113, Amboina, by original designation). Diagnosis Distinguished by following combination of characters. Second dorsal fin rays 1,6-10; anal fin rays 1,5-8; number of second dorsal and anal rays equal except in one species; first dorsal spine often longest and may be elongate; pectoral rays 13-20; 17 (9/8) segmented caudal rays; predorsal scales medium to large, 5-13, extending up to behind eyes; 22-27 scales in lateral series; TRB 6-12; 12-13 circumpeduncular scales; headpores present in three species, absent in two; preopercular pores present in three species; papillae rows a and c consisting of large, widely spaced papillae, as does row p if present (otherwise replaced by headpores), rows b and d always consisting of small close-set papillae, row f of one or two papillae only, two or three s rows on snout of one papilla each; dorsal surface of head without fine villi; gill opening usually extending past pectoral base to under opercle, bony or fleshy flange or ridge, or fleshy knobs and flaps usually present on pectoral girdle; jaws enlarged in adult males, sometimes greatly so; anterior nostril tubular, placed at or just behind preorbital edge; genital papilla flattened and usually elongate in males, rounded and bulbous in females; gut simple, 5-bend shape; colour pattern usually pale with brown midlateral blotches or spots, and black line and I or spots along ventral midline of caudal peduncle. Pterygiophore formula 3-12210; 26 vertebrae, modally 10+16; two epurals; two or three anal fin pterygiophores present before first caudal haemal spine; anterior end of preopercular bone blunt or pointed; ridge and I or posteriorly facing groove present along rear edge of preopercle; neural spines of first few vertebrae slender, pointed; metapterygoid short, relatively small, may be broadened dorsally or with short dorsal process, but not contacting or forming bridge to quadrate; palatine and pterygoid relatively slender and nearly equal in length, palatine not reaching quadrate; quadrate somewhat forked; fifth ceratobranchial stout, triangular, with high flange ventrally; four to nine ossified gill-rakers; scapula usually unossified but tip above foramen may be ossified. Remarks There are five valid species, restricted to fresh water in the Indo-Malay Archipelago (Figure 59), which I recognise so far. Several species of this genus have been described a number of times (e.g. males as one species, females as another). Several museum lots of specimens exist which may represent an additional one or more species, but this cannot be confirmed until more specimens are obtained. The genus as it currently stands may be para- or polyphyletic (Figures 23-28). The problem may not be resolved until revisions of Calamiana, Redigobius and Rhinogobius, its apparent closest relatives, are carried out. These are under way as separate projects outside the scope of this paper (Calamiana and Redigobius by myself and Rhinogobius by 1-5. Chen). PRELIMINARY LIST OF VALID SPECIES Eugnathogobius sp. novo Figure 59 Calamiana sp. nov.: Kottelat and Lim 1995: 247. :xl ｾ <: 0;' I' 140· I 120· 100· 80· 60· :RYUKYU IS ... . - :' o t; (7 .(J o' ::l l " 0- ',' ARABIAN SEA Cl ;r. ::: "/{ 20·' D ｾ SOUTH CHINA 0, I,' LACCADIVE • ", ISLANDS , ' . 10' O<:l o SEA , ( '• • I., , " Maldives " " ［Ｇｾ , • PHILIPPINES ｾ " 'j .. Ｎｾ ,, .. O' * n, Sp. • • SEYCHELLES • microps • oligactis • paludosus T siamensis .- .* . I. - S' .... " Ｎｾ 10· CELEBES SEA ' ＼ｾ O· '" " ,-10; Ｍｾ ........ AA Of" • OC§:> ｾ .. .68 c::::::>Ozj;;'<? 0' " 0 -<..,<::, "" ..: 60· Figure 59 80' Distribution of Ellgnathogobills species, 100' 120' 140' '" '1 H.K. Larson 68 Figure 60 Eugnatlwgobius microps Smith, holotype, 26 mm SL, USNM 90316, Bangpakong River, Thailand. Material Examined Malaysia: Sarawak: ZRC 40279, 34.5 mm SL male, 8.6 km after turnoff to Sungei Cina Matang after entrance to Matang Reserve, near Kuching, RH. Tan, 4 September 1995; ZRC 40280, 6(20.5-27), same data as holotype; NTM S.14299-001, 7(19.529.5), same data as holotype; ZRC 27842-3, 2(14.524.4), 7 km on Kuching-Batu-Kawa Road, M. Kottelat and K Lim, 3 July 1992; ZRC 29082-3, 2(1920.5), 7 km on Kuching-Batu-Kawa Road, M. Kottelat and K Lim, 3 July 1992; ZRC unregistered, 9(18-33), stream I, Ulu Assam, Bako National Park, Kuching, K Lim, 30 June 1994; ZSM\KEW 538, 5(16-24), NW of Bau, Batang Kayan River basin, Sungai Stunggang, C. Kettner, K-E. Witte and R Krumenacher, 24 March 1988; CMK 8401,26(11-27), Sungei Bejit, road from Balai Ringin to Simunjan, M. Kottelat, K Lim and P. Ng, 2 July 1992. Eugnathogobius microps Smith, 1931 Figures 59-63 ?Glossogobius mas Hora, 1923: 742-743, figure 23 (Chilka Lake: off Samal Island, Rambha Bay, off Barkul). Eugnathogobius microps Smith, 1931: 37, figure 18 (Bangpakong River, Siam); Koumans 1931: 6869; Koumans 1940: 129, 200; Smith 1945: 520; Suvatti 1950: 421 (not seen); Suvatti 1981: 202; Kottelat 1989a: 19; Eschmeyer and Bailey 1990: 145. Eugnathogobius macrops: Herre 1940a: 24 (lapsus). Material Examined Holotype of Eugnathogobius microps USNM 90316, 1(26), lower Bangpakong River, central Thailand, H.M. Smith, 1 July 1923. Other Material Thailand: USNM 11951, 1(26), same data as holotype; USNM 119593, 3(20-26.5), near Pitrieu, Bangpakong River, H.M. Smith, 4 June 1928. KUMF unregistered, 2(17.5-17.5) Bang Nara River, Narathiwat Province, D. Tanwilai, 23 December 1983; NTM S.13953-013, 7(18-25.5), small mangrove creek, Klong Bang Sai, Phuket Island, H.K Larson, D.F. Hoese and PMBC staff, 8 December 1993. Other material examined (but not used) Thailand: Two specimens, KUMF 1853, same locality data as USNM 119593 (fish very fragile, heads separated from bodies). Figure 61 Jaws and suspensoria of Eugnathogobius microps, male, ex NTM 5.13953-013, Phuket, Thailand. Scale bar = 1 mm. Figure 62 Eugnathogobius microps papillae pattern. NTM 5.13953-013. Scale bar = 1 mm. Revision of Mugilogobius Figure 63 69 Eugnathogobius microps. 25.5 mm SL, NTM 5.13953-013, Phuket, Thailand. Eugnathogobius oligactis (Bleeker, 1875) Figures 4, 59, 64-Q7 187, figure 14l. Gobiopsis oligactis Bleeker, 1875: 113-114 (Amooina, Indonesia); Koumans 1931: 67; Bleeker 1983: plate 433, figure 4. Glossogobius campbellianus Jordan and Seale, 1908: 542, figure 2 (Buytenzorg, Java). Mugilogobius perakensis: Kottelat 1989a: 19. ?Genus and species undet; Roberts 1993: 44, figure 51. Pseudogobiopsis campbellianus: Kottelat et al., 1993: 150, plate 70 (in part). ?Glossogobius mas Hora, 1923: 742-743, figure 23 (Chilka Lake: off Samal Island, Rambha Bay, off Barkul). Pseudogobius neglectus Koumans, 1931: 102 (nomen nudum; Bleeker museum name); Bleeker 1983: plate 438, figure 13. Pseudogobiopsis neglectus: Kottelat et al., 1993: 150, plate 70. Material Examined Holotype of Glossogobius oligactis RMNH 4459, 1(25.5), Java, Indonesia. S tigmatogobius neglectus Koumans, 1932: 3, 5 (western Java). Pseudogobiopsis oligactis: Koumans 1935: 131-132, figure 4; Fowler 1937: 251; Koumans 1940: 126; Smith 1945: 521-522; Kottelat 1989a: 19; Kottelat et al., 1993: 150, pL70; Lim and Larson 1994: 260; Kottelat and Lim 1995: 247. Vaimosa perakensis Herre, 1940a: 21, plate 16 (Lake above Chenderoh Peninsula). Stigmatogobius isognathus: Mohsin and Ambak 1983: Darn, Perak, Malay Stigmatogobius oligactis: Koumans 1953: 110, 116- Holotype ofGlossogobius campbellianus USNM 61051, 1(26.5), Buitenzorg (= Bogor Botanical Gardens), Java, D.H. Campbell, Indonesia. Paratype ofGlossogobius campbellianus Indonesia: CAS 22498, 1(23), same data as holotype. Holotype ofVaimosa perakensis CAS 32975, 1(25.5), Chanderoh Darn, Perak, Malaysia, A.W. Herre, 9 March 1937. 117, figure 27; Chatterjee 1980: 229,230. Stigmatogobius poicilosoma (in part): Alfred 1966: 47. Figure 64 Jaws and suspensorium of Eugnathogobius oligactis, female, ex CMK 10713, Trat Province, Thailand. Scale bar = 1 mm. Paratypes ofVairnosa perakensis Malaysia: CAS 32977, 17(13.5-28.5), 2 miles N of Sauk, Perak, A.W. Herre, 18 April 1937; ZRC 255, Figure 65 Eugnathogobius oligactis, papillae pattern. 36 mm SL male, CMK 5385, Ranong, Thailand. Scale bar = 1 mm. H.K. Larson 70 . ' : . : / (5).' J'.-I/', • "0 ｾＮＬ . ｾ＿ ) , ' : , -, . cp ＭｾＬＺ Figure 66 I ＭＮＬ｟ｾ ..... .;ｾＮｊｦｴＺＧ ' . ... ｾ ---- - , ｾ '--- , ". J "':,' c ... " " .. -J ｾ ": .. :., ｾ . " " ｾｾＭ Eugnathogobius oligactis, papillae pattern. 32 mm SL female, NTM 5.14239-001, Belalong River, Brunei. Scale bar = 1 mm. 47(12-24.8), lake above Chanderoh Dam, Perak, AW. Herre and M.F. Tweedie, 19 March 1937. Road, Penang, E.R. Alfred, 21 October 1961; FMNH 68465, 3(27.5-29), Sungai Tawan, Sungai Tibas Camp, Kalabakan, Tawau, Sabah, R. Inger, 8 June 1956; NTM S.14240-001, 4(23-26), 7 miles S of Kuching, Kampong Pangkalan Kuap, Sarawak, B.L. Lim, 1 January 1969. Brunei: NTM S.14239-001, 3(32-36), Sungai Belalong, KBSFC, S. Choy, 27 July 1992. Singapore: ZRC 1049i-iv, 4(24-28), MacRitchie Reservoir, S.H. Chung, 4 November 1959. Indonesia: CMK 9621, 6(15-32), clear stream E of Mentok on road from Pangkalpinang, Banka, Sumatra, M. Kottelat et al., 4 March 1993. Eugnathogobiu5 paludo5u5 (Herre, 1940) Figure 59 Ctenogobius paludosus Herre, 1940a: 23, plate 18 (north of Kota Tinggi, Johore, Malay Peninsula). Calamiana sp. undet.: Roberts 1989: 168. Other Material India: CAS 61960, 3(15-20.5), sandy beach at Hampi, Tungabahdra River, Bellary District, Karnataka State, Mysore, T. Roberts, 3 February 1985. Thailand: CMK 5385, 22(17-36), stream on road from Ranong to Kra Buri, Ranong Province, M. Kottelat, 24 April 1985; KUMF uncatalogued, 3(37.5-41), Tha Lad Waterfall, Samui, Surathani Province, C. Vidthayanon, 22 July 1983; NIFI 1230, 3(33.5-41), Chiew Larn Dam, Klong Saeng, Surat Thani Province, J. Karnasutra, 20 March 1983. Cambodia: CMK 4806, 1(35), Stung Tong Hong, 77 km on road from Phnom Penh to Sihanoukville, d'Aubenton, 13 January 1963. Malaysia: CMK 4722, 5(24-30), NE foothills of Gunung Panti, N of Kota Tinggi, Johor, M. Kottelat and party, 1991; ex ZRC 19323-19325, 1(34), Desaru Road, Kota Tinggi, Johor, P. Ng and M. Kottelat, 14 August 1991; ZRC 20457-61, 5(25.5-37.5), Rembia, Malacca, D.S. Johnson, 26 February 1958; ZRC 1651i-iii, 3(32-40.5), Sungei Telok Bahang at 14.5 miles on Telok Bahang Figure 67 Rhinogobius paludosus: Kottelat 1989a: 19. Material Examined Halo type CAS 32998, 1(30.5), 5 miles N of Kota Tinggi, Johore, Malaysia, AW. Herre, May 1937. Other Material Malaysia: ZRC 8411, 1(30), Sungai Sedili, Johore, 22 February 1968; CMK 7384, 10(12.5-29), Sungai Mupor, Johor, M. Kottelat, P.K.L. Ng and K. Lim, 22 January 1991. Indonesia: USNM 230332, 3(18-23), West Kalimantan, Sungai Gentu, near confluence with Kapuas River, T. Roberts, 16 August 1976; CAS 49462, 58(7.5-28), Sungai Mandai Kecil, near confluence with Kapuas River, 18 km WSW of Putissibau, T. Roberts and S. Woerjoatmodj0, 1 August 1976; CMK 9009, 5(13-21), Sumatra, Riau district, Sungai Kalesa, N of Seberida, AJ. Whitten, 22 February 1992. Eugnathogobius oligactis. CMK 10713, 28.2 and 32 mm SL, Khlong Huai Paeng, Trat, Thailand. From colour slide by Maurice Kottelat. Revision of Mugilogobius Eug1lathogobius siamc1lsis (Fowler, 1934) Figure 59 Vailllosa simnensis Fowler, 1934: 157, figure 125 (Silom Canal, Bangkok, Thailand); Fowler 1935: 161; Smith 1945: 538,540-541. Vaimosa mawaia Herre, 1936a: 9, plate 6 (Mawai district, Johore, north of Singapore); Herre and Myers 1937: 40; Fowler 1938: 267; Koumans 1940: 152. Vaimosa jurongensis Herre, 1940a: 18, plate 13 aurong, Singapore); Koumans 1953: 386-387. VainlOsa oratai Herre, 1940a: 20, plate 15 (Brook at Tawau, north Borneo); Koumans 1953: 389. Pseudogobiopsis oligactis: Koumans 1940: 135. Vaimosa singapurensis: Tweedie 1940: 75. Stigmatogobius oligactis: Koumans 1953: 116-117; Suvatti 1981: 204. Stigmatogobius poicilosoma (in part): Alfred 1966: 47. Calamiana siamensis: Hoese in Bbhlke 1984: 110. Pseudogobiopsis jurongensis: Roberts 1989: 169-170. Mugilogobius jurongensis: Kottelat 1989a: 19. Mugilogobius mawaia: Kottelat 1989a: 19. Pseudogobiopsis siamensis: Kottelat 1989a: 19; Tan and Tan 1994: 357; Lim and Larson 1994: 260; Kottelat and Lim 1995: 247. Pseudogobiopsis campbellianus: Kottelat ct al., 1993: 150, plate 70 (in part). Pseudogobius orotai: Kottelat et al., 1993: 150, figure 306. Pseudogobiopsis wuhanlini Zhong and Chen, 1997: 79--81 (Min River, Fujian Province, China). Material Examined Holotype ofVaimosa siamensis ANSP 60025, 1(31.5), Silom Canal, Bangkok, Thailand, RM. de Schauensee, 18 December 1932. Holotype ofVaimosa mawaia CAS 29080, 1(24), Mawai district, Johore, Malaysia, A.W. Bene, 22 March 1934. Figure 68 71 Holotype ofVaimosa oratai CAS 32988, 1(18), Tawau, British North Borneo, Sabah, Malaysia, A.W. Herre, 17 January 1937. Holotype ofVaimosa jurongensis CAS 32982, 1(36), Jurong, Singapore, A.W. Herre, 8 May 1937. Paratypes ofVaimosa jurongensis Singapore: CAS 32983, 22(19.5-34), same data as holotype; BMNH 1938.12.1.215-7, 3(24.5-28.5), same data as holotype. Other Material Thailand: ANSP 87453, 13(23-32.5), Bangkok, R de Schauensee, 1936; ANSP 63126, 2(27.5-28), Bangkok, R. de Schauensee, May 1934. Malaysia: CMK 8485, 11(19-30.5), blackwater stream, 70 km on road to Kuantan-Pekan-Mersing, Pahang, M. Kottelat and party, 24 July 1992; ZRC 19323-19324, 2(33.5-34.5), stream by Desaru Road, Kota Tinggi, Johor, P. Ng and M. Kottelat, 14 August 1991; ZRC 17051-6, 6(23.5-31), freshwater stream on MawaiTanjung Sedili Road, Johore, P. Ng and M. Kottelat, 14 August 1991. Brunei: NTM S.14244-001, 8(2328.5), small stream near Kampong Lempong, Temburong Baru, S. Choy, 9 January 1992. Indonesia: CMK 7304, 24(11.5-21), tributary of Sungei Siak, Riau Province, Sumatra, M. Kottelat, 13 February 1991. 11lccrtac scdis Gobius tigrcllus Nichols, 1951 Figure 68 Gobius tigrcllus Nichols, 1951: 3, figure 2 (Bemhard Camp, Idenburg River, West New Guinea). Ctcnogobius tigrellus: Allen 1996: 19. Material Examined Holotype AMNH 18574, 1(21.5), 75 m altitude, Bemhard Camp, Idenburg River, West New Guinea, W.B. Richardson, April 1939. GobillS tigrelllls Nichols, Idenburg River, West New Guinea [Irian Jaya]. Reproduced with permission from the American Museum of Natural History. H.K. Larson 72 Paratypes AMNH 15096, 9(16-22), same data as holotype. Remarks This species appears to be a gobionelline, having 3-12210 pterygiophore formula, two epurals and one or two pre-anal pterygiophores. It somewhat resembles Eugnathogobius due to fin ray and scale counts; in having a relatively complete oculoscapular canal, including paired anterior interorbital pores, infraorbital pores, and three preopercular pores, and anterior nasal pores absent; and in the remnants of large c or cp papillae across the cheek, contrasting to the small papillae in row d and the preorbital portion of row c. The holotype and nine paratypes, the only known specimens to date, have been dehydrated at some stage and are not in the best condition. Resolution of the status of this species will require additional material. Vaimosa rivalis Herre, 1927 Figure 69 Vaimosa rivalis Herre, 1927: 149, plate 11, figure 1 (Talakop Creek, at foot of Mount Isarog, Calabanga, Camarines Sur Province, Philippines). Remarks The syntypes, BSM 13061 (16 specimens) and BSM 13602 (eight specimens), were all destroyed in WWII. The description was based on four specimens (21-28 mm long), with" ... the account checked by ..." 11 additional specimens (ranging down to 8 mm), and eight cotypes (16-33 mm long) from Hinagianan River, from the same province (Herre 1927). Koumans (1940: 185) gave the Bureau of Science catalogue numbers of the types and stated that they were"A good species of Stigmatogobius". The Figure 69 species could belong to either Redigobius or Eugnathogobius, judging by Herre's description and figure, although Redigobius may be more likely. The drawing shows 1,8 second dorsal fin rays and 1,7 anal fin rays, a combination more usual in Redigobius; Eugnathogobius usually has equal numbers of soft dorsal and anal fin rays. In the description, Herre gives the fin counts as "VI, 1-7 or 8; anal 1,7". From the other information given, it is difficult to be sure what Herre had. Eugnathogobius sp. Pseudogobiopsis sp.: AlIen 1991: 191, plate 15, no. 12. Material Examined Papua New Guinea: WAM P.27849-008, 8(1824.5), Luap Creek, 6 km N of Bewani on Vanimo Road, Bewani Mountains, G. AlIen and J. Patten, 1 November 1982. Remarks Specimens are known so far only from a small freshwater creek in northern Papua New Guinea, close to the border with Irian Jaya. All but one specimen appears to be male. Radiographs and a cleared and stained specimen show that the species has a dorsal pterygiophore formula of 3-21210, one or two epurals and the top of scapula is ossified around the foramen. The status of this species cannot be resolved without further material and clarification of the relationship between Eugnathogobius and Rhinogobius. Hemigobius Bleeker, 1874 Hemigobius Bleeker, 1874 (type species: Gobius melanurus Bleeker, 1849: 31, Java; = Hemigobius bleekeri Koumans, 1953: 191, replacement name for Gobius melanurus Bleeker, by original designation and monotypy). Vaimosa rivalis Herre, Talakop Creek, Mount Isarog, Calamines Sur Province, Philippines. From Herre (1927): plate 11, figure 1. Revision of Mugilogobius 73 Microgobius Koumans, 1931 (listed as synonym of Stigmatogobius). Preoccupied by Microgobius Poey. Sphenen togobius Fowler, 1940 (type species: Sphenentogobius vanderbilti Fowler, 1940: 396, figures 8-11, Sumatra, by original designation and monotypy). Diagnosis Distinguished by following combination of characters. Dorsal rays 1,6-8, modally 1,7; anal rays 1,6-9, modally 1,7; pectoral rays 13-17; 17 segmented caudal rays in 9/8 pattern; some headpores present with no oculoscapular canal over preoperculum, no preopercular pores, no nasal pores, pairs of interorbital pores not connecting across interorbital space; 25-34 lateral scales; circumpeduncular scales strongly modally 12; predorsal scales 7-12, extending close up to behind eyes, margins of anteriormost scales scalloped; preopercle at least partly scaled; interorbital and top of snout with fine villi; papillose flaps or pads present at insertion of first gill arch onto roof of mouth; jaws small, lower jaw symphysis usually raised; thin, folded lower lip; teeth small and flattened in females (at least), usually conical in males; anterior nostril in short tube oriented down and forward over upper lip, preorbital usually curved outward slightly around base of nostril; genital papilla slender, flattened and pointed in males, conical and blunt-tipped in females; intestine very long and tightly coiled. Dorsal pterygiophore formula 3-12210; two epurals, rarely one; one to three pre-anal pterygiophores; neural spine on first vertebra usually short and broad. Palatine and pterygoid short, with broad, T-shaped heads; palatine larger and more robust than pterygoid. Metapterygoid deep, well separated from quadrate, anterior process extends upward and forward, well above quadrate. Mandibular ramus elevated and curved anteriorly in H. mingi, elevated but angled backward in H. hoevenii. Fifth ceratobranchials triangular, very open and lattice-like in structure. Figure 70 Jaws and suspensorium of Hemigobius Izoevenii, male, ex NTM S.1l06S-D02, Leanyer Swamp, Northern Territory. Scale bar = 1 mm. Stigmatogobius hoevenii: Koumans 1953: 125 (in part); Munro 1958: 272; Munro 1967: 499. Microgobius hoevenii: Bleeker 1983: plate 438, figure 17. Mugilogobius obliquifasciata Wu and Ni, 1985: 93-95 (Haikou, Hainan Island, China); Anonymous, 1986: 272-273. Mugilogobius obliquifasciatus: Zhu 1988: figure 162. Hemigobius crassa: Davis 1988: 164. Pseudogobius hoevenii: Murphy 1990: 155. Mugilogobius latifrons: Nguyen 1991: 334-335, figure 143. Hemigobius lwevenii: Kottelat et al., 1993: 146; Larson 1999b: 25. Material Examined Holotype of Gobius hoevenii RMNH 4457,1(32), in river, Sambas, Borneo. Remarks Larson (1999b) reviewed the species of this genus. LIST OF VALID SPECIES Hemigobius hoevenii (Bleeker, 1851) Figures 70-74 Ｇｾ .. " Gobills Izoevenii Bleeker, 1851b: 426-427 (Sambas, in river, Borneo). Vaimosa crassa Herre, 1945e: 403 (brook near Un Long, Hong Kong). Figure 71 Hemigobius hoevel1ii papillae pattern. CA5/5U 38636, Coron, Busuanga, Philippines. Scale bar = 1 mm. 74 H.K. Larson Figure 72 Hemigobius hoevenii, female, 29 mm SL, ZRC 21872-21906, Singapore. Figure 73 Hemigobius hoevenii, male, 31 mm SL, NTM S.13968-008, Singapore. Holotype ofVaimosa crassa 1(35), CAS/SU 39848, Un Long, New Territories, Hong Kong, AW. Herre, 23 February 1941. Paratype ofVaimosa fusca CAS 32987, 25.5 mm SL male, mangrove swamp, Kranji River, Singapore, A Herre, March 1937. Other Material Thailand: URM P.12662, 2(28-28), mangrove swamp at Phuket, H. Senou and V. Chavalit, 2 November 1983; ex URM P.6677, 3(22.5-25.5), Ranong, 9 March 1982. Malaysia: ZSM 27559, 4(2326.5), mangrove at Bamgangan, SW of Sandakan, Sabah, Kettner, Krumenacher and Witte, 13 March 1988. Singapore: NTM S.14235-004, 11(8.5-27), Sungei Buloh mangroves, K. Urn, 30 January 1991. Brunei: NTM S.12812-002, 2(18-24), Kedalayan River, from Nypa leafaxils, R. Hanley and S. Choy, 7 April 1989. Indonesia: BMNH 1935.5.27.28, 1(36), (possibly from Sulawesi), Arnold. Philippines: CAS 38636,30(19-33), Coron, Busuanga, 22-30 June 1940, AW. Herre. Papua New Guinea: WAM P.26751006, 2(21.5-37), Tureture village, Binaturi, G. AlIen, 29 September 1979. Australia: Queensland: AMS 1.23262-001, 8(18-32), The Esplanade, Cairns, D. Hoese and D. Rennis, 2 October 1982; Northern Territory: NTM S.11065-o02, 13(13-21.5), Leanyer Swamp, Darwin, T. Davis, 5 March 1980; Western Australia: AMS 1.25521-009, 1(27.5), Crab Creek, Broome, D. Hoese, D. Rennis, 20 September 1985. Hemigobius melanurus: Koumans 1931: 78; Bleeker 1983: plate 433, figure 9. Gnatholepis mingi Herre, 1936a: 8-9, plate IV (Pulau Ubin, Singapore); Fowler 1938: 266; Koumans 1940: 151. Sphenentogobius vanderbilti Fowler, 1940: 396-397, figures 8-11 (Medan, Sumatra); B6hlke 1984: 111. Stigmatogobius mingi: Koumans 1953: 118-119. Hemigobius bleekeri Koumans, 1953: 191-192, figure 47 (replacement name for Gobius melanurus Bleeker, 1849, not Gobius melanurus Bloch and Schneider, 1801); Kottelat et al., 1993: 146, plate 67. Hemigobius mingi: Larson 1999b: 34. Material Examined Lectotype of Gobius melanurus and Hemigobius bleekeri RMNH 4501, 1(55), Java, Indonesia, in sea (Sunda Archipelago on jar label). Paralectotypes of Gobius melanurus and Hemigobius bleekeri ex RMNH 4501, 2(40.5-55), same data as lectotype. Holotype of Gnatholepis mingi CAS 30960, 1(43), Pulau Ubin, Singapore, AW. Herre, 1934. Hemigobius mingi (Herre, 1936) Figures 11, 12, 20, 21, 74-77 Gobius me1anurus Bleeker, 1849: 31 (Java) [not Gobius melanurus Gmelin); Giinther 1961: 33. Holotype ofSphenentogobius vanderbilti ANSP 68714, 1(40.5), Medan, Sumatra, Indonesia, Vanderbilt Expedition, 23 May 1939. " ｾ <: ::n' o' 80' 60' ::l 0 lOO' ｾ C§, 3O'-l 30' :... RYUKYU IS. '. C"' E' ＮＢｾ .' I.' Ｏｾ ',' McldivGS ｾＬ :,' .. ':. .,' MARSHAU IS. 10' ' CAROUNE IS. V1-. .• ＯｾＮ .. 10' :1;7 , " "',0" <>"0 ":::> ?<?<I>O '.-I>' ",'" ',' O' ＧｾＺＮ c::::::!J •ｾ ." ',' ."••.• SOLOMON 1$, '.' <>4\ '. .. '. 10' ｓａｾﾷ <:l", ｾＭ Figure 74 FIJI'::::>, , <:l'" , .., ". '<:::\" • ;,j 20' :OVALTY IS. 30' 30' 60' " COi'A,{ SEA *• hoevenii mingi Mauritlus '20' • "'of KIRlaATl ｉ｜ｾ '" ; MAl?tANA IS. 0' ｾＮ I 20' Kosroe " { 80' Distribution of Hemigobius species, 100' 120' '80' 'I U1 76 H.K. Larson Figure 75 Hemigobius mingi. Holotype of Gnatholepis mingi Herre, 44 mm SL, CAS 30960, Pulau Ubin, Singapore. Other Material Singapore: ZRC 20263-72, 10(28-47), Sungai Punggol, 22 March 1966; ZRC 20192-37, 46(11.341.2), Sungei Punggol, c.K. Quek and M. Dali, 19 October 1965; CMK 8322, 2(39-41), Kranji mangroves near Sungei Buloh, M. Kottelat and D. Murphy, 8 April 1992; NTM S.14235-003, 4(14.5-22), Sungei Buloh mangroves, K. Lim, 30 January 1992; ZMH 19308, 2(26.5-35), aquarium import, Reichelt, 1 December 1090. Thailand: URM P.6677, 4(33-40), Ranong, 9 March 1982; NTM S.14288-OO1, 1(46), in ponds within research station, Ta-Chaluab, Chantaburi Province, Mahidolia Project, NIFI, Chulalongkorn University and Mahidol University parties, 2 June 1990; KUMF uncatalogued, 1(40.5), Tak Bai canal, Narathiwat Province, D. Tanwilai, 25 September 1984; CMK 5419, 1(32.5), Ban Pliu, near Chantaburi, M. Kottelat, 21 March 1980. No locality: RMNH 12580, 2(28-35.5). Mugilogobius Smitt, 1899 Mugilogobius Smitt, 1899: 552, India and Japan (type species: Ctenogobius abei Jordan and Snyder, 1901, by subsequent designation). Vaimosa Jordan and Seale, 1906: 395, Samoa (type species: Vaimosa jontinalis Jordan and Seale, 1906, by original designation). 24-65; predorsal scales 0-36, all evenly sized and relatively small or with anteriormost nape scale enlarged and placed close behind eyes; TRB 6-22; circumpeduncular scales 9-25; headpores always absent. Head papillae pattern longitudinal, rows composed of small, evenly sized papillae, papillae in rows a, c, cp and p small and closely spaced; long p row on top of snout and around eye forming characteristic "raised eyebrow" shape; on side of head, papilla row c broken under eye, rear portion consisting of two or more small papillae; three 5 rows present on snout, modally of two or more papillae, anteriormost row just above upper lip. Dorsal surface of head (strongly modally) with fine villi, mostly in interorbital space and on top of snout, sometimes extending into preorbital area. Several small fleshy knobs, lobes or fleshy ridge (at least) modally present on pectoral girdle. Mouth subterminal to terminal; jaws often enlarged in sexually mature males, sometimes considerably so. Snout usually broad, rounded and overhanging tips of jaws. Anterior nostril in tube and oriented down and forward over upper lip, preorbital usually curving outward slightly around base of nostril. Genital papilla generally slender, often elongate, flattened and pointed in males; round, bulbous, short to conical in females. Simple short gut with one loop present. Pterygiophore formula 3-12210; modally two Waiteopsis Whitley, 1930: 122, Gundamaian, New South Wales (type species: Waiteopsis paludis Whitley 1930, by original designation). Ellogobius Whitley, 1933: 92, Bateman's Bay, New South Wales (type species: Gobius stigmaticus De Vis, 1884, by original designation). Weberogobius Koumans, 1953: 172, Celebes (type species: Gobius amadi Weber, 1913, by original designation and monotypy). Diagnosis Distinguished by combination of characters. Small to moderately large gobies with second dorsal rays 1,6-10; anal rays 1,6-12, pectoral rays 12-20, 16 segmented caudal rays, in 9/7 pattern, lateral scales Figure 76 Jaws and suspensorium of Hemigobius mingi, female, ex URM P.6677, Ranong, Thailand. Revision of Mugilogobius Figure 77 Hemigobius mingi, head pores and papillae pattern. ZRC 20263-72, Sungei Punggol, Singapore. Scale bar = 1 mm. epurals; two to three pre-anal pterygiophores. Vertebrae 26--27, 10-11 precaudal and 15-16 caudal. Neural spines of first three vertebrae modally stout, and often broadly expanded, flanged or bifid at tip; in few species, spines slender with pointed tips. Palatine and pterygoid usually slender, nearly equal in length, with palatine slightly longer, palatine reaching quadrate; quadrate somewhat forked, dorsal arm broad; metapterygoid low and slender or broad and expanded dorsally (greatest expansion in mature males), upper limb usually forming bridge, overlapping quadrate; metapterygoid narrow and without bridge in one species-complex. Fifth ceratobranchial stout, triangular to nearly straight, with curved sharp teeth and short triangular or pointed flange on ventral surface. Scapula modally unossified (partial ossification observed in two species), fused to cleithrum. Six to no rakers ossified on first gill arch. Glossohyal broadly spatulate or forked. Remarks As defined above, the genus is distinguished by a combination of synapomorphies: small lobes or flaps on the pectoral girdle, fine villi present on dorsal surface of the head, sensory papillae on head all small and evenly sized, papilla row p long, forming a characteristic eyebrow shape over the eye, papilla row c broken under the eye with the rear portion consisting of several papillae, mouth enlarged in males and headpores always absent. No single defining synapomorphy for the genus has been found so far. Jordan (1920) first recognised that " Mugilogobius is substantially the same as Vaimosa Jordan & Seale, which name it may replace". 77 Herre (1927) separated his new genus Tamanka (now considered to be monotypic) from Vaimosa Jordan and Seale, due to the" much smaller and more numerous scales (38 to 54 in a longitudinal series), and by having many small cycloid scales on the opercles instead of a few large ctenoid ones" (Herre, 1927). Vaimosa (a junior synonym of Mugilogobius) was once used as a catch-all name for most of the genera currently considered to be related to Mugilogobius. Most of the species described by Herre as Vaimosa are Redigobius or Pseudogobius, genera which, unlike Mugilogobius, possess headpores. Another synonym of Mugilogobius, Waiteopsis Whitley, 1930, was created to include only paludis Whitley (= Mugilogobius platynotus). Whitley (1930: 123) indicated that he recognised the genus Mugilogobius, in which he placed Mugilogobius stigmaticus (De Vis), but he did not provide characters distinguishing Mugilogobius and Waiteopsis. Whitley (1933) later considered that Australian species previously referred to as Mugilogobius did not belong to that genus, so he created Ellogobius, including two species (paludis and stigmaticus) in the subgenus Ellogobius, with a third species (galwayi McCulloch and Waite (= Pseudogobius olorum)) placed in a separate sub genus (Lizagobius; = Pseudogobius). Aurich (1938) described as new Tamanka maculata and Vaimosa cagayanel1sis. The type specimens of Tamanka maculata could not be located at ZMH in 1988, and it is likely that they were lost during WWII. From the original description, T. maculata may have been a species of Mugilogobius. The syntypes of Vaimosa cagayanensis were found, and, although in poor condition, were observed to be a valid species of Mugilogobius. Aurich (1938) was the first, and one of few authors, to publish the observation that this group of gobies very often possesses papillae, knobs or lobes on the edge of the cleithrum (pectoral girdle). These knobs or lobes are characteristic of most Mugilogobius species, and are present in some species of Brachygobius, Eugnathogobius, Pseudogobius and Redigobius. The genera Stenogobius and Awaous also possess lobes (usually elongate in these two genera) which are mentioned in most works in which these genera are discussed (e.g. Koumans, 1953; Akihito et al., 1988; Watson, 1991; Kottelat et al., 1993). Koumans (1953: 155) mentioned the absence of "flaps" in his diagnosis of the genus TamankLl (Koumans included three valid genera within Tamanka). However, it is apparent from the subsequent text that he did not actually see the material. Koumans characterised Tamanka as having " ... edge of shouldergirdle without fleshy flaps, or with some small flaps" (1953: 155). This observation must be based on Aurich's description, as Koumans H.K. Larson 78 only mentions the flaps in his diagnosis of "Tamanka latifrons" but does not mention the flaps in the diagnoses of any other species, several of which do possess fleshy lobes (= flaps). The genera Tamanka and Weberogobius had previously been considered to be valid genera (Larson and Kottelat, 1992). However, Weberogobius is here treated as a junior synonym of Mugilogobius. 1. lA. 2. 2A. 3. 3A. Key to species of Mugilogobius Preoperculum with small scales at least ventrally; gill rakers with short papillose rakers on outer face of first arch, inner and outer faces of other gill arches with oval papillose pads; body with three or four narrow dusky horizontal lines ... M.lepidotus sp. novo (freshwater, Lake Towuti, Sulawesi) Preoperculum always naked; gill rakers short, stubby or slender, but not with spiny or papillose oval pads; colour pattern variable, but not with distinct narrow horizontal lines 2 Five or six short transverse rows of papillae almost directly under eye; f rows on either side of chin oriented longitudinally; relatively plain yellow to greyish when live ..... M. rexi sp. novo (freshwater, Towuti and Mahalona Lakes, Sulawesi) Papillae rows longitudinal, rows b, d and e may be doubled but not forming transverse rows; f rows oriented transversely across 3 chin; body colour variable Second dorsal and anal rays 1,9-12 (modally 1,10); predorsal scales small, 22-36; adult fish large (up to 124 mm); plain black to brown in colour with narrow head, terminal mouth and distinctly protruding chin ..... M. amadi (Weber, 1913) (freshwater, Lake Poso, Sulawesi) Second dorsal and anal fin rays 1,7-9; predorsal scales 0-36; adult fish reaching only to 65 mm; mouth usually subterminal, if terminal, then other characters not as above 4 4. Vertebral pattern 11+15-16; pectoral girdle smooth-edged, with no knobs or flaps; narrow vertical bars or series of rounded diffuse blotches along side .. .................. M. sarasinorum (Boulenger, 1897) (freshwater, Lake Poso, Sulawesi) 4A. Vertebral pattern 10+15-16; pectoral girdle nearly always with small knobs, lobes, flaps or fleshy ridge; colour not as above 5 5. Dorsal and anal rays modally 1,7 (if 1,8, then anteriormost nape scales enlarged); SA. 6. 6A. 7. 7A. 8. SA. 9. 9A. predorsal scales reaching to close behind eyes and almost entering interorbital space, anteriormost one to few scales on nape enlarged, with scale in centre of nape immediately behind eyes usually largest (few species may have anterior nape scale only slightly larger than those around it) ... 6 Dorsal and anal rays modally 1,8 or more (rarely 1,7, if so, then scales on nape small, none enlarged); extent of scales on nape variable, reaching to above preopercular margin or less, never entering interorbital space, usually all scales small and about equal in size (few species may have first nape scale somewhat enlarged in some specimens), or absent from predorsal midline 12 Body with four to seven distinct encircling blackish bands; 24-28 lateral scales present; first dorsal spine may be elongate 7 Body with short bars or blotches, if distinct bands encircle body, then lateral scales 2635 (average 30); first dorsal spine variable ... .......................................................................... 8 Body bands wide, five usually encircle the body; first body band begins at first dorsal fin origin; second to fourth first dorsal spines may be free and elongate . M. fasciatus sp. novo (Thailand, Singapore, estuarine) Body bands narrow, four usually encircle the body; first body band begins above pectoral base; first or second first dorsal fin spine elongate ......... M. tignnus sp. novo (Thailand, Malaysia, Singapore, estuarine) Strongly marked species with dusky greyish body having six to nine distinct dark to black diagonal narrow bars or cross-hatched blotches, a pair of vertically aligned, dense black spots at base of caudal and very distinct black spot in first dorsal fin 9 If pair of black spots present at base of caudal, brown to greyish body with only indistinct narrow diagonal bars, first dorsal fin markings variable 10 Second and third (at least) dorsal spines usually filamentous; black spot on first dorsal fin placed to rear of fin, anterior edge of spot beginning at third spine ..... M. chulae (Smith, 1932) (Ryukyus, Hong Kong, Thailand, Philippines, Sri Lanka, Singapore, Sulawesi, estuarine) Third first dorsal spine longest, but no spines filamentous; black spot on first dorsal fin placed forward on fin, with anterior edge of spot beginning at second dorsal spine ..... M. Revision of Mugilogobius 10. wilsoni sp. nov. (Northern Australia, estuarine) 13. First dorsal spine longest, often elongate (if 14 so, others are not elongate) Ctenoid scales on body extend to above pectoral base or further, predorsal and opercular scales usually ctenoid; lateral scales 28-32; scales on body with spot in centre giving reticulate appearance, body dark with indistinct pair of spots at caudal base; predorsal scales 9-13; head blunt in large adults M. fuscus (Herre, 1940) (Sri Lanka, Philippines, Papua New Guinea, Ryukyus, estuarine to freshwater) 13. Third to fourth dorsal spine longest; if first dorsal spine is elongate, then so are other spines 16 14. Distinctly marked with four black bands which may encircle body (bands may be partly broken up into black blotches; predorsal scales 11-15; pectoral rays 1415 M. adeia Larson and Kottelat, 1992 (Sulawesi, freshwater) lOA. Ctenoid scales on body extend to behind pectoral fin, predorsal and opercular scales cycloid; lateral scales 26-40; predorsal scales 11 12-22; colour not as above 11. 79 Second or third dorsal spine usually longest (sometimes first spine); predorsal scales 1016 (modally 13), anteriormost few enlarged; pectorals modally 15; side of head with two longitudinal streaks, one from behind eye, and one from rear of upper jaw (in banded form only, a bar extends from below eye to meet streak from jaw); body with distinctive "chequered" (alternating light and dark blotches) colour pattern, nine indistinct body bars underlying this, two or three indistinct spots or marks on caudal base ...... ............. M. mertoni (Weber, 1911) (Northern Australia, western Pacific to western Indian Ocean, South Africa, estuarine to freshwater) 11A. First dorsal spine usually longest, often elongate; predorsal scales 13-22 (modally 19), anteriormost scales often not much larger than others; pectorals modally 16; side of head with distinct reticulated pattern, two or three bars extend from eye to meet streak from jaw; body with oblique dark bars and blotches, rarely "chequered", broad dusky band usually crosses branchiostegals . ......... M. filifer sp. novo (Northern Australia, Solomon Islands, estuarine to freshwater). 12A. Gill opening rather oblique, small knobs on pectoral girdle usually present; mouth terminal to subterminal; body plain dark brown in males, females with four to seven indistinct oblique bars or X-shaped blotches along mid side, caudal fin brown, often with rows of small dark spots; head with three to five faint oblique stripes from eye across preopercle . M. cagayal1ensis (Aurich, 1938) (freshwater, Cagayan Sulu Archipelago, Philippines) 12A Gill opening usually vertical; colour not as above; mouth subterminal 13 14A. Body without distinct bands, colour almost uniform, although up to nine indistinct bars or cross-hatched blotches may be present; predorsal scales 14-22; pectoral rays 1516 15 15. Circumpeduncular scales 12-14; predorsal scales 18-22; side of head with indistinct stripes or simple reticulate pattern, shoulder bar indistinct, single blackish spot on upper caudal fin base ... M. fusculus (Nichols, 1951) (Papua New Guinea, freshwater to estuarine) 15A Circumpeduncular scales always 12; predorsal scales 14-16; side of head with finely reticulate or ocellate pattern, dark shoulder bar conspicuous, two small blackish spots at caudal fin base .. M. rivulus sp. novo (Northern Territory, estuarine) 16. Caudal fin with two to four dark longitudinal streaks originating at base, central two streaks may extend forward onto caudal peduncle as partial or distinct streaks, body with short bars . M. abei Oordan and Snyder, 1901) (Ryukyu Islands, mainland Japan and China, freshwater) 16A Caudal fin variably marked, but without two to four dark longitudinal streaks; an indistinct dark streak may appear along mid-lateral line of body 17 17. Circumpeduncular scales modally 12; caudal fin plain, or with two dark spots at base, or with several rows of small spots (not bands) ........................................................................ 18 17. Circumpeduncular scales 13-20 (rarely 12); two or more dusky diagonal bands may be present on caudal fin 20 18. Caudal, soft dorsal and anal fins with several rows of small dark spots; distinct dark diagonal shoulder bar and second, less distinct, bar extending downward from first dorsal fin; scale margins clearly outlined forming "network" pattern, reticulate pattern variably developed on head .. H.K. Larson 80 ............ M. rambaiae (Smith, 1945) (Thailand, Singapore, Borneo, Sulawesi, Sri Lanka, freshwater to estuarine) 18A. Unpaired fins not marked as above, if shoulder bar present, then six to nine other body bars also present 19 Predorsal scales 16-18; body with light and dark "chequered" pattern (especially in females), six to nine body bars visible, often indicated by cross-hatched blotches, head with reticulated pattern often welldeveloped, two dark spots at base of caudal fin M. notospilus (Giinther, 1877) (Queensland, Papua New Guinea, western Pacific islands, freshwater) 19A. Predorsal scales 18-23; body with indistinct narrow bars and no marks at base of caudal fin . ...... M. latifrons (Boulenger, 1897) (Sulawesi, freshwater) 19. 20. Three or more diagonal dark bands cross caudal fin, bands may be indistinct or broken (but present), upper dark spot on caudal base usually visible; lower spot indistinct; ctenoid scales on side of body usually extend without interruption to behind pectoral fin M. cavifrons (Weber, 1909) (Ryukyus, Taiwan, Philippines, western Pacific islands, introduced into Hawaii, Indonesia, Kei Islands, Papua New Guinea, freshwater to estuarine) 20A. Caudal fin with indistinct spots or streaks, not forming diagonal bands, often two dark spots at base of caudal; ctenoid scales on side of body may be in two sections: a patch behind pectoral fin and a wedge extending forward from caudal to below the gap between the dorsal fins 21 21. Predorsal scales 14-30, never absent; ctenoid scales on side of body extending forward to below fifth second dorsal ray or further; dorsal and anal fin rays usually 1,7-8 22 21A. Predorsal scales 0-21 (modally 0 in males, 3 in females), sides of nape may have small scales present if midline naked; ctenoid scales on side of body restricted to caudal peduncle and patch under pectoral fin; . dorsal and anal fin rays often 1,9 M. platynotus (Giinther, 1861) (Queensland to Victoria, marine to estuarine) 22. Ctenoid scales on side of body extending continuously up to behind pectoral base, not broken into two areas; predorsal scales 1725, sometimes anteriormost few scales may be larger than those behind; all dorsal spines short; pair of variably developed dark spots at caudal base; about nine indistinct narrow bars or square brown blotches along midside of body . M. stigmaticus (De Vis, 1884) (east coast of Australia, estuarine) 22A. Usually a distinct gap in ctenoid scales on side of body, or ctenoid scales form very narrow wedge up to behind pectoral fin; predorsal scales 8-21, all small; some dorsal spines may be elongate 23 23. Peritoneum dense black, less so ventrally; single dusky spot at centre of caudal base, caudal fin plain; body with indistinct mostly vertical bars and spots along midside (males may be nearly uniform in colour) . M. littoralis sp. novo (Northern Territory, Western Australia, estuarine to marine) 23A. Peritoneum brown dorsally, pale ventrally; colouring not as above 24 24. Caudal fin with spotting and small blotches forming vertically oriented rows; body with indistinct oblique bars and blotches across upper sides ..... M. myxodermus (Herre, 1935) (China, freshwater) 24A. One dark spot at caudal base and two distinct oval dark spots on base of caudal fin, caudal fin plain or with dusky streaks; body often with short diagonal dark bars which may form rounded blotches M. platystomus (Giinther, 1872) (north-eastern Australia, Indonesia, Belau, Singapore; freshwater to estuarine) Mugilogobius abei (Jordan and Snyder, 1901) Figures llA, 12F, 78-83; Tables 5-9 Ctenogobius abei Jordan and Snyder, 1901: 55, figure 5 (Wakanoura, Kii, Japan). Mugilogobius abei: Jordan et al., 1913: 345; Tanaka 1918a-b: 514-516, plate 136, figure 381; Koumans 1931: 90; Whitley 1931: 156; Akihito 1963: 11, 13, 16; Chu and Wu 1965: 132; Arai and Kobayasi 1973: 1-2,4, figure 13; Shao 1980: 178; Kanabashira et al., 1980: 191-198; Akihito et al., 1988: 268, plate 247M; Kim and Lee 1986: 21-24, figures 1-3; Kim et al., 1986: 400; Tzeng 1986: 131; Anonymous, 1986: 271-272; Kim et al., 1987: 533,536; Miller 1987: 688, 694-6, 699, 702; Takagi 1989: 564. Tamanka bivittata Herre, 1927: 224, plate 17, figure 4 (Hoihow, Hainan, China); Koumans 1940: 156. Gobius abei: Tomiyama 1936: 74; Kamohara 1950: 252; Aoyagi 1957: 229. Gobius (Tamanka) bivittata: Nichols 1943: 265. Revision of Mugilogobius Figure 78 81 Mugilogobius abei. Holotype of Ctenogobius abei Jordan and Snyder, 35 mm SL, CAS 6447, Wakanura, Kii Peninsula, Japan. Mugilogobius fontinalis: Hayashi and Itoh 1978: 71, plate IS, figure 42; Akihito et al., 1988: 268, plate Other material examined (no data taken) China: CAS/SU 28182, I, Tai Ping, Kwangtung 247N; Miller 1987: 688; Kawanabe and Mizuno 1989: 563, 576. Province; Hong Kong: CAS 53892, I, Castle Peak; AMS 1.19319-001, I, Deep Bay; CAS/SU 61119, 30, Keui Island; ANSP 76734, 2. Japan: FMNH 57196, I, Shirnbara; ANSP 26371, 5, Wakanoura, Kii; URM P.5307, I, Saigo River, Nagasaki; YCM P.1245, 2, Abukuma River, Miyagi; YCM P.6936, 5, Iida River, Kashirna; AMS 1.20371-001, 5, Izumi, Kochi; NSMT unregistered, I, Iriomotejima. Tamanka bittata Kim and Lee, 1986: 21 (lapsus for bivittata). ?Mugilogobius abei: Nguyen 1991: 335-337, figure 144. Ctenogobius abei: Ding 1994: 519. Material Examined Holotype of Ctenogobius abei CAS/SU 6447, 1 (35), Wakanoura, Kii, Japan, Jordan and Snyder, Apri11939. Paratype of Ctenogobius abei USNM 49892, 1(35), same data as holotype. Other Material Japan: MNHN 1967-566, 3(32-40.5), Hayama, Shirnoyama system, Kanagawa, Hirohito, July 1963; YCM 7055, 3(36-40), Yodo River, Hidegashidogawa Ward, Ohsaka Prefecture, 20 June 1977; BLIH 1984213, 13(15.5-35), Kamoba, Shinhama, Chiba Prefecture, 15 November 1984; YCM 908, 7(30-40), Nojima, Yokohama, Kamagawa Prefecture, 7 August 1973; URM P.7053, 66(11-39), Manko, Naha City, Okinawa, 31 January 1983; AMS 1.20372-001, 5(25.5-29), Kiyonna, Kagoshima Prefecture, 28 March 1968; URM P.4099, 1(41), Shiira River, Iriomotejima, Ryukyu Islands, T. Yoshino, 30 January 1982; USNM 132780, 11(11-33), Okinawa, J.R. Simon, 7 September 1945; BLIH 1984144, 8(1534); mouth of Adake River, Tanegashimajima, Kagoshima Prefecture, 25 October 1984; URM P.3507, 1(28.5), Teima River, Okinawa, 6 June 1982. Taiwan: USNM 316176, 1(36), Tan-Hai, 18 June 1969. China: ex AMNH 37029, 3(30-34), Foochow, Fukien Province, E.C. Pope, March-September 1926; CAS/SU 30266, 2(31.5-35), Kwangtung Province, A. Herre, 1934. Hong Kong: CAS/SU 39616,8(25-37), Un Long, New Territories, A. Herre, 23 February 1941. Korea: USNM 143027, 1(25.5), I.B. Bernadon, 1886. Diagnosis A moderate-sized Mugilogobius with second dorsal and anal rays 1,7-9; pectoral rays 12-17; longitudinal scales 33-44; TRB 10-15; 12-16 circumpeduncular scales; predorsal scales small, usually reaching halfway between preopercular margin and eyes; scales on body mostly ctenoid; second and third spines of dorsal fin often filamentous; distinctive pair of dark stripes along side of caudal peduncle, extending onto caudal fin, short dark bars and irregular blotches along anterior half of body; restricted to southern Japan, Hong Kong, Korea, Taiwan and Fukien Province, China; barred colour form known from Ryukyus, Okinawa, Tanegashima and southern Kagoshima. Figure 79 Jaws and suspensorium of Mugilogobius abei, male, ex YCM 908, Yokohama, Japan. Scale bar= 1 mm. H.K. Larson 82 Table 5 Frequency distribution of second dorsal and anal fin ray counts in Mugilogobius species. Species abei adeia amadi cagayanensis cavifrons chulae fasciatus sp. novo filifer sp. novo fuscus fusculus latifrons lepidotus sp. novo littoralis sp. novo mertoni myxodermus notospilus platynotus platystomus rambaiae rexi sp. novo rivulus sp. novo sarasinorum stigmaticus tigrinus sp. novo wilsoni sp. novo 6 Second dorsal rays 8 9 7 6 50 22 2 14 55 23 54 31 90 3 11 1 29 2 29 38 20 23 55 2 37 45 42 27 28 24 4 5 1 1 1 2 1 73 1 4 1 2 3 1 1 6 34 39 Anal rays 10 6 7 8 9 1 4 6 51 16 3 2 4 55 25 54 29 100 3 1 16 10 1 28 3 1 35 16 14 1 1 1 35 1 2 1 1 8 1 7 34 4 6 74 2 50 1 2 2 2 2 3 2 1 2 Description Based on 60 specimens, 15-41 mm SL. Counts of holotype of Ctenogobius abei (Figure 78) indicated by asterisk. First dorsal V (in two), V1* (in 57), VII (in one); second dorsal 1,7-9 (mean 1,8*); anal 1,7-9 (mean 1,8*); pectoral rays 12-17 (mean 16*); segmented caudal rays 15-16 (mean 16*); caudal ray pattern 7/ 6 to 9/7 (modally 9/7, 8/7* in holotype); branched caudal rays 13-16 (15*; mean 16); unsegmented (procurrent) caudal rays 7/7 to 10/9; longitudinal scale count 33-44 (36*; mean 37); TRB 10-15 (14*; mean 12); predorsal scale count 15-23 (22*; mean 18); circumpeduncular scales 12*-16 (mean 13). Gill rakers on outer face of first arch 3+7 to 5+7* (modally 3+7). Pterygiophore formula 3-12210* (in seven), 3-12211 (in one). Vertebrae 10+16* (in 14), 10+17 (in two). Neural spine on first and/or second vertebra broad, expanded at tip (in nine) or pointed (in one). Two (in 13) or one (in two) epurals. Two (in 14) or three (in one) anal pterygiophores before haemal spine of first caudal vertebra. Head usually somewhat wider than deep, length 3.1-3.8 (mean 3.4) in SL, in some specimens head nearly square in cross-section; large specimens with head considerably wider than deep, cheeks inflated and muscular in males. Depth at posterior preopercular margin 1.1-2.0 (mean 1.8) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.4) in HL. Mouth usually subterminal, may 1 2 10 34 36 10 11 12 17 3 1 1 35 29 13 17 2 4 39 44 42 27 28 21 3 31 2 1 1 1 2 2 4 be barely terminal, slightly oblique, forming angle of about 20-25° with body axis; jaws reaching to below anterior half of eye or to below mid-eye. Upper jaw 2.0-3.1 (mean 2.4) in HL, mouth not enlarged in mature males. Lips smooth, upper lip often with densely fimbriate tissue on inner edge, inner edge of lower lip slightly fimbriate. Lower lip free at sides, fused to underside of head well before mandibular symphysis. Eyes lateral, 3.3-4.8 (mean 4.0) in HL, set relatively high on head, upper margin of eye often forming part of dorsal profile. Snout usually rounded, often plump, 2.8-4.2 (mean 3.5) in HL. Interorbital broad, fleshy skin over eye quite thick, 2.4-4.4 (mean 3.2) in HL. Upper surface of head, especially interorbital, dorsal surface of snout, upper part of opercle and preopercle, often covered with small closely-spaced villi (usually visible when mucous coat scraped away). Body relatively compressed, especially posteriorly, depth at anal origin 4.2-6.5 (mean 5.3) in SL. Caudal peduncle compressed, length 3.7-4.8 (mean 4.1) in SL. Caudal peduncle depth 6.4-8.4 (mean 7.2) in SL. First dorsal fin with second to fourth spines elongate in both sexes, second or third spine longest, 2.7-7.9 (mean 5.4) in SL. First dorsal spines reaching to third to fifth second dorsal fin element when depressed. Second dorsal and anal fins low, anterior rays slightly shorter than posterior rays, not reaching caudal fin when depressed. Pectoral fin rounded, 3.8-9.1 (mean 6.6) in SL, central rays Revision of Mugilogobius longest; rays all branched in adults except for uppermost (lowermost also often unbranched). Pelvic fins short, rounded, usually reaching halfway or less to anus; 3.0-7.5 (mean 5.6) in SL. Caudal fin relatively short, rounded, 4.2-10.4 (mean 7.8) in SL. No mental fold or fraenum. Anterior nostril tubular, tube placed just behind upper lip, oriented downward or forward and down over lip; rear margin of tube sometimes expanded. Posterior nostril oval, without raised rim, placed near centre of anterior margin of eye. Gill opening usually extending forward to below opercle. Inner edge of pectoral girdle smooth (in 11), with low irregular fleshy ridge or raised flange (in 15) or with one to four fleshy knobs or bumps (in 24). Gill rakers on outer face of first arch generally smooth and very short, two or three rakers closest to angle of arch longest, rakers on upper limb very stubby; rakers on inner face of first arch short and stout, rakers on other arches somewhat longer. Tongue tip usually blunt or concave. Outer row teeth in upper jaw stout, curved and pointed, largest at front of jaw (outer row teeth larger in males than in females), figure 80 MIIgilogobius abei, papillae pattern. Composite, based on holotype of Ctenogobius abei (CAS 6407) and YCM 908. A, lateral view; B, ventral view of chin area (not to scale). Scalation suggested only. Scale bar = 1 mIn. 83 followed by two to three rows of much smaller pointed teeth; only one or two rows of teeth on rear half of jaw (Figure 79). Teeth in lower jaw pointed, curved, in four to five rows across front, one or two rows at sides; innermost row of large curved teeth across front of jaw only, posteriormost tooth in this row sometimes enlarged in males. Predorsal scales small, evenly sized, usually reaching about halfway between eyes and preopercular margin but sometimes reaching up to behind eyes. Operculum with small cycloid scales, usually upper third to half scaled. Cheek always naked. Pectoral base covered with small cycloid scales. Prepelvic area fully covered by small cycloid scales. Belly usually with patch of ctenoid scales under pelvic fins; up to anterior half of belly with ctenoid scales (separated from ctenoid scales on side of body by wide band of cycloid scales). Ctenoid scales on side extending forward in wedge nearly to base of pectoral fin or to below origin of first dorsal fin; scales largest posteriorly. Genital papilla in male elongate, flattened, narrowing to pointed tip; in female, short, rounded and bulbous. Head pores absent, as in all Mugilogobius species. Sensory papilla pattern longitudinal, as shown in Figure 80. Papillae rows typical, composed of small, evenly sized, closely spaced papillae; long prow along snout and around eye forming characteristic "raised eyebrow" shape; on side of head, row c broken below eye, with several papillae in rear portion; three s rows present, usually of two or more papillae; mental f row of at least six papillae. Coloration of fresh material The following description is based on photographs in Akihito et al. (1988), Sakurai (1981) and Kawanabe and Mizuno (1989). There are two colour forms of this species. Typical M. abei are found mainly in southern Japan and China and have two distinct stripes on the caudal peduncle. The other form, in which the peduncle stripes are partly obscured by barring, is known mainly from the Ryukyu Islands. "Typical" M. abei with basic colour pattern of overall pinkish-brown to yellowish-brown, with two broad brown stripes extending from caudal base forward to at least below second dorsal fin origin, and anterior half of body with five to seven slightly curved short brown bars. Head similar, pinkishbrown to yellowish-brown with brown to reddish brown markings. Outer margin of lower lip reddish. Iris dark brownish gold, with narrow gold rim around pupil. About six brown dorsal saddles present on rear half of body, beginning behind posteriormost brown bar; saddles usually extending down to reach uppermost peduncle stripe, sometimes extending further ventrally. Caudal peduncle stripes extending posteriorly onto H.K. Larson 84 Table 6 Frequency distribution of pectoral ray counts in Mugilogobius species. Species 12 13 abei adeia amadi cagayanensis cavifrons chulae fasciatus sp. noy. ftlifer sp. noy. fuscus fusculus latifrons lepidotus sp. noy. littoralis sp. noy. mertoni myxodermus notospilus platynotus platystomus rambaiae reri sp. noy. rivulus sp. noy. sarasinorum stigmaticus tigrinus sp. noY. wilsoni sp. noy. 1 1 4 1 4 14 15 16 17 4 17 5 28 8 7 28 21 15 4 7 3 28 1 27 45 10 43 24 63 3 25 33 2 26 4 12 12 2 34 9 3 12 1 4 17 7 8 1 1 2 40 2 10 29 6 38 15 14 20 27 12 19 1 1 22 15 9 2 5 1 11 1 2 3 1 1 8 24 translucent yellowish or pinkish caudal fin, and becoming blackish; stripes extending at least halfway onto fin, rear end of each stripe often radiating outward into several narrow black streaks along fin rays. Black streak over upper procurrent caudal rays, often similar brown streak posterior to it; upper margin of fin sometimes white. First dorsal fin striped: uppermost, narrow brownish stripe running along margin (membrane of elongate spines brown to pinkish-brown); translucent yellow band below this; median brownish stripe forming upper border of pink to brownish lower half of fin, stripe intensified posteriorly into distinct dense irregularly shaped black spot at rear edge of fin, black spot usually with small bright yellow area immediately above; third stripe brownish, running just above base of fin. Brownish stripes, especially lowermost, often intensified into blotches or spots over fin spines. Second dorsal fin similar in colour, with broad brownish marginal stripe, broad translucent yellowish submarginal stripe, and lower two-thirds of fin pinkish-brown; upper border of this region usually with blotches or short oblique marks anterior to each fin ray; at base of fin, three or more dark brown oblique blotches or streaks often present. Anal fin plain translucent brownish to pinkish, with posteriormost rays tipped with white. Pectorals hyaline or pale translucent yellowish. Pelvics brown to translucent pinkish brown. Ryukyu colour form with basically similar colouring, but for increased number of lateral 18 19 20 5 15 1 1 1 4 1 5 3 3 10 16 23 4 4 brown bars, less distinct caudal peduncle stripes; also, lowermost black caudal streak often less pronounced, though always present. Good colour photographs of living fish can be found in Kawanabe and Mizuno (1989: 576). Colour photographs of freshly dead specimens appear in Akihito et al. (1988: 257, 268, plate 247), Tzeng (1986: 131), Sakurai (1981) and Shao (1980: 178). Coloration of preserved material Head and body light greyish yellow, paler ventrally, with brown to dark brown bars and stripes. Anterior half of body usually with five short brown bars on sides; posterior half of body with two brown stripes along midside, stripes beginning at level of origins of second dorsal and anal fins respectively, and extending onto caudal fin as black streaks (Figure 81). Head with brown stripes or irregular network of markings, top of head with darker vermiculation and spots. Side of head with three to four oblique stripes, often coalescing at intervals, forming an irregular network. First stripe beginning halfway between end of jaw and anterior nostril tube, usually running along ventral margin of eye (sometimes ending at eye and reappearing at rear margin of eye to join second stripe), then bending back and meeting second stripe on upper preopercle behind eye. Second stripe beginning as horizontal mark from rear of jaw, often curved or hook-shaped, running obliquely across cheek to meet first stripe Revision of Mugilogobius Figure 81 85 Mugilogobius abei, female, 34 mm SL, BLIH 1984213, Shinhama, Chiba, Japan. indistinct bands. Top of snout and interorbital with short venniculate lines and spots, becoming blotchy toward nape; nape with indistinct square brownish blotches, posteriormost extending down behind pectoral fin and forming first brown body bar ("shoulder" bar). Body bars about equal to eye width, bars not extending to belly, but ending at point just below rnid-lateralline. Second bar beginning below origin, and third bar below rear, of first dorsal fin. Fourth bar beginning at origin of second dorsal fin, bar joining two caudal peduncle stripes; two or more short lines often joining stripes anteriorly, making ladder-like pattern. All four body bars sometimes broken up into series of indistinct blotches. Dorsal midline with about eight evenly spaced brown patches coinciding with body bars. Breast evenly dusky brownish. Ventral half of body pale, unmarked. Pectoral base brownish on anterior half, behind eye. First two stripes often joined anteriorly by brown line running along preorbital area just above upper lip. Third stripe beginning below head behind chin, running along ventral margin of preopercle, ending at centre of rear margin of preopercle and meeting fourth stripe. Fourth stripe broadest, beginning below head on branchiostegal membranes, curving up over branchiostegal rays and vertically onto opercle and meeting with preopercular margin stripe. Fifth partial stripe extending partly onto cheek from upper lip between first two stripes. Brown line running from anterior nostril tube (entire tube brown), below posterior nostril to anterior edge of eye; rear counterpart of line running obliquely from upper rear margin of eye onto dorsal surface of head becoming diffuse or venniculate. Broad diffuse brown line running from edge of eye around snout tip, just above posterior nostril. Underside of head dusky, with two Table 7 Frequency distribution of transverse backward scale counts in Mugilogobius species. Species abei adeia amadi cagayan. cavifrons chulae fasci. sp. novo ftlifer sp. novo fuscus fusculus latifrons lepid. sp. novo littor. sp. novo mertoni myxodermus notospilus platynotus platystomus rambaiae rexi sp. novo rivul. sp. novo sarasinorum stigmaticus tigr. sp. novo wils. sp. novo 6 7 17 18 155 788 3 20 19 3 2 8 1 8 2 5 15 12 2 4 4 1 2 11 11 11 4 4 5 2 5 2 12 10 455 5 6 1 2 8 9 10 11 12 13 14 15 3 8 16 24 7 2 5 8 7 2 16 19 20 21 22 3 1 1 1 1 1 1 1 3 23 1 17 18 20 21 3 7 6 2 4 2 25 2 7 12 1 19 9 6 14 4 H.K. Larson 86 Table 8 Frequency distribution of lateral scale counts in Mugilogobius species. Species abei adeia amadi cagayanensis cavifrons chulae fasciatus sp. novo filifer sp. novo fuscus fuseulus latifrons lepidotus sp. novo littoralis sp. novo mertoni myxodermus notospilus platynotus platystomus rambaiae rexi sp. novo rivulus sp. novo sarasinorum stigmaticus tigrinus sp. novo wilsoni sp. novo 24 25 2 5 5 26 27 1 Species 39 40 abei adeia amadi cagayanensis cavifrons chulae fasciatus sp. novo filifer sp. novo fuscus fusculus latifr.ons lepidotus sp. novo littoralis sp. novo mertoni myxodermus notospilus platynotus platystomus rambaiae rexi sp. novo rivulus sp. novo sarasinoru m stigmaticus tigrinus sp. novo wilsoni sp. novo 4 4 2 14 3 14 34 35 36 37 38 5 8 9 11 10 6 1 2 2 1 1 1 5 3 6 9 16 4 8 6 2 7 9 12 9 4 4 5 6 6 5 5 4 1 4 3 3 10 3 4 5 1 1 4 35 2 2 7 7 3 1 1 29 30 31 2 5 10 1 10 1 6 2 2 1 8 1 3 10 32 15 7 19 3 11 1 10 3 1 1 4 1 6 10 20 18 10 12 1 13 6 16 10 11 1 5 3 7 5 5 14 13 12 2 9 11 3 3 10 2 1 33 28 1 24 16 1 14 3 2 20 12 6 1 41 42 43 44 45 46 1 1 6 47 48 49 7 4 4 5 1 1 4 1 1 1 1 1 3 3 50 51 52 53 1 2 1 2 1 2 7 10 1 1 1 1 8 11 5 7 3 2 1 3 2 4 4 5 9 1 4 6 2 3 2 4 3 3 3 3 5 4 2 1 2 6 1 3 3 3 posteriorly pale; brown horizontal mark at midbase, often extending back onto pectoral rays. Brown stripes on caudal peduncle extending onto caudal fin as black streaks stopping short of rear edge of fin; streaks often bifurcating, following fin ray branches. Dorsalmost procurrent rays of caudal 3 1 5 5 1 2 2 3 1 1 2 2 covered by intense black streak from upper base to anterior quarter of upper fin margin. Behind black streak, two to four short black streaks along bases of uppermost caudal rays. First dorsal fin with three dusky to blackish stripes: first along margin, elongate spines dusky; second stripe oblique, placed Revision of Mugilogobius 87 Table 8 (cant.) Species abei adeia anuuii cagayanensis cavifrons chulae jasciatus sp. novo filifer sp. novo fuscus fusculus latifrons lepidot. sp. novo littoralis sp. novo mertoni myxodermus notospilus platynotus platystomus rambaiae rexi sp. novo nvulus sp. novo 54 55 3 3 3 6 56 3 57 58 59 3 1 4 4 60 61 62 63 64 3 65 1 3 1 sarasinorum stigmaticus tigrinus sp. novo wilsoni sp. novo halfway across fin; third stripe oblique, beginning on lower part of first spine and ending at midbase of fin; dense black spot at rear of fin. Second dorsal fin with wide dusky margin; translucent submarginal band of equal width; proximal twothirds of fin dusky, darkest dorsally. Two or more dark blotches or streaks along base of fin (usually two elongate blotches). Barred colour form similar, but caudal peduncle stripes mostly indistinct, lowermost stripe usually discernible (Figure 82). Ladder-like lines joining two stripes in "typical" M. abei proliferated and broadened, with five or more short bars or blotches along side of caudal peduncle (in addition to five or six broad body bars). In one lot from Kagoshima, both lateral stripes completely obscured and "ladder-bars" on caudal peduncle present as short, broad, X-shaped blotches (similar markings present in M. notospilus; possibly why Akihito et al., 1988 Figure 82 referred to this form of M. abei as 'jontinalis", a junior synonym of notospilus). In specimens from Kagoshima, blackish caudal streaks not so pronounced. Aoyagi (1957: figure 189) shows a drawing of M. abei with "barred form" colour pattern, but no provenance indicated. Comparisons The distinctive caudal fin colour separates this species from others of the genus. Mugilogobius notospilus (Giinther) shares similarities in colour pattern and it is possible to derive the abei colour (stripes on peduncle) from the bars on the side of M. notospilus. The spots on caudal base and pectoral base are very similar in these two species, as is the reticulate pattern on the head. Differences between the two species are: M. abei often has filamentous first dorsal fin spines and has higher scale counts (lateral scales 33-44, mean 37, Mugilogobius abei, barred colour form, 37.5 mm SL, URM P.7053, Naha, Okinawa, Japan. H.K. Larson 88 Table 9 Morphometries as percentages of SL or HL, as indicated, of Mugilogobius abei Oordan and Snyder, 1901). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral !. in SL Pelvic l. in SL Caudall. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 28.6 58.0 75.0 19.7 27.1 49.0 62.5 15.3 9.7 21.0 11.9 25.0 20.8 32.3 25.5 18.3 15.1 23.9 12.6 31.9 65.3 85.1 21.0 16.8 26.2 15.3 32.5 30.4 50.4 42.1 24.8 21.9 29.2 36.4 30.0 56.7 72.8 18.8 12.9 24.1 14.0 29.3 25.1 44.2 31.9 21.6 18.3 26.2 22.4 26.3 50.3 62.9 15.4 9.7 21.1 12.5 24.1 21.9 33.9 22.9 18.5 15.4 19.3 13.2 31.2 90.1 82.1 23.7 15.7 26.8 15.7 35.5 29.6 43.0 40.9 26.8 22.5 29.7 26.8 28.8 59.3 72.5 19.5 12.9 24.3 14.0 28.2 25.7 38.5 32.0 20.9 17.7 23.9 18.2 24.0 14.0 27.0 25.0 38.0 30.0 20.3 18.0 24.3 16.0 versus 28-36, mean 31, in M. notospilus; circumpeduncular scales 12-16, mean 13, versus always 12; TRB IQ-IS, mean 12, versus 9-12, mean 10 in M. notospilus). Distribution South-central to southern Japan, Hong Kong, Korea, northern Viet Nam, Taiwan and Fukien Province, China (Figure 83). Nichols (1943) refers to a specimen from a "tiny brook at Tai Ping". Kanabashira et al. (1980) record it from the Kanto district in middle Honshu southward to the Ryukyu Islands and the southern part of China including Hainan Island; they do not refer to any colour variants. The barred colour form is known from the Ryukyus, Okinawa, Tanegashima and southern Kagoshima. The "typical" striped M. abei also apparently occurs on Okinawa (Tomiyama, 1936), and one badly dehydrated CAS/SU specimen is known from Naha which has lost most of its colour, but still shows the two peduncular stripes. Chu and Wu (1965) list M. abei as being found in the South China Sea, Korea, Japan and possibly the East China Sea. Kim and Lee (1986) record M. abei from Mokpo, Puan and Poryong, on the west coast of the Korean Peninsula. Ecology This species inhabits sandy and muddy substrates in estuaries and coastal fresh water streams not far from the stream mouth (Kim and Lee 1986). It is probably somewhat fresh water tolerant. Hayashi and Itoh (1978) indicate that specimens they collected from the Shiiugawa River, Ishigakijima, were feeding on tanaidaceans. Kanabashira et al. (1980) report M. abei from sandy and muddy estuarine substrates, and describe the spawning season (April to October) of a population from the Takahama Channel in Tokyo Bay. They also successfully hatched and reared larvae (up to about 40 mm total length) in captivity. The larvae remained planktonic for about 40 days. It was not clear from their study whether larvae remained planktonic for less than this length of time in the field, or indeed, were able to delay settling even further. Kim and Lee (1986) refer to sexual dichromatism observed during the breeding season: " ... the body color of the male darkened and the margin of the dorsal fin became yellow, the female's body colour faded". Remarks This is the type species of the genus Mugilogobius, as designated by Jordan (1920: 487). In his original description of the genus Mugilogobius, Smitt (1899) neglected to designate a type species. Jordan (1920) indicated that Smitt had written to him in 1903, pointing out his intention " ...that his type species was the one named Ctenogobius abei Jordan and Snyder". Jordan accepted this intention, hence his designation. Smith (1945), however, disapproved, considering that it was" ...unjustifiable to attempt to validate the genus as of the date of its establishment by accepting as the genotype a species that was undescribed at the time the genus was set up ... ", and believed that the validity of the genus Mugilogobius " ... should date from the first published correlation of the name with a genotype, in 1913". As was indicated in the Introduction, Jordan's designation of abei as the genotype is acceptable under the International Code for Zoological Nomenclature (1999). The holotype of Tamanka bivittata Herre was registered as BSM 13194, and was 29 mm SL, 35 mm TL, according to Koumans (1940: 188); he recognised that bivittata was identical to abei. The type was destroyed in WWII. ｾ /'I) :::. en o ｾｬＮｷﾷｾ 80' O· ::l ［ｰＺＮｾＴ o ..., 40' fA/ CHNA MA 30" tｾ :::: { AAAPAN <:r ;,;. 30" • !. Ｎｾ " "' .' ' . 20' 20' ＿ｾｓ LACCAOIVE IS.• : .. 10' r Moldfve. :;. 'I 's :,' ..; 10' 20' .. (j MADAGASCAR 30' • ＭＧＺｾｬｏｍｎ SEYCHEUES (.q..r ••• ｴｾＮ '.. Ｌｾｏｾ • * • T ... *• 0' 10' , SAMOA ｃｊｾＧ［ FUI .... COfitAl SEA ＧｾＬ AU$TRAUA .... , '.",; , Mauritius '}OS ... '. o ＼ｲｾ I(IRlaAn IS. .\,. abei eavifrons chu/ae fasciatus n. sp. fiBfer n. sp. fuses 10' .': MARSHAllIS. Kosrae 20' -LOYALTY 1$. 30' v .wo l.w' Figure 83 • : '", CAROUNE IS: 0' ) iiii HAWAIIAN ISLANOS {) .l MAlnANA IS. Distribution of Mugilogobius abei, M. cavifrons, M. chulae, M. fasciatus n. sp., M. filifer n. sp. and M. fUscus. T.4SMANSEA 160' ､ｾ 40' o 180' 160' 00 \C H.K. Larson 90 Hayashi and Itoh (1978: 71) refer to the barred form of M. abei as M. fontinolis, without explanation as to why they use the name. Kim and Lee's (1986) name Tamanka bittata is clearly a typographic error for bivittata. Mugilogobius abei is called abehaze in Japan, and the barred colour form is known as izumihaze (Akihito et al., 1988). In Korea, it is called mochi-mang-dug (Kim and Lee 1986). Nguyen's (1991) record of M. abei from Viet Nam (based on other literature which I have not seen) could be something else entirely, because the illustration is not consistent with M. abei. The drawing is reminiscent of Herre's original illustration (1940a) of Vaimosa kabilia (= Eugnathogobius kabilia) and is likely a composite from previously published drawings. Akihito (1963) documented the partial ossification of the scapula in M. abei. Part of the portion above the foramen, nearest the radials, becomes ossified in some specimens, beginning at 26 mm SL. The scapula is cartilaginous in smaller specimens, and in some specimens up to 40 mm SL. Nogusa (1957; not seen, quoted in Arai and Kobayasi 1973) and Arai and Kobayasi (1973: figure 13) described the chromosomes of Mugilogobius abei, finding 46 diploid chromosomes, all with monoarms. Mugilogobius adeia Larson and Kottelat, 1992 Figures 16C, 84-87; Tables 5-8, 10 Mugilogobius adeia Larson and Kottelat, 1992: 225234 (Lake Matano, Sulawesi). Material Examined Holotype MZB 5891, 1(27.5), east of Talu, north margin of Lake Matano, Sulawesi, Indonesia, 19 March 1989, M. Kottelat and A. Werner. Paratypes Indonesia: Sulawesi: ZSM/CMK 6513, 4(16-22), same data as holotype; MZB 5892, 2(16-23), same data as holotype; NTM 5.13067-001, 4(13-28), same data as holotype; ZSM/CMK 6468, 4(15-19), Mengonuwai, eastern end of Lake Matano, 12 March 1989, M. Kottelat and A. Werner; NTM 5.13068-001, 3(21-26), southern margin of Lake Matano, east of Tandjung Potipuaha, 1 February 1991, M. Kottelat; ZSM/CMK 9063, 4(20.5-27.5), same data as preceding. Diagnosis A robust Mugilogobius with second dorsal fin rays 1,8; anal rays 1,7-8; pectoral rays 12-15; longitudinal scales 28-32; TRB 8-11; 11-13 circumpeduncular scales; predorsal scales small, reaching to above preopercular margin; scales on body mostly ctenoid; first spine of dorsal fin longest and often filamentous; at least five irregular broad black bands across sides, bands interspersed with large spots and irregular markings; known only from Lake Matano, central Sulawesi. Description Following description largely taken from Larson and Kottelat (1992). Based on 22 specimens, 12.527.5 mm SL. An asterisk indicates counts of holotype. First dorsal VI*; second dorsal 1,8*; anal 1,7*-8 (mean 1,8); pectoral rays 12-15 (mean 14*); segmented caudal rays 16*-17 (mean 16); caudal ray pattern modally 9/7; branched caudal rays 10-13* (mean 12); unsegmented (procurrent) caudal rays 6/6 (in two); longitudinal scale count 28-32 (29*; mean 30); TRB 8-11 (10*; mean 9); transverse scale count forward 9-13 (14*; mean 12); predorsal scale count 11-18 (15*; mean 14); circumpeduncular scales 11-13* (mean 12). Gill rakers on outer face of first arch 3+6 to 4+6 (no mode; mean total rakers 10). Pterygiophore formula 3-12210* (in three). Vertebrae 10+16 (in one), 11+16* (in three). Neural spine on first few vertebrae pointed (in one). Two epurals (in three). Three (in three) anal pterygiophores before haemal spine of first caudal vertebra. Head somewhat compressed, almost square in cross-section, 2.8-3.1 in SL (mean 3.0). Head depth at posterior preopercular margin 1.6-1.9 in SL (mean 1.8). Head width at posterior preopercular margin 1.4-1.7 in SL (mean 1.5). Mouth slightly oblique, forming angle of about 30° with body axis; jaws ending under mid-eye or anterior half of eye. Mouth 2.4-3.1 in HL (mean 3.0), terminal, upper jaw length same in males and females. Eyes lateral, set high on sides of head, forming part of dorsal profile, 2.9-4.0 in HL (mean 3.6). Snout slightly pointed, 3.4-4.3 in HL (mean 4.3). Interorbital moderately broad, 3.7-{).8 in HL (mean 5.2). Fine villi often scattered on interorbital (present on cheeks and preorbital of 45 mm SL specimen). Body robust, mean depth at anal origin 5.0 in SL. Body width above anal fin origin 6.7-9.5 (mean 7.7). Caudal peduncle compressed, length 3.5-4.6 in SL (mean 4.0). Caudal peduncle depth 7.3-10.3 in SL (mean 8.2). First dorsal fin generally triangular, with first dorsal spine usually elongate. First dorsal fin spine usually longest, maximum 5.7 in SL in males, 5.9 in females; extending beyond other spines when fin depressed and sometimes reaching to fifth or sixth second dorsal fin element. Second dorsal spine sometimes subequal to first. Second dorsal fin low, not much higher anteriorly than posteriorly. Anal fin low, with posteriormost rays longest. Pectoral fins narrow, roughly rectangular, 3.5-4.5 in SL (mean 3.9); rays usually branched (uppermost one Revision of Mugilogobius Table 10 91 Morphometries as percentages of SL or HL, as indicated, of Mugilogobius adeia Larson and Kottelat, 1992. Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic!. in SL Caudal!. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 33.1 59.3 72.5 21.1 14.9 24.0 12.7 27.5 25.3 35.2 18.7 26.2 25.1 27.6 27.3 32.5 52.1 58.6 18.1 10.3 21.6 11.0 24.1 25.8 32.8 14.8 22.4 21.8 24.7 17.4 36.3 59.7 68.5 22.7 15.0 28.3 13.6 29.7 33.3 41.0 26.9 28.9 28.6 31.6 42.9 33.8 56.2 65.7 20.4 13.6 24.8 12.7 26.7 29.6 36.3 20.1 25.7 24.9 28.4 31.2 32.3 51.5 59.1 17.0 10.5 21.8 9.7 23.3 25.3 34.3 16.4 22.9 20.6 24.6 17.0 35.4 61.2 72.5 22.3 14.9 27.2 13.0 29.2 34.9 38.1 22.2 28.8 25.6 30.4 38.7 33.7 55.5 65.1 19.8 13.1 25.1 11.9 26.4 30.8 35.6 19.0 25.5 23.6 27.5 27.7 to three rays often unbranched). Pelvic fins long, narrow, reaching to anus, origin below pectoral base, mean 4.0 in SL in males, 4.3 in females. Caudal fin rounded, upper rays slightly longer than lower. No mental fold or fraenum. Anterior nostril in short tube directed forward and down over upper lip. Posterior nostril with low rim, closer to eye than midway between eye and tip of snout. Gill opening extending forward to below opercle. Gill rakers on outer face of first arch short, stubby; rakers on inner face of first arch with very short spiny papillae (resembling raker teeth). Outer rakers on second arch short, stout; rakers on inner face of second arch with fine short spiny papillae. Pectoral girdle smooth in three specimens (including holotype), others with one to four low bumps or short lobes on one or both sides of body. Tongue blunt to slightly trilobed. Inner margin of both lips finely fimbriate. Teeth in outer row of upper jaw enlarged, caniniform, widely spaced; four to five inner rows of smaller, sharp conical teeth. Outer and innermost rows of teeth in lower jaw larger than remainder, outer row widely spaced, followed by two to three rows smaller teeth. Teeth similar in males and females. Predorsal scales cycloid, small and evenly sized, none enlarged, scales reaching forward to above preopercular margin, occasionally further forward, but never up to behind eyes. Operculum with small cycloid scales, lower third of opercle often unscaled. Cheek usually naked; two specimens, including holotype, with few scales behind eye. Pectoral base scales smalt cycloid. Prepelvic area scales cycloid, usually fully scaled. Belly with ctenoid scales anteriorly, anterior quarter to at least 3/4 of belly with ctenoid scales, remainder cycloid; scales close to anus always cycloid. Genital papilla in male slender, elongate and flattened, narrowing toward tip; papilla in female short, rounded and stout, with no lobes at tip. Head pores absent as in all Mugilogobius species. Sensory papillae on head in longitudinal pattern, as in Figure 84. Three s rows on snout, of three or more papillae. Mental f row of five or more papillae across chin. Coloration of fresh material Based on notes and colour slides of captive specimens. Bod y pale greyish white, yellowish, light brownish grey, or light bluish grey, overlaid by black to dark brown markings. Posterior half of body tending to light blue-grey. Peritoneum white to pinkish, showing through body waIt abdominal region lighter than rest of body. Four distinct broad black slightly diagonal bars crossing body; bands may be broken into several blotches, especially those posteriorly; usually irregular black spots present on upper back, interspersed with black bands. Conspicuous large irregularly rounded Figure 84 Mugilogobius adeia, papillae pattern. Holotype, MZB 5891. Scalation suggested only. Scale bar = 1 mm. Reproduced with permission from Larson and Kottelat (1992). H.K. Larson 92 shoulder blotch above pectoral fin base always present; blotch sometimes continuing diagonally backward behind pectoral base to belly, but nearly always broken up into rounded shoulder blotch and separate bar or blotch behind pectoral base. First distinct black band originating from base of third or fourth first dorsal fin spine, extending nearly vertically to belly; band broad, regular and straight-sided, or irregular and partly broken-up. Indistinct, rounded, irregular black spot on upper back between first band and shoulder mark sometimes partly connected to first band. Second black band beginning behind second element of second dorsal fin and extending slightly diagonally backward, ending behind third or fourth anal fin element. Upper portion of second band often wide or forming separate blotch partly connected to spot on upper back between second and first black bands. Anterior to second black band, triangular black blotch beginning at origin of anal fin and extending up to midside of body. Third black band beginning approximately behind sixth second dorsal fin element, and extending diagonally backward to behind rear of anal fin; band usually broken into upper oval blotch and partly to wholly separated irregular diagonal band. Fourth black band beginning at middle of upper caudal peduncle, extending diagonally back to just before lower caudal fin rays. Fourth band broad and continuous, or broken into two separate diagonallyoriented black blocks. Square to rounded dense black blotch at centre of caudal fin base. Slightly smaller, but similarly shaped, black blotch anterior to central caudal blotch on upper side of body; two blotches sometimes partly joined, forming irregular bar. Lower side of head light bluish grey, with upper surface of head, snout and predorsal region pale brownish to brownish grey. Distinct broad black streak running diagonally from upper rear corner of preopercle across lower cheek, ending at middle of lower preopercular margin. Similar but generally less distinct streak extending from upper rear corner of opercle diagonally forward to subopercle. Opercular streak quite narrow and distinct or broad and diffuse; sometimes joined to large shoulder blotch. Branchiostegal membranes dense black, Figure 85 iridescent blue markings sometimes overlying black. Both lips black, with short black line on head just above rictus. Intense black blotch or rounded spot extending from behind each eye onto nape. Top of head and snout with irregular black or dark brown vermiculate markings. Underside of head banded with black and whitish grey, markings following outline of branchiostegal membranes, preopercular margins, and folds of isthmus. Iris pale golden dorsally, remainder red gold, with wide black rim. First dorsal fin with whitish triangular area anteriorly (approximately covering bases of first four or five spines), and broad black diagonal band occupying most of middle of fin; distal to black band, fin dull whitish or yellowish, with bright white area posteriorly; white mark sometimes extending forward and forming very narrow white margin to broad black band. First dorsal fin spines brownish; fin with narrow brownish margin, or brownish markings on membrane only. Second dorsal fin with proximal two-thirds dusky to pale brownish, with fin rays brown to blackish; three evenly-spaced pale yellowish patches at base of fin, and short upright brown mark between each fin ray, on membrane near base; outermost quarter of fin whitish or yellowish; fin margin sometimes with very narrow dusky edge. Anal fin black, overlaid by dark iridescent blue spots and short streaks (streaks usually near fin base). Caudal fin brown to reddish brown, with some blue markings on lower half. Pelvics dusky to blackish. Pectorals translucent, with blackish fin rays. Pectorals and pelvics often with blue markings over dark background. Females similar to males, but second dorsal and anal fins paler in colour, caudal almost translucent and without blue markings, and white patch at rear of first dorsal fin quite dull and inconspicuous. Captive fish observed (by Maurice Kottelat) changing background colour to dull brown, with black bars and spots becoming brown, blending into main body colour and becoming difficult to see. Fin spines and rays blackish, caudal fin sometimes reddish brown. During courtship (in captivity), displaying males darken entire head and body, black markings MlIgilogobills adeia, paratype, 27 mm SL, CMK 9063, Lake Matano, Sulawesi. Revision of Mugilogobius Figure 86 93 Mugilogobius adeia, paratype, 26 mm 5L, NTM 5.13068-001, Lake Matano, 5ulawesi. almost indiscernible, pale bands on first and second dorsal fins becoming intense yellow with very narrow brown outer margin. White area at rear edge of first dorsal fin becoming bright pale lemon yellow. Lower dusky portions of dorsal fins and entire caudal fin becoming brown in colour, almost reddish. Dorsal fin margins in spawning females clear, not yellow as in males. Females not changing colour noticeably during spawning. Coloration of preserved material Body light brown to pale fawn, black bands and spots black or brown, depending on intensity of markings (Figures 85-86). Fin markings black or brown; unpaired fins' background colour pale dusky to hyaline, with brownish rays; pectoral fin membranes translucent or light dusky, with dark rays; pelvic fins usually quite dark. Comparisons The only similar Mugilogobius in Lake Matano and surrounding lakes is M. latifrons (Boulenger, 1897). Mugilogobius adeia differs from M. latifrons by colour pattern (compare Figures 85-86 and 125, 127), predorsal scale arrangement and number (in M. latifrons, the predorsal scales number 16-21, mean 18, and extend forward to halfway between eyes and rear of preopercular margin, versus 11-15 scales, mean 14, usually only reaching to above preopercular margin), lateral scale counts (average 32 in M. latifrons versus 30 in M. adeia), first dorsal fin shape (second or third spine longest, first longest in one specimen; versus first spine always longest and often elongate in m. adeia), and head length (M. latifrons head length averages 3.2 in SL, while M. adeia averages 3.0 in SL). However, five specimens (CMK 11430) recently examined from a tributary entering Lake Matano NE of Matano village, appear to share characters of M. adeia and M. latifrons. The colour resembles that of M. latifrons in four specimens and that of M. adeia in one. All specimens have 15 pectoral rays, 31-33 lateral scales, and 15-18 predorsal scales which extend to just before the rear preopercular margin. Two specimens have the first dorsal spine longest and the others have the third spine longest. These five specimens cannot be distinguished as either adeia or latifrons based on the combination of characters outlined in the previous paragraph. Based on body colour and first dorsal fin shape, four are M. latifrons and one is M. adeia. However, the extent of scalation, scale numbers and head proportions are the same. Maurice Kottelat advised me (in litt.) that the area in which the fish were collected was disturbed due to road construction. It is possible that the local populations of these two species may have hybridised due to habitat disturbance. Data from these specimens is not included in this paper and no decision made as to their status (awaiting further specimens and information about the area). Distribution Restricted to Lake Matano, central Sulawesi (Figure 87). Ecology The following information is from Larson and Kottelat (1992). Lake Matano is a very deep, steepsided, graben lake situated along the Matano Fault line, with its surface at an elevation of 382 meters. It is the deepest lake in Sulawesi (590 meters, according to Abendanon, 1910, 1915), and has three main, intermittently flowing rivulets draining into it, and an outflowing river, the Patea, which flows into Lake Mahalona to the south-east. The tectonic lakes of central Sulawesi are inhabited by Anguilla, hemiramphids, telmatherinids, adrianichthyids, oryziids, and gobiids (Weber, 1913; Kottelat, 1990ac, 1991). Due to the isolation of these lakes, most fish species are restricted to particular lakes (either a single lake, or a group of lakes). Recent collections in the region have provided a considerable increase in the number of endemic species known (Kottelat 1990a-<:, 1991). A number of exotic species also exist in most of the lakes, including several which were probably introduced by early settlers to the region (Whitten et al., 1987; Kottelat et al., 1993). These introductions represent a considerable threat to the native fauna, part of which may already be extinct (Whitten et al., 1988; Kottelat, 1990a). In sheltered, shallow bays of Lake Matano, the substrate is sand and gravel (seldom mud), often with quite large rocks and boulders interspersed. Aquatic vegetation is restricted to the edges of the shallow bays. At the type locality, east of Talu on H.K. Larson 94 which occurs throughout the lake. About half the specimens were collected by shaking Brotia shells in a bucket or dipnet. Captive M. adeia have been observed entering empty shells head first. MugiIogobius adeia has also been observed, but not collected, at Teluk Nuha, a long beach at the northern shore of the lake. Here the lake is very shallow, with a pebbly substrate shelving at an approximately 10% slope to a depth of about 2 rn, the substrate abruptly becoming sand at 2-2.5 m depth. Mugilogobius adeia was observed among the pebbles, close to the pebble/sand borderline, but never on the sand. Empty Brotia shells were abundant, and M. Iatifrons was also present, in 0-50 cm depth. Specimens of M. latifrons I collected in 1989 were all from depths of less than 0.5 rn, often from only a few centimetres depth (M. adeia was not observed). There is apparently some separation of preferred habitat between Mugilogobius adeia and M. latifrons, especially in terms of depth and exposure to wave action. Mugilogobius Iatifrons seems to prefer shallower water, which can be exposed to considerable wave activity. Further field work would be required to confirm this, however. Some X-rayed specimens of M. adeia had a few whole or fragmented tiny short-spired gastropods in the gut. Mugilogobius amadi (Weber, 1913) Figures 5, 16B, 87-88; Tables 5-8, 11 Gobius amadi Weber 1913: 211-212, figure 8 (Posso Lake). Figure 87 Location of central Sulawesi lakes, in which are found Mugilogobius adeia, M. amadi, M. 1atifrons, M. lepidotus sp. nov, M. rexi n. sp. and M. sarasinorum. the northern shore, the lake shore is steep and heavily forested, with overhanging branches reaching the water's surface. The lake bottom is moderately steep, and consists mostly of large boulders (0.5-1 m in diameter). All specimens of Mugilogobius adeia were observed and collected below 1.5-2 m depth. Their body colour blended in well with the thick algae cover on the nearby boulders. The fish were regularly seen to stay close by empty shells of the gastropod Brotia gemmifera (Sarasin and Sarasin), which is endemic to Lake Matano. At Mengonuwai, a shallow bay along the southeastern shore, M. adeia was collected and observed mostly in very shallow water (20-50 cm). The substrate was pebbles and dead leaves, in thick groves of submerged Pandanus roots. The fish were observed to seek shelter in empty Brotia shells. The shells were also used as shelter by M. Iatifrons, Weberogobius amadi: Koumans, 1953: 172-174; Whitten et al., 1987: 43, 46; Miller 1987: 697, figure 7; Whitten et al., 1988: 295; Kottelat 1990a: 49, 64-65; Larson and Kottelat 1992: 226, 233; Nijssen et al., 1993: 232; Kottelat et al., 1993: 154, plate 73. Material Examined Lectotype ZMA 112.664, 1(122), Lake Poso, Sulawesi, Indonesia, E.C. Abendanon, 1909. Paralectotypes Indonesia: Sulawesi: ZMA 121.293, 8(93-124), same data as lectotype; RMNH 14365, 1(111), same data as lectotype. Other Material ZMH 7579, 11(60-88), Lake Poso, Sulawesi Selatan, M. Brembach, 1978; CMK 5774, 4(69-84), same data as preceding. Diagnosis A striking, large species with second dorsal rays Revision of Mugilogobius Figure 88 95 Mugilogobius amadi. Lectotype of Gobius amadi Weber, 129 mm SL, ZMA 112.664, Lake Poso, Sulawesi. 1,9-10; anal rays 1,10-12; pectoral rays 18-20; longitudinal scales 52--65; TRB 14-20; predorsal scales small, 22-36, extending past preopercular margin but not reaching eyes; scales on body mostly ctenoid; third or fourth spine of dorsal fin longest; body relatively compressed and slender; snout and jaws long, mouth terminal; gill opening oblique, pectoral girdle smooth; body and fins plain dark brown to blackish in colour; known only from Lake Poso in central Sulawesi; very rare and possibly extinct. Description Based on 21 specimens, 60-124 mm SL. An asterisk indicates counts of lectotype (Figure 88). First dorsal VI*; second dorsal 1,9-10 (mean 1,10*); anal 1,10-12 (mean 1,10*), pectoral rays 18-20 (mean 19*), segmented caudal rays 14-16 (mean 16, 15 in lectotype); caudal ray pattern modally 9/7; branched caudal rays 12-15 (mean 13*); unsegmented (procurrent) caudal rays 10/9 to 10/ 10; longitudinal scale count 52--65 (mean 57, 59 in lectotype); TRB 14-20 (mean 16, 18 in lectotype); predorsal scale count 22-36 (mean 28, 26 in lectotype); circumpeduncular scales 21-25* (mean 22). Gill rakers on outer face of first arch 6+17 to 8+18 (modally 7+17). Pterygiophore formula 312210* (in 14). Vertebrae 11+15* (in 15). Neural spine of first to third vertebra may be broad, or expanded at tip (in 13). Two epurals (in 15). Three anal pterygiophores before haemal spine of first caudal vertebra (in 15). Metapterygoid slender, with narrow bridge across to quadrate. Fifth ceratobranchial very slender, nearly straight. Body relatively compressed, slender. Head long, deeper than wide, HL 2.7-3.1 (mean 2.9) in SL. Depth at posterior preopercular margin 1.7-2.5 (mean 2.0) in HL. Width at posterior preopercular margin 1.9-2.6 (mean 2.2) in HL. Mouth terminal, chin distinctly protruding in most specimens, often sharply pointed, mouth slightly oblique, forming angle of 20-25° with body axis; jaws reaching to below anterior half of eye or nearly to below mideye. Lips with very fine small fimbriae present along edges of both lips, fimbriae usually more noticeable on lower lip; lower lip free at sides, fused on either side of chin. Upper jaw 2.4-3.3 (3.3 in single female specimen, mean 2.8 in males) in HL. Eyes large, lateral, top of eye sometimes forming part of dorsal profile, 4.0--6.9 (mean 5.2) in HL. Snout long, somewhat flattened, with pronounced bump in centre formed by premaxillary ascending process, 3.3-4.0 (mean 3.6) in HL. Interorbital broad, flattened to slightly convex, 3.4-4.9 (mean 4.1) in HL. Top of head smooth, without fine villi on interorbital and snout region. Body depth at anal origin 3.7-5.8 (mean 4.9) in SL. Caudal peduncle compressed, length 4.5-5.5 (mean 4.9) in SL. Caudal peduncle depth 8.3-10.0 (mean 8.9) in SL. First dorsal fin pointed, no spines filamentous or with free tips; third or fourth spines longest or subequal (fifth spine longest in one specimen); spines reaching to first element of second dorsal fin when depressed. First dorsal spine always shorter than next three. Third dorsal spine length 6.4-7.0 (mean 6.7) in SL. Fourth dorsal spine length 5.7--6.9 (mean 6.3) in SL. Second dorsal and anal fins shortbased, rather pointed, posteriormost rays longest, rays just falling short of caudal fin base when depressed. Pectoral fin narrow, oval to pointed, central rays longest, 4.4-5.6 (mean 4.8) in SL; rays usually all branched but for uppermost. Pelvic fins oval and rather pointed posteriorly, nearly reaching anus, 4.5-6.2 (mean 5.3) in SL. Caudal fin rectangular, truncate, 3.7-5.0 (mean 4.4) in SL. No mental fraenum, chin smooth, usually pointed. Anterior nostril in small tube, placed just at preorbital edge, tube short, oriented down and forward. Posterior nostril small, oval, placed halfway between anterior margin of eye and preorbital edge. Gill opening wide, extending forward to under preopercle, or to just anterior to rear preopercular margin; gill opening oblique, with lower edge of pectoral girdle well anterior to upper edge. Inner margin of pectoral girdle smooth with no bony ridge or fleshy flange or lobes present. Gill rakers on outer face of first arch very short, rather rounded, smooth, two or three longest rakers near angle of arch; rakers on inner face of first arch nearly twice length of those on outer face and equal in size to outer rakers on second and all other arches. Tongue large, flat, blunt to slightly concave, I H.K. Larson Table 11 Character Morphometries as percentages of SL or HL, as indicated, of male Mugilogobius amadi (Weber, 1913) (only one female known). Lectotype Males Males Males Minimum Maximum Mean 35.4 Head length in SL 49.8 Head depth in HL 52.3 Head width in HL 19.7 Body depth in SL Body width in SL 20.9 Caud. ped. 1. in SL Caud. ped. d. in SL 10.7 Snout length in HL 30.5 14.9 Eye width in HL 38.3 Jaw length in HL Interorbital 1. in HL 26.4 17.7 Pectoral 1. in SL 18.7 Pelvic 1. in SL 21.3 Caudal 1. in SL Longest D1 spine in SL 17.3 32.5 40.0 39.0 17.3 10.1 18.3 10.0 25.2 14.5 30.8 20.3 17.7 16.0 16.9 14.6 37.7 57.9 52.3 26.8 13.6 22.3 12.0 30.6 25.2 42.5 29.7 22.9 22.4 27.3 17.5 34.9 50.1 45.5 20.5 11.4 20.7 11.2 27.9 19.5 36.2 24.7 20.8 18.9 22.8 16.0 sometimes trilobed; tongue often sharply folded along longitudinal axis, forming V-shape in crosssection. Outer teeth in upper jaw largest, straight and sharply pointed, three to five rows of very small sharp teeth behind this row; one or two rows at side of jaw. Lower jaw with four or five rows of pointed teeth across front, outermost two rows largest, straight and pointed; lower jaw teeth all pointing posteriorly; usually only one or two rows of teeth at side of jaw. Predorsal scales small, cycloid and evenly sized, usually reaching forward to past preopercular margin, scales never reaching up to behind eyes, usually ending not far in front of rear preopercular margin. Operculum covered with small cycloid scales, area below horizontal papilla row oi naked. Cheek usually naked, lectotype and two other specimens with scattered small cycloid scales on lower rear corner of preopercle and just behind eye respectively. Pectoral base and prepelvic area covered with small cycloid scales. Belly scales all cycloid or weakly ctenoid; several specimens with isolated area of ctenoid scales under pelvics (anterior IA to ｾ of belly), rest of belly scales cycloid. Ctenoid scales on side of body extending forward in narrow wedge up to behind pectoral base or to below first dorsal fin; two specimens with ctenoid scales extending narrowly forward only as far as below second dorsal fin origin (some specimens with ctenii missing from most scales). Male genital papilla long, slender and flattened, with pointed tip. Female genital papilla fairly short, rounded, with slight groove on ventral surface close to opening at tip. Swimbladder present; membrane thin and fragile. Gut short, muscular; 5-bend shape. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 5. Three to four s rows on snout, composed of many small papillae; rows may be wavy or broken up. Broken row c below eye composed of many small papillae (most Mugilogobius species usually with few papillae in rear portion of row c). Papilla rows / on chin formed into two patches of many papillae. Coloration of fresh material No information available, probably plain dark brown with black fins. Coloration of preserved material Head and body very dark brown, interorbital and snout nearly black. Several specimens somewhat lighter brown, with about seven or eight indistinct dark brown mid-dorsal blotches and some irregular dark brown blotches along upper side of body; markings clearest on upper half of body. Dorsals, anal, pelvics and caudal fins entirely black. Pectoral fins with black or dark brown rays; membranes clear at least on distal half, proximal part of membrane dark brown to black. Peritoneum dense dark brown, slightly paler ventrally. Comparisons Mugilogobius amadi differs from other species in the genus by its high dorsal fin counts (modally 1,10 versus 1,7-8), long snout and protruding chin (versus short snout and subterminal mouth) and oblique gill opening with smooth pectoral girdle (versus nearly somewhat oblique to vertical gill opening, usually with lobes or bony flange on the pectoral girdle). Mugilogobius amadi is most similar in appearance to M. sarasinorum, which also inhabits Lake Poso, in that the two species share 11+15 vertebrae, an oblique gill opening and low slender metapterygoid. However, M. sarasinorum has lobes on the pectoral girdle, 1,8 dorsal and anal fin rays, a subterminal to sometimes just barely terminal mouth and a relatively broad, sometimes slightly pointed, snout. Male M. sarasinorum in breeding condition are very dark plain brown (and nonbreeding males and females are fairly plain). Mugilogobius adeia and M. lati/rons from neighbouring Lake Matano also have 11+15 vertebrae and low slender metapterygoids, but are "typical" Mugilogobius in always having subterminal mouths, lobes on the pectoral girdle and smaller size. The ancestor of the Sulawesi freshwater Mugilogobius has apparently undergone three reversals, resulting in species usually having a terminal mouth (character 11), the higher number of pre-anal pterygiophores and low narrow metapterygoid (characters 12 and 13). The vertebral pattern of 11+15-16, as opposed to the more typical pattern of 10+16, also separates this group from other species of Mugilogobius. Revision of Mugilogobius Distribution Specimens are known only from Lake Poso, central Sulawesi, Indonesia (Figure 87). It has not been reported from any of the neighbouring lakes (Whitten et al., 1987; Whitten et al., 1988; Kottelat et al., 1993). Poso is a very deep (450 m) graben lake, formed by tectonic activity (Weber 1913; Whitten et al., 1988). Remarks Weberogobius was established by Koumans (1953), who designated the type species Gobius amadi Weber, 1913. A lectotype is chosen here, from among nine syntypes, a 122 mm SL specimen in good condition. The same specimen was also selected as a potential lectotype by Peter Miller (in a letter to ZMA in 1987), but the designation was not published. Weberogobius was previously considered to be a valid (monotypic) genus by Larson and Kottelat (1992). Weber (1913) discussed the fish species of Lake Poso, the names given to them by fishermen, and the methods of catching them. Mugilogobius amadi was once abundant in Lake Poso and formed an important local fishery (Kottelat, 1990a). Kottelat noted that the species was called bungu by the fishermen of Dulumai village, who formerly subsisted almost exclusively on a bungu fishery. This species has been reported as being nearly extinct (Whitten et aI., 1987; Whitten et al., 1988; Kottelat 1990a; Larson and Kottelat 1992) since about 1985. Whitten et al. (1987) discussed extinctions of various species in Sulawesi, but did not propose a cause for the fish extinctions in Lake Poso. However, they mentioned that fishermen from lakeside villages attributed the decline in W. amadi and two of the endemic ricefish species to the Mount Colo eruption (on Una Una Island in Tomini Bay, in 1983). Whitten et al. (1987) considered this to be unlikely because volcanic ash from the eruption blew to the west rather than south over Lake Poso and so could not have fallen over the lake area, causing fish deaths. Kottelat (1990a) agreed, and considered that the introduction of exotic fishes and associated parasites and diseases was much more likely, based on information given to him by villagers at Pendolo (southern end of Lake Poso. These villagers indicated that the native fish populations began to decline after the introduction in the 1980s of ikan lele (the catfish Clarias sp.). However, in 1989, at Tentena on the northern end of Lake Poso, a villager named Wan told me about the bungu hitam ("black goby" = M. amadi) fishery which used to exist in the lake, and his observations of the species' decline. In the past, the fish could be found in the centre of the lake (in very deep water) during the dry season, and were fished for using very long lines. In the wet season, large numbers of bungu hitam would come in to the shoreline to 97 spawn, where they could be caught easily by everyone, using a variety of nets. The fish were reported as being good and fat. According to Wan, the fish disappeared after the 1983 Mount Colo volcanic eruption and that after the eruption there were a great many earthquakes and earth tremors around Lake Poso. Thousands of bungu hitam (and other fishes) washed ashore in large drifts, and the fish populations have apparently never recovered. It is likely that toxic gases such as sulphur dioxide were released within the lake as a result of the tectonic activity (Krauskopf, 1979; Weigelt, 1989; Megirian personal communication, 1995), which would have quickly killed aquatic organisms (Lake Poso sits across a major fault line). Wan made no mention of any ash cloud. During six days' searching around Lake Poso in 1989, I saw no M. amadi, despite dipnetting, scoop-netting, seining and snorkelling. However, M. amadi's close relative, M. sarasinorum, was observed to be common in the shallows at night. If M. amadi still exists, it probably does so as small scattered populations in deeper water. Lake Poso, like all the tectonic lakes of central Sulawesi, is very deep and steep-sided and almost nothing is known of the fish fauna in deeper waters (Kottelat 1989b). It is very likely that the populations of M. amadi and several species and genera of endemic buntingi (ricefish), damaged by the seismic activity, are having difficulty re-establishing due to the additional pressure of the introduced predatory fishes (Clarias, Channa and Osphronemus) and their parasites. Mugilogobius cagayanensis (Aurich, 1938) Figures 89-92; Tables 5-8, 12 Vaimosa cagayanensis Aurich, 1938: 169-170 (Cagayan Sulu, Singuan Lake and Sulu Lake); Koumans 1953: 156 (as synonym of Tamanka latifrons); Ladiges et aI., 1958: 166 (as synonym of Tamanka latifrons). Tamanka cagayanensis: Herre 1953b: 766. Material Examined Lectotype ZMH 420a, 1(25), Cagayan Sulu, Philippines, Woltereck, Sta.K173, Wallacea Expedition, 14 May 1932. Paralectotypes Philippines: ZMH 420b, 2(18.5-20), same data as lectotype. Other Material Philippines: ZMH 746, 3(22-26), Singuan Lake, Cagayan Sulu, Wallacea Expedition, 1932; USNM H.K. Larson 98 122921, 99(18.5-62), Ernestine Lake, Cagayan Sulu, RV Albatross Philippine Expedition 1907-9, 8 January 1909. Diagnosis A large Mugilogobius with adult males mostly plain dark brown in colour and females with indistinct mottling and oblique bars posteriorly on light brown background; soft dorsal rays 1,7-8, anal rays 1,7-9, pectoral rays 15-18, lateral scales 32-42, TRB 10-16, 14-24 small predorsal scales and 12-19 circumpeduncular scales; gill opening oblique, usually with few small round knobs on edge of pectoral girdle; restricted to freshwater lakes (Lakes Ernestine, Singuan, Sulu) on Cagayan Sulu Island, Cagayan Sulu Archipelago, southern Philippines. Description Based on 33 specimens, 18.5-62.0 mm SL. Counts of lectotype (Figure 89) indicated by asterisk. First dorsal VI*; second dorsal 1,7-8* (mean 1,8); anal 1,7-9 (mean 1,8*); pectoral rays 15*-18 (mean 16); segmented caudal rays 15 to 18 (mean 16*); caudal ray pattern modally 9/7; branched caudal rays 7/6 to 10/7 (modally 8/7); unsegmented (procurrent) caudal rays 7/8 to 8/8; longitudinal scale count 32-42 (33 in lectotype, mean 37); TRB 10-16 (12 in lectotype, mean 14); predorsal scales 14-24 (21 in lectotype, mean 19); circumpeduncular scales 12-19 (12 in lectotype, mean 15). Gill rakers on outer face of first arch 3+8 to 5+9 (modally 4+9). Dorsal pterygiophore formula 3-12210 (in 20), 311310 (in three). Vertebrae 10+15 (in four), 10+16 (in 32). Neural spines of first three vertebra broad and expanded dorsally. Metapterygoid broad, expanded dorsally, forming distinct bridge to quadrate. Fifth ceratobranchial long and slender; tall, rounded flange present on ventral surface. Two epurals (in 32), one in one specimen. Two (in 27) or three (in eight) anal pterygiophores before haemal spine of first caudal vertebra. Body rather compressed, more so posteriorly. Head broad, rounded, somewhat depressed Figure 89 anteriorly, always wider than deep, HL 2.6-3.3 (mean 2.9) in SL; profile bluntly rounded. Depth at posterior preopercular margin 1.8-2.2 (mean 2.0) in HL. Width at posterior preopercular margin 1.6-2.0 (mean 1.8) in HL, preopercular area sometimes slightly inflated in males. Mouth moderately large, terminal to slightly subterminal (upper jaw anterior to lower), slightly oblique, forming angle of about 25-30 0 with body axis; jaws reaching from below anterior half (females) to posterior half of eye (males), jaws averaging longer in males than females. Lips fleshy, narrow, smooth, with fine fimbriae on inner edges of both lips; lower lip mostly free, fused near tip of jaw. Upper jaw 2.6-3.0 (mean 2.8 in females, 2.5 in males) in HL. Eyes large, placed dorsolaterally, sometimes forming part of dorsal profile; eye width 3.6-5.6 (mean 4.6) in HL. Snout broad, rounded in dorsal view; snout profile rounded to somewhat pointed; some specimens (usually female) with concave interorbital and snout with raised bump in centre (formed by ascending premaxillary process); males often with rounded to somewhat blunt snout; snout 2.0-4.9 (mean 4.0) in HL. Interorbital broad, flat to concave, 2.9-6.3 (mean 4.1) in HL. Fine dense villi present on naked areas on top of head (interorbital and upper part of snout). Body depth at anal origin 4.7-5.8 (mean 5.2) in SL. Caudal peduncle compressed, length 3.8-4.8 (mean 4.3) in SL. Caudal peduncle depth 7.4-9.0 (mean 8.2) in SL. First dorsal fin rounded, low, usually fourth spine longest; fin often not reaching second dorsal origin when depressed, falling well short of fin origin. Third dorsal spine length 6.5-8.2 (mean 7.8) in SL. Fourth dorsal spine length 6.5-8.4 (mean 7.6) in SL. Second dorsal fin usually taller than first dorsal fin; posteriormost rays usually longer than anteriormost; fin falling well short of caudal fin base when depressed. Anal fin similar to second dorsal fin, posteriormost rays longest, rays falling short of caudal fin base when depressed. Pectoral fin oval, central rays longest, 4.2-6.5 (mean 4.8) in SL; all rays branched but for uppermost. Pelvic fins oval, MlIgilogobills cagayanensis. Lectotype of Vaimosa cagayanensis Aurich, 25 mm SL, ZMH 420a, Cagayan Sulu, Philippines. Revision of Mugilogobius B Figure 90 Mugilogobius cagayanensis, papillae pattern. Specimen from USNM 122921. A, lateral view; H, ventral view of chin area (not to scale). Scales omitted. Scale bar = 1 mm. rays not reaching anus (often only reaching halfway to anus), 4.7-6.6 (mean 5.4) in SL. Caudal fin moderate in size, oval, rounded posteriorly, 3.8-4.7 (mean 4.1) in SL. Chin smooth, without mental fraenum. Anterior nostril in very short tube, placed at edge of upper lip, usually pointing down and forward over lip, preorbital slightly curved outward near nostril. Posterior nostril oval, sometimes with slightly raised rim, placed nearly halfway between preorbital edge and eye, slightly closer to eye. Gill opening somewhat oblique, usually extending forward to under opercle. Inner edge of pectoral girdle with small rounded knobs or bumps (in 21) or smooth (in nine); one specimen with low flange on one side. Gill rakers on outer face of first and second arch rather short, usually with fine spiny papillae on inner faces of rakers on lower limb, rakers longer toward angle of arch; rakers on inner face of first arch slightly longer than those on outer face, similar in size to those of lower limb of second arch, fine spiny papillae at tips; irmer rakers on second arch, outer and inner rakers of third and fourth arches longer, usually finely papillose at tips. Tongue tip concave to blunt. Outer teeth in upper jaw largest, but not particularly enlarged, straight and sharp, evenly spaced along length of jaw; 99 largest teeth across front; behind this row, two to five rows of small sharp curved teeth. Lower jaw with four to six rows of sharp curved teeth across front, two to three rows at sides; outermost row teeth largest and stoutest, oriented nearly upright; innermost teeth evenly sized, tending to point posteriorly. Predorsal scales cycloid, mostly evenly sized, reaching forward to past preopercular margin, extending forward to about halfway between rear of eyes and preopercular margin; scales near nape midline extending further forward than those on sides of nape. Operculum covered with small cycloid scales on upper third to half. Cheek always naked. Pectoral base covered with small cycloid scales. Prepelvic area covered with small cycloid scales. Belly scales cycloid, with patch of weak ctenoid scales anteriorly, close to base of pelvics (ctenoid scales extending third to halfway down belly). Ctenoid scales present posteriorly, extending forward on side of body in wedge (occasionally quite narrow anteriorly) up to behind pectoral base. Genital papilla in male slender, elongate and flattened, broad at base and narrowing toward tip; papilla in female bulbous and stout, narrow toward tip and broad at base, groove present along ventral surface. Head pores absent, as in all Mugilogobius species. Sensory papillae on head in longitudinal pattern, as in Figure 90. All papillae small, close-set, equal in size. Three or four s rows present, each consisting of three or more papillae. Mental f rows of four or five papillae each, rows parallel to row i. Coloration of fresh material Aurich (1938) did not mention live colour of this species. A small watercolour sketch is held at USNM, of a formalin preserved Mugilogobius specimen, with collecting locality given as Singuan Lake, Cagayan Sulu, dated 8 January 1909. The specimen is probably part of USNM 122921, which has the same collection date, but a collection locality of Ernestine Lake, Cagayan Sulu. Aurich (1938) gave Singuan Lake and Sulu Lake as type localities for Vaimosa cagayanensis. The sketch shows a blackish brown fish with white submarginal band on each dorsal fin. The notes next to the sketch say "Body nearly uniform bright blue black. D. white, tip of last with tawny tip ... Vairnosa singuanensis". Coloration of preserved material Aurich (1938) described colour in alcohol as yellowish to reddish brown, head darker, with dark or black markings. Head and body plain light brown to dark brown (darkest specimens usually males); darker dorsally and lighter brown ventrally (Figure 91). Dorsal surface of head, nape and upper part of body with indistinct brown splotches and H.K. Larson 100 Figure 91 Mugilogobius cagayanensis, 54 mm SL, USNM 122921, Ernestine Lake, Cagayan Sulu, Philippines. mottling. Females with four to seven indistinct brown oblique bars or cross-hatched blotches along midside, especially on rear half of body (barring and mid side blotches visible on lectotype). Small specimens (less than 30 mm SL) usually more distinctly marked than adults. Largest mark, if present, brown blotch on middle of caudal fin base. Some specimens with about seven indistinct brown saddles across dorsal midline; saddles sometimes partly connecting with faint oblique bars or blotches on upper half of body. Head with three to five faint, oblique brown stripes extending from eye across preopercle; most distinct stripes running from lower rear edge of eye and along lowermost margin of eye (head markings illustrated by Aurich 1938: 169, figure 22). Opercle and top of head with indistinct brown spotting and vermiculation. All fins brown to dark brown (almost black in males), pectoral fins usually paler than others. Small specimens (30-40 mm SL) with very light dusky fins. First dorsal fin with narrow submarginal white band, tips of fin spines dusky to dark brown; females with fin paler and often with blackish blotch on rear half. Second dorsal fin with similar Table 12 narrow submarginal white or translucent band; some pale blotches anteriorly in middle of fin; vertically elongate brown streaks forming band along middle of fin. Anal fin plain dark brown. Caudal and pelvic fins with translucent to whitish rear margins, or entire fin plain brown; in mature fish, especially males, caudal fin often with vertically aligned rows of brown spots, often coalescing and forming broken-up rows. Peritoneum dark brown, its ventral midline pale with scattered melanophores. Comparisons Mugilogobius cagayanenslS IS very similar in appearance to Tamanka siitensis Herre, 1927, being large and robust, with adult males plain brown in colour. The two can be distinguished by the number of segmented caudal rays (always 17 in M. siitensis and modally 16 (15-18 observed) in M. cagayanensis), and colour pattern on the head (dark brown indistinct moUling on the top of the head only in T. siitensis and three to five dark oblique stripes from eye across cheek in M. cagayanensis). Additionally, Tamanka siitensis always has a smooth pectoral girdle, while M. cagayanensis modally has fleshy lobes or a flange present. Morphometries as percentages of SL or HL, as indicated, of Mugilogobius cagayanensis (Aurich, 1938). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. 1. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital 1. in HL Pectoral 1. in SL Pelvic 1. in SL Caudal 1. in SL Longest D1 spine in SL Lectotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 35.2 30.6 48.7 50.8 17.1 12.5 21.1 11.6 20.3 17.9 36.3 15.9 19.4 16.1 21.3 15.3 38.0 57.1 63.3 21.4 14.1 25.9 13.4 28.6 28.1 45.2 34.0 23.7 21.1 26.3 15.3 35.0 52.7 57.2 19.5 13.4 23.6 12.4 25.4 22.2 40.1 26.6 21.7 18.9 24.9 14.0 32.2 46.3 51.1 17.3 12.4 21.0 11.1 23.5 18.7 33.3 21.4 15.4 15.3 22.8 11.9 36.5 53.7 59.8 21.2 15.3 26.3 13.1 27.6 25.8 38.8 26.3 22.1 19.7 26.5 13.3 34.9 50.3 55.5 19.1 13.8 23.5 12.1 25.0 22.3 35.8 24.0 20.0 18.2 24.2 12.6 21.6 21.6 37.5 19.9 ttau Revision of Mugilogobius 101 . '- I SOUTH CHINA SEA " CELEBES SEA BORNEO .. Ｎｾ Q' Figure 92 The Philippines, showing location of Cagayan Sulu and Jolo Islands in the Sulu Sea, the only known localities of MlIgilogobills cagayanensis and Tamanka siitensis respectively. Distribution Known only from three freshwater lakes on Cagayan Sulu Island, Cagayan Sulu Archipelago, Philippines (Figure 92). Remarks Aurich (1938) considered that his new species more closely resembled Mugilogobius latifrons Boulenger, and he placed it in Vaimosa. Aurich used the phrase "Typ.: la tifrons ", possibly implying that he considered M. latifrons to be similar to his new species, as he used "Typ." also for Vaimosa (Typ. latifrons) karatunensis (= Mugilogobius parvus (Oshima, 1919)). However, it is possible that what Aurich meant by "typ." was his way of distinguishing between the two groups of Mugilogobius: Typ. latifrons being those with small evenly sized predorsal scales and higher numbers of anal and second dorsal fin rays, and Typ. zebra being those with an enlarged predorsal scale and 102 H.K. Larson lower numbers of anal and second dorsal fin rays. Under Vaimosa (Typ. zebra), he described V. zebra (= M. chulae), noting the predorsal scale arrangement. Aurich's definition of Typ. latifrons (1938: 165) is not consistent with that given for Typ. zebra (1938: 170), however. The types are in poor condition. They appear to have become dried out and attacked by fungus at some stage. The largest specimen had been placed in a vial with a label saying "lectotype", but with no indication as to who was responsible for this designation. However, it is likely that the authors of this note were Ladiges et al. (1958) who state that a lectotype and paralectotypes are present in ZMH 420, but they did not state which specimen was the lectotype. To end any confusion the largest specimen, ZMH 420a, 25 mm SL, is hereby designated as lectotype of Vaimosa cagayanensis, and ZMH 420b, two specimens, 18.5-20 mm SL, are paralectotypes. Mugilogobius cavifrons (Weber, 1909) Figures 9E, 18A, 83, 93-101; Tables 5-8,13 ?Mugilogobius villa: Myers 1988: 164; Donaldson et al., 1994: 330. Mugilogobius parous: RandalI et al., 1993: 222, 232233, plate 1H; Fang et al., 1996: 28, 174, plate 102. Mugilogobius karatunensis: Kottelat et al., 1993: 146. Mugilogobius cavifrons: Donaldson et al., 1994: 330. Material Examined Lectotype of Gobius cavifrons ZMA 112.616, 1(27.5), Soela Takoemi di bawa, freshwater, Temate, Indonesia, M. Weber, 1903. Paralectotypes of Gobius cavifrons ZMA 123.468, 38(8.5-28.5), same data as lectotype. Holotype of Glossogobius parvus FMNH 59138, 1(33.5), Kizanto, Taiwan, M. Oshima. Holotype ofVaimosa karatunensis ZMH 421, 1(31), Karatun, Talaud Archipelago, Indonesia, Woltereck, June 1923. Gobius cavifrons Weber, 1909: 152 (Sula Takomi di bawah, Temate, Indonesia); Weber 1911: 36; Weber 1913: 462-463, figure 89; Nijssen et al., 1993: 232. Holotype ofTamanka talavera CAS 36824, 1(34), Nypa swamp, Capiz, Panay, Philippines, A.W. Herre, 3 August 1940. Glossogobius parvus Oshima, 1919: 305, plate 3, figure 3 (Kizanto near Giran, Formosa). ?Tamanka tagala Herre, 1927: 222 (Malabon, Rizal Province, Philippines). Vaimosa karatunensis Aurich, 1938: 168 (Karatun, Talaud Archipelago, Dutch East Indies). Para types ofTamanka talavera CAS 36825, 10(11.5-32), same data as holotype. Holotype of Tamanka philippina CAS 39884, 1(30), Hacienda Waterous, Mangarin, Mindoro, Philippines, A.W. Herre, 20-22 July 1940. Gobius tagara Tomiyama 1936: 74 (lapsus). Tamanka tagala: Koumans 1940: 188. Tamanka talavera Herre, 1945b: 4 (Capiz, Panay, Philippines); Herre 1953b: 766. Tamanka philippina Herre, 1945c: 75 (Hacienda Waterous, Mangarin, Mindoro, Philippines); Herre 1953b: 766. Tamanka cavifrons: Koumans 1953: 155-158, figure 38. Vaimosa karatunensis: Ladiges et al., 1958: 166 (as questionable synonym of V. cavifrons Weber). ?Vaimosa tagala: Kami 1975: 118. ?Vaimosa villa: Kami 1975: 118. Mugilogobius tagara: Hayashi et al., 1981: 10, plate 7, figure 127 (lapsus). Mugilogobius tagala: Akihito et al., 1988: 268. ?Mugilogobius tagala: Myers 1988: 164; Donaldson et al., 1994: 330. Other Material Taiwan: LICCP 1967194, 3(40.5-44), Tangchiang, Ping Tung Prefecture, 15 June 1967; ZMH 7992, 18(19-40), Takao, Schwinghammer, Jan.-Feb. 1908. Japan: Iriomotejima: URM P.6686, 37(18-38), Hoshidate, H. Senou and Suzuki, 28 December 1978; YCM P.8579, 47(20-37), Sumiyoshi, 25 December 1978. Philippines: CAS 38632, 8(26-35), Hacienda Waterous, Mangarin, Mindoro, A.W. Herre, 20 July 1940; USNM 316190, 4(22-31), SEAFDEC Fishery Station, Lleganes, E of Hoilo City, Panay, T. Roberts, 6 April 1976; ZMH 7578, 2(28-34.5), Lake Danao, Camotes Is., Cebu, J. N. Jumalin, October 1962. Guam: NTM 5.13626-001, 14(12-33), mangroves at Laguas River, H.K. Larson, 7 July 1992. Micronesia: NTM 5.13482-001, 22(10-32), Insrefusr Stream, Kosrae, H.K. and J.A. Larson, 2 July 1992. Hawaii, Oahu: BPBM 33930, 25(23-46), drainage canal, Ala Moana Park, B. Nagada and R. Kam, 10 December 1989; BPBM unregistered, 2(14-31), mangroves at Hawaii Institute of Marine Revision of Mugilogobius Figure 93 103 Mugilogobius cavifrons. Lectotype of Gobius cavifrons Weber, 27.5 mm SL, ZMA 112.616, Ternate, Indonesia. Biology, Coconut Island, Kaneohe Bay, J. Randall and J. Smith, 25 August 1989. Indonesia: ZMK unregistered, 34(11-24), Ohoithiet, Kai Dulah east coast, Kai Islands, 3 May 1922; USNM 99609, 1(43), Limbe Island [Sulawesi], RV Albatross, 12 November 1909; CMK 8968,9(15-24), mangroves, Benoa Bay, Bali, A.J. Whitten, 1 January 1992; CMK 8988, 2(17.5-22), Sukamade estuary, Meru Betiri, SE Java, A.J. Whitten, 13 November 1991; ZMA 113.001, 1(40), from Buitenzorg Museum, no date; ZMA 112.617, 2(25.5-36), freshwater near Ampenam, Lombok, P.N. van Kampen, 26 January 1907. Papua New Guinea: ZMK unregistered, 21(7-20), Papitalei Lagoon, SE of Lombrum, Manus Island, Bismarck Archipelago, Noona Dan station 70, 29 June 1962; ZMK unregistered, 16(11-32.5), dammed lake near Lombrum, Manus Island, Bismarck Archipelago, Noona Dan expedition 1961-2, 29 June 1962. Other material eXilmined (but not used in description) Japan: ZUMT 10987, 1, Ryukyu Islands; URM P.4857, 1, Sumiyoshi, Iriomotejima; URM P.4420, 9, Sumiyoshi, Iriomotejima; URM P.6688, 5, Sumiyoshi, Iriomotejima; URM P.6685, 1, Yonada River, lriomotejima; NTM 5.12136-002, 13, Sumiyoshi, Iriomotejima; URM P.6689, 2, Yonada River, Iriomotejima. Philippines: USNM 112842, 116, Hoilo, Panay; USNM 99598, 1, Iliolo, Panay; USNM 99589, 1, Nabatas Luzon fishpond, Luzon; CAS 46517, 1, Siluag Island, Sulu; USNM 99591, 2, Uanjan River, Mindoro; CAS 46509, 1, Dumaguete, Negros Oriental; CAS 38625, 1, Capiz, Panay; USNM 316187, 1, Maribago Mactan, Cebu; AMS 1.30309-003, 3, Mangarin, Mindoro; CAS 46512, 1, Siluag Island, Sulu. Indonesia: ZMA 112.618, 1, Dobo, Aru Islands; USNM 99610, 3, Limbe Island [Sulawesi]; BMNH 1913.12.9.154-5, 1, Aru Islands; NTM 5.14147-009, 1, Sejorong River, Sumbawa. Papua New Guinea: USNM 143658, 3, Los Negros, Admiralty Islands; USNM 143651, 7, Los Negros, Admiralty Islands. NO DATA: RMNH 14347, 1. Diagnosis A moderately sized Mugilogobius with second dorsal rays 1,6-9; anal rays 1,7-9; pectoral rays 1318; longitudinal scales 30-52; TRB 9-20; 12-20 circumpeduncular scales; predorsal scales 12-27, small, usually reaching forward part-way between preopercular margin and rear of eyes; scales on body mostly ctenoid; body light brown or grey, and distinctively marked with nine or more narrow oblique or vertical bars on upper sides, bars interspersed with blotches and chevron-shaped markings, and at least three oblique to vertical dark bars across caudal fin, bar closest to fin base always darkest; distributed from southern Japan throughout the SE Asian archipelago to insular Papua New Guinea. Description Based on 123 specimens, 14-44 mm SL. An asterisk indicates counts of lectotype of Gobius cavifrons (Figure 93). First dorsal V (in nine), VI* (in 112), VII (in two); second dorsal rays 1,6-1,9 (mean 1,8*); anal rays 1,79 (mean 1,8*), pectoral rays 13-18 (mean 16*), segmented caudal rays 15*-17 (mean 16); caudal ray pattern 7/6 to 10/7 (8/7 in lectotype); branched caudal rays 12-17 (mean 15*); unsegmented (procurrent) caudal rays 5/6 to 7/8; longitudinal scale count 3D-52 (mean 39, 35 in lectotype); TRB 920 (mean 13, 11 in lectotype); predorsal scale count 12-27 (mean 19, 17 in lectotype); circumpeduncular scales 12-20 (mean 15*). Gill rakers on outer face of first arch 2+6 to 5+6 (modally 3+61). Pterygiophore formula 3-12210 (in 26), 3-12211 (in one), 3-11310 (in two), 3-11311 (in one). Vertebrae 9+16 to 11+16 (modally 10+16). Neural spine of second and third vertebra (rarely first) expanded at tip (in 11), or narrow and pointed (in 16). Two epurals (in 27). Two (in 23) or three (in 10) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid broad, expanded dorsally (Figure 94). Body compressed posteriorly, rounded anteriorly. Head always wider than deep, HL 2.8-4.6 (mean 3.3) in SL; cheeks sometimes inflated in adult males. Depth at posterior preopercular margin 1.5-2.0 (mean 1.7) in HL. Width at posterior preopercular margin 1.1-1.6 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of 2530° with body axis; jaws generally reaching to below posterior half of eye in males and to below anterior half to mid-eye in females (to below 104 Figure 94 H.K. Larson Jaws and suspensorium of Mugilogobius cavifrons, male, ex BPBM 33930, Ala Moana Park, Oahu, Hawaii. Scale bar = 1 mm. posterior half of eye in lectotype). Lips externally smooth, fleshy fimbriae often present on inner edges of upper lip and front of lower lip (fimbriae sometimes very close to outer edge); lower lip free at sides, fused across front. Upper jaw 1.8-3.7 (mean 2.7 in females, 2.3 in males) in HL. Eyes lateral, high on head, top usually forming part of dorsal profile, 2.5-5.4 (mean 3.9) in HL. Snout rounded, 3.0-5.2 (mean 3.8) in HL. Interorbital broad, flat, 1.4-4.8 (mean 3.3) in HL. Top of head from unscaled area behind eyes to interorbital and snout usually densely covered with fine villi (sometimes tips bifid), villi occasionally present on upper part of preopercle and on suborbital. Body depth at anal origin ＴＮｾＸ (mean 5.3) in SL. Caudal peduncle compressed, length 3.6-5.5 (mean 4.3) in SL. Caudal peduncle depth 6.0-9.7 (mean 7.4) in SL. First dorsal fin low, rounded, tips of spines usually free, third and fourth spines longest or subequal; spines slightly longer in males than females; spines always falling short of second dorsal origin when depressed (gap of one to three scales usually present). First dorsal spine usually shorter than next three; if longest spine, its length 3.6-7.2 (mean 5.4) in SL in females, 5.1 in SL in one male. Second dorsal spine length 6.0-8.4 (mean 7.5 in females, 6.5 in males) in SL. Third dorsal spine length 6.2-9.4 (mean 7.4) in SL. Fourth dorsal spine length 6.2-9.3 (mean 7.6) in SL. Second dorsal and anal fins low, posteriormost rays longest, rays not reaching caudal fin when depressed. Pectoral fin rounded, central rays longest, 3.7-6.3 (mean 4.6) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, reaching about halfway to anus, 4.2-8.0 (mean 5.4) in SL. Caudal fin rounded to oval, 3.1-4.6 (mean 3.9) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, placed at edge of upper lip, tube short, oriented down and forward, preorbital curved forward to accommodate nostril. Posterior nostril oval, placed close to anterior centre margin of eye. Gill opening reaching to lower pectoral base or extending forward to under opercle. Inner edge of pectoral girdle smooth with no knobs or flange (in 11), with low irregular fleshy ridge or flange (in 17), or with distinct fleshy knobs or flaps (in 74). Gill rakers on outer face of first arch short and smooth, longest rakers near angle of arch; rakers on inner face of first arch not much longer than those on outer face; inner rakers on other arches nearly twice the length of first arch inner rakers. Tongue tip blunt to concave, occasionally rounded. Outer teeth in upper jaw largest, evenly spaced, stout and curved, three to four rows of small sharp teeth behind this row; usually only two rows present at side of jaw, outermost row teeth usually largest in males. Lower jaw with five or six rows of small pointed teeth anteriorly, outermost row oriented upright, inner rows all pointing posteriorly; innermost row teeth largest and stoutest, no individual teeth particularly enlarged; usually one or two rows of teeth at side of jaw. Predorsal scales small, evenly sized, very rarely with one scale somewhat larger than others, usually reaching forward to part-way between preopercular margin and rear of eyes; rarely, scales reaching only to above preopercular margin. Operculum with small cycloid scales on upper half to two-thirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1,4 to 1h of belly), remaining scales cycloid. Ctenoid scales on side of body in broad wedge extending forward to up behind pectoral fin; rarely in narrow wedge of ctenoid scales or with wedge broken, with separate patch of ctenoid scales behind pectoral fin (as in holotype of Glossogobius parous). Genital papilla in male elongate, flattened, narrowing toward tip; in female, short, rounded, bulbous to conical. ···········@····O·······1····:· 0·.. . ; '. .',' Figure 95 .'8 . Mugilogobius cavifrons, papillae pattern. URM P.6686. Scalation omitted. Scale bar = 1 mm. Revision of Mugilogobius 105 A '. Q / ｜ｾ @j).\ , \.... . ". I \ ,- . \ \ ... .. .. 8 '.' '" f . \ .. c , ·f ｾ ... , ... , -'f I ( Figure 96 \ \ MlIgilogobills cavifrons. A, papillae pattern on dorsal surface of snout and interorbital (YCM P.6686); B, mental papillae (YCM P.6686); C, mental papillae (ZMK unregistered), Not to scale. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 95. Papillae on head all small, evenly sized. Three s rows on snout, of three to nine papillae each; mandibular f row present on each side of chin, of about three papillae each; broken cheek row c with wide gap (Figure 96). Coloration of fresh material Live specimens grey to light grey to yellowish or light brown, depending on substrate on which they were found, with dark grey to blackish bands and fin markings, with dense black, white-bordered spot on first dorsal fin. An illustration in Akihito et al. (1988: plate 247R) shows a pale grey fish with black and grey markings. Freshly killed specimen from Taiwan (from slide by 1-5. Chen) pale yellowish grey with grey to dark grey bars and vermiculation, dorsal surface of head and body dull yellow, abdominal region whitish. First dorsal fin with broad black central band, widest posteriorly, lower part of fin yellowish white, outer third of fin bluish white with several yellow spots posteriorly. Second dorsal fin yellowish grey with bluish white submarginal band (yellow at rear tip of fin), dense black vertical streak between each fin spine along centre of fin. Anal fin yellow with broad submarginal grey band, margin white. Caudal fin grey with centre of fin yellow alternating with grey irregular vertical bars, margin of fin narrowly bordered with white. Pelvic fins white. Pectoral fins translucent light grey. Rather dark colour photograph in Randall et al. (1993: plate I, figure H) shows a grey fish with narrow, diffuse black bars on body, about five irregular, vertically oriented rows of black lines on caudal fin, distinct black spot on rear half of dusky grey first dorsal fin with yellowish white submarginal band present and dusky grey second dorsal fin with yellowish white submarginal band increasing in width posteriorly and vertically oriented black streak on membrane, in centre of fin, between each fin ray (no head, nape or shoulder bar pattern discernible from this photograph). Coloration of preserved material Head and body yellowish grey to light brown (depending on history of preservation); side of body with nine or more dark brown to blackish oblique bars, elongate blotches or chevrons across upper half of body; alternate bars sometimes joining across dorsal midline (Figures 97-98). Lower half of body relatively plain. No obvious sexual dichromatism, other than some males (probably in breeding colour) being generally darker. Anteriormost dark bar short, sometimes darker than others, extending obliquely from close behind pectoral base up to rear edge of opercle. Second bar beginning at midbase of second dorsal fin, extending down and forward obliquely, ending midlaterally. Remainder of dark bars, blotches and chevrons equally spaced along side of body; bars oriented vertically or angled backward (or alternating obliquely forward and backward). Short upright brown streaks and/or variably shaped blotches often present between all body bars; margins of scales between bars, especially toward midlateral part of body, sometimes narrowly outlined with brown, giving diffuse reticulate 106 H.K. Larson Figure 97 Mugilogobius cavifrons, 27 mm SL, BPBM 33930, Ala Moana Park, Oahu, Hawaii. Figure 98 MlIgilogobius cavifrons, 32.5 mm SL, NTM 5.13626-001, Laguas River, Guam. pattern to background. At caudal base, dark brown chevron or oblique to vertical bar present just before hypural crease, apex of chevron facing anteriorly; bar extending dorsally over upper procurrent rays of caudal fin. Top of head and snout with irregular vermiculate network of light to dark brown lines or blotches; lines often joining across interorbital space. Side of head with two to four horizontal to oblique diffuse to dark brown and dense stripes. Uppennost stripe short, oblique, running from rear of eye up to nape, joining vermiculation on top of head. From lower rear margin of eye, short dark brown stripe present, running obliquely across rear of cheek, usually ending at point halfway down rear preopercular margin. Occasionally, diffuse brown short stripe visible at lower edge of eye, curving back to end on cheek. Lowermost head stripe usually diffuse and broad, or partly broken up, running from corner of mouth to reach edge of preopercle. Opercle with horizontal to slightly oblique streak on upper half, lower half plain dusky. Lips plain dusky, or with pencil-thin dark line along edges. Underside of head plain dusky; sometimes thin or diffuse brown lines visible following lower lip margin, preopercular margin and skin folds across isthmus, one or two lines may cross branchiostegal membranes. Pectoral base light brown, with diffuse to distinct, horizontally elongate brown spot on upper half. Lower half of body, breast and ventral edge of caudal peduncle light brown to yellowish brown or whitish. First dorsal fin with lower two-thirds dusky grey to brownish grey, with large dense black round to oval spot on central rear half of fin; black spot sometimes present on widest and darkest reannost portion of dark grey streak running along centre of fin. Outer third of first dorsal fin white to translucent, with brownish to dusky grey narrow margin; tips of fin spines usually dusky. Second dorsal fin dusky grey with narrow dark margin, submarginal translucent to whitish band present, and row of short vertical blackish streaks and oval spots on each membrane between spines; streaks and spots often fanning rows of black blotches. Anal fin evenly dusky brown to grey, usually with narrow translucent to whitish margin; sometimes indistinct vertical blackish streaks on membranes visible. Pectoral and pelvic fins evenly dusky grey; pelvics often darker than pectorals. Caudal fin pale to dusky, with three to five vertically to obliquely oriented broad brown to blackish bars occasionally formed by rows of vertically oriented elongate blotches; very dark bar closest to fin base most conspicuous marking on caudal fin, occasionally bar no darker than others. Comparisons This species may possibly be confused with M. platystomus and they can be most easily separated by caudal fin colouring. Mugilogobius cavifrons has three to many vertical or oblique dark bars crossing the caudal fin, with the bar closest to the fin base the darkest, but not fonning two spots, while M. platystomus has two distinct dark oval spots on the caudal base and the caudal fin is relatively plain, with indistinct narrow dark streaks along the fin rays. Revision of Mugilogobius Table 13 107 Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius cavifrons (Weber, 1909). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped.l. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbitall. in HL Pectorall. in SL Pelvic I. in SL Caudall. in SL Longest D1 spine in SL Lectotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 30.5 54.8 72.6 18.2 10.5 21.5 13.5 23.8 28.6 47.6 27.4 19.3 19.3 27.3 14.2 21.7 51.1 64.4 14.7 6.8 18.3 10.3 21.1 18.4 27.3 25.4 15.8 12.5 22.3 11.6 35.4 68.2 86.2 22.3 17.3 28.1 16.7 33.6 39.5 56.9 69.7 26.3 23.0 32.8 16.0 31.2 58.0 74.1 19.3 12.6 23.3 14.1 27.2 25.1 43.7 32.9 21.8 18.9 26.4 13.6 27.3 53.3 63.6 15.1 9.3 19.6 11.4 19.3 23.1 34.2 21.1 18.1 16.0 26.6 11.8 35.0 62.8 92.2 21.6 14.8 27.4 15.0 33.6 31.8 45.1 35.4 26.8 23.7 23.3 30.6 29.9 58.3 73.0 18.9 12.3 23.4 13.5 26.1 26.8 37.8 30.0 21.9 19.0 25.4 28.2 Distribution Remarks Specimens are known from the Yaeyama Islands, southern Japan and Taiwan throughout the Philippines and Micronesia to Indonesia and islands off Papua New Guinea; and has been introduced in Hawaii (Figure 83). There are two forms represented here, differing in distribution and mean scale counts: cavifrons from Indonesia, Micronesia, Sulawesi, and the Admiralty, Kei and Am Islands, having 31-40 lateral scales (mean and mode of 36); and parvus from Taiwan, Iriomotejima and the Philippine Islands (Figure 99), with 35-52 lateral scales (mean 41, mode 42). The number of predorsal, TRB and circumpeduncular scales differ slightly between the two forms, as do first dorsal fin spine lengths, all with considerable overlap: Mugilogobius cavifrons: circumpeduncular scales 12-17 (mode 15), predorsal scales 12-23 (mode 18), TRB 9-15 (mode 12), longest first dorsal spine is second to fourth spine; Mugilogobius parvus: circumpeduncular scales 1420 (mode 16), predorsal scales 17-25 (mode 19), TRB 11-18 (mode 13), longest first dorsal spine is third to fourth spine. Initially, it was considered that the two forms represented different species. However, there do not appear to be any other consistent differences between the two such as colour pattern or dorsal Ecology This species is known mostly from brackish to freshwater habitats: mangroves, Nypa swamp, coastal lakes and streams. The specimens from Guam came from muddy pools left among broken coral rock at the mouth of a mangrove-lined river. The Kosrae specimens were from isolated salty (to taste) pools in a sandy-bottomed coastal stream running through mangrove forest. Hawaiian specimens were first found at Coconut Island, Kaneohe Bay, Oahu, in mangrove habitat (Randall et al., 1993). The species also occurs in muddy aquaculture ponds (varying from fresh to brackish) and in a concrete drainage canal at Ala Moana, Honolulu. About two out of ten fish survived being placed directly into a freshwater aquarium (B. Alexander, in Randall et al., 1993). Figure 99 MlIgilogobius cavifrons. Holotype of Glossogobius parous Oshima, 34.5 mm SL, FMNH 59138, Taiwan. Light area at mouth is air bubble. H.K. Larson 108 Figure 100 Mugilogobius cavifrons. Holotype of Vaimosa karatunensis Aurich, 31 mm SL, ZMH 421, Karatun, Talaud Archipelago, Indonesia. spine length etc., that can be used to separate the populations. Without knowledge of its provenance, it is often difficult to assign a given specimen to one fonn or the other, due to the overlap in scale counts. Therefore, the two fonns are placed together here under a single species, albeit with some misgivings. The original description of Gobius cavifrons Weber, 1909, was based on more than one specimen, but the number of fish examined was not stated by Weber, and he did not designate a holotype. Weber (1913) later stated that he had 44 specimens. Of the 44 syntypes that apparently were originally in ZMA 112.616, four were loaned to Peter Miller in May 1987 from ZMA (these four were not examined for this paper). A syntype was figured in Weber (1913: 462, figure 89) and reproduced in Koumans (1953: 155, figure 38, as Tamanka cavifrons). None of the available types presently look particularly like the figured specimen. The lectotype of Gobius cavifrons Weber, 27.5 mm SL (Figure 93; ZMA 112.616), here designated, was chosen because it was a relatively large adult in good condition. The holotype of Glossogobius parous Oshima, 1919 (Figure 99), has been partly dehydrated at some time, but considering this, it is reasonable condition. Oshima's illustration (1919: plate 3, figure 3) shows the characteristic caudal fin barring and two oblique cheek stripes; the head markings are not visible now on the specimen. Vaimosa karatunensis Aurich was possibly based on more than one specimen, as Aurich gave a range of lateral scale counts (42-43; which could possibly refer just to left and right sides of the body). Only one type specimen is currently present in ZMH 421 (Figure 100). The holotype of Tamanka tagala Herre, 1927 (BSM 820; 31 mm SL), and paratype (BSM 804; 33 mm SL), were destroyed in WWII. The original description was based on the type and cotype, with "two tiny specimens, 13.5 and 21 millirneters long" mentioned as an addition to, but not part of, the description (Herre 1927). Koumans (1940: 188) listed BSM catalogue numbers and the total lengths of the types and stated that they"Agree with the description" (referring to Herre's original description). Tamanka tagala may be Mugilogobius cavifrons or similar species of the genus, as it had high lateral (52) and predorsal (25 to 30) scale counts, unlike members of other related genera with large mouths and broad flattened heads, such as Eugnathogobius. Tomiyama (1936) decided that Tamanka tagala Herre, 1927, was the same species as Glossogobius parous Oshima. He placed it in the genus Gobius, but as this action would have made parvus a homonym of "an American species" (he was referring to Gobius parvus Meek, 1902, a junior synonym of Evorthodus lyricus (Girard, 1858)), he used Herre's name, but misspelled it as Gobius tagara, for what is now recognised as Mugilogobius cavifrons. A specimen from Malabon (CAS/SU 33160), in poor condition, shares the same collection data with the lost types of Tamanka tagala Herre, although it is Figure 101 Mugilogobius cavifrons. Holotype of Tamanka talavera Herre, 34 mm SL, CAS 36824, Capiz, Panay, Philippines. Revision of Mugilogobius 109 too large to be either of them (45 mm SL) and may be mis-Iabelled. This specimen has been tentatively identified as M IlgilogobillS cavifrons. Tamanka talavera Herre (Figure 101) is a typical M. cavifrons. The specimen referred to as M. cavifrons by Akihito et al. (1988: plate 2470) is MllgilogobillS mertoni. The material from Guam, identified as Vaimosa tagala and V. villa in Kami (1975), was not available for study. Mizuno 1989: 563, 576-577; Tan and Tan 1994: 356. MugilogobillS sp.: Bright and June 1981: 109. Mugilogobius tagara: Hayashi et al., 1981: 10, plate 7, figure 127 (lapsus). MugilogobillS hoevenii: Horsthemke 1987a: 22 (misidentifica tion). Mugilogobius zebra: Kottelat et al., 1993: 147, figure 297. Material Examined MllgilogobillS cllUlae (Smith, 1932) Figures 21F, 83, 102-108; Tables 5-8,14 Vaimosa dlUlae Smith, 1932: 260, plate 23 (Koh Samui, Gulf of Thailand); Fowler 1937: 251-252, figure 262 (dubious record); Smith 1945: 540; Koumans 1953: 125 (as synonym of Stigmatogobius llOevenii); Suvatti 1981: 306, figure 127. Tamanka sinensis Herre, 1935b: 288 (Hong Kong); Koumans 1940: 149 (as synonym of Stigmatogobills llOevenii Bleeker); Koumans 1953: 125. VainlOsa valigollva Deraniyagala, 1936: 219, figure 1 (Pt. Pedro, Ceylon); Pethiyagoda 1991: 24-25, 336. Vaimosa zebra Aurich, 1938: 171, figure 23 (Boloang, north Celebes); Koumans 1953: 125 (as synonym of Stigmatogobills hoevenii). Holotype ofVaimosa chulae KUMF 1889, 1(29), brackish pool, Koh Samui, Gulf of Siam, Thailand, L.M. Chitrakam and N.B. Chuay, 21 July 1931. ｳ･ｐｬｾｴ｡ｲ ofVaimosa chulae KUMF 1889, 1(20.5), in same jar as holotype, same data as holotype; USNM 119645, 2(26-32.5), same data as holotype; KUMF 1888, 2(24.3-27.7), rocky pools, Lem Sing shore, estuary of Chantabun River, south-east Siam, L.M. Chitrakam, 28 June 1931. sinensis Holotype ｡ｫｮｭｔｦｾ CAS/SU 31518, 1(29.5), Hong Kong, A.W. Herre, 2 March 1934. Holotype ofVaimosa valigouva BMNH 1936.4.23.1, 1(19.5), Pt. Pedro, N.P. Ceylon (Sri Lanka). Vainzosa hoevenii: Herre 1939: 348. ?Mugilogobius lllzonensis Roxas and Ablan: 1940: 307, plate 6 (Luzon, Philippines). Gobius notospilus: Koumans 1953: 125 (as possible synonym of Stignzatogobills hoevenii). Vaimosa fllsca: Koumans 1953: 125 (as possible synonym of Stigmatogobills hoevenii Bleeker). Paratypes ofVaimosa fusca BMNH 1938.12.1.213-4, 2(19-24), Dumaguete, Negros Oriental, Philippines, A.W. Herre, 24 December 1936; CAS 32985, 12(13.5-30), same data as previous. Stigmatogobius bomeensis (in part): Koumans 1953: 127. Lectotype ofVaimosa zebra ZMH 727, 1(20), creek at Boloang, north Celebes, Woltereck, Indonesia, Wallacea Expedition K.167, 1 July 1932. Stigmatogobius hoevenii: Koumans 1953: 125 (in part); Fowler 1961: 218-219, figure 59; Sterba 1973: 765, figure 941; Suvatti 1981: 203. Paralectotypes ofVaimosa zebra ex ZMH 727, 3(13.5-19.5), same data as lectotype. ?Mugilogobius valigolwa: Koumans 1941: 271. Mugilogobius valigollva: Munro 1955: 241. Mugilogobius abei: Masuda et aI., 1975: 276, plate 91J (misidentified M. chulae). Mllgilogobius elmlae: Hayashi and Itoh 1978: 71, plate 15, figure 41; Akihito et al., 1988: 268, plate 247P; Horsthemke 1986a: 492-496; Horsthemke 1986b: 543-545; Horsthemke 1987a: 22-26; Horsthemke 1987b: 59-60; Kottelat 1989a: 19; Kawanabe and Other Material Thailand: CMK 4830, 3 (29.5-30.5), Ko Similan, Phang Ngna Province, Ottinger, January 1985. Japan: BLlH 1978045, 15(14-24.5), mouth of Miyara River, Ishigakijima, 13 August 1978; YCM 9324, 96(13-38.5), Ishigakijima, 26-27 July 1981. Philippines: ROM 48526, 1(23), closed estuary, Tobuan Creek, Lingayan Bay, D.F. Hoese, 15 April 1980; CMK 10035, 1(34), Calbiga Creek, Baybay, Leyte, J. Margraf, 1991; CMK 10054, 4(18-24), I b H.K. Larson 110 Calbiga Creek, Baybay, Leyte, J. Margraf, 5 May 1991; CMK 10005, 1(22.5), 32 km 5 of Tacloban, Dulag, San Jose, Leyte, M. Kottelat and J. Margraf, 9 July 1993. Malaysia: ZRC 22757-63, 7(28-38), Trengganu, Pulau Redang, freshwater stream about 100 m inland of Pasir Chagar Huntang, K. Lim et al., 25 June 1992. Singapore: ZMH 1344,2(35.5-38), imported aquarium specimens, Meincken, March 1961; CMK 7426, 1(29), K. Lim and P. Ng, 1988. Indonesia: ANSP 64935, 1(32.5), Sumatra, F. Mayer, December 1933; ZMH 1246, 1(35), imported aquarium specimen, Borneo, 1959; CMK 8921, 11(18-30.5), Bali, Lovina, stream at Nirwana Beach Cottages, M. Kottelat et al., 27 March 1992. Micronesia: CAS 76144, 1(31), Aebukuru Village, bomb crater near mangroves, Koror, Palau, A. Fehlmann et al., 1 October 1955; AMS 1.23388-001, 1(20.5), taro swamp, Koror, Palau, G. Bright, 28 July 1977. Other material examined (no data taken) Hong Kong: CAS 60978, 3, Plover Cove, Tolo Channel. Japan: Iriomotejima: URM P.4893, 1, Shiira River; NSMT unregistered, 6; NTM 5.12731014, 5, Yonada River; YCM P.8664, 16, Yonada River; AMS 1.27369-003, 2, Amitori Bay; NTM 5.12110-014, 3, Udara River. Thailand: CAS 51229, 7, Songkhla Channel; CAS 53915, 9, Rayong; CAS 209620, I, Klong Ban Phe, Rayong; URM P.13304, 10, Khung Kraben Bay; URM P.12661, 7, Phuket mangrove. Malaysia: ZSM/KEW 680, 1, Banganan mangrove, Sabah; FMNH 100853, 5, Sandakan, Sabah; FMNH 44933, I, Tenah Merah, Sandakan, Sabah; FMNH 44934, 14, Fish Pond, Mile 2, Sandakan; Sabah; CMK 8486, 4, Pahang. Singapore: CAS 33872, 20, Serangoon; CMK 6010, 3, west coast; NTM 5.13957-012, 16, Sungei Pandan. Philippines: Paratype of Vaimosa jusca, CAS 32986, I, Guinlo, Malampaya Sound, Palawan. CMK 9944, I, Calbiga Creek, Baybay, Leyte; USNM 204946, 4, Ayuquitan Daku River, Negros Oriental; USNM 316188, 4, Calasiao River, Pangasinan, Luzon; USNM 264949, 7, Mojon Creek, near Dumaguete Airport, Negros Oriental; USNM 354445, 7, Siquijor Island, Negros Oriental; CAS 38638, 144, Zamboanga, Mindanao; CAS 69657, 6, Hacienda Waterous, Mangarin; CAS 38631,3, Dumaguete; CAS 38634, 2, Old Ayukitan, Dumaguete, Negros Oriental; AMS 1.30309-002, 1, Mangarin, Mindoro; CAS 38633, 9, Estancia, Panay; CAS 69806, 21, Coron, Busuanga; CAS 46385, 4, Batangas, Luzon; AMS 1.21896-001, 5, Tobuan Creek, Lingayan Bay; USNM 57928, 1. Indonesia: RMNH 16805, 1, Batavia; CMK 15113, I, Bungus Bay, Sumatra. Micronesia: CAS 54739, 16, Airai, Babelthuap, Palau. Diagnosis A moderate sized Mugilogobius with second dorsal and anal rays 1,6-8; pectoral rays 12-16; longitudinal scales 25-32; TRB 8-13; circumpeduncular scales 9-13; predorsal scales 11-15, extending up to behind eyes, with anteriormost scale or two distinctly larger than remainder; scales on body mostly ctenoid; some dorsal fin spines elongate and filamentous in males, less so in females, first spine longest in females, second to third spines longest in males; body with at least six oblique black bars across greyish background and pair of black spots at caudal base, first dorsal fin with dense black spot on rear half; widely distributed in Indo-west Pacific, in fresh to brackish waters. Description Based on 59 specimens, 14-38.5 mm SL. An asterisk indicates counts of holotype of Vaimosa chulae. First dorsal V (in three), VI* (in 56); second dorsal 1,6-1,8 (mean 1,7*); anal 1,6-8 (mean 1,7*), pectoral rays 12-16 (mean 14, 15 in holotype), segmented caudal rays 14-17 (mean 16, 15 in holotype); caudal ray pattern modally 9/7 (8/7 in holotype); branched caudal rays 13-16 (mean 15*); unsegmented (procurrent) caudal rays 6/6 to 7/8 (modally 7/7); longitudinal scale count 25-32 (mean 29*); TRB 8-13 (mean 10*); predorsal scale count 11-15 (mean 12*); circumpeduncular scales 9-13 (mean 12*). Gill rakers on outer face of first arch 3+8 to 5+9 (modally 4+9). Pterygiophore formula 312210 (in 14), 3-11310 (in one). Vertebrae 10+15 to 11+16 (modally 10+16). Neural spine of second and third vertebra expanded or thickened at tip (in four), or stout and pointed (in seven). Two (in 18) or one (in two) epurals. Two (in 13) or three anal (in eight) pterygiophores before haemal spine of first caudal vertebra. Body compressed, less so anteriorly. Head wider than deep, often broad and depressed in males, HL 2.9-3.7 (mean 3.4) in SL; cheeks sometimes inflated and muscular. Depth at posterior preopercular margin 1.0-1.9 (mean 1.5) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.4) in HL. Mouth subterminal to barely terminal (upper jaw extending just over lower), oblique, forming angle of about 25-30° with body axis; jaws generally reaching to below posterior half of eye in males and to below anterior half of eye in females. Lips usually smooth, fleshy fimbriae present on inner edges of upper lip and front of lower lip (fimbriae sometimes very close to outer edge); lower lip free at sides, fused across front. Upper jaw 1.8-3.0 (mean 2.8 in females, 2.1 in males) in HL. Eyes lateral, high on head, usually forming part of dorsal profile, 2.8-4.4 (mean 4.2) in HL. Snout rounded, 3.0-4.2 (mean 3.6) in HL. Interorbital broad, usually flat, sometimes concave in large males, 2.2-4.6 (mean 3.8) in HL. Top of head usually covered with fine villi from just behind eyes up to tip of snout (villi density Revision of Mugilogobius variable; often not visible in specimens without well-preserved mucous coat). Body depth at anal origin 4.5-5.6 (mean 5.0) in SL. Caudal peduncle compressed, length 3.3-5.8 (mean 3.8) in SL. Caudal peduncle depth 6.2-8.2 (mean 7.1) in SL. First dorsal fin with second to fourth spines free and often greatly elongate in males, first three spines sometimes elongate in females but always comparatively much shorter; in males, spines sometimes reaching past end of second dorsal fin base when depressed. First dorsal spine always shorter than next three; second and/ or third spine usually longest. Second dorsal spine length in males 1.8-5.9 (mean 3.4) in SL; in females, second dorsal spine length 3.1-6.7 (mean 4.6) in SL. Third dorsal spine length in males 2.0-6.1 (mean 3.6) in SL; in females, third dorsal spine length 3.2-7.2 (mean 5.0) in SL. Second dorsal and anal fins with posteriormost rays longest, dorsal rays longer than anal rays, and often reaching onto caudal fin when depressed; fins more pointed and posterior rays longer in males than females. Pectoral fin oval, central rays longest, 3.7-5.7 (mean 5.6) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, may reach half to two-thirds of distance to anus, 4.3-6.1 (mean 5.1) in SL. Caudal fin rounded, somewhat rectangular in shape, 3.34.5 (mean 3.8) in SL. No mental fraenum, chin smooth. Anterior nostril A Figure 102 Jaws and suspensoria of Mugilogobius clzulac, ex URM P.9324, lshigakijima, Japan. A, male; B, female. Scale bar = 1 mm. 111 tubular, placed just behind upper lip, tube short, oriented down and forward, preorbital curved forward slightly to accommodate nostril. Posterior nostril round to oval, placed close to just above centre anterior margin of eye. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle rarely smooth with no ridge or flange (in four) or with low irregular to smooth fleshy ridge or flange (in three); most specimens (in 41) with distinct fleshy knobs and flaps present. Gill rakers on outer face of first arch very short, longest one or two rakers by angle of arch; outer rakers on other arches short, with few spiny papillae; rakers on inner face of first arch slightly longer than those of outer; inner rakers on other arches usually longer than first arch inner rakers, with small spiny papillae on sides. Tongue tip blunt to concave. Teeth sexually dimorphic (Figure 102). In males, upper jaw with three to four rows of stout, curved, sharp teeth, outer row largest; two or three rows at sides of jaw. In females, outer row in upper jaw largest, consisting of curved sharp teeth; three to four rows of tiny sharp teeth behind this row; two to three rows of small teeth at end of jaw. In males, lower jaw with four to six rows of stout, curved teeth anteriorly; teeth of innermost two rows tending to be larger and stouter (no individual teeth particularly enlarged); usually only one row of teeth along side of jaw. In females, three to five rows of small, evenly-sized pointed teeth in lower jaw; teeth tending to curve inward; usually only one row of teeth along side of jaw. Anteriormost predorsal scale enlarged, placed at rear of interorbital space; sometimes several anterior scales enlarged. Operculum with small cycloid scales on upper half at least. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated area of ctenoid scales under pelvics (covering anterior 1,4 to Yz of belly), rest of belly scales cycloid; sometimes all belly scales cycloid. Ctenoid scales on side of body extending forward in narrow wedge to behind pectoral fin, sometimes ctenoid scales extending above pectoral base, but never to over opercle. Genital papilla in male elongate, flattened, narrowing to pointed tip; in female, short, rounded with blunt tip. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 103. Papilla row p in characteristic "eyebrow" shape; row may be set in groove. Cheek papillae rows all composed of small, close-set papillae; row c broken, with wide gap under eye. Preopercular margin papilla row e not extending far up vertical limb of preopercle. Three 5 rows on snout, consisting of three to six papillae each. Four to six f row papillae present behind mandibular symphysis. H.K. Larson 112 A @ . . . : . . . 0 .:. \ ｾｾｾ［ :' . : ' : : ｾ Singapore mangroves. Horsthemke (1986a-b, 1987a-b) shows live captive fishes displaying. Head and body greyish yellow or pale greenish grey to greyish. Oblique shoulder bar and other body markings very dark brown to dense black; shoulder bar and spot on first dorsal fin always black; paired black spots on caudal fin base always conspicuous. Margins of soft dorsal, anal and lower edge of caudal usually bright translucent bluish white or white. Area directly above spot on first dorsal fin yellowish; bright yellow in fish shown in Kawanabe and Mizuno (1989). Peritoneum white. Large, heavily blotched male (identified as M. hoevenii), shown in Horsthemke (1987a), with deep orange-yellow abdomen, anal fin blue below submarginal dusky stripe and lower half of first dorsal fin orange-white and light-coloured body scales edged with golden yellow. Smith (1932), in his original description of the species, describes his specimens as having blackishbrown markings over a light yellowish-green background, with a dull orange belly and the "muzzle, opercles, and lower jaw dull plumbeous". 0 ::" ." :'.. :." 1: ｾｾ［ . c" 'cp' '. \', \: :'. ......... . ｾＮＮ .... :/ .•..,• .. ...... ';A .f • • B .. • Ｎｾ ... . '.:' , ."0' " '::\•• . , . . ｲｾ ".: " ｟ｾ •• • t .. Figure 103 Mugilogobius chulae, papillae pattern. Paratype of Vaimosa chulae, USNM 119645. A, lateral view; H, ventral view of chin area (not to scale). Scales omitted. Scale bar = 1 mm. Coloration of fresh material A clear photograph of a live specimen is in Kawanabe and Mizuno (1989: 577); the colour in this photograph is very close to that which I observed in live specimens in Thailand and Table 14 Coloration of preserved material Head and body pale yellowish to greyish white (depending on history of preservation), with scale margins usually outlined with dark brown to blackish, giving overall reticulate appearance; side of body with six or seven dark brown to blackish oblique bars, usually joining across dorsal midline (Figure 104). No obvious sexual dichromatism other than males being more heavily marked. Anteriormost bar on side darkest, usually black, running obliquely forward behind pectoral base up to above opercle; sometimes zigzag line or series of small blotches meeting bar on top of head and extending forward into interorbital space. Second bar beginning at midbase of second dorsal fin and Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius chulae (Smith, 1932). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. I. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital I. in HL Pectoral I. in SL Pelvic I. in SL Caudal I. in SL Longest Dl spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 29.3 60.0 80.0 20.3 28.7 52.9 63.5 17.7 10.2 17.3 13.5 23.8 22.8 39.6 28.6 17.6 16.8 22.4 24.9 34.0 67.6 85.7 22.4 16.1 30.0 16.2 33.6 32.7 55.2 44.8 24.4 23.3 30.6 54.5 30.8 58.4 73.0 19.7 13.2 25.6 14.5 29.6 26.9 47.6 35.1 21.5 19.5 27.2 32.0 27.1 52.9 62.3 19.1 11.3 24.2 12.2 23.6 25.3 33.3 21.8 18.6 16.3 22.5 14.9 30.0 67.6 78.1 22.5 16.2 28.4 15.7 32.4 36.4 41.7 37.7 27.2 22.3 30.0 32.1 28.7 58.8 70.2 20.7 13.6 26.4 14.0 27.1 30.1 36.5 30.8 22.2 20.2 26.6 22.4 24.8 16.2 31.8 25.9 54.1 35.3 21.0 17.2 27.6 36.2 Revision of Mugilogobius Figure 104 113 Mugilogobius chulae, CMK 6010, 22.7 and 25 mm SL, Singapore. From colour slide by Maurice Kottelat. extending down and forward obliquely, ending midlaterally. Remaining four to five bars equally spaced along sides of body, usually oriented vertically; lower half of bars often sharply bent posteriorly. Between all body bars, usually short upright brown streaks and/ or variably shaped blotches; margins of scales between bars outlined with dark brown, giving overall reticulate background pattern (pattern may include chevrons and Xs and Ws). At caudal base, dark brown chevron or vertical bar present just before hypural crease (apex of chevron facing anteriorly). Over scales on base of caudal fin, vertically aligned pair of distinct black round spots present; area immediately surrounding spots usually pale, giving ocellate appearance; black spots occasionally indistinct or diffuse. Top of head with irregular network of dark brown lines; network variable, sometimes forming vermiculate blotches or wavy lines joining across interorbital space. Side of head with two horizontal stripes. Uppermost (broader) stripe running from midupper lip around lower edge of eye and across upper preopercle, then abruptly turning obliquely downward, ending on opercle. Lowermost head stripe very narrow and slightly wavy, sometimes partly broken up; stripe running from corner of mouth to reach edge of preopercle. Some indistinct vermiculation and blotches sometimes present on cheek between head stripes. Opercle often with two irregular oblique streaks on upper half (often quite obscure); lower half plain dusky. Snout tip with dark brown vermiculation and spotting. Lips plain dusky, or with pencil-thin darker lines along edge. In males, interorbital space, snout and lips often distinctly dark brownish grey, contrasting with rest of body. Underside of head plain dusky; sometimes thin dark lines visible following lower lip margin, preopercular margin and skin folds across isthmus; several lines sometimes crossing branchiostegal membranes. Most distinct marking wide, diffuse but dense dark grey to blackish band from lower half of each opercle, crossing branchiostegal membranes to join its opposite; band most diffuse on anterior part of breast. Pectoral base light brown, with distinct rounded brown spot on centre of upper half. Abdomen and ventral edge of caudal peduncle pale yellowish to whitish. Breast dusky; duskiness extending up to below lower pectoral base. First dorsal fin with dark grey margin (including fin spines) and two narrow blackish bands. First blackish band approximately median, with large dense black round spot at rear of band; spot placed between third to sixth spines. Second blackish dorsal fin band slightly curved, oblique and incomplete; extending along bases of first three spines. Area below median blackish band dusky grey; area above band translucent to whitish. Second dorsal fin dusky grey with submarginal translucent to whitish band and short vertical blackish streaks and spots on membranes between spines. Streaks and spots may be very intense, forming rows of black blotches. Anal fin evenly dusky brown to grey; sometimes indistinct vertical blackish streaks on membranes visible. Pectoral and pelvic fins evenly dusky grey; centre of pelvics dusky only in pale specimens. Caudal fin usually dusky with vertically oriented rows of fine blackish spots; near fin base, spots often coalescing, forming two to three narrow irregular lines; spots becoming more diffuse toward rear third of fin. Two specimens from the island of Ko Similan, Thailand, have an unusual colour pattern with minimal barring: of the body bars, only the shoulder bar remains, with only one row of blackish blotches along mid-lateral line on one specimen, and no such markings on the other. Both fish retain paired black blotches on the caudal fin base and the overall grey reticulate pattern. A cleared and stained specimen H.K. Larson 114 was aberrant skeletally. These isolated island specimens are quite atypical (the habitat was a landlocked freshwater lake). Comparisons This species is very similar to M. wilsoni sp. novo of northern Australia; see under Comparisons for that species. It is also similar to M. fuscus (see under Comparisons for that species). Distribution This is a relatively abundant species, mostly restricted to the south-east Asian archipelago; ranging from estuaries in the southernmost islands of Japan, Hong Kong, Thailand, Sri Lanka, Singapore, Malaysia, Borneo, Philippines and Indonesia, with Palau being its easternmost point of distribution (Figure 83). In northern Australia, it is replaced by the closely related species, M. wilsoni sp. novo Ecology This species is apparently tolerant of a variety of conditions; specimens have been obtained from a turbid, brackish bomb crater in Palau to a clear freshwater stream in Bali. Hayashi and Itoh (1978: table 1) give some infonnation on food habits of this species from the Shiiugawa River, Ishigakijima; their specimens were feeding on annelids. Horsthemke (1986a-b, 1987a-b) gave detailed infonnation on captive husbandry and breeding, illustrating egg and larval development. He was unable to discover the original "wild" locality of his captive specimens. He found the species easy to breed in seawater and was able to successfully raise the larvae to adulthood, with the fish reaching sexual maturity by five months (Horsthemke, 1987a). There are undoubtedly more photographs, infonnation and anecdotes about this species in the "aquarium" literature (where this species is likely to be referred to as 5tigmatogobius hoevenii). Remarks Vaimosa chulae Smith, 1932, was originally described from seven specimens (three males, of which one was the holotype, and four females) (Figure 105). Smith's notes kept at Kasetsart University (KUMF) show that he collected seven specimens: three males and four females. Upon examining the specimens held at KUMF and USNM, it was found that Smith actually had four males and two females. The fate of the seventh specimen is not known, and it was not found in December, 1993, when the holotype and the three KUMF paratypes were re-discovered. The male holotype was collected at Koh Samui along with four females. The holotype was retained at the Department of Fisheries, Thailand, and two of these females are the paratypes deposited in USNM 119645. Smith (1945) stated that "a paratype" had been sent to USNM (USNM 119645, which consists of two females). Smith's fish collections were moved to different buildings several times, as the Fisheries Department moved, until they ended up at Kasetsart University, Bangkok (P. Wongrat: personal communication). Two male paratypes (KUMF 1888) were collected at Lem Sing Shore, Chantabun River estuary; these are the " ... 2 male specimens" referred to in Smith's description. Figure 105 Mugilogobius chulae. Paratype of Vaimosa chulae Smith, 32.5 mm SL, USNM 119645, Koh Samui, Gulf of Thailand. Figure 106 Mugilogobius chulae. Holotype of Tamanka sinensis Herre, 29 mm SL, CAS 31518, Hong Kong. Revision of Mugilogobiu5 Herre's (1935b) Tanumka sinensis (Figure 106) was recognised by Koumans (1940, 1953) as being the same as Smith's species (although Koumans referred to it as Stigmatogobius hoevenii). In his description of Vaimosa valigouva, Deraniyagala (1936) stated that holotype was sent to the BMNH (where it is catalogued as BMNH 1936.4.23.1). He did not indicate the location of the paratype (and only other type specimen). Pethiyagoda (1991), in his Appendix 3, listed the type specimens held at the National Museum of Sri Lanka (NMSL), and indicated that a specimen of Vaimosa valigouva was held there, uncatalogued. This is probably the paratype. Vaimosa zebra was described by Aurich (1938), from Boloang, north Celebes. The type specimens are typical M. chulae (Figure 107). The largest specimen in reasonable condition from among his four syntypes (ZMH 727) is a 20 mm SL female, which is hereby designated as lectotype of Vaimosa zebra Aurich. Fowler (1937) described a single specimen of M. chulae from Tachin, but his drawing and description do not clearly identify the species (it looks more like a Pseudogobius). Herre (1939) reported Stigmatogobius hoevenii from Long Island in the Andamans, giving a very brief description of the specimen. However, he gave a count of 13 predorsal scales, which allows an identification of M. chulae with this record (Hemigobius hoevenii has 8-10 predorsal scales). A number of paratypes of Vaimosa fusca Herre are 115 M. chulae (Figure 108); see under Remarks for M. fuscus. The problem of the identity of Mugilogobius luzonensis Roxas and Ablan (1940) is discussed under Remarks for M. fuscus. Mugilogobius luzonensis could be either of these two species or another species or genus entirely. Koumans (1953) confused M. chulae and Hemigobius hoevenii, referring to it as Stigmatogobius hoevenii, and others, following Koumans, have made similar errors. Koumans did recognise that Tamanka sinensis and Vaimosa zebra were the same species as chulae (his "hoevenii"). Koumans' 1953 colour description of "Stigmatogobius hoevenii" (Koumans, 1953: 126) clearly was of M. chulae, not H. hoevenii. Fowler (1961) considered that Gobius notospilus Giinther was a synonym of Stigmatogobius hoevenii and re-drew Giinther's figure. Fowler's (1961: 219) description of S. hoevenii was clearly of M. chulae. Munro (1958, 1967) records Stigmatogobius hoevenii from West New Guinea. From the information given in his 1967 key to New Guinea gobies, the fish could be M. chulae, M. wilsoni sp. novo or Hemigobius hoevenii. Bright and June's (1981) Mugilogobius sp. was identified here as M. chulae on the basis of one of their 14 specimens, deposited at AMS (AMS 1.23388-001); the remainder are probably still in Koror, as indicated in their paper. Hayashi et al. (1981) discussed this species' similarity to M. parous (citing Tomiyama 1936), but stated that scale counts of the two species differed, indicating that further work was necessary. Figure 107 Mugilogobius chulae. Lectotype of Vaimosa zebra Aurich, 20 mm SL, ZMH 727, Boloang, Sulawesi. Figure 108 Mugilogobius chulae. Paratype of Vaimosa fusca Herre, 29 mm SL, CAS 32985, Dumaguete, Philippines. H.K. Larson 116 Mugilogobius fasciatus sp. novo Figures 83,109-111; Tables 5-8,15 Material Examined Holotype 20 mm SL male, ZRC 17099, Sungei Buloh mangroves, Singapore, P. Ng, 23 February 1991. Paratypes Thailand: NTM S.13953-016, 9(8-20.5), small muddy mangrove creek, Klong Bang Sai, Phuket, H. Larson, D. Hoese and PMBC staff, 8 December 1993; NTM S.13954-042, 1(12), sandy mangrove creek, Ao Nam Bor, Phuket, H. Larson, D. Hoese and PMBC staff, 9 December 1993; ROM 68199, 13(14-24), mangroce channel, E coast Phuket, between Cape Phanwa and Phuket City, R. Winterbottom, R. Mooi, W. Holleman and U. Satapoomin, 22 November 1993. Singapore: CMK 8316, 2(22-24), Kranji mangrove near Sungei Buloh, M. Kottelat and D. Murphy, 8 April 1992; ZRC 36706-7,2(11.5-18), Sungei Buloh mangroves, P. Ng, K. Lim, N. Sivasothi, 14 January 1994; ZRC 29698, 1(12), Sungei Buloh mangroves, A. Wong et al., 8 September 1992; AMS 1.37570-001, 1(11), Sungei Pandan, D. Hoese and K. Lim, 22 December 1993; NTM S.13957-011, 1(10.5), small mangrove stream, Sungei Pandan, H. Larson, P. Ng, D. Murphy, 29 December 1993; NTM S.14303-OO1, 1(19.5), site 1, mangroves at Sungei Buloh, K. Tai, 17 July 1992. Diagnosis A rather small Mugilogobius with second dorsal rays 1,6-7, anal rays 1,7; pectoral rays 14-15; longitudinal scales 25-28; TRB 7-9; circumpeduncular scales 12; predorsal scales 8-11, beginning close behind eyes, anteriormost scale enlarged; scales on body mostly ctenoid; second to Table 15 fourth spines of dorsal fin longest, often elongate; head and body whitish to light brown, with seven black bands, at least five of which encircle body; rounded to elongate black spot on centre of caudal fin base; black-bordered whitish oblique streak from eye to corner of mouth; known only from mangrove habitats in peninsular Thailand and Singapore. Description Based on 25 specimens, 10.5-24 mm SL. Counts of holotype (Figure 109) indicated with asterisk. First dorsal V (in one), VI* (in 24); second dorsal 1,6-7 (mean 1,7*); anal rays always 1,7*; pectoral rays 14-15*; segmented caudal rays 15-16 (mean 16*); caudal ray pattern modally 9/7*; branched caudal rays 6/4 to 9/6 (7/6 in holotype); unsegmented (procurrent) caudal rays 5/7 to 7/7; longitudinal scale count 25-28 (mean 26*); TRB 7-10 (mean 8*); predorsal scale count 8-12 (mean 10*); circumpeduncular scales 12*. Gill rakers on outer face of first arch 3+6 to 4+8 (modally 3+7). Pterygiophore formula 3-12210 (in 11). Vertebrae 10+16 (in 14). Neural spine of first and/or second vertebra broad at tip or with flange (in five) or pointed (in one). Two epurals (in 11), possibly one epural in one specimen (X-ray unclear). Two (in 11) or three (in one) anal pterygiophores before haemal spine of first caudal vertebra. Body slender, rather rounded anteriorly, compressed posteriorly. Head wider than deep, but not greatly so, HL 3.2-3.7 (mean 3.5) in SL; somewhat rectangular in cross-section, with rounded cheeks. Depth at posterior preopercular margin 1.4-1.8 (mean 1.6) in HL. Width at posterior preopercular margin 1.3-1.7 (mean 1.4) in HL. Mouth subterminal to barely terminal, oblique, forming angle of about 25-35° with body axis; jaws reaching to below anterior half of eye in both sexes Oaws slightly longer in some males). Lips usually Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius fasciatus sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pedoral!. in SL Pelvic !. in SL Caudal!. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 29.0 65.5 74.1 19.0 13.5 28.5 13.5 25.9 29.3 34.5 32.8 19.0 20.0 23.5 28.5 27.9 55.6 63.9 18.1 10.4 25.4 12.4 22.2 26.5 27.8 28.8 19.0 20.0 23.5 22.9 31.3 69.5 77.0 20.9 13.7 30.4 14.1 25.9 32.3 37.1 32.8 23.8 26.1 30.4 32.7 29.2 62.1 71.3 19.5 12.6 28.2 13.4 24.2 29.0 33.9 30.5 21.6 21.7 26.6 28.3 28.2 63.6 72.1 20.2 13.7 27.4 13.1 23.7 30.0 33.2 29.4 21.0 20.8 25.1 20.8 30.8 66.2 78.6 21.5 20.4 30.6 13.8 27.6 32.0 36.0 32.7 24.1 22.1 26.7 28.3 28.2 63.6 72.1 20.2 13.7 27.4 13.1 23.7 30.0 33.2 29.4 21.0 20.8 25.1 20.8 Revision of Mugilogobius 117 Figure 109 Mugilogobius fasciatus n. sp., holotype, 21.5 mm SL, ZRC 17099, Singapore. smooth, small fleshy fimbriae may be present on inner front edges of lips; lower lip free at sides, fused across front. Upper jaw length same in males and females, 2.7-3.6 in HL (mean 3.0). Eyes lateral, high on head, top usually forming part of dorsal profile, 3.1-3.8 (mean 3.4) in HL; eye width greater than snout length. Snout short, profile rounded to slightly pointed, 3.6-4.6 (mean 4.3) in HL. Interorbital broad, flat, 3.1-4.0 (mean 3.4) in HL. Top of head from anterior portion of interorbital forward to snout tip often with sparsely scattered fine villi. Body depth at anal origin 4.6-5.5 (mean 5.1) in SL. Caudal peduncle long, compressed, length 3.3-3.9 (mean 3.7) in SL. Caudal peduncle depth 4.9-9.6 (mean 7.7) in SL. First dorsal fin approximately triangular, tips of second to fourth spines free and elongate in males, in females, tips free but spines not as elongate, second or third spines longest; elongate spines often reaching to middle of second dorsal fin, rest of fin barely reaching second dorsal fin origin when depressed. First dorsal spine longest in one female (never in males), 5.3 in SL. Second dorsal spine length in males 3.1-5.3 (mean 3.8) in SL; length in females 3.5-5.4 (mean 4.9) in SL. Third dorsal spine length in males 3.1-4.4 (mean 3.6); 5.6 in SL in one female. Second dorsal and anal fins short-based, about equal in height to first dorsal fin, posteriormost rays longest, rays falling short of caudal fin when depressed. Pectoral fin oval to slightly pointed, central rays longest, 4.1-5.3 (mean 4.7) in SL; rays usually all branched but for uppermost. Pelvic fins rather slender, oval, reaching about two-thirds of distance to anus, 3.8-5.3 (mean 4.7) in SL. Caudal fin oval, rounded posteriorly, 3.34.3 (mean 3.9) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, short, placed on edge of upper lip, tube oriented down and forward, preorbital may be curved to accommodate nostril. Posterior nostril oval to rounded, placed close to anterior centre margin of eye (one specimen with nostrils fused together, forming single oval nostril on each side of snout). Gill opening extending forward to under opercle. Inner edge of pectoral girdle with low fleshy to bony flange bearing several to many small flattened lobes or bumps; or entire flange irregular to "frilly" to partly smooth. Gill rakers on outer face of first arch slender, short, with or without tiny spiny papillae at tip, longest rakers near angle of arch; rakers on inner face of first arch usually longer than those of outer face (sometimes equal to longest raker); outer rakers on second (and remaining arches) arch short, tiny papillae sometimes absent from tips, and inner face rakers relatively long and slender, as are inner face rakers on third and fourth arches. Tongue tip usually blunt. Outer teeth in upper jaw largest, curved and pointed, three or four rows of very small sharp teeth behind this row; one or two rows of small teeth at side of jaw; outer teeth in upper jaw larger in males than in females. In females, lower jaw with five or six rows of evenly sized, small pointed teeth anteriorly, innermost teeth usually largest; one or two rows of teeth at sides. In males, four or five rows of curved, pointed teeth anteriorly, innermost row teeth abruptly larger, stout and curved; usually only one row of teeth at side of jaw. Predorsal scales about same size as body scales but for anteriormost, this scale enlarged, placed close behind eyes and partly entering interorbital area. Operculum with cycloid scales on upper half to two-thirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area completely covered with small cycloid scales. Belly usually with isolated area of ctenoid scales under pelvics (covering anterior 1,4 to V2 of belly), rest of scales cycloid; some specimens with all belly scales cycloid or few ctenoid scales under pelvic disk. Ctenoid scales on side of body extending well forward to behind pectoral fin. Genital papilla in male elongate, flattened, with pointed tip; in female, short, blunt-tipped, rounded to bulbous. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 110. lhree s rows present, of one to three papillae each; central row often of one papilla only. F row on chin consisting of two papillae. Coloration of fresh material No information available, other than my field notes indicating that fresh specimens were light greyish with black bands. 118 Figure 110 Mugilogobius fasciatus n. sp., papillae pattern. Paratype, CMK 8316. Scale bar = 1 mm. Scalation suggested only. Coloration of preserved material Head and body whitish to very light brown, with dark brown saddle or bar across head and six dark brown bars present, five encircling body; bars usually three scales wide (Figures 109, 111). Bars usually wider dorsally. Dark brown band or saddle crossing head behind anteriormost (enlarged) scale, rear margin of brown band ending at rear corner of operc1e; band continuing onto upper half of operc1e as triangular blotch. Second dark brown band beginning at first dorsal fin origin and nearly meeting its counterpart at centre of belly; bands separated by about two whitish scales at belly midline. Some specimens with small dark brown blotch on nape before first dorsal fin. Third dark brown band encircling body, beginning at rear of first dorsal fin and crossing anus or slightly before anus. Fourth dark brown band beginning below anterior half of second dorsal fin and extending down to anterior part of anal fin. Fifth dark brown band beginning at rear of second dorsal fin and ending at rear of anal fin. Sixth dark brown band encircling mid-point of caudal peduncle. Posteriormost dark brown band encircling base of caudal fin, crossing upper and lower procurrent rays. H.K. Larson Snout and top of head between eyes brown to blackish, blackish bar or broad line usually present from upper edge of each eye across interorbital space; second blackish bar or line usually discernible, extending from eye to eye just behind posterior nostril. Two dense blackish bars or broad lines extending from front of eye to upper jaw, with relatively conspicuous whitish area between them (forming indistinct black-bordered whitish stripe); posteriormost blackish line extending from rictus up along ventral margin of eye, ending on preoperc1e behind eye. Indistinct brown to dark brown blotchy line running from top of operc1e obliquely forward onto middle of cheek. Blackish narrow line following lower edge of preoperc1e to underside of chin; line sometimes extending posteriorly onto upper part of operc1e; similar line sometimes present along rear margin of opercle. Underside of head plain dusky to blackish. Lips dark brown to blackish. Pectoral base pale with irregular to rounded brown blotch or spot on upper half; spot sometimes extending onto pectoral ray bases. Breast brown to blackish. Belly pale to light brown. First dorsal fin dusky brownish anteroventrally, with dense black band occupying middle third of fin; broad translucent white submarginal band present; fin margin itself dark brown to black; elongate spines translucent white edged with brown, or entirely brown. Second dorsal fin with lower two-thirds light brownish, with broad translucent white marginal band; brown portion of fin usually with elongate darker brown streaks along centre; streaks horizontal or oblique. Anal fin plain dusky or brown with narrow whitish margin. Caudal fin plain dusky, often with dark brown to blackish upper and lower edges near fin base; distinct dark brown to (more usual) black spot on centre base of fin; spot round to vertically elongate and sometimes partly joined to centre of brown body band at base of caudal fin by short dark brown line. Pectoral fin translucent to dusky, fin rays outlined with dusky pigment. Pelvic fins brownish to blackish, including fraenum, latter whitish in some specimens. Figure 111 Mugilogobius fasciatus n. sp., paratype, 22.8 mm SL, CMK 8316, Kranji, Singapore. From colour slide by Maurice Kottelat. Revision of Mugilogobius Comparisons This species looks very similar to M. tigrinus sp. novo and is syntopic with it in Thailand and Singapore. The two species are distinguished under Comparisons for M. tigrinus sp. novo Distribution Specimens are so far known only from peninsular Thailand and Singapore (Figure 83). Ecology Mugilogobius fasciatus has been found in shallow small mangrove creeks and pools, often concealed among leaf litter. Etymology From the Latin fascia, band or stripe, referring to the encircling black bands on the body. Mugilogobius flli/er sp. novo Figures 22B, 83, 112-115; Tables 5-8, 16 Material Examined Holotype 32 mm SL male, NTM S. 12358-001, mangroves, Darwin Harbour, Northern Territory, Australia, R. Hanley, 1986. Para types Australia: Western Australia: WAM P.25668-014, 12(17-32), Port Warrender, Admiralty Gulf, 4 m depth, J.B. Hutchins and party, 22 October 1976; AMS 1.25511-004, 2(23-27), mangroves at road crossing, Cossack, D. Hoese and D. Rennis, 15 September 1986. QM 1.22552, 1(25), Pago Ruins Creek, Napier Broome Bay, N of Kalumburu, R. Leggett,l1 August 1986; Northern Territory: NTM S.12836-001, 2(30-33), Charlotte Rover, road crossing on Finniss River Station road, D. Wilson, August 1990; NTM S.14348-001, 1(39), Tumbling Waters, Blackmore River, D. Wilson, 26 December 1994; NTM S.10472-022, 9(14-24), NW side Channel Island, Darwin Harbour, B. Russell and R. Williams, 119 15 July 1982; NTM S.10694-013, 33(13-36), mangrove creek at Gunn Point, H. Larson and R. Williams, 20 September 1982; USNM 31686, 1(31), desiccating mudpool, cut off from creek at Yirrkala, Arnhem Land, R.R. Miller, 30 July 1948; Queensland: AMS 1.23279-008, 66(15-38.5), creek at Embley River, D. Hoese and D. Rennis, 10 October 1982; AMS 1.23281-017, 69(15.5-39), Mission River mouth, SW of Wallaby Island, Weipa, D. Hoese and D. Rennis, 11 October 1982; AMS 1.19356-013, 1(21.5), N side of Terry Beach, Prince of Wales Island, Torres Strait, W. Ponder and D. Brown, 2 July 1976; AMS 1.20927-010, 2(27.5-34), creek at Smith Point, Prince of Wales Island, Torres Strait, joint Australian Museum - AIMS Expedition, 16 February 1979; AMS 1.20927-012, 1(34), same data as preceding. Papua New Guinea: USNM 316179, 1(34), brackish lagoon cut off from ocean by sandspit, PukPuk Island opposite Kieta, Bougainville Island, Te Vega Cruise 6, 9 March 1965. Other material examined (but not used in description) Australia: Western Australia: AMS 1.25522-002, 4, King Sound; Northern Territory: AMS 1.24685005,18, Ludmilla Creek, Darwin; NTM S.13518-005, 1, Bing Bong Creek, Gulf of Carpentaria; NTM S.14046-016, 1, Roper River; NTM S.11575-001, 1, Ludmilla Creek, Darwin; AMS 1.24686-006, 3, Ludmilla Creek, Darwin; NTM S.14292-002, 2, golf links canal, Darwin; AMS 1.24692-003, 1, Leader's Creek, Gunn Point; AMS 1.24683-002, 1, Blackmore River; NTM S.12740-002, 1, Darwin River; Queensland: AMS 1.23278-001, 2, Mission River; AMS 1.23281-019, 4, Mission River; AMS 1.23287011, 17, Embley River. Diagnosis A moderately large Mugilogobius with second dorsal and anal rays always 1,7; pectoral rays 15-17; longitudinal scales 29-40; TRB 9-13; circumpeduncular scales 11-15; predorsal scales 1321, small, reaching close up behind eyes, anteriormost scale somewhat larger than rest; scales on body mostly ctenoid; first spine of dorsal fin usually longest, filamentous (usually white and Figure 112 Mugilogobius filifer n. sp., holotype, 32 mm 5L, NTM 5.12358-001, Darwin Harbour, Northern Territory. H.K. Larson 120 conspicuous) in adult males; light brown to grey body with seven to nine short oblique dark blotches, bars or chevrons on sides, bars usually broken up into off-set pairs of blotches, anteriormost body marking a dark oblique shoulder bar, head with radiate or reticulate pattern of brown lines, broad brown band crossing branchiostegal membranes on underside of head; known from northern Australia and eastern Papua New Guinea. Description Based on 54 specimens, 17-39 mm SL. An asterisk indicates counts of holotype (Figure 112). First dorsal Vl*; second dorsal always 1,7*; anal always 1,7*, pectoral rays 15-17* (mean 16), segmented caudal rays 16*; caudal ray pattern modally 9/7; branched caudal rays 7/6 to 9/7 (modally 8/7*); unsegmented (procurrent) caudal rays 7/7 to 8/9; longitudinal scale count 29-40 (mean 35, 36 in holotype); TRB 9-13 (mean 12*); predorsal scale count 13-21 (mean 19*); circumpeduncular scales 11-15 (mean 13, 12 in holotype). Gill rakers on outer face of first arch 3+6 to 5+7 (modally 4+7). Pterygiophore formula 312210 (in six). Vertebrae 10+15 (in one; deformed vertebra), 10+16 (in five). Neural spine of second vertebra expanded at tip (in two). Two epurals (in two). Two (in five) or three (in one) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid broad, expanded, reaching across to, but not overlapping, quadrate. Body robust, compressed (less so anteriorly), large specimens rather rounded anteriorly. Head broad, wider than deep, HL 3.1-3.6 (mean 3.3) in SL; cheeks sometimes inflated in large males. Depth at posterior preopercular margin 1.5-2.0 (mean 1.8) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of 20-25° with body axis; Table 16 jaws generally reaching to below posterior half to mid-eye in males and to below anterior half or mideye females (to below mid-eye in holotype). Lips smooth, fleshy fimbriae usually present on inner edge of upper lip, mostly across front; lower lip free at sides, broadly fused across front. Upper jaw 2.23.0 (mean 2.7 in females, 2.5 in males) in HL. Eyes lateral, high on head, top usually forming part of dorsal profile, 3.1-4.3 (mean 3.7) in HL. Snout rounded, sometimes slightly inflated, 3.3-4.5 (mean 4.0) in HL. Interorbital broad, flat, 2.7-4.7 (mean 3.5) in HL. Top of head from rear of interorbital up to snout usually with dense or relatively sparse, fine villi (sometimes difficult to see, depending on preservation). Body depth at anal origin 4.4-6.0 (mean 5.2) in SL. Caudal peduncle compressed, length 3.4-4.3 (mean 3.8) in SL. Caudal peduncle depth 6.5-8.3 (mean 7.3) in SL. First dorsal fin triangular, tip of first spine always free; second to sixth spines usually falling short of second dorsal fin origin when depressed; spines nearly always longer in males than females. First dorsal spine usually longest, often elongate and filamentous in males (spine reaching back to last few second dorsal fin elements); 3.8-7.7 (mean 5.4) in SL in females, 2.2-5.6 (mean 3.8) in SL in males. Second dorsal spine length 6.1-7.1 (mean 6.7) in SL in females and 3.0-7.0 (mean 4.3) in SL in males. Third dorsal spine longest in only two specimens, spine lengths 7.2 and 7.4 in SL. Second dorsal and anal fins low, short-based, posteriormost rays longest, rays falling well short of caudal fin when depressed. Pectoral fin rounded, central rays longest, 4.1-5.0 (mean 4.6) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, reaching half to two-thirds distance to anus, 4.7-6.6 (mean 5.4) in SL. Caudal fin rounded, 3.3-4.5 (mean 3.9) in SL. No mental fraenum, chin smooth. Anterior nostril Morphometries as percentages of SL or HL, as indicated, of Mugilogobius filifer sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic !. in SL Caudal!. in SL Longest Dl spine in SL Holotype 31.9 60.8 78.4 20.6 13.4 28.1 14.1 28.4 26.5 40.2 34.3 22.5 19.7 27.2 35.0 Males Males Minimum Maximum 28.1 50.0 63.1 16.9 9.8 24.1 13.2 22.3 23.2 33.9 23.2 20.0 15.2 22.4 17.7 31.9 62.1 79.8 22.7 15.8 29.4 15.5 30.1 32.2 45.1 36.9 24.4 21.2 30.5 45.6 Males Mean 30.2 57.1 71.9 19.7 12.3 26.1 14.1 25.9 27.2 40.1 30.3 22.1 18.6 26.8 28,8 Females Minimum 28.4 52.5 65.6 16.8 9.4 23.5 12.1 22.0 23.0 33.3 ' 21.4 20,0 17.3 23.4 13.0 Females Maximum Females Mean 32.3 66.0 83.0 21.3 15.8 28.8 15.0 27.8 29.8 40.7 35.6 23.5 20.8 30.0 26,2 30.1 58.3 72.5 19.5 12.6 26.5 13.5 25.3 27.6 36.5 29.3 21.6 18.9 25.9 19.2 Revision of Mugilogobius 121 tubular, placed just at preorbital edge, tube short, oriented down and forward, preorbital curved forward to accommodate nostril. Posterior nostril rounded, placed close to upper anterior margin to anterior centre of eye. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth (in two), with low irregular fleshy flange (in 13) or with small fleshy knobs (in 23); often only one rounded knob present in combination with fleshy ridge. Gill rakers on outer face of first arch very low, almost rudimentary, smooth, longest rakers near angle of arch; rakers on inner face of first arch slightly longer, about equal to rakers near angle; outer rakers on second arch similar to those of first arch; inner rakers on other three arches short and slender. Tongue tip usually concave, may be blunt. Outer teeth in upper jaw largest, stout and curved, three to four rows of evenly sized small sharp teeth behind this row; one or two rows at side of jaw. Lower jaw with three or four rows of small pointed teeth anteriorly, outermost row oriented upright, inner rows tending to point rearward; innermost row teeth largest, stout and curved; usually only one row of teeth at side of jaw (innermost row); males generally with larger teeth than females (especially outer row teeth). Predorsal scales small, reaching forward to close behind eyes, anteriormost scale usually larger than remainder, and placed on nape midline; scale not as enlarged as is usual for this species-group. Operculum with small cycloid scales on upper half to two-thirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior % to 1h of belly), rest of scales cycloid. Ctenoid scales on side of body extending forward in wedge to behind pectoral fin. Genital papilla in male elongate, flattened, narrowing to pointed tip; in female, short, short and rounded. ｾ \ Figure 113 MlIgilogobills filifer n. sp., papillae pattern. Paratype, NTM 5.10472-022. 5cale bar = 1 mm. Scalation suggested only. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 113. Three s rows on snout present, middle row consisting of one to three papillae. Mandibular f row in two groups of about three papillae each. Papillae in rows on side of head small, evenly sized and close-set; rear portion of row a behind eye curved posteriorly. Coloration of fresh material Information taken from colour slides taken by Doug Hoese. Head and body light grey, yellowish white or dark brownish grey (latter in males in breeding coloration; non-breeding males and females similar, with dark blotches and markings on head and body more distinct than in males exhibiting dark breeding coloration), with brown to purplish brown bars, blotches and chevrons on side of body, and brown reticulate pattern blotches and lines on head; underside of head and body plain light grey to whitish. Side of body with eight midlateral brown to purplish brown X-shaped blotches or chevrons; above these blotches, about nine dorsal saddles or oblique bars (often reaching down to, and blending with, midlateral blotches); anteriormost bar a brownish oblique shoulder bar directly above pectoral base. Pectoral base pale with rounded dense to diffuse dark brown spot on base of fin rays, placed just above centre of fin base. Side of head with five short dark brown curved lines radiating from eye; lines diffuse, broad or narrow, or partly joining to form reticulate pattern. Iris pale gold to gold marbled with dark brown. Lower half of head and lower third of body distinctly paler than remainder. Breast and branchiostegal membranes usually dusky brown to blackish. Fin colour most intense in males. Broad submarginal band on dorsal and anal fins bright yellow to yellowish white, remainder of fins yellowish grey to greyish yellow with black to brownish grey markings and narrow dusky grey margin. Caudal fin purplish brown to brownish grey, with scattered indistinct brown spots and streaks near base; most distinct markings usually two square blackish blotches vertically oriented near fin base, each blotch joined by diffuse blackish brown bar to blackish blotch on centre of hypural crease; lowermost blotch on caudal fin often indistinct; all three blotches often barely darker than blotches and bars on side of body. Coloration of preserved material Preserved colour variable, basically similar to live colour. Head and body light brown to yellowish brown or greyish with dark brown oblique bars, chevrons, elongate blotches and X-shaped markings along sides; in some specimens, underlying lighter background colour visible on side mainly as series 122 H.K. Larson Figure 114 Mugilogobius filifer n. sp., paratype, 32 mm SL, NTM S.10694-013, Gunn Point, Northern Territory. of light blotches or as two or three alternating rows of light blotches in between dark markings, giving "chequered" appearance. Seven to nine oblique narrow dark bars, chevrons, rounded blotches or Xshaped markings present; anteriormost bar a short, oblique "shoulder" bar not always distinguishable from other markings but sometimes more densely pigmented than other body markings; posteriormost dark marking a rounded spot or < shaped blotch on middle of caudal base (across hypural crease), extending onto lower part of caudal fin base and sometimes ventrally onto lower part of caudal fin. Upper and lower parts of posteriormost body bar sometimes intensified as dark brown to blackish spot or streak, extending onto procurrent rays of caudal fin. Pectoral base pale brown to whitish with narrow brown horizontal bar placed just above midpoint, diffuse brown curved streak sometimes covering upper pectoral ray bases. On head, five to seven short straight or curved lines radiating outward from eye (Figures 112, 114), lines entering interorbital area short, lines crossing cheek long, often reticulate, joining up with brown wide line or blotch on lower cheek behind rictus (brown cheek blotch variable in shape and intensity). Snout and interorbital area dark brown spotted or with dark vermiculate markings. Rearmost line from eye extending horizontally onto upper part of opercle, meeting brown or blackish blotch or diffuse reticulate pattern; rear edge of opercle with narrow brown margin (usually confluent with dark brown band crossing branchiostegals). Underside of head pale or dusky brown, crossed by series of about seven thin curved brown lines; lines occasionally fuse, forming marbled or ocellate pattern; broad dark brown band (most distinct marking on underside of head) crossing branchiostegal membranes and breast; usually branchiostegal bands heavily pigmented and distinct, breast paler. Gill arches dusky to brownish; roof of branchial chamber pale to light dusky. First dorsal fin with lower half dusky grey to brownish grey, with large dense black spot on central rear half of fin. Outer third of first dorsal fin white to translucent whitish, often with narrow brownish to dusky grey marginal band; tips of fin spines usually white; elongate first spine white and conspicuous (Figure 115). Second dorsal fin with proximal two-thirds dusky brownish to grey, narrow grey to brown fin margin present, broad white to translucent whitish stripe below marginal band. Dusky lower part of fin with series of short, vertically elongate brown streaks on fin membrane, and two dark brown irregular blotches on fin base, one near anterior end of fin and one near mid-point. Anal fin plain dusky to brownish, with narrow white to translucent margin. Caudal fin mostly plain dusky grey to brownish grey, with some brown spots and indistinct streaks near base, two small dark brown to blackish vertically oriented distinct spots at base, lower spot sometimes replaced by diffuse oblique brown bar; upper spot always distinct. Pectoral fin translucent, fin rays usually narrowly outlined with brown. Pelvic fins quite dusky to dark brown but for narrow translucent margin on fraenum. Figure 115 Mugilogobius filifer n. sp., 34 mm SL, AMS 1.20927-010, Prince of Wales Island, Torres Strait, Queensland. From colour slide by Doug Hoese. Revision of Mugilogobius Comparisons See Comparisons under M. mertoni. Distribution Specimens are known only from tropical northern Australia and eastern Papua New Guinea (Bougainville) (Figure 83). Ecology Found in mangrove creeks and estuaries, sometimes well into freshwater influence; for example, the N1M specimen from the Blackmore River was taken from among vegetation close to the riverbank in fresh water. Remarks Mission River (Queensland) specimens were syntopic with M. mertoni. This species is sometimes identified in museum collections as "DFH sp. 7", in reference to the numbering system used by Doug Hoese of AMS. Etymology From the Latin filum, thread or filament, and fer, suffix meaning to bear or carry; in reference to the elongate first dorsal fin present in adult males. Mugilogobius fuscus (Herre, 1940) Figures 83,116-119; Tables 5-8,17 Vaimosa fusca Herre, 1940b: 359, plate 3 (Dumaguete, Oriental Negros Province, Philippines); Herre 1950: 75. ?Mugilogobius luzonensis Roxas and Ablan: 1940: 307, plate 6 (Luzon, Philippines). Stigmatogobius hoevenii: Koumans 1953: 125 (as questionable synonym of hoevenii). Mugilogobius sp.: Hayashi et al., 1981: 10, plate 7, figure 129. Mugilogobius luzonensis: Akihito et al., 1988: 268, plate 247Q figure 148. 123 Material Examined Holotype ofVaimosa fusca CAS 32984, 32.5 mm female, Dumaguete, Negros Oriental, Philippines, A. Herre, December 1936. Other Material Philippines: USNM 99611, 1 (35.5), brackish water, Port Dupon, Leyte, Albatross Expedition, 17 March 1909. Japan: YCM 9323, 1 (39), Nagura River, Ishigakijima, M. Hayashi and party, 26-27 July 1981; YCM P.9167P, 1 (42), Yonada River, Iriomotejima, 30 August 1978; URM P.4840, 1 (42), Shiira River, Iriomotejima, T. Yoshino, 14 September 1982. Papua New Guinea: USNM 260525, 2(21-33), mangrove creek at Gona, Popondetta, T. Roberts, 8 August 1975. Sri Lanka: USNM 316185, 1 (22), estuary about 3 miles N of Mahaweli River mouth, W side of road, Trincomalee, C. Koenig, 9 April 1970. Seychelles: USNM 265058, 2(33-47), Beau Valley, freshwater stream draining into sea at Beau Vallon Bay, W. Starck, J. Tyler, 26 February 1964; R.G. Mus. Africain Centrale 188771-3, 2(23-26), Les Canelles, Mahe south, P. Benoit and J.J. von Mol, 20 June 1972. Diagnosis A robust, blunt-headed Mugilogobius with second dorsal rays 1,7-8; anal rays 1,6-8; pectoral rays 1416; longitudinal scales 28-30; TRB 9-11; circumpeduncular scales 12; predorsal scales 9-11, reaching to behind eyes, anteriormost scale enlarged; scales on body ctenoid, extending forward to at least over top of opercle and often onto predorsal area, opercle usually with ctenoid scales; dorsal fin low, without filamentous spines; body with reticulate dark brown pattern, diffuse crosshatched barring along midside of body, more distinct posteriorly, and pair of brown spots at caudal fin base; known from Japan, Philippines, Papua New Guinea, Sri Lanka and the Seychelles. Description Based on 12 specimens, 21-47 mm SL. An asterisk indicates counts of holotype of Vaimosa fusca (Figure 116). Figure 116 Mugilogobius fuSCllS, holotype of Vaimosa fusca Herre, 32 mm SL, CAS 32984, Dumaguete, Philippines. 124 Table 17 H.K. Larson Morphometries as percentages of SL or HL, as indicated, of Mugilogobius fuscus (Herre, 1940). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. l. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbitall. in HL Pectorall. in SL Pelvic l. in SL Caudall. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 31.4 61.8 75.5 22.5 14.8 26.2 13.5 27.5 30.4 32.4 27.5 24.3 22.5 28.5 54.5 67.6 20.0 12.3 26.7 12.9 26.9 24.6 31.3 29.9 22.2 21.7 23.6 16.1 31.9 64.9 80.6 22.1 14.5 29.6 15.5 35.8 32.8 45.5 47.0 26.1 24.8 31.5 17.1 29.6 57.9 70.8 20.6 13.2 27.8 14.0 28.9 28.4 36.0 37.2 24.2 23.3 28.8 16.6 27.6 55.7 66.4 20.6 11.5 25.0 13.0 22.3 26.7 32.4 27.5 20.0 19.7 25.7 14.0 31.4 63.8 75.5 23.3 15.5 28.5 14.8 30.2 30.4 35.9 39.7 24.3 22.5 27.6 15.5 29.4 60.5 71.6 22.3 13.9 26.2 14.0 26.5 28.2 33.7 33.8 22.7 21.2 26.4 15.2 First dorsal VI*; second dorsal 1,7*-1,8 (mean 1,7); anal 1,6-8 (mean 1,7*), pectoral rays 14-16 (mean 15*), segmented caudal rays always 16*; caudal ray pattern modally 9/7; branched caudal rays 14-16 (mean 15*); unsegmented (procurrent) caudal rays 6/6* to 7/7; longitudinal scale count 28-30 (mean 29*); TRB 9-11* (mean 10); predorsal scale count 911 (mean 10*); circumpeduncular scales always 12*. Gill rakers on outer face of first arch 3+10 (in one), 4+10 (in one), 4+11 (in one). Pterygiophore formula 3-12210* (in six). Vertebrae 10+16* (in seven). First and second neural spines short and pointed (in two) or broad, with flanges (in three). Two epurals (in six). Two (in six) or three (in one) anal pterygiophores before haemal spine of first caudal vertebra. Body generally compressed, rounded anteriorly (some specimens approximately square in crosssection anteriorly). Head wider than deep, HL 3.13.6 (mean 3.4) in SL; top of head flattened, but head not depressed. Depth at posterior preopercular margin 1.6-1.8 (mean 1.7) in HL. Width at posterior preopercular margin 1.2-1.5 (mean 1.4) in HL. Mouth subterminal, slightly oblique to nearly horizontal, forming angle of about 12-15° with body axis; jaws generally reaching to below anterior half of eye. Lips smooth, relatively narrow, rarely with fleshy fimbriae on inner edges; lower lip thin, not fleshy, free at sides, fused across front. Upper jaw 2.2-3.2 (mean 2.8 in males, 3.0 in females) in HL, jaws in adult males reaching to below mid-eye; largest male specimen with jaws just reaching to below rear portion of eye. Eyes relatively large, lateral, high on head, forming part of dorsal profile, 3.0-4.1 (mean 3.6) in HL. Snout blunt to rounded, 2.8-4.5 (mean 3.7) in HL, overhanging tips of both jaws in large specimens. Interorbital broad, flat to slightly concave, 2.1-3.6 (mean 2.9) in HL. Top of head, on unsealed portion of interorbital up to snout tip, with scattered fine villi. Body depth at anal origin 4.3-5.0 (mean 4.7) in SL. Caudal peduncle compressed, length 3.4-4.0 (mean 3.7) in SL. Caudal peduncle depth 6.4-7.8 (mean 7.2) in SL. First dorsal fin rounded, tips of second to third spines free, but no spines elongate; spines slightly longer in males than females; spines just reaching second dorsal fin origin when depressed. First dorsal spine always shorter than next three. Second dorsal spine length 6.5-7.1 (mean 6.6) in SL. Third dorsal spine length 5.8-7.1 (mean 6.0 in males, 6.6 in females) in SL. Fourth dorsal spine longest in one male. Second dorsal and anal fins low, posteriormost rays longest, rays falling well short of caudal fin when depressed; rear margin of anal fin somewhat rounded. Pectoral fin oval, central rays longest, 3.8-5.0 (mean 4.2) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, reaching up to two-thirds of distance to anus, 4.0-5.1 (mean 4.5) in SL. Caudal fin broad, rounded, 3.2-4.2 (mean 3.7) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, placed just behind upper lip, tube short, oriented down and forward, preorbital curved forward to accommodate nostril. Posterior nostril oval, slightly closer to anterior centre margin of eye than to halfway to lip. Gill opening usually extending forward, just past lower pectoral base, to under opercle. Inner edge of pectoral girdle with smooth raised flange (in one) or with low bumpy ridge or fleshy flange (in 11), flange may have flaps or bumps present. Gill rakers on outer face of first arch very short and smooth, longest two rakers near angle of arch; rakers on inner face of first arch also stubby, but slightly longer than outer rakers; inner rakers on other arches twice length of first arch inner rakers. Tongue tip usually blunt (absent in holotype). Teeth quite small, similar in immature males and females, larger in mature males. Outer Revision of Mugilogobius teeth in upper jaw largest, especially across front, but not particularly enlarged, curved; two to three rows of very small sharp teeth behind outer row; two or three rows on side of jaw. Lower jaw with four or five rows of small, stout, pointed teeth, teeth gently curved, most teeth evenly sized, teeth at side of jaw tending to be upright; innermost row teeth sometimes larger and stouter U1an others but no individual teeth particularly enlarged; usually only one or two rows of teeth at side of jaw. Predorsal scales with anterior scale enlarged, entering rear of interorbital space; scales often ctenoid, ctenoid scales extending forward to above rear of preopercle in three specimens; ctenoid scales present on sides of nape only in five specimens. Operculum mostly covered with ctenoid scales in Japanese specimens; otherwise ctenoid scales present on upper half of opercle, cycloid scales below (small male from Sri Lanka with all opercular scales cycloid). Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly scales ctenoid; may be few cycloid scales before anus. Ctenoid scales on side of body extending forward to over top of pectoral base and over top of opercle then up onto nape. Genital papilla in male elongate, flattened, with pointed tip; in female, short, rounded and bulbous. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figures 117-118. Papillae illustrated in Akihito et al. (1984: figure 148). Papillae in cheek rows small, evenly sized; three 5 rows on snout, with three or more papillae in each row; d row fairly short;f rows with two to three papillae in each; broken c row with sections very short (consisting of three to four papillae each). ·@'b . C' . cp···· . d Figure 117 MugilogobillS !USCllS, papillae pattern, female, lshigakijima, Japan, YCM 9323. Scale bar = 1 mm. 125 Figure 118 Mugilogobius juSCllS, papillae pattern, male, USNM 265058, Beau Vallon, Seychelles. Scale bar = 1 mm. Coloration of fresh material This species is illustrated in colour in Akihito et al. (1988: Plate 247Q), where it appears to be generally dark brown. Coloration of preserved material Head and body light brown with dark brown spot on each scale, outer margin of spot darkest, body appearing finely reticulated with dark brown; toward abdominal region, spot placed closer toward centre of scale, abdomen appearing finely spotted (Figures 116, 119). Row of indistinct small brown blotches along midside, blotches darkest on pos terior half of body, sometimes forming incomplete oblique cross-hatched dark bars, especially toward ventral part of caudal peduncle; posteriormost blotch just before hypural crease. Upper half of body with indistinct brown small blotches or spots, often indiscernible from reticulate background; brown spots sometimes forming short irregular horizontal line backward from above pectoral fin base. Most conspicuous markings are two square to oval brown spots (one above the other) on base of caudal fin. Scaled portion of head similarly coloured to side of body; interorbital and snout with few indistinct brown vermiculate lines; side of head with brown mottling and small brown spots, markings sometimes forming short dark lines radiating from eye. Lips plain dusky. Underside of head and breast plain dark brown; belly light brown. First dorsal fin light dusky with narrow dusky margin and transparent submarginal band; central band on fin dark grey to blackish, intensified into black blotch at about middle of fin, lower part of fin plain dusky. Second dorsal fin with narrow dusky margin and translucent submarginal band, proximal two-thirds of fin dusky to translucent grey, with vertically oriented dark grey to brown H.K. Larson 126 Figure 119 Mugilogobius fuscus, 40 mm SL, YCM 9323, Ishigakijima, Japan. streak or blotch on each membrane between fin rays, forming row along middle of fin; usually dark grey to brown blotches present along fin ray bases. Anal, pectoral and pelvic fins plain dusky. Caudal fin dusky greyish, with four to six vertically oriented irregular rows of indistinct small brown spots on proximal half of fin; spots indistinct or absent on distal half of fin. Comparisons This species is similar to M. chulae and M. mertoni. After examination of the holotype of Vaimosa fusca, I initially considered this species to be conspecific with M. chulae (as an aberrant form). The consistent differences between the two species, in colour pattern (conspicuous oblique bars in M. chulae versus plain reticulate brown body in fuscus) and arrangement of ctenoid scales (nape and opercle with cycloid scales in M. chulae versus ctenoid scales on nape and usually on opercle in M. fuscus), do indicate that the species are separate. Additionally, M. chulae usually has long, free to filamentous dorsal spines; in M. fuscus specimens, the dorsal spines are low and never filamentous. Some specimens of Mugilogobius fuscus are similar in colouring to M. mertoni, but have a greater covering of ctenoid scales (at least to opercle versus a wedge of ctenoid scales reaching to behind pectoral fin but not extending over the top of the fin base), and has marginally fewer lateral scales (2830, mean 29; versus 26-37, mean 31). Herre (1950: 75) considered that Vaimosa zebrinus (= Calamiana mindora Herre) was related to M. fuscus " ... but differs markedly in the predorsal and opercular scalation". Distribution Specimens are known from the Philippines, Sri Lanka, Japan, Papua New Guinea and the Seychelles (Figure 83). It appears to be rare. Ecology Most specimens come from mangrove creeks or river estuaries. This species can occur syntopically with M. chulae and M. mertoni. Hayashi et al. (1981) collected M. fuscus with M. cavifrons on a mud bottom in the Yonadagawa River estuary, Iriomotejima. Remarks The holotype, CAS 32984, is a large female (32 mm long) (Figure 116). Herre described the species based on the large female "type" and 16 paratypes (12.5-29 mm in length); he described the differently coloured paratypes as variations from the "type". Plate 3 in Herre (1940) is of one of the paratypes (= M. chulae), as can be seen from the characteristic colour pattern and dorsal fin spine form. None of the other paratypes are the same species as the holotype. Paratypes CAS 32985 and 32986 are all specimens of Mugilogobius chulae, as are the two paratypes in BMNH 1938.12.1.213-214. The paratype CAS 32987, from a mangrove swamp at Kranji, Singapore, is a Hemigobius hoevenii. Akihito et al. (1988) identified their Japanese specimens of M. fuscus as M. luzonensis Roxas and Ablan, 1940, undoubtedly due to the appearance of M. luzonensis as illustrated in the original description (there are no extant types). This species was described from one specimen (as far as can be determined from the description), housed in the Bureau of Science, Manila (BSM 31950), and was subsequently lost during WWII. Even if Roxas and Ablan's species is conspecific with Herre's, the question is somewhat academic, as Herre's description of Vaimosa fusca has priority, being published in August, 1940, while Roxas and Ablan's description appeared in September/December, 1940 (both papers in different issues of The Philippine Journal of Science). Mugilogobius luzonensis could be the same species as M. fuscus, M. chulae or another taxon altogether. The predorsal and transverse scale counts given by Roxas and Ablan are very similar among the three species (TRB 8-13 in M. chulae, 9-11 in M. fuscus, 89 in M. luzonensis; predorsal scales 11-15 in M. chulae, 10-11 in M. fuscus, 13-16 in M. luzonensis). Roxas and Ablan state there are three spots at the base of the caudal fin, and their illustration (1940: plate 6) does look rather like a Mugilogobius. Revision of Mugilogobius It is difficult to be sure what M. luzonensis really is, as in the original description it is not clear how Roxas and Ablan measured interorbital width, as the eye is given as being " ... 2 times interorbital" on p. 307, and as 2 mm wide compared to the 4 mm wide interorbital on p. 308. Roxas and Ablan compare their species to "M. dispar Peters", which is a Redigobius, a genus with a narrow interorbital. Another genus with a narrow interorbital is Pseudogobius, a genus which is also similar in appearance to Roxas and Ablan's illustration. Mugilogobius fusculus (Nichols, 1951) Figures 120-124, Tables 5-8,18 Gobius (Tamanka) fusculus Nichols, 1951: 5-6, figure 3 (New Guinea). Tamanka fusculus: Munro 1958: 276. Acentrogobius fusculus: Munro 1964: 147; Munro 1967: 505. Mugilogobius fusculus: AlIen and Boeseman 1982: 102; AlIen and Coates 1990: 33, 110-111; Alien 1990: 11; AlIen 1991: 188-189, figure 35. Mugilogobius sp.: Parenti and AlIen 1991: 318. Material Examined Holotype AMNH 16887, 30 mm SL male, New Guinea, E. Ruda. Para types AMNH 19520,4(21-27), same data as holotype. Other Material Papua New Guinea: WAM P.28206-009, 1(21), Pagwi, Sepik River, D. Coates, 1 September 1983; N1M S.13689-005, 33(11.5-34), creek entering Biges River, N of Madang, H.K. Larson, J. Mizeu, M. Balem, 23 October 1992; N1M S.13698-001, 1(32), Biges River just S of Alexishafen bridge, Madang, H.K. Larson, J. Mizeu, M. Balem, 29 October 1992; NTM 5.13674-002, 2(9-25), Biges River close to entrance into Sek Harbour, H.K. Larson, J. Mizeu, M. Balem, 12 October 1992; CAS 63250, 9(8.5-23), pond NW of Riwo village, Madang, L. Parenti and B. Buddemeier, 26 September 1987; CAS 63580, 24(10-21), pond NW of Riwo village, Madang, L. Parenti and J. Mizeu, 3 November 1987; USNM 316193, 1(23.5), Popondetta, mangrove creek entering Oro Bay, T. Roberts, 4 August 1975; ZMH 19344, 2(27-34), Admiralty Islands, N coast Seeadler-hafen, Papitalai, Hamburg SudseeExpedition, Duncker, 18-20 October 1908. Other material eX11mined (but not used in description) Papua New Guinea: NTM S.13678-003, 10, Kokon village, Madang. 127 Diagnosis A rather slender Mugilogobius with second dorsal rays t8-9; anal rays C7-9; pectoral rays 14-17; longitudinal scales 28-41; TRB 8-13; circumpeduncular scales 12-15; predorsal scales 14-22, smalt evenly sized, reaching up to close behind eyes; scales on side of body ctenoid; first spine of dorsal fin longest white and usually filamentous in both sexes; plain light to dark grey or brownish grey with indistinct bars and X-shaped markings along side, and most distinct marking a blackish spot on upper caudal fin base; known only from fresh water and estuaries of northern Papua New Guinea. Description Based on 30 specimens, 15-34 mm SL. An asterisk indicates counts of holotype. First dorsal V (in one), VI* (in 29); second dorsal t8*-9 (mean C8); anal C7-9 (mean t8*), pectoral rays 14-17 (mean 16*), segmented caudal rays 1517 (mean 16*); caudal ray pattern modally 9/7*; branched caudal rays 7/6-9/7 (mean 8/7*); unsegmented (procurrent) caudal rays 8/8 (in two); longitudinal scale count 28-41 * (mean 35); TRB 813* (mean 11); predorsal scale count 14-22 (mean 18,21 in holotype); circumpeduncular scales 12-15* (mean 13). Gill rakers on outer face of first arch 3+7 to 5+9 (modally 4+7). Pterygiophore formula 312210 (in two). Vertebrae 10+16 (in two). Neural spine of second vertebra blunt and expanded at tip (in two). Two epurals (in two). Two anal pterygiophores before haemal spine of first caudal vertebra (in two). Body slender, compressed, somewhat rounded anteriorly. Head depressed, wider than deep, but not greatly so, HL 3.1-3.5 (mean 3.4) in SL; cheeks sometimes inflated in large males. Depth at posterior preopercular margin 1.4-2.0 (mean 1.7) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.4) in HL. Mouth subterminat slightly oblique, forming angle of about 25° with body axis; jaws generally reaching to below posterior half of eye in adult males and to below mid-eye or anterior half of eye in females. Lips mostly smooth, fleshy fimbriae may be present on inner edges of upper lip (sometimes very close to outer edge); lower lip free at sides, fused across front. Upper jaw 2.1-3.0 (mean 2.7 in females, 2.3 in males) in HL. Eyes laterat set high on head, top usually forming part of dorsal profile, 3.3-4.2 (mean 3.8) in HL. Snout rounded, 1.1-4.6 (mean 3.8) in HL. Interorbital broad, flat to slightly concave, 2.9-3.9 (mean 3.4) in HL. Top of head from behind eyes forward to tip of snout covered with fine villi, most easily seen in specimens with well-preserved mucous coat. Body depth at anal origin 5.1-6.6 (mean 5.6) in SL. Caudal peduncle compressed, length 3.7-4.4 (mean 4.1) in SL. Caudal peduncle depth 6.9-9.6 (mean 8.0) in SL. First dorsal fin triangular, tips of all spines free; H.K. Larson 128 first dorsal spine longest (only one male with all spines short, fourth spine longest), spine length 2.36.9 (mean 3.7) in SL in males and 3.7-7.1 (mean 5.5) in SL in females; other spines rarely reaching second dorsal origin when depressed. Second dorsal and anal fins low, posteriormost rays longest, pointed; rays just reaching caudal fin base when depressed; second dorsal fin reaching further back than does anal fin. Pectoral fin oval, central rays longest, pointed, 4.0-6.3 (mean 4.7) in SL; rays usually all branched (uppermost ray sometimes unbranched). Pelvic fins short, oval, reaching half to two-thirds of distance to anus, 4.7-6.8 (mean 5.4) in SL. Caudal fin oval, rounded posteriorly, 3.3-4.6 (mean 3.8) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, placed at edge of preorbital, tube oriented down and forward, preorbital distinctly curved forward to accommodate nostril. Posterior nostril rounded, small, placed closer to anterior centre margin of eye than to halfway between eye and preorbital edge. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in three), with low irregular fleshy ridge (in one) or with fleshy flaps or knobs (in 26). Gill rakers on outer face of first arch short, slender, longest rakers near angle of arch; rakers on inner face of first arch longer, but short and slender; inner rakers on other arches twice as long as first arch inner rakers. Tongue tip blunt or concave, A B ,:'; \" Figure 120 Mugilogobius fusculus, papillae pattern, USNM 316193, Popondetta, Papua New Guinea. A, lateral view, scale bar = 1 mm; B, ventral view of chin (not to scale). Scalation omitted. absent in four specimens. Outer teeth in upper jaw largest, stout and curved, three to four rows of small sharp teeth behind this row; one or two rows at side of jaw; teeth in both jaws considerably larger and stouter in males. Lower jaw with about five rows of small sharp teeth across front, outermost row often oriented upright, inner row teeth all pointing posteriorly; innermost teeth largest and stoutest; usually only one row (innermost) of teeth at side of jaw. Predorsal scales small, evenly sized, usually reaching forward to close behind eyes, or at least to above preopercular margin. Operculum with small cycloid scales on upper half to two-thirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly usually with isolated patch under pelvics of weakly ctenoid scales (covering anterior 1,4 to lfz of belly), rest of scales cycloid. Ctenoid scales on side of body extending forward in wedge to behind pectoral fin. Genital papilla in male elongate, flattened, with pointed tip; in female, rounded and bulbous, blunttipped. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 120. Three s rows on snout, of three to six papillae each. Papillae in rows on cheek small, evenly-sized. Mental f row of four to eight papillae; anterior to this, row i extending forward in U -shape onto chin. Coloration of fresh material Few live colour notes available; I recorded fish as being dull greyish brown with yellow streak above black blotch on first dorsal fin. Notes below based on rather poor slide of freshly dead specimen from the Biges River, Madang. Fish dark brownish grey with dark brown mottling and stripes on head, and dark brown cross-hatching, indistinct oblique bars and X-shaped markings on posterior half of body and dark brown oblique shoulder bar; most distinctive marking a black spot on upper part of caudal fin base. First dorsal fin dark grey with broad pinkish yellow submarginal stripe; dense black oval spot on rear half of fin immediately below this stripe; elongate first spine dark pinkish yellow. Second dorsal fin dark grey with broad pinkish yellow submarginal stripe placed closer to dark grey fin margin than counterpart on first dorsal fin (i.e. more than lower half of fin dark grey); short blackish vertical streak on membrane on lower half of fin, next to each fin ray. Anal fin plain, very dark brown, with dark brownish grey margin. Caudal fin plain dark brownish grey with single round black spot on upper half of fin base. Coloration of preserved material Head and body light brown to yellowish brown Revision of Mugilogobius 129 Figure 121 Mugilogobius fusculus, 33 mm SL female, NTM 5.13689-005, Biges River, Papua New Guinea. Figure 122 Mugilogobius fusculus, 33 mm SL male, NTM 5.13689-005, Biges River, Papua New Guinea. with seven to nine dark brown dorsal saddles, oblique bars, chevrons and X-shaped markings along side; anteriormost short, oblique "shoulder" bar usually not distinguishable from other markings; posteriormost dark bar or chevron at caudal base not intensified, sometimes extending onto upper and lower parts of caudal fin and onto procurrent rays (Figures 121-122). Nine to ten indistinct dorsal saddles often joining lateral oblique bars or chevrons, giving banded appearance to body. Pectoral base pale brown to whitish with narrow brown horizontal streak or Table 18 rounded blotch just above midbase, extending onto fin as diffuse brown curved streak over pectoral ray bases. Head pattern variable, often indistinct. Most distinct marking (always present to some degree) a dusky brown oblique stripe from rear of eye to midopercle, joining brownish blotch on anterior part of opercle. Up to seven indistinct brownish curved or straight lines radiating outward from eye, shortest lines entering interorbital area, lines crossing preorbital and cheek often reticulate and diffuse. Snout and interorbital area plain, dark brown Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius fusculus (Nichols, 1951). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. 1. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital 1. in HL Pectoral 1. in SL Pelvic 1. in SL Caudal 1. in SL Longest Dl spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 28.7 64.0 83.7 17.0 28.7 53.3 61.2 15.2 9.6 21.6 11.9 23.9 23.2 39.7 25.4 16.0 14.7 21.8 14.6 32.5 66.7 85.6 19.5 14.1 25.1 13.8 29.9 30.8 48.8 34.6 24.8 21.4 30.5 44.2 30.5 59.0 74.5 17.9 12.1 24.1 13.1 27.0 26.3 43.3 31.1 21.9 18.9 26.5 28.9 28.4 50.0 60.9 16.1 10.0 22.5 10.4 21.7 24.2 33.3 26.0 19.3 15.2 22.2 14.0 32.0 69.2 79.5 19.6 15.6 26.9 14.4 27.5 30.0 39.4 32.2 22.5 19.7 30.0 26.8 29.7 57.9 71.4 18.1 12.0 24.9 12.4 25.1 27.0 36.9 28.4 20.9 18.4 26.6 18.9 24.0 13.3 29.1 23.3 47.7 26.7 16.0 14.7 23.3 34.7 ... ｾｾ o ＢＧＯＨＬｾＱ Aa!JJAPAN rJ?!' :;# , . 30' 'RYUKYU IS, . , '. .' HAWAIIAN ISLANO(Cl 20", 20" ft' LACCADIV( IS, ': .. 10" 1: MoIdlve. :'; ';f , : MARlANA IS, ',' :,. .... ". CAROUN( IS: .,i 'iSEYCHELlEs ＴｹｩＧＯｾ Ko'rae . O' ., 10' ..... ,': MAIlSHALL IS, ;: 10" KIRIIlATI .. ,.: 20" '0" MADAGASCAR 30' a , Mauritius • . 10' ;;':J;A .,e,"'" ;:', ", \ * IJI" • ｾ fuscu/us littora/is n. sp. • mertoni • myxodermus ｾ notospilus ... p/atynotus * LOYALTY IS. '. ',' 20' ". 30"· .' ｾＧ V AO' 0' ' ' , " '. ....'"ｾＮｲ 120" _.. ｾ､Ｇｾｉ Figure 123 Distribution of Mugilogobius jusculus, M. littoralis n, sp., M. mertoni, M. myxodermus, M. notospilus and M. platynotus, 160' ｾＧ " 180' 160' :t ｾ rUl o ::l 131 Revision of Mugilogobius spotted or with dark vermiculate markings. Rearmost radiating line from eye (if visible) extending horizontally onto upper part of opercle. Underside of head and branchiostegal membranes pale or dusky brown, or crossed by about four curved brown lines; posteriormost curved line crossing branchiostegal membranes and extending up onto rear part of opercle; usually branchiostegal area plain dark brownish grey, without curved lines; breast pale. Gill arches dusky; roof of branchial chamber pale to light dusky. Peritoneum brown dorsally, pale ventrally and laterally. First dorsal fin with lower half dusky grey to brownish grey, with large dense black spot on central rear half of fin; white to translucent whitish broad submarginal band, marginal band narrow, brownish to dusky grey; tips of elongate fin spines white, usually first dorsal spine mostly white. Second dorsal fin mostly dusky brownish to grey, with narrow grey to brown marginal band and broad white to translucent submarginal band; dusky lower part of fin with series of indistinct vertically elongate brown streaks on fin membrane. Anal fin plain dusky to brownish, margin slightly darker. Caudal fin mostly plain dusky grey to brownish grey with narrow translucent to whitish margin, sometimes diffuse brown spotting and indistinct streaks near base, most distinct marking a single small dark brown to blackish rounded spot close to fin base just dorsal to mid-point. Pectoral fin translucent, fin rays usually narrowly outlined with brown. Pelvic fins dusky to dark brown, sometimes with translucent to white margin, fraenum usually paler than rest of fin. Mature males display an indistinct colour pattern, with head considerably darker than pale body and chevrons and bars on side usually faded; spot on caudal fin always visible. Comparisons In morphology and general appearance, M. fusculus is very like M. rivulus sp. novo from northern Australia. It can be differentiated most easily by colour: M. fusculus has several indistinct stripes on the head, some of which may be curved (M. rivulus sp. novo has a finely reticulate and spotted head pattern, giving it an ocellate appearance); M. fusculus has a relatively plain to barred dark body with the single blackish caudal fin base spot being the most distinctive feature (versus a distinct dark oblique shoulder bar, body scale margins all narrowly edged with brown and body barred anteriorly or only dorsal saddles present, in M. rivulus sp. nov.). Mugilogobius fusculus also resembles M. filifer sp. novo from northern Australia and eastern New Guinea, which differs in having 1,7 dorsal and anal rays, usually has the anteriormost few predorsal scales larger than others and, although it has similar basic colouring, is more distinctly marked. Mugilogobius fusculus is generally plainer, never looks "chequered", with its most distinct marking being the small blackish spot on the upper part of caudal fin base. Distribution Specimens are known only from northern Papua New Guinea (Figure 123). Ecology Known from freshwater to brackish habitats, from mangrove creeks to a lily-covered still lake. A salinity range of 1-33 %0 was recorded in the Biges River, N of Madang, from three localities inhabited by this species. Remarks In the original description of M. fusculus, Nichols (1951) did not state exactly where his specimens were from, only that they were from "new Guinea". No subsequent information has clarified the locality. Alien and Coates (1990) state that their Sepik specimen was the "second" to have been collected. Strictly speaking, this is inaccurate, as there are four AMNH paratypes (Figure 124). I have examined Parenti and Allen's (1991) "Mugilogobius sp." specimens held at CAS; hence the confidence in placing their name in the synonymy. Figure 124 Mugilogobius fusculus. Paratype of Ta mankn fusculus Nichols, 28 mm SL, AMNH 19520, New Guinea. H.K. Larson 132 Mugilogobius latifrons (Boulenger, 1897) Figures 18B, 87, 125-127; Tables 5-8, 19 Gobius latifrons Boulenger, 1897: 427, plate 28, figure 2 (Lake Matanna and Kalaena River, Celebes). NTM 5.12707-003,1 Lake Towuti; CMK 6470, 4, Lake Towuti; CMK 6489, 1, Lake Towuti; CMK 6227, 1, Lake Towuti. CMK 6179, 1, Lake Matano; NMBA 2735, 1, Celebes; NMBA 2736, 1, Celebes. No locality given: RMNH 14358, 1. Gobius latifrons: Weber 1913: 198, 200, 212; Kottelat and Sutter 1988: 56. Vaimosa latifrons: Aurich 1938: 165-167. Tamanka latifrons: Koumans 1953: 155-156; Whitten et al., 1988: 295. Mugilogobius latifrons: Kottelat 1991: 343; Larson and Kottelat 1992: 226, 231-232; Kottelat et al., 1993: xxviii, 146, plate 68. Material Examined Lectotype NMBA 1847, 38.5 mm SL female, Lake Matanna and Kabaena River, Sulawesi, Indonesia, P. and F. Sarasin, 1895-1896. l'aralectotypes NMBA 1848-52, 2734, 6(18.5-29), same data as lectotype; BMNH 1897.3.8.5-7, 3(26-33), south-east Celebes, Sarasin. Other Material Indonesia: Sulawesi: ZMA 113.627, 2(22.5-29.5), Soroako, Matano Lake, E.C. Abendanon, 3 Sept. 1909; ZSM/CMK 6274, 3(17-19), about 13 km west of Soroako, Lake Matano, M. Kottelat, 5 July 1988; ZSM/CMK 6221, 7(18.5-29.5), 6 km south of Timampu, Lake Towuti, M. Kottelat, 22 June 1988; NTM 5.12706-005, 4(23-29.5), Lake Towuti, H. Larson and R. Williams, 15 September 1989; NTM 5.12704-005, 5(10-19), rice padi margin, Lake Matano, H. Larson and R. Williams, 14 September 1989; NTM 5.12702-002, 34(8-25), old buffalo wallow W of Nuha village, Lake Matano, H. Larson, 12 September 1989; CMK 11396, 1(45), tributary entering Lake Larona from south, M. Kottelat, 12 February 1995. Other material examined (but not used in description) Indonesia: Sulawesi: CMK 6206, 20, Lake Towuti; Diagnosis A small Mugilogobius with second dorsal rays 1,79, modally 1,8; anal rays 1,7-8, modally 1,8; pectoral rays 14-16; longitudinal scales 28-35; TRB 9-13; circumpeduncular scales 11-13; predorsal scales small, 16-23, usually reaching halfway between preopercular margin and eyes; ctenoid scales on sides of body usually extending forward in wedge to behind pectoral fin; third spine of dorsal fin longest, no spines elongate; body plain brownish, mottled with irregular dark brown markings; known only from freshwater lakes and tributaries of central Sulawesi. Description Based on 41 specimens, 17-45 mm SL. An asterisk indicates counts of lectotype of Gobius latifrons (Figure 125). First dorsal VI* (V in one); second dorsal 1,7-1,9 (mean 1,8*); anal 1,7-8 (mean 1,8*), pectoral rays 1416* (mean 15), segmented caudal rays 16-17 (mean 16*); caudal ray pattern modally 7/6 (8/6*); total branched caudal rays 12-15 (mean 14*); unsegmented (procurrent) caudal rays 7/7 (in four); longitudinal scale count 28-35 (31*; mean 32); TRB 9-13 (12*; mean 11); predorsal scale count 16-23* (mean 18); circumpeduncular scales 11-13 (mean 12). Gill rakers on outer face of first arch 2+7 to 4+7 (modally 2+7). Pterygiophore formula 3-12210 (in nine). Vertebrae 11+16* (in 13). Neural spines of anterior vertebrae with tip narrow and pointed (in seven). Two epurals present (in 11). Three anal pterygiophores before haemal spine of first caudal vertebra (in 11). Head and anterior part of body roughly rounded, body compressed posteriorly. Head bluntly rounded, may be slightly depressed anteriorly, HL 2.8-3.4 (mean 3.2) in SL. Head depth often equal to or slightly greater than depth at posterior preopercular margin; depth 1.6-2.0 (mean 1.8) in Figure 125 Mugilogobius latifrons. Lectotype of Gobius latifrons Boulenger, 38.5 mm SL, NMBA 1847, Lake Matano, Sulawesi. Revision of Mugilogobius HL. Width at posterior preopercular margin 1.3-1.6 (mean 1.4) in HL; cheeks not particularly enlarged or fleshy. Mouth subterminal, sometimes barely tenninal, slightly oblique, fonning angle of about 15-20° with body axis; jaws generally reaching to below anterior half of eye or to mid-eye (to below mid-eye in lectotype, a large female); jaws not much enlarged in adult males. Upper jaw 2.3-3.6 (mean 3.0 in females, 2.8 in males) in HL. Lips smooth, not thick or fleshy; row of fine low fimbriae often present on inner edges of both lips; lower lip free at sides, fused across front. Eyes relatively small, lateral, high on head, top usually forming part of dorsal profile, 3.0-4.4 (mean 3.7) in HL. Snout rounded, not fleshy or overhanging, 3.2-4.4 (mean 3.9) in HL. Interorbital moderately broad, flat to very slightly concave between bulges of eyes, 1.45.5 (mean 4.0) in HL. Top of head anterior to nape scales sometimes with small, sparsely scattered villi, usually present on interorbital area (villi often only visible if mucous coat preserved and undamaged). Body depth at anal origin 4.3-5.9 (mean 5.1) in SL. Caudal peduncle compressed, length 3.4-4.6 (mean 4.3) in SL. Caudal peduncle depth 7.2-9.1 (mean 8.3) in SL. First dorsal fin low, rounded, tips of spines free, second to fourth spines often longest, but none ever much longer than others and never filamentous; fin not reaching second dorsal origin when depressed. First dorsal spine longest in one specimen, 3.1 in SL. Second dorsal spine length 5.9-7.7 (mean 6.8) in SL. Third dorsal spine length 6.2-8.3 (mean 7.1) in SL. Fourth dorsal spine length 6.0-7.3 (mean 6.9) in SL. Second dorsal and anal fins low, rounded, posterior rays not much longer than anterior; rays falling well short of caudal fin when depressed. Pectoral fin rounded to slightly pointed, central rays longest, 3.7-4.9 (mean 4.1) in SL; uppermost ray usually short and unbranched. Pelvic fins short, round to oval; reaching up to about half distance to anus, or slightly more, 3.8-5.4 (mean 4.7) in SL. Caudal fin rounded to roughly rectangular, 3.3-4.2 (mean 3.7) in SL. One apparently sexually mature male with all fin rays relatively longer than usual and depressed first dorsal fin reaching back to base of second element of second dorsal fin; posterionnost rays of second dorsal and anal fins much longer than anterior, fully depressed rays still falling short of caudal base; pelvics long, nearly reaching anus. No mental fraenum, chin smooth. Anterior nostril short, tubular, placed just behind upper lip, and oriented down and forward; preorbital produced forward to acconunodate nostril. Posterior nostril small, round to oval, placed just before upper half or anterior centre margin of eye. Gill opening extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in one), with low irregular fleshy ridge or raised bumpy flange (in 17), or with one to several distinct 133 fleshy knobs and bumps, often on thickened ridge (in 22). Gill rakers on outer face of first arch very low, stubby and smooth, longest rakers near angle of arch; rakers on inner face of first arch fat and stubby; outer rakers on other arches stubby, inner rakers on other arches more slender, nearly twice length of first arch inner rakers. Tongue tip usually blunt, sometimes slightly concave (nearly bilobed in one). Outer row teeth in upper jaw largest, sharp and curved, inner two to three rows of very small sharp teeth present. Lower jaw with about four rows (five in lectotype) of small sharp, curved teeth, evenly sized; inner row teeth pointing posteriorly. Teeth size not greatly dissimilar between male and female. Predorsal scales small, evenly sized, usually reaching forward about halfway between preopercular margin and rear of eyes. Operculum covered with small cycloid scales at least on upper two-thirds; rarely only upper half scaled. Cheek always naked. Pectoral base covered with small cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated area close to pelvic base with weakly ctenoid scales (covering anterior 112 to 1/3 of belly), rest of scales cycloid. Ctenoid scales on side of body in wedge extending up to behind pectoral fin, wedge narrowing approximately below second dorsal fin origin. Genital papilla in male elongate, flattened, with pointed tip; in female, rounded and bulbous, with blunt tip. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 126. Cheek row a extending posteriorly just past row p in one large male specimen (in CMK 6221); three 5 rows on snout, composed of three to five papillae each; two to three papillae on each side of mandibular symphysis (anterior part of row i) and short row f of about six papillae immediately behind these. Figure 126 Mugilogobius latifrons, papillae pattern. Composite of two specimens from CMK 6221. Scales omitted. Scale bar = 1 mm. 134 Table 19 H.K. Larson Morphometries as percentages of SL or HL, as indicated, of Mugilogobius latifrons (Boulenger, 1897). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic !. in SL Caudal!. in SL Longest D1 spine in SL Lectotype 29.9 59.1 65.2 22.3 14.8 24.9 24.3 24.3 37.4 28.7 19.7 23.9 12.5 Males Males Minimum Maximum 29.2 50.0 63.3 17.1 10.2 22.8 11.0 22.8 22.6 32.3 21.5 22.2 20.4 24.9 14.1 35.3 64.3 77.8 21.9 14.3 28.6 13.7 28.6 29.7 44.3 32.9 27.1 24.7 30.6 16.7 Coloration of fresh material A good colour photograph of large (50 mm SL) captive specimen in Larson and Kottelat (1992: 232), shows pale body colour (very light grey) with brown scale margins forming characteristic reticulate pattern and pale yellow-gold ventral area, and very small reticulations on side of head. Freshly dead specimen shown in Kottelat et al. (1993: plate 68). From my field notes on live specimens from Lake Towuti: fish very dark with body scales outlined by narrow blackish edges and irregular, just visible, dusky marbling and barring across the back. Some specimens very pale, but barring still visible. Side of head marbled with darker brown in large specimens, with blackish trim along edges of opercle and preopercle; both lips nearly black. Underside of head pinkish to yellowish with fine curved black lines forming U-shapes across width of head. Pectoral base with broad dusky vertical bar across fin ray bases, with yellowish bar before and behind dark bar. Area above black blotch on first dorsal fin bright yellow. Distalrnost stripe on second dorsal fin pinkish; base of fin with yellowish stripe. Males Mean Females Minimum Females Maximum Females Mean 31.9 57.3 70.1 19.5 12.4 25.8 12.1 25.8 26.7 36.0 26.1 24.8 21.6 27.6 15.4 29.6 51.6 61.4 16.9 10.8 21.7 11.0 22.2 24.2 29.1 20.4 18.6 23.9 16.8 12.0 33.7 66.2 82.4 23.6 16.6 29.7 13.8 30.9 31.1 38.2 34.6 26.8 25.3 29.5 16.9 31.6 58.6 70.7 19.9 13.3 24.8 12.0 26.1 26.8 33.7 25.1 24.1 20.9 26.9 14.8 Coloration of preserved material Head and body light yellowish brown, brown on posterior half of scales on most of body, giving overall finely reticulate pattern; underside of head, belly and ventral edge of body relatively pale (Figure 127). Body faintly and very irregularly barred and mottled with brown, bars indistinct, usually partly joined by mottling, markings on anterior upper half of body often more distinct; about nine narrow brown dorsal saddles or crossbars sometimes visible. Unscaled portion of top of head (interorbital and snout) with indistinct brown reticulation and vermiculation. Side of head with two to four brown bars extending from eye (bars often obscure), arranged almost exactly as in M. notospilus, and cheek sometimes with dark spotting and reticulation enclosing lighter areas. Opercle with obscure brown mottling or plain brown. Lips brown to very dark brown, edges of lips darkest. Underside of head pale, with up to seven narrow brown curved lines running up onto side of head in U-shape; first line crossing isthmus at beginning of preopercular limb, last line on branchiostegal membranes; these lines Figure 127 Mugilogobius latifrons, 29.5 mm SL, CMK 6221, Lake Towuti, Sulawesi. Revision of Mugilogobius may be indistinct, broken-up, or one or more missing, but present even in pale specimens. Caudal base usually without any dark spots; occasionally small dark brown spot at centre of base, and another small brown spot at base of upper procurrent rays. First dorsal fin with dusky to blackish distal stripe (stripe width about one quarter to one third of dorsal fin height); tips of spines unpigmented. Below this stripe, narrow unpigmented area present, remainder of fin dusky to blackish, with central area Gust below unpigmented area) usually more intense, widening posteriorly to dense black spot between membranes of fifth to sixth rays, sometimes scattered pale areas on fin in front of spot. Second dorsal fin with pale to unpigmented margin, remainder of fin dusky, especially toward centre, where vertical dark streaks present on membranes between rays; bases of fin rays often very dark, with adjacent membrane paler than dusky area above. Anal fin plain dusky with translucent margin. Caudal fin translucent to dusky, with small brown spots or broken-up wavy lines, spots or lines often forming irregular vertical rows, especially toward midbase of fin; fin-rays usually brownish, especially near bases; usually translucent margin around fin. Pectoral fin with ray bases dusky to dark brown, membranes sometimes dusky towards bases. Pelvic fins dusky with translucent margin; fraenum may be unpigmented. Peritoneum brown dorsally, pale ventrally and on lower sides. Comparisons This species is closely related to M. adeia and is discussed under Comparisons for that species. Distribution This species is known only from three freshwater lakes (Lakes Matano, Towuti and Mahalona) in central Sulawesi, Indonesia (Figure 87). Ecology This species is restricted to freshwater, in three of the tectonic lakes of central Sulawesi. These are very steep-sided, deep lakes (Matano is 590 m deep, Towuti 203 m and Mahalona, the smallest of these three, is 73 m deep), with aquatic vegetation and native fish fauna generally restricted to the shallow sides (Kottelat, 1989b, 1990a). The lakes and their habitats are discussed further under the Ecology section for M. adeia. In Lake Matano, Mugilogobius latifrons is usually found in very shallow water (less than 50 cm depth), typically among pebbles (Larson and Kottelat, 1992). It also occurs in shallow, muddy bays protected from wind and wave disturbance, or where vegetation is thick such as rice padi or Charafilled water buffalo wallows. In general terms, Mugilogobius species favour leaf litter over aquatic 135 vegetation for concealment. However, field observations indicate that, where the Chara was thick, M. latifrons would be found among it and not among the leaf litter in neighbouring areas without Chara. Remarks In 1897, Boulenger described a species of Mugilogobius (as Gobius latifrons), which was collected from Lake Matano by the Swiss naturalists Fritz and Paul Sarasin in 1896. Weber (1913) described several freshwater fishes from Sulawesi, such as the now very rare Mugilogobius amadi (Weber) from Lake Poso (Whitten et al., 1987; Kottelat, 1990a), and Glossogobius matanensis (Weber) from Lake Matano. Koumans (1953) included Vaimosa cagayanensis as a synonym of this species, without giving reasons, although he placed a question mark next to the locality. Ladiges et al. (1958), in their type catalogue of Hamburg museum specimens, erroneously included Vaimosa cagayanensis as a synonym of Tamanka latifrons and established a lectotype for cagayanensis (see further discussion in Remarks under Mugilogobius cagayanensis). The largest NMBA syntype, NMBA 1847 (Figure 125), is hereby designated lectotype of Gobius latifrons, as it is in reasonable condition and resembles the specimen illustrated by Boulenger (1897: plate 28, figure 2). As Larson and Kottelat (1992: 233) indicated, one 19.5 mm female syntype (now paralectotype) (NMBA 1847-52) does not belong to this species; it is now identified as a specimen of Redigobius penango (Popta, 1922). Weber (1913) gave the Buginese name of this fish (based on information from E.C. Abendanon) as piniponro. During my visit to Lake Towuti in 1989, local villagers (from Nuha and Subario villages) used the name bontini for small lake gobies of the genera Mugilogobius and Glossogobius. Mugilogobius lepidotus sp. novo Figures 87, 128-132; Tables 5-8, 20 "Sandgrundel": Kottelat 1989b: 684, figure 17. Material Examined Holotype MZB 5946, 24.5 mm SL male, Tandjung Subalaote, Lake Towuti, Sulawesi, Indonesia, M. Kottelat, 30 January 1993. Pa ra types Indonesia: Sulawesi: MZB 5948, 3(18-20.5), about 3 km S of Timampu, Tandjung Posombuwang, M. Kottelat, 29 June 1988; CMK 6251,3(16.5-22.5), same 136 Figure 128 H.K. Larson Mugilogobius lepidotus n. sp., holotype, MZB 5946, 24.5 mm SL male, Tandjung Subalaote, Lake Towuti, Sulawesi (elongate object at front of mouth is paper tag). data as preceding; CMK 6491, 5(18.5-24.5), Watidi, 4-7 km E of Timampu, M. Kottelat and A. Werner, 15 March 1989; N1M 5.14700-001, 5(19.5-26; ), same data as preceding; MZB 5947, 4(19.5-25.5), same data as holotype; N1M 5.14701-001, 6(21-26), same data as holotype; CMK 9752, 7(22-26), same data as holotype. Diagnosis A small Mugilogobius distinguished by combination of characters: second dorsal rays 1,8-9; anal rays 1,8-9; pectoral rays 14-16; lateral scales 25-28; TRB 6-8; circumpeduncular scales 11-12; predorsal scales 8-12; preoperculum with small cycloid scales at least ventrally; profile rather pointed; upper jaw somewhat protrusible, mouth tending to open down and forward; third and fourth first dorsal spines very long but usually not free from membrane, gill arches papillose with short papillose rakers on outer face of first arch, inner and outer faces of other gill arches with oval papillose pads; colour pattern including three or four narrow dusky stripes along upper side of body; restricted to sandy habitats in Lake Towuti, central Sulawesi, Indonesia. Description Based on 30 specimens, 16.5-26 mm SL. Counts of holotype (Figure 128) indicated by asterisk. First dorsal VI*; second dorsal 1,8*-9 (mean 1,8); anal 1,8-9* (mean 1,9); pectoral rays 14*-16 (mean 15); segmented caudal rays 15-16 (mean 16*); caudal ray pattern modally 9/7; branched caudal rays 5/5 to 7/6 (modally 6/5); unsegmented (procurrent) caudal rays 8/7 (in one); longitudinal scale count 25*-28 (mean 27); TRB 6-8* (mean 7); predorsal scale count 8-12 (mean 11, 10 in holotype); circumpeduncular scales 11-12* (mean 12). Gill rakers on outer face of first arch 2+5 to 4+6 (modally 3+5). Dorsal pterygiophore formula 312210 (in 14). Vertebrae 11+16 (in 15). Neural spines of first few vertebrae slender and pointed. Premaxilla with long ascending process. Quadrate low, nearly triangular, not forked, joined to metapterygoid by cartilage (Figure 129). Metapterygoid low, not expanded dorsally, without bony bridge to quadrate. Fifth ceratobranchial rather slender; narrow high flange present on ventral surface. One or two rakers ossified on first gill arch. Pectoral radials partly unossified. Two epurals (in two). Three (in 15) anal pterygiophores before haemal spine of first caudal vertebra. Body compressed, more so posteriorly. Head rather compressed, square to somewhat triangular in cross-section, HL 3.1-3.6 (mean 3.4) in SL. Head width usually slightly greater than depth, although depth and width often equal; mean head depth at posterior preopercular margin 1.4-1.8 (mean 1.7) in SL. Mean head width at posterior preopercular margin 1.4-2.0 (1.6) in SL. Profile pointed to slightly rounded. Mouth terminal to slightly subterminal, slightly oblique, forming angle of about 15° with body axis; upper jaw fairly protrusible, jaws tending to open downward and forward; jaws ending under anterior third of eye. Mouth 2.6-3.5 (mean 3.1) in HL, upper jaw length similar in males and females, 2.7-3.5 (mean 3.2) in females, 2.6-3.5 (mean 3.0) in males in HL. Lips narrow, relatively thin, smooth, usually with fine fimbriae on inner edges of both lips; lower lip mostly free, fused across front of jaw. Eyes lateral, set high on sides of head, forming part of dorsal profile, 2.7-3.5 (mean 3.2) in HL. Snout relatively steep, slightly pointed, 3.1-3.9 (mean 3.6) in HL. Interorbital rather narrow, 5.4-8.5 (mean 6.8) in HL. Usually without any fine fleshy villi on naked areas of head. Body slender, Figure 129 Jaws and suspensorium of Mugilogobius lepidotus, male, ex CMK 6491, Lake Towuti, Sulawesi. Stippling indicates poorly ossified bone. Scale bar = 1 mm. Revision of MlIgilogobills Figure 130 137 MlIgilogobills lepidotlls n. sp., 26 mm SL female paratype, ex CMK 9752. Upper, first gill arch showing papillose rakers; lower, second gill arch showing fleshy papillose pads. Scale bar = 1 mm. depth at anal origin 4.9-5.7 (5.4) in SL. Body width above anal fin origin 7.3-9.2 (mean 8.3) in SL. Caudal peduncle long, compressed, length 3.3-4.2 (3.7) in SL. Caudal peduncle depth 8.6-10.3 (mean 9.2) in SL. First dorsal fin triangular, with second to fourth dorsal spines elongate, but generally not free from fin membrane. Third dorsal fin spine longest, Table 20 maximum 2.6-4.4 (mean 3.2) in SL in males, and 3.3-4.7 (mean 4.2) in females; extending well beyond other spines and sometimes reaching to base of last second dorsal fin element. Fourth dorsal spine longest in one male specimen only. Second dorsal fin roughly triangular, low, usually higher anteriorly than posteriorly. Anal fin low, with posteriormost rays longest. Depressed second dorsal and anal fin rays usually only reaching halfway along caudal peduncle. Pectoral fins narrow, central rays longest, 3.8-5.1 (mean 4.4) in SL; most pectoral rays branched, upper and lower two or three often unbranched. Pelvic fins long, narrow, reaching to anus and slightly beyond, 3.74.8 (mean 4.3) in SL. Caudal fin rounded to slightly rectangular in shape, upper rays sometimes slightly longer than lower, 3.5-4.3 (mean 4.0) in SL. No mental fold or fraenum. Anterior nostril in short tube placed on preorbital edge, directed mostly downward over upper lip. Posterior nostril with low rim, closer to eye than midway between eye and tip of snout. Gill opening extending to below opercle. Gill rakers on outer face of first arch very short, round and robust, with papillae on tips and outer face; rakers on inner face of first arch reduced to small round pads covered with papillae; anteriormost portion of arch covered with papillae (Figure 130). On second arch, outer rakers are oval pads covered with papillae; rakers on inner face of second arch short, resembling those on outer face of first arch, as do inner and outer rakers on remaining arches. Pectoral girdle smooth (in 15, including holotype), with low irregular flange and no distinct bumps (in six), or with one to three low bumps (in 14). Tongue short, blunt to slightly concave, usually very reduced, absent in two specimens. Teeth in outer row of upper jaw somewhat enlarged, curved, evenly spaced, followed by four or five rows of smaller, sharp conical teeth; two rows at sides of Morphometrics as percentages of SL or HL, as indicated, of MlIgilogobills lepidotlls sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. L in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital L in HL Pectoral L in SL Pelvic L in SL Caudal L in SL Longest Dl spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 298 63.0 671 18.8 122 269 11.4 288 315 370 178 23.3 24.1 26.5 36.3 28.8 54.8 53.4 17.6 10.9 24.7 9.7 255 288 28.3 118 22.2 22.5 24.3 22.7 32.3 67.1 68.5 19.6 13.3 29.7 11.7 31.9 338 385 185 25.1 27.2 28.3 38.0 30.3 60.1 61.3 18.6 12.1 27.2 10.9 28.4 31.6 34.0 15.1 23.8 24.1 25.8 36.3 27.5 54.8 49.3 17.4 11.1 24.1 10.2 25.4 29.9 28.8 11.9 19.6 21.0 23.1 21.2 30.9 70.0 729 20.4 13.8 30.0 116 30.0 36.8 37.1 17.9 26.7 26.7 26.7 30.0 29.4 61.3 61.8 18.9 12.2 27.3 11.0 27.6 31.6 31.8 14.7 22.2 22.7 24.4 24.2 H.K. Larson 138 flattened, narrowing toward tip; papilla in female short, rounded and stout, with no lobes at tip. Gut simple S-bend, although rather long and slender. Head pores absent. Sensory papillae on head in longitudinal pattern, all papillae small, evenly sized (Figure 131). Three s rows on snout, of three or four papillae; often only one or two papillae in middle s row. Row f usually of one or two papillae behind chin, oriented transversely. B Figure 131 Mugilogobius lepidotus n. sp., papillae pattern. Para type, from CMK 6491. A, lateral view; B, ventral view of chin area (not to scale). Scales omitted. Scale bar = 1 mm. jaw. Teeth in lower jaw small, sharp and curved, about four rows anteriorly, and two at sides. Teeth often slightly larger in males. Predorsal scales cycloid, small and even, none enlarged, scales reaching forward to close behind eyes. Operculum with small cycloid scales, lower third or more of opercle often unscaled. Cheek scaled, usually ventrally, with two to seven rows of small cycloid scales. Pectoral base scales small, cycloid. Prepelvic area scales cycloid, area usually fully scaled. Belly scales ctenoid, small scales close to anus cycloid. Genital papilla in male slender, elongate and Figure 132 Coloration of fresh material Based on colour slides. Head and body translucent yellowish grey with six evenly spaced, brownish grey, square blotches or saddles crossing dorsal midline (first two on nape, last two on caudal peduncle); scales on upper half of body with slightly darker centres; and four dull yellow stripes along side of body; upper two stripes beginning behind head, lower two behind pectoral base; stripes ending at caudal base; dark brown mark on head behind upper rear edge of eye; lips speckled brownish; iris mottled dark brown and gold; fins translucent with dusky rays; first dorsal fir with rounded dark grey blotch crossing upper portions of fifth and sixth spines. In freshly dead specimen, yellow stripes appear light brown; two central body stripes forming small brown spots at caudal base; dorsal saddles reaching down to uppermost body stripe, each saddle forming dark spot where it meets stripe; head with three oblique light brown lines: darkest extending from upper rear margin of eye (continuation of uppermost body stripe); remaining two oblique lines crossing upper part of preopercle and opercle, running forward obliquely. Kottelat (1989b) described live fish as grey with a few longitudinal stripes and some dark saddles along the back. Coloration of preserved material Very similar to that given above (Figure 132). On head, most distinct mark a short brown line from lower front margin of eye to upper lip; scattered brown spots and mottling present on top of head and nape; stripes on sides and dorsal saddles sometimes faint, but nearly always visible. Mugilogobius lepidotus n. sp., 23.5 mm SL, CMK 6251, Lake Towuti, Sulawesi. Revision of Mugilogobius Comparisons This species resembles M. rexi sp. nov., the other small slender species found in Lake Towuti. However, the papillose gill raker pads, striped colouring and scaled preoperculum set this species apart from all other species of the genus. Distribution Restricted to Lake Towuti, central 5ulawesi, Indonesia (Figure 87). Ecology Kottelat (1989b) described this species as being found exclusively on sandy bottoms. Mugilogobius littoralis sp. novo Figures 123, 133-136; Tables 5-8, 21 Material Examined Holotype NfM 5.14293-001, 27.5 mm 5L female, rock pools to left of Nightcliff Boat Ramp, Darwin, Northern Territory, Australia, C Jones, 23 July 1995. Paratypes Australia: Western Australia: WAM P.25668-018, 1(20.5), Port Warrender, Admiralty Gulf, J.B. Hutchins, 22 October 1976; WAM P.30303-017, 2(12.5-20.5), Mission Bay, Napier Broome Bay, G.R. Alien, 14 August 1991; AM5 1.25498-004, 1(22.5), New Beach, 5hark Bay, 45 km W of Camarvon, D.F. Hoese and D. Rennis, 8 September 1985; Northern Territory: NTM 5.12443-001, 8(6-24), beach rock pools, beach at Galiwinku, W side of Elcho Island, H.K. Larson, 1 February 1988; NTM 5.14296-001, 10(30.5-39), freshwater spring behind beach at Galiwinku, Elcho Island, K. Aland, November 1995, kept in aquarium until 5 March 1996; NTM 5.10439001, 4(22.5-25.5), mangrove lagoon at Danger Point, Cobourg Peninsula, H. Larson and R. Williams, 30 April 1982; NfM 5.14293-002, 14(20-28), rock pools to left of Nightcliff Boat Ramp, Darwin, C Jones, 23 July 1995; NfM 5.14294-001, 4(16.5-23.5), creek by 5ki Club, Darwin, M. 5elway, 23 April 1992; AM5 Figure 133 139 1.23929-003, 1(24.5), East Point, Darwin, D. Rennis, 1 August 1983; NfM 5.10452-022, 16(9-23), sandy mangrove at mouth of Cairnan Creek, Port Essington, H.K. Larson, B.C Russell and R.5. Williams, 4 May 1982; NTM 5.13513-010, 1(20.5), mouth of Bing Bong Creek, W of McArthur River, H. Larson, 4 September 1992; NTM 5.10430-002, 2(18.5-26), Ludmilla Creek, Darwin, R. Hanley, 21 January 1982; AM5 IA.4387, 1(20.5), Port Darwin, L. Wilson. Other material examined (but not used in description) Australia: Western Australia: WAM P.30919-001, 6, Macleay Island; AM5 1.25514-005, 1, Port Hedland; Northern Territory: NfM 5.10408-006, 6, Rapid Creek, Darwin; NTM 5.14295-001, 15, Vestey's Beach, Darwin; NTM 5.11931-001, 3, Vestey's Beach, Darwin; NTM 5.12990-002, 1, Vestey's Beach, Darwin; NTM 5.12871-001, 2, Groote Eylandt; NTM 5.10036-001, 14, Lee Point Reef. Diagnosis A moderately slender Mugilogobius with second dorsal rays 1,7-9, modally 1,8; anal rays 1,7-9, modally 1,8; pectoral rays 13-16; longitudinal scales 36-46; TRB 13-18; circumpeduncular scales 14-19; predorsal scales 12-21, small, reaching forward of preopercular margin but not up to behind eyes; ctenoid scales on side of body separated into patch behind pectoral fin and area from below second dorsal fin origin back to caudal base; body colour dusky grey with irregular vertical bars, blotches and small spots, often almost plain grey, first dorsal fin grey, black posteriorly, bright yellow submarginal band present; inhabiting high intertidal pools or lagoons; known from northwestern Australia. Description Based on 43 specimens, 12.5-39 mm 5L. An asterisk indicates counts of holotype (Figure 133). First dorsal V (in one), VI* (in 40); second dorsal 1,7-9 (mean 1,8*); anal 1,7-9 (mean 1,8*), pectoral rays 13-16* (mean 15), segmented caudal rays 15- Mugilogobius littoralis n. sp. Holotype, NTM 5.14293-001, Nightcliff, Darwin, NT. H.K. Larson 140 16* (mean 16); caudal ray pattern 7/6 to 9/7, modally 8/7*; branched caudal rays 13-16* (mean 15); unsegmented (procurrent) caudal rays 7/6 to 8/8; longitudinal scale count 36-46 (mean 41, 39 in holotype); TRB 13-18 (mean 16*); predorsal scale count 12-21 (mean 17, 15 in holotype); circumpeduncular scales 14-19 (mean 16, 15 in holotype). Gill rakers on outer face of first arch 3+8 to 4+9 (modally 3+8, 4+8 in holotype). Pterygiophore formula 3-12210 (in two). Vertebrae 10+16 (in 15), 10+17 (in one), 11+15 (in one). Neural spine of second and/or third vertebra thickened or rounded at tip (in five), or with tip expanded (in four). Two (in 16) or one (in one) epurals. Two (in 14) or three (in three) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid relatively slender, forming bridge to quadrate, not expanded dorsally. Body slender, compressed (less so anteriorly). Head somewhat depressed, always wider than deep, but not greatly so, HL 3.0-4.3 (mean 3.6) in SL. Depth at posterior preopercular margin 1.7-2.1 (mean 1.8) in HL. Width at posterior preopercular margin 1.1-1.5 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of about 20-25° with body axis; jaws generally reaching about to below middle of eye or slightly more anteriorly (as in holotype; not much difference in jaw length between males and females). Lips usually smooth, fleshy fimbriae often present across front of inner edge of upper lip and less often, on inner edge of lower lip; lower lip free at sides, fused across front. Upper jaw 2.3-3.6 (mean 2.9 in females, 2.6 in males) in HL. Eyes dorsolateral, high on head, top forming part of dorsal profile, 2.7-4.4 (mean 3.7) in HL. Snout slightly pointed to rounded, tip slightly inflated but not overhanging upper lip, 3.3-4.3 (mean 3.8) in HL. Interorbital broad, flat to slightly concave, 1.3-6.3 (mean 3.4) in Table 21 HL. Top of head, mostly in interorbital region, occasionally covered with very small, sparsely scattered villi. Body depth at anal origin 4.5-6.8 (mean 5.6) in SL. Caudal peduncle compressed, length 3.5-7.2 (mean 4.3) in SL. Caudal peduncle depth 6.6-13.4 (mean 7.7) in SL. First dorsal fin low, triangular to somewhat rounded, spines not free of membrane at tips, second to fourth spines longest or subequal; spines slightly longer in males than females; spines falling short of second dorsal fin origin when depressed; gap or three to four scales between dorsal fins. First dorsal spine always shorter than next three. Second dorsal spine length 6.5-10.6 (mean 7.7 in males, 8.6 in females) in SL. Third dorsal spine length 6.5-9.4 (mean 7.5 in males, 8.4 in females) in SL. Fourth dorsal spine length 7.5-8.7 (mean 8.0 in males, 8.1 in females) in SL. Second dorsal fin taller than or equal in height to first dorsal fin; anal fin lower than dorsals, posteriormost rays of second dorsal and anal fins longest, rays falling well short of caudal fin base when depressed. Pectoral fin broad, rounded, central rays longest, 4.1-6.1 (mean 4.9) in SL; rays all branched but for uppermost. Pelvic fins short, rounded to oval, reaching about half distance to anus, 4.7-6.5 (mean 5.5) in SL. Caudal fin short, rounded, 3.7-5.5 (mean 4.4) in SL. No mental fraenum, chin smooth. Anterior nostril in short tube, placed at edge of upper lip, tube oriented down and forward, preorbital curved forward to accommodate nostril. Posterior nostril round to oval, placed close to anterior centre margin of eye. Gill opening restricted to pectoral base or extending forward to just under opercle. Inner edge of pectoral girdle with low knobby or fleshy ridge or narrow irregular flange (in 11); or with distinct fleshy knobs or flaps (in 29). Gill rakers on outer face of first arch short, longest rakers near angle of arch; rakers on inner face of first arch same Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius littoralis sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. 1. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital 1. in HL Pectoral 1. in SL Pelvic 1. in SL Caudal 1. in SL Longest Dl spine in SL Holotype 26.2 56.9 76.4 15.6 9.8 23.3 12.4 27.8 23.6 34.7 26.4 19.6 16.0 20.4 11.6 Males Males Minimum Maximum 26.8 47.4 67.1 14.6 8.8 22.5 12.1 23.6 24.7 32.7 26.2 18.9 16.4 21.4 11.9 32.9 58.7 80.0 20.0 14.1 28.8 15.1 30.6 32.7 43.1 35.3 24.6 20.6 27.1 15.3 Males Mean Females Minimum Females Maximum Females Mean 29.2 54.2 72.6 17.5 10.9 25.0 13.8 26.7 27.9 38.3 29.6 21.3 18.6 24.8 13.4 23.3 50.0 66.0 15.6 9.3 20.8 11.9 24.0 22.7 28.0 15.9 16.9 15.3 18.2 10.7 30.2 68.4 87.4 22.5 17.0 26.1 14.8 29.5 30.2 37.4 36.4 22.6 21.2 26.8 13.2 27.3 59.5 77.6 19.0 13.6 23.7 13.7 27.1 26.4 34.9 29.3 20.1 17.8 22.1 12.4 Revision of Mugilogobius '. ....:: 0 . :.....q,:. ',' . : Figure 134 141 @ ..-: ＺＮｾＭ .," "";- ..... .. -' . - Mugilogobius littoralis n. sp., papillae pattern. Paratype, NTM 5.10439-001. Scales omitted. Scale bar = 1 mm. length as those on outer face; inner rakers on other arches slightly longer than those on first arch. Tongue tip blunt to slightly concave. Outer teeth across front of upper jaw largest, curved and pointed, three to four rows of small sharp teeth behind this row; one or two rows at side of jaw; teeth in females smaller than those of males. In males, lower jaw with four to five rows of small, curved teeth across front, rows generally pointing posteriorly; teeth toward midside of jaw largest and stoutest (but no individual teeth particularly enlarged), usually only one row of teeth at side of jaw; teeth in females similar, but smaller and less curved. Predorsal scales small, evenly sized, usually reaching forward to above preopercular margin or further; if further, usually scales near nape midline extending slightly forward (as in holotype); scales not reaching up to behind eyes. Operculum with small cycloid scales on upper two-thirds. Cheek always naked. Pectoral base covered with small cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anteriormost 1/4 to 1/3), rest of scales cycloid; some specimens only with few ctenoid scales close to pelvic fin base. Ctenoid scales on side of body extending forward in narrow wedge from caudal base to just below Figure 135 fourth dorsal fin ray or to gap between dorsals, and isolated oval patch of ctenoid scales behind pectoral fin (in three specimens, two areas of ctenoid scales barely joining together below dorsal fin gap). Genital papilla in male elongate, flattened, with pointed tip; in female, papilla short, rounded and bulbous. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 134. Broken cheek row c with two or three papillae in posterior portion. Three s rows on snout, central row consists of one papilla only. Mental f row of four, occasionally two, papillae. Gut simple, in long looped 5-bend. Coloration of fresh material From notes based on living specimens (NTM 5.14293-002 and NTM 5.14296-001). Head and body translucent whitish grey to iliac grey or greyish yellow, with pale grey bars and blotches, and scattered pale yellowish white or pearly white patches which can quickly be intensified or turned off. About seven saddles or pairs of grey blotches on either side of mid-dorsal line in one specimen. Markings along sides variable, usually rounded large dusky patches or blotches. Some fish without clear markings, others with dusky grey nebulous reticulation or spotting. On large fish, narrow grey scale margins visible. Nape, shoulder and top of head with dusky irregular anastomosing pattern; dark oblique shoulder bar present, very short, oval to rounded, shoulder bar may be indistinct or nearly horizontal. Small black spot crossing top of caudal peduncle, at base of uppermost few caudal rays. Side of head pale greyish with three distinct oblique blackish stripes crossing preorbital and cheek. Top of head and snout with short dusky vermiculate lines and spots. Dorsal surface of abdominal cavity, visible through musculature, whitish, with blackish dorsal midline and black blotches. First dorsal fin with grey to dark grey margin, with dull orange to chrome yellow submarginal stripe bordered above and below with dark grey to black; lower third of fin translucent pale grey. Second dorsal fin translucent to greyish with very narrow MlIgilogobills littoralis n. sp. Paratype, NTM 5.10439-001, Danger Point, Cobourg Peninsula, NT. H.K. Larson 142 Figure 136 Mugilogobius littoralis n. sp. Captive specimen from Darwin, NI. From colour slide by Neil Armstrong. dark grey margin and pinkish orange submarginal stripe. Caudal fin plain translucent greyish to whitish, with diffuse dusky spot at fin base, short vertical bar on fin close to base in large specimens. Pelvic fins dark grey. Caudal fin plain grey to brownish. Peritoneum dense black, slightly lighter ventrally in small specimens. Coloration of preserved material Head and body brownish yellow to grey or yellowish grey (depending on preservation), with variable light to dark grey spots, irregular crosshatched blotches and 7-10 vertically oriented, short, dark grey bars or blotches along sides, and about seven dark blotches or saddles across dorsal midline; males often almost plain dark brown or grey (Figures 133, 135). Most consistent and distinctive markings: two oblique lines crossing cheek and preorbital region (other less distinct lines sometimes present) (Figure 136) and small dark grey spot at centre of caudal base (on hypural crease). Belly and lower third of body generally plain greyish. Nape and top of head with dark grey vermiculation and dusky spots, vermiculation more pronounced on snout; side of head pale with dark spotting and blotches. All specimens with short dark grey line from lower anterior margin of eye to upper lip (midway between anterior nostril and rictus) and similarly coloured line from posterior lower margin of eye ending just behind rictus. In recently preserved material, often dark grey line from front of eye connecting both nostrils, an almost vertical dark grey line down rear preopercular margin, and less defined dark line along rear margin of opercle. Underside of head and branchiostegal membranes usually plain dusky grey to dark grey (aquarium-raised specimens from Galiwinku with five to seven irregular narrow dark lines crossing underside of head). First dorsal fin with proximal two-thirds dusky grey, becoming black posteriorly, broad whitish black-bordered submarginal stripe present, margin of fin dark grey to blackish. Second dorsal fin similarly coloured, but without black area posteriorly. Anal fin grey to dusky with translucent to whitish margin. Pectorals brownish to greyish. Comparisons The dense black peritoneum distinguishes this species from M. platystomus, which is sometimes similar in external appearance and may occur syntopically. Distribution Specimens are known from north-western Australia, from Shark Bay, WA, to Bing Bong Creek, Gulf of Carpentaria, NT (Figure 123). Ecology This species occurs intertidally on marine shores, typically in beach-rock pools (near fresh water seepages), in brackish pools behind beach dunes, or in sandy mangrove lagoons along beaches or headlands. Northern Territory beach-rock pools inhabited by this species were often influenced by fresh water seepage, or near a potential fresh water source (storm drain). Mugilogobius littoralis were usually present at high tide level, in very small pools in which they were often the only fish species present (Omobranchus rotundiceps and Istigobius ornatus also occurred, but less commonly). Remarks The 30-39 mm SL aquarium-raised specimens from Galiwinku (NTM 5.14296-001) are about 10 mm longer than any of the wild-caught fish, and they are much more stoutly built (almost cylindrical anteriorly). All are females, most filled with ripening eggs and a layer of abdominal fat. Etymology From the Latin litoralis, in reference to the seashore habitat in which this species is found. Revision of Mugilogobius Mugilogobius mertoni (Weber, 1911) Figures 123, 137-146; Tables 5-8, 22 Gobius mertoni Weber, 1911: 37, figures 5-6 (Panua Bori River near Sungei Manumbai, Aru Island, Indonesia) (in part). Gobius durbanensis Barnard, 1927a: 70-71 (Durban Bay, South Africa); Barnard 1927b: 815-816; Smith 1965: 333, figure 915. Tamanka mindora Herre, 1945c: 75 (Hacienda Waterous, Mangarin, Mindoro, Philippines); Herre 1953b: 766. Vaimosa layia Herre, 1953a: 13 (Layia, Batangas Province, Luzon, Philippines); Herre 1953b: 769. Tamanka mertoni (in part): Koumans 1953: 160. Stigmatogobius inhaeae Smith, 1959: 198, plate 9G (Inhaca, Mozambique); Smith 1961: 570; Smith 1965: 570; Smith and Smith 1969: 50. Stigmatogobius durbanensis: Smith 1960: 306; Smith 1961: 570. Mugilogobius inhacae: Hoese and Winterbottom 1979: 4; Hoese in Smith and Heemstra 1986: 795, figure 240.63. Mugilogobius durbanensis: Hoese and Winterbottom 1979: 4; Hoese in Smith and Heemstra 1986: 795, figure 240.62; Mauge 1986: 376; Smith and Heemstra 1986: 795. Mugilogobius valigouva: Hoda 1980: 476, figures lOII (misidentification). Mugilogobius sp.: Hayashi et al., 1981: 10, figure 128; AlIen and Boeseman 1982: 87; AlIen 1989: 165, plate 59. Mugilogobius in/wcae: Smith and Heemstra 1986: 795. Mugilogobius cavifrons: Akihito et al., 1988: 268, plate 2470. Material Examined Leetotype of Gobius mertoni SMF 6699, 1(15), Panua Bori River, Sungei Manumbai, Wokam, Aru Islands, Indonesia, H. Merton, 14 March 1909. Holotype of Gobius durbanensis SAM 17356, 1(35), Durban Bay, Natal, SS Pieter Faure. Holotype of Tamanka mindora CAS 39885, 1(24.5), Hacienda Waterous, Mangarin, Mindoro, Philippines, Herre 1940 Oriental Expedition, 20-22 July 1940. Holotype ofVaimosa layia USNM 202503, 1(31), Layia, Batangas Province, Luzon, Philippines, AW. Herre, 30 June 1948. 143 Paratypes ofVaimosa layia USNM 202573, 3(23-28), same data as holotype. Holotype of Stigmatogobius inhacae 1(27), RUSI 207, Inhaca Island, Seychelles. Paratypes of Stigmatogobius inhacae RUSI 7247, 4(22-30), Mahe, Seychelles, M.M. and J.L. Smith, September 1954. Other Material South Africa: ANSP 73284, 1(21), freshwaters of False Bay, North Zululand, Natal, H.W. BellMarIey, 1925; ANSP 96760, 1(31), False Bay, North Zululand, H.W. Bell MarIey, 1931. Mozambique: AMS 1.23637-001, 1(22.5), Benguerua Island, off Inhassoro. Madagascar: ZMH 7990, 1(49), freshwater, Oswald, March 1895. Seychelles: USNM 316046, 2(25.5-25.5), Rochon River, about a mile SE of Victoria, Mahe, Anton Bruun cruise 9, H.A. Fehlmann, 10 December 1964; USNM 316192, 1(30), same data as preceding; R.G. Mus. Africain Centrale 188771-3, 1(22), Les Canelies, Mahe south, P. Benoit and J.J. von Mol, 20 June 1972. Djibouti: USNM 99592,1(29), Djibouti, French SomaIiland, B. Brown, June 1920. Sri Lanka: USNM 316194, 3(2023), estuaries NW of western mouth of Mahaweli River, C. Koenig, 9 April 1970; USNM 316185, 6(1622.5), same data as preceding. India: ZMH 7994, 4(20-22.5), Bandra Rocks, near Bombay, Muller, April 1926; ZMH 7564, 81(10.5-22), Alibag, Arabian Sea, S of Bombay, German Indian Expedition, V. Maydell, 25 November 1955. Japan: YCM P.8719, 3(28.5-34.5), Sukuji River, Ishigakijima, 5 July 1979; URM P.4387, 7(25-36), Shiira River, Iriomotejima, T. Yoshino, 14 September 1982; URM P.1650, 9(27.538.5), same data as preceding. Philippines: AMS 1.23024-002, 7(23.5-29.5), Mactan Island, Cebu, E. Murdy and C. Ferraris, 6 July 1979; AMS 1.30309001, 1(22), Mindoro, Mangarin, AW. Herre, 20 July 1940. Indonesia: CMK 4545, 6(18.5-25), mangrove and small creek entering sea, Padang, Bungus Bay, Sumatera Barat, M. Kottelat and Bianco, 29 November 1984; CMK 8877, 1(25.5), Benoa mangroves, S of Sanur, BaIi, M. Kottelat and A Whitten, 23 March 1992. Australia: Northern Territory: NTM S.11276-002, 1(17.5), small mangrove at East Woody Island, near Gove, H. Larson and D. Percival, 22 February 1984; Queensland: AMS 1.22717-010, 7(19-24.5), Saunder's Beach, N of TownsviIIe, D. Hoese and R. Winterbottom, 4 October 1981; AMS 1.20978-012, 32(16-29), Mrs Watson's mangrove, Lizard Island, D.F. Hoese and party, 25 November 1978; ex AMS IA6560, 3(17-29.5), Plantation Beach, Lindeman Island, G. Whitley, 2 September 1935; AMS 1.22706003, 1(21), channel near mouth of Daintree River, D. Hoese and R. Winterbottom, 27 September 1981; WAM P.27780-D21, 1(22), mangroves near mouth of 144 H.K. Larson Daintree River, G. Allen and R. Steene, 2 September 1982; ex AMS 1.23281-017, 6(21-27), Mission River mouth, SW of Wallaby Island, Weipa, D. Hoese and D. Rennis, 11 October 1982. New Caledonia: USNM 316173, 6(18-26.5), Baie de Pecheurs, Noumea Harbour, Chapman, Cheyne and Smith, 28 January 1944. Santa Cruz Islands: ZMK P.781384, 7(9.5-22), Te Roto (brackish lake), Tikopia, T. Wolff, 20 April 1965. Other material examined (but not used in description) South Africa: RUSI 37917, 4, St Lucia, Natal; RUSI 17042, 2, Umtata River, Transkei; RUSI 32617, 8, Richard's Bay, Natal. Seychelles: USNM 270714,11, Victoria, Mahe. Pakistan: University of Karachi unregistered, 1, Kakkarpir, Karachi coast. Thailand: URM P.14771, 1, Phuket; URM P.12663, 6, Phuket; URM P.14863, 1, Phuket; CAS 51224, 23, Klong Ban Phe, Rayong; CAS 76142, 10, Trat Bay, Goh Mak Island; USNM 316177, 1, Lam Goh Peninsula, Satul; NTM 5.13953-015, 6, Klong Bang Sai, Phuket; ROM 68716, 1, creek between Cape Phanwa and Phuket City. Brunei: NTM unregistered, 28, Kuala Belait, Sungei Dalit; Japan: URM P.1678, 3, Shiira River, Iriomotejima; URM P.4839, 1, same data as preceding; URM P.1654, 7, same data as preceding; YCM P.8182, 7, Kabira Bay, Ishigakijima; YCM P.8296, 4, Saji River, Iriomotejima; YCM P.89OO, 4, same data as preceding; N. Oseko private collection 7036, 7012, 2, Honera River, Iriomotejima. Philippines: CAS 69680, 1, same data as holotype of Tamanka mindora; CAS 69655, 3, Hacienda Waterous, Mangarin; AMS 1.21938-013, 5, Mactan Island, Cebu; USNM 264948, 30, Cuyo Island, Palawan; USNM 260641, 29, Marigondon, Cebu; USNM 264945, 4, Oriental Negros; USNM 268197, 37, Siquijor Island; CAS 69815, 2, Coron, Busuanga; USNM 264947, 1, Negros Island; ROM 53342, 1, Negros Island, Oriental Negros; CAS 38629, 1, Dumaguete, Oriental Negros; AMS 1.23027-003, 1, Mactan Island, Cebu; CMK 9811, 5, Leyte, N of Baybay. Singapore: NTM 5.13957-010, 4, Sungei Pandan; NTM 5.13968-009, 3, Sungei Pandan; CAS 69814, 2, Serangoon. Indonesia: SMF 6738, 2, Kobroor, Aru Islands; USNM 297033, 2, Batanta Island, Irian Jaya. Papua New Guinea: NTM 5.13662-009, 1, Nagada River; USNM 316191, 2, Popondetta; Australia: Western Australia: WAM Figure 137 P.23193-001, 3, Onslow. Northern Territory: NTM 5.10472-022, 4, Channel Island, Darwin; Queensland: AMS 1.21263-004, 3, Yule Point; QM 1.21851, 3, Holloway's Beach; QM 1.31234, 66, Red Bream Creek; QM 1.31236, 3, Red Bream Creek; ROM 38806, 3, Saunder's Beach; AMS 1.22055-015, 2, Cape Tribulation; AMS 1.23262-003, 1, Cairns; AMS 1.22051-019, 1, Daintree River; AMS 1.23281020, 1, Mission River; WAM P.26962-009, 1, Daintree River; AMS 1.17158-002, 2, Bohle River, near Townsville; AMS 1.22041-016, 1, Mowbray River; AMS 1.20780-054, 4, Nymph Island; AMS 1.19102-012, 2, Lizard Island; AMS 1.22724-022, 3, Lizard Island; AMS 1.19468-042, 10, Lizard Island; AMS 1.19468-059, 1, Lizard Island; AMS 1.21273004, 13, Esk River; AMS 1.20785-041, 12, Hartley's Creek, Cairns; AMS 1.21259-005, 33, Cape Tribulation. New Caledonia: MNHN 1980-384, 1; No Data: RMNH 21069, 6. Diagnosis A moderate to large Mugilogobius with second dorsal and anal rays 1,6-8, modally 1,7; pectoral rays 14-17; longitudinal scales 26-37; TRB 8-13; 12-16 circumpeduncular scales; predorsal scales 9-19, mostly small, central anteriormost scale immediately behind eyes always largest, anteriormost one to four scales larger than those posterior to them; scales on body mostly ctenoid; third spine of first dorsal fin most often longest; body brownish with 7-11 darker narrow diagonal bars, chevrons or X-shaped markings along sides, interspaces between chevrons or bars pale, forming chequered or banded pattern, two to three dark spots or short diagonal bars on caudal base; widely distributed, from South Africa, the western Indian Ocean to Pakistan, Japan, the Indo-Malayan Archipelago, Papua New Guinea, northern Australia and the western Pacific. Description Based on 82 specimens, 15-49 mm SL. An asterisk indicates counts of lectotype of Gobius mertoni (Figure 137). First dorsal VI* (in 79), VII (in one); second dorsal 1,6-8 (mean 1,7*); anal 1,6-8 (mean 1,7*), pectoral rays 14-17 (mean 15, lectotype with 15 on right, 16 on left), segmented caudal rays 15-17 (mean 16*); Mugilogobius mertoni. Lectotype of Gobius mertoni Weber, SMF 6699, 15 mm SL, Wokam, Aru Islands. 145 Revision of Mugilogobius caudal ray pattern modally 9/7; branched caudal rays 7/6 to 9/7 (mean 8/7*); unsegmented (procurrent) caudal rays 5/6 to 7/7*; longitudinal scale count 26-37 (mean 31, 29 in lectotype); TRB 814 (mean 11, nine in lectotype); predorsal scale count 9-19 (mean 14*); circumpeduncular scales 1216 (mean 12*). Gill rakers on outer face of first arch 2+7 to 4+8 (modally 4+7). Pterygiophore formula 312210 (in 37), 3-122110 (in one). Vertebrae 10+15 (in two), 10+16 (in 44),10+17 (in two). Neural spines of first three, or first and second vertebra expanded or bifid at tip (in 19), or pointed and slender (in 15). Two epurals (in 43). One (in one), two (in 28) or three (in three) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid wide, but not expanded upward greatly; forms broad bridge to quadrate. Body relatively compressed (especially posteriorly). Head broad, rounded, wider than deep, HL 3.1-3.6 (mean 3.3) in SL; cheeks sometimes slightly inflated. Depth at posterior preopercular margin 1.4-2.0 (mean 1.7) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of about 20-25° with body axis; jaws generally reaching to below mid-eye (to posterior half of eye in large specimens) in males and to below anterior half of eye in females (to below mid-eye in large females). Lips usually smooth, small fleshy fimbriae may be present on inner edges of upper lip; lower lip free at sides, fused narrowly across front. Upper jaw 2.3-3.1 (mean 2.7 in females, 2.6 in males) in HL. Eyes rather small, lateral, high on head, top usually forming part of dorsal profile, 2.8-4.3 (mean 3.6) in HL. Snout rounded, 2.8-4.5 (mean 3.8) in HL. Interorbital broad, flat, 2.1-4.5 (mean 3.4) in HL. Top of head from rear of interorbital space forward to snout tip covered with fine villi (their tips Table 22 sometimes branched), usually visible in specimens with well-preserved mucous coat; scattered villi on preorbital and below eye in some specimens. Body depth at anal origin 4.4-5.9 (mean 5.0) in SL. Caudal peduncle compressed, length 3.4-4.2 (mean 3.8) in SL. Caudal peduncle depth 6.4-8.4 (mean 7.1) in SL. First dorsal fin low, rounded, tips of first three spines often free, third spine most often longest; first spine sometimes longest and filamentous; spines always falling short of second dorsal fin origin when depressed, usually by gap of one scale width (unless first spine elongate). First dorsal spine 3.0-8.2 (mean 6.3) in SL. Second dorsal spine length 6.4-8.8 (mean 7.2) in SL. Third dorsal spine length 5.9-8.8 (mean 7.4) in SL. Fourth dorsal spine length 5.9-8.3 (mean 7.5) in SL. Second dorsal and anal fins low, angled anteriorly, pointed posteriorly, posteriormost rays longest, rays falling well short of caudal fin when depressed. Pectoral fin rounded, central rays longest, 3.4-5.4 (mean 4.5) in SL; rays all branched but for uppermost. Pelvic fins short, rounded to oval, reaching half to two-thirds distance to anus, 3.8-5.8 (mean 5.1) in SL. Caudal fin rounded, 3.2-4.2 (mean 3.8) in SL. No mental fraenum, chin smooth. Anterior nostril in short tube, placed just at edge of preorbital, tube oriented down and forward, preorbital usually curved forward to accommodate nostril. Posterior nostril oval, often close to anterior centre margin of eye or closer to mid-point between eye and preorbital edge. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in four), with low irregular flat or fleshy flange (in 17), or distinct fleshy knobs and/or flaps on edge (in 43). Gill rakers on outer face of first arch short, longest rakers near angle of arch; rakers on inner face of first arch also stubby but with fine spiny papillae at tips; outer rakers on remaining three arches similar Morphometries as percentages of SL or HL, as indicated, of Mugilogobius mertoni (Weber, 1911). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. I. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital I. in HL Pectoral I. in SL Pelvic I. in SL Caudal!. in SL Longest D1 spine in SL Lectotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 32.0 54.2 64.6 18.7 13.3 29.3 13.3 22.9 31.3 33.3 25.0 27.4 50.0 64.1 16.8 10.0 23.9 12.2 22.4 23.5 33.3 22.1 18.4 17.4 24.1 11.2 32.7 69.5 87.0 22.1 14.6 29.6 15.6 358 36.2 47.8 47.0 26.1 24.0 31.5 33.7 30.0 58.7 73.2 19.7 12.7 26.4 14.0 270 27.4 38.9 30.5 22.3 19.9 27.0 18.5 27.6 52.6 65.8 18.0 11.4 23.6 11.9 22.4 24.5 32.7 22.7 18.5 17.6 23.9 11.4 32.0 65.0 83.9 22.7 19.6 29.3 15.7 31.3 32.1 40.0 32.7 29.1 26.3 30.9 20.5 30.3 59.7 73.6 20.7 18.8 26.2 14.1 26.5 28.5 36.5 28.5 22.5 20.1 26.4 15.9 20.7 12.0 -------- 146 H.K. Larsan A ...... :.' '. . .......... C":' :.: . cp' . ' .. -. "" .. 0.. .0 ＺｾＢ I \ ) , )" .... } ' \ , ' - ' B ..;:. ｾ .-" ｾ t / ＢＧｾＺＬＮＧＺｾＮＧＢｾ｜ "", ＧｾＭｾＮｾ "///fｾ -,.. /.'" ｾＮ Figure 138 Mugilogobius mertoni, papillae pattern. A, holotype of Vaimosa layia, USNM 202503, Luzon, Philippines; B, banded form, Weipa, Queensland, AMS 1.23281-017. Scalation suggested only. Scale bar = 1 mm. to first arch inner rakers; inner rakers on other arches twice length of first arch inner rakers. Tongue tip blunt or concave. In males, outer teeth across front of upper jaw largest, stout and curved, three to four rows of small sharp teeth behind this; one or two rows at side of jaw. Lower jaw with five or six rows of small sharp curved teeth across front, outermost row oriented nearly upright, teeth in inner rows all pointing inward; innermost row teeth largest and stoutest, no individual teeth particularly enlarged; usually only one row of teeth at side of jaw. Teeth in females similar, but teeth tending to Figure 139 be slightly smaller, especially innermost row of lower jaw. Most predorsal scales small, evenly sized, reaching to close behind eyes, with anteriormost scale enlarged, or anteriormost two to four scales larger than remainder of nape scales; scale on nape midline immediately behind eyes always larger than other three scales around it, scale never as greatly enlarged as usual in this species-group (the chulae-group). Operculum with small cycloid scales on upper two-thirds to half. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1,4 to 1;2 of belly), rest of scales cycloid. Ctenoid scales on side of body extending forward in broad wedge to behind pectoral fin. Genital papilla in male elongate, flattened, narrowing to pointed tip; in female, short to moderate, resembling blunt-tipped cone. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 138. Three 5 rows present, middle 5 row sometimes consisting of only one or two papillae. Two to four mandibular f row papillae. One specimen from Tikopia (ZMK P.781384) with extra papillae arranged in row between rows cp and d. Coloration of fresh material Two colour forms present in this species: typical "chequered" form and less common "banded" form. From colour photographs of male and female from Iriomotejima. Head and anterior quarter of body grey to violet grey with darker grey lines, blotches and mottling; rest of body dull greyish yellow; side of body with two alternating rows of dull violet to dark grey cross-hatched bars and blotches, lower row along midside of body; second dorsal fin with two rows of elongate dark grey spots; caudal fin grey with curved, somewhat comma-shaped dark grey blotch across bases of upper rays, base of fin dull yellow. Colour slide of "banded form" specimen from near Weipa (AMS 1.23281-020) with head and body yellowish white with purplish brown markings and first dorsal fin with dense black spot (Figure 139). Side of specimen from Lizard Island (AMS 1.20978012) showing light brown body overlaid with dense Mugilogobius mertoni, banded colour form, ex AMS 1.23281-017, Mission River, Weipa, Queensland. Revision of Mugilogobius mottling and banding in dark brown, forming "chequered" pattern; fin pigmentation translucent white and grey. Colour photograph in AlIen (1989), locality not given, of "banded form" with nine wide purplish brown bands alternating with yellowish background, whitish abdominal wall showing through musculature; caudal fin plain pinkish but for square black blotch on upper base; anal fin and pelvics plain purplish; pale yellowish band above dense black first dorsal fin spot, pale yellowish submarginal band in second dorsal fin. From colour photograph of banded specimen from Iriomotejima. Head and body light violet grey with dull violet to dusky violet grey lines and vermiculation on head; eight violet grey bars on body alternating with dusky to dull yellow interspaces; posteriormost bar darkest and extending over upper procurrent caudal fin rays; abdomen light pinkish; dorsal fins dusky with grey margin and translucent submarginal streak; yellow patches along fin bases; anal fin grey with dull yellow basally; caudal fin dull yellow with very broad violet grey margin and two diffuse curved grey bars near base; pelvics mostly grey. Hoda (in litt.) described a specimen from Pakistan as having the head and body brown with black markings, the upper part of both dorsal fins yellow with black margins and a black band along centre of fins. This is the specimen shown in Hoda (1980: figures 10-11). Coloration of preserved material Head and body light brown to yellowish brown with dark brown oblique bars, chevrons and Xshaped markings along sides; combination of dark markings and underlying lighter background colour usually giving reticulate or "chequered" appearance (Figures 140-143); alternately, with light background showing through as two or three alternating rows of light blotches. Seven to nine oblique narrow dark bars, chevrons or X-shaped markings visible, anteriormost an oblique "shoulder" bar (sometimes more intense than others); posteriormost dark bar crossing caudal base and extending ventrally onto lower part of caudal fin. Upper and lower parts of posteriormost body bar sometimes intensified as dark brown to blackish spot or streak, extending onto procurrent rays of caudal fin; blackish streak often somewhat commashaped, with "dot" of comma uppermost. Banded colour form with eight vertical diffuse dark bands crossing body (bands replacing narrow bars and chevrons); four specimens from Sumatra with pattern intermediate between broad bands and wide X-shaped markings; and two lots from India (near Bombay) with nine very narrow dark brown bands somewhat pointed ventrally, giving distinctive appearance. 147 Side of head with two brown horizontal stripes: upper extending from middle of upper lip, running along lower margin of eye across preopercle and ending on opercle as large diffuse or rounded dark brown blotch; lower brown stripe from rictus just above lower preopercular edge, ending on lower part of opercle either abruptly or as dark brown blotch. Lower streak often with one or more curved lines or hooks extending dorsally, sometimes forming reticulate pattern on cheek (holotype with remnants of reticulate pattern on side of head); banded specimens from Queensland always with reticulate pattern present. Top of head and predorsal area light brown, usually with scale margins narrowly outlined in darker brown, and irregular brown mottling. Some specimens with light "chequer" blotches along dorsal midline alternating with narrow brown bars. Snout and interorbital area plain brown, indistinctly mottled with dark brown, or light brown with dark brown reticulate pattern. Lips, underside of head and branchiostegal membranes brown to dark brown; dark brown blotch on lower part of opercle usually extending ventrally over branchiostegal rays as curved brown streak; some specimens with indistinct curved dark brown lines crossing isthmus and running behind lips. Gill arches and roof of branchial cavity brown to blackish. Pectoral base light brown with horizontal dark brown elongate blotch or streak on centre, extending part-way onto pectoral fin. Underside of body plain yellowish to light brown; breast dark brown and area immediately behind pelvic fin origin usually dark brown. Peritoneum mostly dark brown with abruptly pale ventral area. First dorsal fin with lower half dusky grey to brownish grey, with large dense black spot on central rear half of fin; black spot sometimes only visible as widest and darkest rearmost portion of dark grey streak running along centre of fin. Outer third of first dorsal fin white to translucent whitish, often narrow brownish to dusky grey marginal band present; tips of fin spines usually white (elongate spine sometimes dusky). Second dorsal fin with lower two-thirds dusky brownish to grey, narrow grey to brown fin margin, broad white to translucent whitish stripe below marginal band. Dusky lower part of fin with series of short, vertically elongate brown streaks on fin membrane, and usually two or three dark brown irregular blotches evenly spaced along fin base. Anal fin plain dusky to brownish, often with narrow white to translucent margin. Caudal fin mostly plain, membrane between rays often quite dark, forming streaks; most distinct marking an oblique to nearly vertical dark brown to blackish streak just posterior to fin base, lower part of streak often bending posteriorly, as does posteriormost body bar crossing H.K. Larson 148 Figure 140 Mugilogobius mertoni, AMS 1.20978-012, Mrs Watson's mangrove creek, Lizard Island, Queensland. Figure 141 Mugilogobius mertoni, 22.5 mm SL, AMS 1.20978-012, Mrs Watson's mangrove creek, Lizard Island, Queensland. Figure 142 Mugilogobius mertoni, 31 mm SL, AMS 1.23024-002, Mactan Island, Philippines. Figure 143 Mugilogobius mertoni, Coffee Bay, South Africa. From colour slide by Doug Hoese. caudal fin base; upper part of caudal fin streak usually intensified as comma-like black spot. Pectoral fin translucent, fin rays usually narrowly outlined with brown. Pelvic fins quite dusky to dark brown but for narrow translucent margin on fraenum. Comparisons This species is very similar to Mugilogobius filifer sp. novo and can be confused with it. Mugilogobius mertoni has a lateral scale count of 26-35, with a mean of 30 (versus 29-40, and mean of 35, in M. filifer sp. nov.), 10-16 predorsal scales, with a mean of 13 (versus 13-21, mean 19, in M. filifer sp. nov.), and usually has the first to fourth dorsal fins spine longest (versus the first spine usually longest and often elongate). Most frequently the first spine is longest. in M. filifer sp. novo and the third spine longest in M. mertoni. The predorsal scales are mostly small with the anteriormost scale enlarged in M. mertoni; in M. filifer sp. novo all the predorsal scales are small, with the anteriormost scale larger but not enlarged in comparison with the remainder. Body colour is quite similar in these two species. On the side of the head, however, M. mertoni usually has two longitudinal streaks but may have a partial or complete reticulate cheek pattern, while in M. filifer sp. novo this is always a reticulate or radiate pattern. Distribution A very widely distributed species, with specimens Revision of Mugilogobius Figure 144 Mugilogobius mertoni. Holotype of Gobius durbanensis Bamard, 35.5 mm SL, SAM 17356, Durban Bay, South Africa. known from the Seychelles, India, Pakistan, throughout the Indo-Malayan Archipelago, Japan, Philippines, Indonesia, Papua New Guinea, northern Australia and the western Pacific (New Caledonia and the Santa Cruz Islands) (Figure 123). Ecology Most specimens are recorded as being from mangroves, estuaries, or "brackish water". The Indian (Bombay) material came from "rocks" and "rocky pools". The USNM Seychelles specimens came from coastal freshwater streams. Remarks Among the syntypes of Gobius mertoni Weber, 1911, only one specimen, SMF 6699, is M. mertoni (the remainder are six M. platystomus and one Pandaka; see also Remarks under M. platystomus). Koumans (1953: 160) referred to this specimen (SMF 6699), recognising it as not the same species as the others in the lot, and considered that it " ... probably belongs to Stigmatogobius". In the jar containing SMF 6692 (another syntype of G. mertoni) is a label designating it as lectotype, a designation which is unpublished, as far as I can determine. This specimen is shown in Weber (1911: figure 5), and is M. platystomus. Although the syntypes of Gobius mertoni include two species of Mugilogobius, the name mertoni is still available (International Code of Zoological Nomenclature, Article 17), and is thus the oldest name applicable to the present species. The illustration of M. mertoni in Weber (1911: figure 6, which is M. mertoni; while figure 5 is of M. platystomus) exaggerates the triangular form of the fin, but it does show the alternating dark and light Figure 145 149 "chequered" colour pattern on the body. The specimen in SMF 6699 (Figure 137) is here designated lectotype of Gobius mertoni Weber. The holotype of Gobius durbanensis (Figure 144) has the left side of the head cut open, has had a tag (missing) tied to the caudal peduncle at one time, and most of its colour is washed out. Barnard (1927a-b) described the species as new in two publications (1927a appeared in July, 1927b in October). Stigmatogobius inhacae was described from the ("35 mm") holotype from Inhaca (Figure 145), and five putative paratypes from Mahe (RUSI 7247). The holotype is relatively plain in colour, as shown in the photograph in Smith (1959: plate 9G), and does not show any of the markings typical of M. mertoni. The 22 mm female paratype still retains some typical colour pattern. The five paratypes, which include a 15 mm SL Eleotris, are not in very good condition. Two of the M. mertoni are very flabby and bleached, while two are darker, but "crispy", as though they have been partly dehydrated at some stage. Tamanka mindora Herre, 1945, is based on the holotype male (given as 23 mm long) (Figure 146). A " ... juvenile male taken with the type was not used in writing the description" (Herre, 1945c: 75), being specifically excluded from the description, is thus not a type. There were two specimens present in CAS 39885: the 24.5 mm SL holotype and the non-type 20.5 mm SL specimen (the smaller not very "juvenile"). The non-type specimen is now catalogued as CAS 69680. The four types of Vaimosa layia Herre, 1953a (holotype USNM 202503, 1(29); three paratypes, Mugilogobius mertoni. Holotype of Stigmatogobius inhacae Smith, 27 mm SL, RUSI 207, Inhaca Island, Seychelles. 150 H.K. Larson Figure 146 Mugilogobius mertoni. Holotype of Tamankn mindora Herre, 25 mm. SL, CAS 39885, Mangarin, Philippines. USNM 202573) were formerly University of Washington specimens, UW 18959. Hoda (1980) briefly described and illustrated a specimen of what he identified as M. valigouva from Pakistan. After correspondence, in 1993 the specimen was sent to me and I identified it as M. mertoni. The Mugilogobius sp. listed in AlIen and Boeseman February 1934; FMNH 47229, 7(11-24), same data as preceding; FMNH 47058, 3(23-30), same data as holotype; ZMH H.415 (previously ZMH 17234), 2(31-32) same data as holotype; ZMH H.416 (previously ZMH 17238), 6(15-22), Kwangsi University campus, Kwangsi Province, Wu Chow, A. Herre, 12 February 1934. (1982) was examined and found to belong to M. Other Material mertoni. China: AMNH 37029, 32(25-34) (42 in lot, 10 are two different species), Foochow, Fukien Province, KC. Pope, March-September 1926; USNM 85967, 1(30.5), Foochow, Fukien, A. de C. Sowerby, 1922; CAS/SU 32579, 1(33.5), Kwangtung Province, A. Herre, 25 June 1929. One lot of seven specimens of this species from Cebu (Philippines), includes five specimens with second dorsal fin counts of 1,8, but agreeing with M. mertoni in other characters. These fish all have an enlarged scale on the nape close behind the eyes, and can therefore be distinguished from the abei species-group which modally has dorsal ray counts of 1,8, and small, evenly sized scales on the nape, which usually do not extend close up behind the eyes. Mugilogobius myxodennus (Herre, 1935) Figures 123, 147-150; Tables 5-8, 23 Ctenogobius myxodermus Herre, 1935c: 395 (Lingnan University, Honam Island, Canton, China); Ladiges et al., 1958: 166; Chu and Wu 1965: 130. Gobius myxodermus: Nichols 1943: 264-265. Rhinogobius myxodermus: Anonymous, 1976: 209, Diagnosis A moderately slender Mugilogobius with second dorsal rays 1,7-9; anal rays 1,7-9; pectoral rays 1417; longitudinal scales 33-40; TRB 10-16; circumpeduncular scales 13-18; predorsal scales 821, small, none enlarged, reaching to above preopercular margin; ctenoid scales on body in narrow wedge anteriorly, or with isolated patch under pectoral fin; third and fourth spines of first dorsal fin longest, subequal, not prolonged; preserved specimens with about nine indistinct narrow bars or X-shaped markings, indistinct oblique shoulder bar present; known only from southern China. figure 185. Mugilogobius myxodermus: Zheng and Wu 1985: 327328; Zheng 1989: 343-344; Ding 1994: 519-521. Material Examined Holotype ofCtenogobius myxoderrnus CAS 29075, 34 mm SL male, Honam Island, Canton (= Guangzhou), China, from claypits, fishponds, puddles and ditches about Lingnan University, A. Herre, 22 February 1934. Paratypes ofCtenogobius myxodermus China: CAS 69679 (formerly CAS/SU 29075), 16(16-31.5), same data as holotype; CAS/SU 29076, 146(10.5-38.5), Kwangsi University campus, Kwangsi Province, Wu Chow, A. Herre, 12 Description Based on 26 specimens, 18-34 mm SL. An asterisk indicates counts of holotype of Ctenogobius myxodermus (Figure 147). First dorsal VI* (in 24), VII (in two); second dorsal 1,7-1,9 (mean 1,8*); anal 1,7-9 (mean 1,8*), pectoral rays 14-17 (mean 16*), segmented caudal rays 1416 (mean 16*); caudal ray pattern modally 9/7; branched caudal rays 13-16 (14*; modally 15); unsegmented (procurrent) caudal rays 7/7 (in two); longitudinal scale count 33*-40 (mean 37); TRB 1016 (12*; mean 14); predorsal scale count 8-21 (13*; mean 14); circumpeduncular scales 13*-18 (mean 15). Gill rakers on outer face of first arch 3+7 (in two). Pterygiophore formula 3-12210 (in eight), 312211 (in one). Vertebrae 10+16 (in 13), 10+17 (in Revision of Mugilogobius Figure 147 151 Mugilogobius myxodermus. Holotype of Ctenogobius myxodermus Herre, 34 mm SL, CAS 29075, Honan Island, Canton (= Guangzhou), China. one). Neural spine of second and third vertebra expanded at tip (in seven), or first spine only slightly thickened (in one). One (in three) or two (in nine) epurals. Two (in seven) or three (in five) anal pterygiophores before haemal spine of first caudal vertebra. Body compressed (less so anteriorly). Head wider than deep, but not greatly so, HL 5.6-10.0 (mean 8.2) in SL; cheeks sometimes quite inflated. Depth at posterior preopercular margin 3.1-6.7 (mean 5.1) in HL. Width at posterior preopercular margin 3.8-8.4 (mean 6.1) in HL. Mouth subterminal, slightly oblique, forming angle of about 25° with body axis; jaws generally reaching to below posterior half of eye in males and to below anterior half of eye in females (to below mid-eye in holotype). Lips usually smooth, fleshy fimbriae present mostly on inner edges of upper lip and front of lower lip (fimbriae sometimes very close to outer edge); lower lip free at sides, fused across front. Upper jaw 1.8-2.5 (mean 2.7 in females, 3.8 in males) in HL. Eyes lateral, high on head, top usually forming part of dorsal profile, 1.6-2.5 (mean 2.2) in HL. Snout rounded, 1.5-2.9 (mean 2.3) in HL. Interorbital broad, flat, 1.3-3.8 Table 23 (mean 2.7) in HL. Top of head from preopercular margin up to anterior interorbital usually covered with fine villi, relatively sparse in interorbital area; in most of type specimens, mucous coat and villi wellpreserved. Body depth at anal origin 3.5-7.9 (mean 5.6) in SL. Caudal peduncle compressed, length 4.28.2 (mean 6.7) in SL. Caudal peduncle depth 2.4-5.0 (mean 3.9) in SL. First dorsal fin low, tips of second to fourth spines free, third and fourth spines longest or subequal; spines slightly longer in males than females; spines barely reaching second dorsal fin origin when depressed. First dorsal spine always shorter than next three. Second dorsal spine length 2.5-7.2 (mean 4.1) in SL. Third dorsal spine length 2.3-5.0 (mean 4.0) in SL. Fourth dorsal spine length 3.2-5.0 (mean 4.1) in SL. Second dorsal and anal fins low, posteriormost rays longest, rays not reaching caudal fin when depressed. Pectoral fin rounded, central rays longest, 3.6-6.5 (mean 5.3) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, reaching half to two-thirds of distance to anus, 3.5-5.7 (mean 4.5) in SL. Caudal fin rounded, 4.8--8.0 (mean 6.5) in SL. Morphometries as percentages of SL or HL, as indicated, of Mugilogobius myxodermus (Herre, 1935). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic !. in SL Caudal!. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 28.8 68.4 85.7 23.2 16.2 22.1 14.7 28.6 25.5 44.9 38.9 18.8 12.6 22.9 14.7 28.7 55.0 68.6 19.1 12.0 20.6 13.2 26.0 23.0 35.7 25.7 15.6 12.6 22.1 8.2 31.8 68.4 85.7 23.2 16.2 25.0 14.8 30.2 27.8 48.4 38.8 21.0 19.0 27.3 21.2 29.8 62.4 76.3 20.2 13.8 23.4 14.1 27.7 25.9 42.8 33.6 18.9 16.2 24.3 15.0 26.0 55.4 67.9 17.8 12.4 23.1 12.7 25.4 25.3 32.1 23.2 16.8 13.7 21.4 10.7 31.1 69.8 78.7 21.5 17.4 26.2 14.7 30.4 28.6 38.5 36.5 20.0 19.4 27.2 13.5 28.9 61.5 72.8 19.8 14.1 24.2 13.8 27.8 26.9 35.4 32.0 18.8 16.1 23.7 12.3 H.K. Larson 152 Figure 148 Mugilogobius myxodermus, papillae pattern. Holotype of Ctenogobius myxodermus, CAS/ SU 29075. Scales omitted. Scale bar = 1 mm. No mental fraenum, chin smooth. Anterior nostril tubular, placed just behind upper lip, tube short, oriented forward and down, preorbital produced forward to accommodate nostril. Posterior nostril without tube, oval, placed very close to anterior centre margin of eye. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in six) or with low irregular fleshy ridge or raised bumpy flange (in 18); one specimen with two distinct fleshy knobs on one side. Gill rakers on outer face of first arch very short and smooth, longest rakers near angle of arch; rakers on inner face of first arch also stubby; inner rakers on other arches twice length of first arch inner rakers. Tongue tip usually blunt, concave in three specimens. Outer teeth in upper jaw largest, stout and curved, three to four rows of small sharp teeth behind this row; one or two rows at side of jaw (no difference between male and female); tips of teeth (especially outer row) often tinted translucent orange to brown. Lower jaw with five or six rows of small pointed teeth across front, outermost row oriented upright, inner rows all pointing posteriorly; innermost row teeth largest and stoutest but no individual teeth particularly enlarged; usually only one row of teeth at side of jaw; tips of teeth often tinted translucent orange or brown. Predorsal scales small, evenly sized, usually reaching forward to above preopercular margin or further; if further, usually scales near nape midline extending further forward than those at sides; predorsal scales in holotype not reaching preopercular edge. Operculum with patch of small cycloid scales on upper third to half; sometimes only few scales present. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1,4 to Yz of belly), rest of scales cycloid. Ctenoid scales on side of body in narrow wedge, often broken into patch of scales behind pectoral fin (in holotype and 13 other specimens), with remaining ctenoid scales beginning below second dorsal fin origin or further back (below end of second dorsal fin in most extreme case observed). Genital papilla in male moderate, flattened, with blunt tip; in female, rounded and bulbous. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 148. Four to eight papillae in s rows on snout, middle row with usually two papillae; row c broken under eye, with three or more papillae in rear portion; row a following curve of eye; four to six f row papillae on chin. Coloration of fresh material A slide of a recently collected (dead) specimen from Guangdong, China, shows the body colour as light yellowish grey with brown to grey broken oblique bars, small blotches and mottling. The lower half of the first dorsal fin is translucent, the upper half pale yellow; fin crossed with three grey stripes and a distinct black spot posteriorly. Other fins are yellowish, with dusky grey or brown markings. In his original description, Herre (1935c) reported the live colour as follows: The colour in life is butter yellow to yellowish gray, with dusky markings along the scale margins on the upper half of the body, which form blotches most conspicuous posteriorly; there is a conspicuous yellow band on the upper part of the first dorsal and a large basal black spot of striking appearance on the posterior part, the rest of the fin dusky or dusky yellow; the second dorsal has two (rarely but one) longitudinal yellow bands, the rest of the fin more or less blackish, or with a median band of black vertical bars; the pectorals are colorless to yellow; the other fins are all more or less yellow, speckled with cross bars of dusky dots, or the anal may be clear yellow. Zheng (1989) showed a line drawing of this species which probably exaggerated the spot in the first dorsal fin; unfortunately a translation of the Chinese text was not available. Another illustration is in Anonymous (1976), showing the vermiculation on the head. Coloration of preserved material Most specimens with colour pattern faded and indistinct. Head and body very pale brownish, lighter ventrally; upper half of body with groups of scales with brownish edges, roughly forming nine bars sometimes joining at midside, forming Revision of Mugilogobiu5 153 Figure 149 Mugilogobius myxodermus. Paratype of Ctenogobius myxodermus Herre, 30.5 mm SL, CAS 69679, Honan Island, Canton (= Guangzhou), China. Figure 150 Mugilogobius myxodermus, 34 mm SL, AMNH 37029, Foochow, Fukien Province, China. indistinct X or V shapes, or scattered small round brown spots especially along midside of caudal peduncle (Figures 149-150). Anteriormost brown bar oblique, running very close behind pectoral base up onto predorsal area, sometimes extending anteriorly to nearly above rear margin of preopercle, or becoming diffuse at predorsal midline. Top of head and nape dusky (predorsal scale margins usually outlined with brown), interorbital and top of snout with faint spotting and short vermiculations. Often, curved line extending from rear of eye onto centre of nape, close to but not meeting oblique shoulder bar. Side of head with indistinct reticulation or vermiculation; in many specimens dark-margined light round spots present in two or three rows over cheek and lower half of opercle. Six thin brown lines crossing underside of head, from rear of opercle, along branchiostegal rays and membranes and joining underneath head, lines not always visible. Pectoral base with large brown blotch at least on anterior half (or entire base), sometimes posterior half of base with white area. Breast and belly usually pale, unmarked; male specimens most clearly marked, usually with brown pigment scattered over underside of head and prepelvic area, and fins darker. Pectorals pale brownish; pelvics pale to dark brown (latter in males). First dorsal fin with dusky margin, free tips of spines dusky; whitish or translucent stripe below this, median brown stripe present, darkest posteriorly, this area often intensified, forming black spot; dusky area along base of fin, usually most developed on anterior quarter of fin. Second dorsal fin with wide brownish margin, equal-sized white to translucent stripe below this, median broad dusky stripe (often darkest along top of stripe) with elongate dark blotches placed in front of each fin ray; below median dark stripe another clear stripe, and ventralmost, an irregular brown stripe sometimes intensified into several blotches usually corresponding with fin rays. Median dusky stripe often dark brown, partly obscuring fin ray blotches. Anal fin plain brownish, darkest posteriorly. Caudal fin dusky, with faint irregular markings often forming many to few irregular rows of spots, or about five narrow wavy bars, often partly broken-up. Small but distinct brown round to rectangular spot always present at centre of caudal fin base, just at hypural crease, and short elongate brown mark over upper procurrent rays. Two brown spots, equal in size to caudal base spot or much smaller, on caudal fin, close to either side of median fin base spot, but never joining it. Caudal fin of holotype with two dark spots on either side of caudal fin base spot; distally, three narrow, irregular vertical bars, partly broken up into spots. Comparisons This is a rather nondescript species, which bears some resemblance to Chlamydogobius ranunculus in colour pattern and general physiognomy. The colour pattern is also reminiscent of that of M. cavifrons, albeit diffuse and broken-up. Distribution Specimens are known from a few localities (Wuchou, Kuang-chou, Fu-chou) in Kwangsi, Kwangtung and Fukien Provinces, China (Figure H.K. Larson 154 123). Chu and Wu (1965) list this species as a Chinese endemic, recorded from the "Basins of Pearl River and Min River (including Hainan and Taiwan)". Ecology In Herre's original description, he refers to the fish being present in apparently shallow clay pits, wayside ditches, small ponds and mud puddles on the Lingnan University campus. Remarks Ladiges et al. (1958) incorrectly designated the 31.5 mm SL specimen in ZMH HA15 as lectotype, and ZMH H.416 as a paralectotype, with no explanation as to why they did so. The holotype's existence has never been in question. ZMH HA15 is labelled with a metal tag SU 29075, since it was part of Herre's (Stanford University) type series. Herre's original description clearly indicates that there was a type, 32 mm long, and 20 paratypes, 20-31 mm long, taken from Lingnan University campus, as well as about 100 paratypes from Kwangsi University campus. The labelled holotype (CAS/SU 29075) is in CAS where it has always been kept. Of the seven paratype specimens of Ctenogobius myxodermus in FMNH 47229, one 23 mm SL specimen is a Rhinogobius sp. In CAS 69679 (SU 29075), one of the 16 paratypes is probably a Favonigobius. In CAS 29076, two of the 146 paratypes are non-gobioid, probably oryziatids. Mugilogobius notospilus (Giinther, 1877) Figures 10D, 123, 151-157; Tables 5-8, 24 Gobius notospilus Giinther, 1877: 173, plate 109, figure B (Namusi, Viti-Levu, Fiji); Koumans 1953: 125 (as possible synonym of Stigmatogobius hoevenii sensu Koumans, = Mugilogobius chulae). Vaimosa fontinaiis Jordan and Seale, 1906: 395, figure 85 (near Apia, Samoa); Jordan 1927: 10; Fowler 1928: 408; Koumans 1931: 87-88; Schultz 1943: 228,239. Mugilogobius notospila: Whitley 1927: 7. Vaimosa notospila: Fowler 1928: 408; Fowler 1934: 443. Mugilogobius fontinalis: Koumans 1940: 124, 161; Fowler 1949: 135-136; Wass 1984: 28. Stigmatogobius duospilus Fowler, 1953: 387, figure 2 (near Voh, New Caledonia); Bohlke 1984: 106. Vaimosa frontinalis Mauge, 1986: 376 (error for fontinalis). Mugilogobius duospilus: AlIen 1991: 188-189. Mugilogobius notospilus: Pusey and Kennard 1994: 27. Material Examined Holotype of Gobius notospilus BMNH 1869.11.12.31 [formerly Godeffroy Museum No. 5618], 1(51), Namusi [= Namosi], Viti Levu, Fiji, Godeffroy. Holotype ofVaimosa fontinalis USNM 51776, 1(38.5), Gasegase River at Vaimosa near Apia, American Samoa; D.S. Jordan and V.L. Kellogg, 1902. Paratypes ofVaimosa fontinalis USNM 171993, 7(19.5-43), Apia, V.L. Kellogg, 1902; USNM 126361, 32(15-33), same data as previous; MCZ 36013 [ex. USNM 51776], 1(31), Samoa; CAS/ SU 8700, 29 (12-45), Gasigasi River near Apia, D.S. Jordan; AMS 1.7379, 3(16.5-38.5), Samoa; BPBM 5321 [ex. USNM 51776],1(33), Samoa. Holotype of Stigmatogobius duospilus ANSP 71970, 1(29), small pool at edge of swamp near Voh, New Caledonia; Dr Marshall Laird, 12 June 1952. Other Material American Samoa: BMNH 1924.12.11.35 and 1925.3.30.6 [in same bottle], 2(38-39), Apia, Buxton and Hopkins; USNM 316183, 1(29.5), Vaipito Stream, near mouth, C. Couret, 6 April 1978. Solomon Islands: USNM 316171, 2(27-34), Bunia Point, NNW of Honiara, Guadalcanal, W.P. Davis and party, Te Vega Cr.6/Stn 250, 20 March 1965. Papua New Guinea: ZMK unregistered, 6(12-28.5), Brook at Liei River, Lorengau, Manus Island, Bismarck Archipelago, T. Wolff, Noona Dan Stn. 64, 24 June 1962; WAM P.27829-007, 1(25.5), small creek 1.5 km SE of Lorengau, Manus Island, rotenone, G.R. AlIen and R. Knight, 10 October 1982; ZMH 19346, 13(14-29.5), strand pools at Langemak Bay, E coast, Hamburg Sud-see Expedition, 12-16 January 1909. Australia: Queensland: WAM P.26985-002, 3(22-27), small creek on N side of Daintree River, 9 km E of ferry crossing, G.R. AlIen and W. Starck, 4 October 1980; AMS 1.22044-001, 111(9-28), backwater creek, eight miles upstream from mouth of Daintree River, 0-1 m, rotenone, G.R. AlIen, D.F. Hoese, H.K. Larson and W. Starck, 14 September 1980; AMS 1.22052009, 4(14.5-29.5), creek at Starck's jetty, four miles upstream of mouth of Daintree River, 0-1 m, rotenone, D.F. Hoese and H.K. Larson, 16 September 1980; ROM 38329, 15(11-24), Barrett Creek, tributary of Daintree River, 1 m, D.F. Hoese and R. Winterbottom, 24 September 1981; WAM P.26955-008, 30(12-23), on road crossing near Mossman, Mossman River, G.R. AlIen and D.F. Hoese, rotenone, 3 September 1980; AMS 1.21419009, 4(21.5-26.5), creek on S side Cairns Harbour, Revision of Mugilogobius D.F. Hoese and R. 5teene, 20 December 1978; N1M 5.14345-001, 1(32.5), Waterfall Creek, near Cardwell, site 090, sample 51, B. Pusey, 1993. Other material examined (but not used in description) Papua New Guinea: CA5 66726, 1, Ramu River, near Madang; ZMH 19345, River at Mejm Bay, south coast New Britain. Australia: Queensland: QM 1.14611, 1, drain on Anderson Road, Cairns; AM5 1.14597, 4, Cairns; QM 1.30621, 1, Missionary Bay, Hinchinbrook Island; QM 1.30625, 1, Deluge Inlet, Hinchinbrook Island; WAM P.269S7-006, 1, tributary on N side Daintree River; AM5. 1.22045003, 101, Daintree River; AM5 1.22703-003, 3, Daintree River; AM5 1.17985-001, 2, Daintree River; AM5 1.22049-008, 3, Daintree River; N1M 5.14234001, 14, North Johnstone River; N1M 5.14233-001, 2, North Johnstone River; NTM 5.14232-001, 13, Mulgrave River. Tonga: U5NM 342067, 1, Vava'u Island. Diagnosis A moderate-sized Mugilogobius with dorsal rays 1,7-8; anal rays 1,7-9; pectoral rays 14-16; 28-36 longitudinal scales; TRB 9-12; circumpeduncular scales 12; 12-20 small predorsal scales reaching up to eyes or at least to above preopercular border; always 12 circumpeduncular scales; second or third dorsal fin spines longest, with none elongate or filamentous; side of body with about seven bars or vertical X-shaped blotches alternating with pale areas, giving chequered appearance; sides of head with pale round spots and dark reticulated lines; two dark rounded spots at base of caudal fin; mostly inhabiting freshwater; known from Fiji, New Caledonia, 5amoa, 5010mon Islands, NE coast of New Guinea and the NE coast of Queensland. Description Based on 59 specimens, 15.5-51.0 mm 51. Counts of holotype of Gobius notospilus (Figure 151) indicated by asterisk. First dorsal VI"; second dorsal 1,7-8 (mean 1,8"); anal 1,7"-9 (mean 1,8); pectoral rays 14-16 (mean 15"); segmented caudal rays 16-17" (mean 16); caudal ray pattern modally 9/7 to 9/8" (modally 9/ 7); branched caudal rays modally 8/7"; Figure 151 155 unsegmented (procurrent) caudal rays 6/7 to 8/8; longitudinal scale count 28-36 (29"; mean 31); TRB 9-12 (mean 10"); predorsal scale count 12"-20 (mean 17); circumpeduncular scale count always 12". Gill rakers on outer face of first arch 3+7 to 4+12 (modally 3+8). Pterygiophore formula 312210" (in eight). Vertebrae 10+16" (in nine), 10+17 (in one). Neural spine of second and sometimes third vertebra somewhat expanded or bifurcate at tip (in five) or slender and rather pointed (in three). Two epurals (in nine). Two (in nine) anal pterygiophores before haemal spine of first caudal vertebra. Body relatively compressed, especially posteriorly; some specimens slender (e.g. holotype of Stigmatogobius duospilus). Head rounded, 2.6-3.9 (mean 3.5) in 5L, somewhat depressed anteriorly, always wider than deep. Depth at posterior preopercular margin 1.5-2.0 (mean 1.7) in 5L. Width at posterior preopercular margin 1.2-1.7 (mean 1.4) 5L. Mouth usually subterminal, sometimes barely terminal, slightly oblique, forming angle of about 25° with body axis; jaws reaching at least to below anterior half of eye and as far as posterior half of eye in adult males, and to anterior half of eye or to mid-eye in females (i.e. not much difference between males and females). Upper jaw length 2.0-3.9 (mean 2.8) in H1. Lips smooth, upper lip often fimbriate on inner edge. Lower lip free at sides, fused to symphysis anteriorly. Eyes lateral, set high on head, forming part of dorsal profile, 3.1-4.2 (mean 3.7) in H1. 5nout rounded in dorsal view, plump, profile often rounded, 3.3-4.5 (mean 3.9) in H1. Interorbital broad, 2.7-4.0 (mean 3.2) in HL, sometimes slightly concave. Interorbital and adjoining unscaled area of predorsal often covered with closely-spaced villi, often extending onto snout past level of posterior nostril. Body depth at anal origin 4.0-6.1 (mean 5.3) in 51. Caudal peduncle length 3.-4.5 (mean 4.0) in 51. Caudal peduncle depth 5.2-8.2 (mean 7.2) in 51. First dorsal fin with second to third dorsal spines longest (never elongate), no spine extending much beyond depressed fin; depressed dorsal often not reaching first element of second dorsal fin, if so, longest spine reaching only to base of second dorsal fin spine; longest spine 6.1-9.8 (mean 7.7) in 51. Mugilogobius notospilus. Holotype of Gobius notospilus Giinther, 51 mm SL, BMNH 1869.11.12.31, Namusi [= Namosi], Viti Levu, Fiji. H.K. Larson 156 Second dorsal fin with posteriormost rays longest, pointed and sometimes up to twice length of anteriormost two fin elements; rays reaching to lower part of caudal fin in large specimens or males in breeding condition. Anal fin pointed posteriorly, anterior few rays nearly half length of posteriormost ray; falling short of caudal fin base except in males in breeding condition, fin then often reaching base. Pectoral fin rounded to rhomboid, rays usually all branched in adults but for uppermost; reaching to above anus, 3.7-5.1 (mean 4.5) in SL. Pelvic fins oval to slender, reaching at least halfway to anus, often nearly to anus when flattened, 4.0-6.2 (mean 4.9) in SL. Caudal fin relatively short, rounded to oval in shape, 3.0-4.8 (mean 3.5) in SL. No mental fraenum. Anterior nostril tubular, placed just behind upper lip, tube often short, and oriented forward and down over upper lip; preorbital sometimes curved outward slightly to accommodate nostril tube. Posterior nostril rounded to oval, placed close to centre of anterior edge of eye. Gill opening extending forward to below opercle. Inner edge of pectoral girdle smooth edged (in eight), with low irregular fleshy ridge or fleshy flange (in 20) or with fleshy knobs, lobes or bumps (in 30); knobs arising from raised ridge or directly from smooth edge of pectoral girdle. Gill rakers on outer face of first arch short, stubby, smooth, as with rakers on inner face of first arch. Rakers on second and succeeding arches with longer rakers (rakers about twice as long as those on first arch). Tongue tip usually blunt or concave. Teeth in outer row in upper jaw largest, stout, curved or nearly conical, followed by three to four inner rows of tiny even sharp teeth; two rows at rear of jaw (Figure 152). Outer row and inner four or five rows across front of lower jaw similar; teeth all small, sharp, evenly-sized, about two rows present along side. Teeth not particularly enlarged in males. Predorsal scales small, cycloid, usually reaching forward to close behind eyes or at least halfway between preopercular margin and rear of eyes. Figure 152 Jaws and suspensorium of Mllgilogobius notospilus, male, ex ZMH 19346, Langemak Bay, Papua New Guinea. Scale bar = 1 mm. . Qit--@ . "0 ...-:. . ..-: ::: c Figure 153 :1 . :: MllgilogobillS notospilus, papillae pattern. Paratype of Vaimosa fontinalis, USNM 171993. Scales omitted. Scale bar = 1 mm. Operculum usually with cycloid scales on upper half. Cheek always naked. Pectoral base fully scaled with small cycloid scales. Prepelvic area covered with small cycloid scales. Belly scales ctenoid (often weakly) under pelvics, with anterior 1/4 to 1/3 of belly scales ctenoid and remainder of scales cycloid. Side of body with broad to narrow wedge of ctenoid scales extending forward to behind pectoral fin. Genital papilla in male elongate, flattened, with rounded tip; in female, papilla short, rounded and bulbous or conical with blunt tip. Head pores absent, as in all Mugilogobius species. Papillae in longitudinal pattern (Figure 153). Three 5 rows on snout, central row with one or two papillae; two to three mental f row papillae in line behind chin; in many specimens, row a forming posteriorly-directed bend (dorsalmost few papillae); cheek rows e, ep and d sometimes with extra papillae above or below rows, extra papillae occasionally forming additional short rows. Coloration of fresh material From colour photograph of freshly dead M. notospilus in AlIen (1991: 188). Head and body pale yellowish above and whitish below; scale margins narrowly outlined with brown, giving cross-hatch effect to seven bars on side of body and indistinct dark blotch at caudal base. Base of caudal fin whitish, with two round black spots at centre of base and black streak along upper procurrent rays; rest of fin translucent whitish with dusky margin (more pronounced ventrally). Stripes on dorsal fins black, lowermost areas pale yellow. Anal fin reddish orange on proximal half, dusky submarginal stripe, and bluish white margin. Pectorals and pelvics whitish. White pectoral base divided in two by blackish bar, orange pigment in front of and behind white area. Iris golden (indistinct in photograph). Revision of Mugilogobiu5 Coloration of preserved material Head and body pale yellowish or yellowish white (depending on state of preservation), with brown to dark brown or blackish markings (Figures 154-156). Head with four or five brown lines radiating from eye and forming reticulate pattern across preopercle and opercle, always less distinct on opercle; often whole side of head brown with round white spots. Posteriormost eye stripe not reticulate; stripe extending from upper rear margin of eye up onto nape, and curving toward midline, or stopping abruptly. Anteriormost stripe eye stripe usually running just below posterior nostril to above anterior nostril tube; next eye stripe reaching to mid-jaw; these two stripes usually joined by brown line running around tip of snout to rictus of jaw. Underside of head with three brown to blackish stripes: one beginning just below lower lip and extending to rictus, one behind chin at beginning of preopercular edge and following this edge, and one across anterior part of branchiostegal membranes and extending up onto opercle; stripes diffuse and barely discernible, or very broad and dense. Unscaled dorsal surface of head with brownish or dusky reticulate or vermiculate pattern, especially 157 on snout, some lines meeting across interorbital space. Side of body with scale margins outlined in brown, forming about seven short bars, often broken up into vertical pairs of slightly offset blotches, with very pale whitish yellow spaces between bars along midside, giving fish distinctive chequered appearance; scale margins within some bars heavily pigmented, giving bars an X-shape. Bars variable in shape and sometimes in width, being short and straight, or diagonal (formed by short midlateral bar joining with slightly offset blotches on upper part of body); tops and bottoms of bars partly or wholly joined by row of brownish pigmented scales, spaces between bars always remaining pale. Just above pectoral base, dark brown blotch always present, this blotch rounded or forming diagonal bar extending obliquely from above pectoral base to up near nape midline. Abdomen and lower sides pale yellowish white. Pectoral base pale yellowish, with brown bar extending from pectoral girdle onto base of pectoral fin, bar closer to upper edge of pectoral base than lower; fin base usually darker on anterior half, paler on posterior half. At centre of caudal fin base, two Figure 154 Mugilogobius notospilus, 29 mm SL, AMS 1.22044-001, Daintree River, Queensland. Figure 155 Mugilogobius notospilus, 29.5 mm SL, ZMH 19346, Langemak Bay, Papua New Guinea. Figure 156 Mugilogobius notospilus. Holotype of Vaimosa fontinalis Jordan and Seale, 38.5 mm SL, USNM 51776, Apia, American Samoa. 158 Table 24 H.K. Larson Morphometries as percentages of SL or HL, as indicated, of Mugilogobius notospilus (Giinther, 1877). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. I. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic I. in SL Caudal I. in SL Longest 01 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 29.0 64.2 70.9 21.6 13.7 25.1 15.7 27.7 26.4 25.7 26.4 20.8 16.1 24.5 12.0 28.1 51.5 63.1 16.5 9.4 23.0 12.5 23.0 23.6 25.7 26.0 20.0 16.1 24.5 11.8 38.1 64.2 84.5 24.8 17.6 31.9 19.5 29.8 32.7 49.2 39.1 26.7 25.2 33.8 16.5 30.3 58.4 74.1 19.6 13.2 25.2 14.7 26.7 26.5 38.0 32.4 22.9 20.9 29.6 14.8 25.9 50.7 59.8 16.8 9.3 22.4 12.1 22.0 25.3 30.0 25.3 19.7 16.5 20.7 10.3 32.3 65.6 79.0 21.8 14.7 28.3 15.3 29.9 31.0 39.8 37.0 27.1 23.3 33.5 22.6 28.4 57.7 72.0 19.0 12.6 25.0 13.6 25.7 28.0 34.9 30.9 22.3 20.6 28.0 15.3 distinct dark brown to blackish, vertically aligned, round or oval spots; third, sometimes slightly less intense, brown blotch or streak on dorsal procurrent rays of fin; uppermost of paired spots usually largest, sometimes forming oblique oval blotch. First dorsal fin translucent to dusky, with narrow brownish marginal stripe and broad brown central stripe usually intensified posteriorly as black spot on membrane between fourth and sixth spines; at base of fin, dusky stripe present at least anteriorly. Second dorsal fin with narrow brown margin, a narrow transparent submarginal band below this, and broad brown stripe on central half of fin, with narrow translucent area below this; base of fin dusky with three dark brown vertically elongate blotches present along base; one at base of first soft ray, one at centre of base, and one at base of second last ray. Some specimens, probably males in breeding colour, with entire fin dusky, but dark margin always separated from rest of fin by narrow transparent stripe. Anal fin brownish to dark brown, usually with pale or translucent margin. Caudal fin dusky, some heavily pigmented specimens with dark lines in centre following rays, and upper and lower margins of fin more intensely pigmented, especially upper margin near dark blotch on procurrent rays; lowermost caudal base spot sometimes less intense and smaller than upper spot. Pectoral fin dusky, but pale area near base present (confluent with pale areas on base itself). Pelvic fins and fraenum dusky, especially near centre. Males, in probable breeding condition, often very dark, heavily pigmented, and head, breast and second dorsal and anal fins almost evenly brown with head stripes and spots often indistinct. In some specimens, such as the holotype of G. notospilus and some specimens from north Queensland, the caudal base spots are indistinct or indiscernible. The holotype is unusual not only because of its great size, but because the body bars and pale interspaces are not exhibited (pattern similar to a specimen from Cairns, Queensland). The holotype is very heavily pigmented and all scale margins are outlined in brown, the reticulation and pale spots on the head are partly visible, the first dorsal fin is plain brownish with darker brown area on membranes between the fourth and sixth spines, and the only markings on the caudal fin are scattered brown spots on dusky membranes near the fin base, a brown bar along the hypural crease and a short brown bar along the bases of the upper caudal rays. Comparisons Living specimens of this species could be confused with M. mertoni, which has a similar "chequered" colour pattern. However, M. mertoni has the anteriormost few nape scales enlarged (unlike the small, evenly sized scales in M. notospilus) and modally 1,7 dorsal and anal fin rays (1,8 in M. notospilus). Some specimens of this species look similar to the "barred" form of M. abei (see Comparisons for that species). Distribution East coast of Queensland, northern Papua New Guinea and islands of the western Pacific (Fiji, Tonga, New Caledonia, Samoa, Solomon Islands) (Figure 123). Ecology Restricted to freshwater streams and brackish areas. Most specimens were recorded as being from Revision of Mugilogobius Figure 157 159 Mugilogobius notospilus. Holotype of Stigmatogobius duospilus Fowler, 30 mm SL, ANSP 71970, New Caledonia. brackish, estuarine, "swamp" or freshwater habitats, in shallow water (usually less than 1 m). The nominal type specimen came from freshwater near Namosi, Viti Levu. The most detailed habitat information is available for material from shallow (0.3-0.5 m) freshwater habitats on the Johnstone and Mulgrave Rivers, Queensland. Here the species inhabits: short tributary streams close to the respective river mouths and fed by drainage from well vegetated rainforest catchments. These streams ... have a riparian vegetation of lowland rainforest with a smattering of mangrove species and palms (Licuala) and closed canopy. They have a dominant substrate of fine gravel and sand with some mud in places. All specimens so far collected have been from areas of dense leaf litter. This seems to be a fairly specific habitat requirement. (8. Pusey, in litt.). Remarks Gobius notospilus Giinther, 1877, was based on a specimen formerly misidentified by Godeffroy as Gobius petrophilus (original Godeffroy catalogue number 5618) (Figure 151). The specimen is unusually large, 51 mm SL, with the next largest known specimen being 43 mm SL. It is also the only specimen available from Fiji, so it is unclear whether its relatively plain colour is due to its preservation history or whether all the Fijian population is coloured similarly. Jordan and Seale (1906) described Vaimosa fontinalis (Figure 156) from 75 specimens from four localities: 23 from Vaisigano River, eight from Vailima Brook and tributaries near Apia, two from Gasegase River at Vaimosa near Apia and 42 from a stream at Pago Pago. The type was designated as USNM 51776, from the Gasegase River near Apia. The collection localities seem to have become confused when the paratype specimens were distributed from USNM to other museums (e.g. the 29 CAS specimens are labelled as being from the Gasegase River, although only two specimens were stated in the description to have been taken from that locality). Most museum labels for these type specimens give "Apia" or "Gasegase River" as the locality. USNM 12631 (formerly U. S. Bureau of Fisheries 1269) contains 32 fish, of which one is a salarine blenny and another is the gobiine Asterropteryx semipunctatus. CAS 8700 includes 29 specimens, of which one is an Eviota (Gobiinae). The holotype of Stigmatogobius duospilus Fowler, 1953 (Figure 157), is rather slender-bodied, but otherwise typical. Fowler (1928) synonymised fontinalis with notospilus with no reason given for the synonymy. Koumans (1940) refers to a 48 mm specimen in the AMS Vaimosa fontinalis paratype series (AMS 1.7379), which he identified as a Stigmatogobius hoevenii (Koumans often identified specimens of Mugilogobius chulae as s. hoevenii). There are three specimens, all M. notospilus, presently in AMS 1.7379. In 1953, Koumans tentatively listed G. notospilus as a synonym of Stigmatogobius hoevenii (= M. chulae). Fowler (1961) appeared to agree, as he illustrated his diagnosis of s. hoevenii with an illustration, redrawn from Giinther's (1877) original figure of G. notospilus. From Fowler's synonymy (he did not include G. notospilus in the synonymy) it appears that he is referring here to Mugilogobius chulae, but he gives no reason why he considers Giinther's species to belong to M. chulae. Mugilogobius chulae is only known from the western Pacific by specimens from Palau. Mugilogobius platynotus (Giinther, 1861) Figures 180, 123, 158-162; Tables 5-8, 25 Gobius platynotus Giinther, 1861: 66 (type locality unknown). Waiteopsis paludis Whitley, 1930: 122-123 (Gundamaian, Port Hacking, Sydney, Australia); Koumans 1931: 162; Koumans 1940: 169; Ivey 1951: 55; Whitley 1951: 405-106. Ellogobius abascantus Whitley, 1937: 17, figure 4 (Bateman's Bay, New South Wales); Whitley 1937: 18; Koumans 1940: 171; Whitley 1951: 406 (as synonym of Waiteopsis paludis). Mugilogobius paludis: Bell et aI., 1984: 37; Morton et al., 1987: 222-223, 227; Morton et al., 1988: 189; Gee and Gee 1991: 19-27; Kuiter 1993: 358; Hoese and Larson 1994: 793. H.K. Larson 160 Material Examined Holotype of Gobius platynotus BMNH 1859.5.7.71, 1(39), no locality data, Sir A. Smith. 106(19-30), Towra Point, Botany Bay, NSW Fisheries, 9 February 1979; AMS 1.30997-001, 2(27.538), beach washup, Harrington Lagoon, Harrington, B. Williams, 20 January 1989. Victoria: NTM S.14346-001, 10(26.5-35), Blind Bight, Westemport Bay, P. Unmack, S. Ballestrin, 25 January 1993. Holotype of Waiteopsis paludis AMS IA.3917, 1(37.5), Gundamaian, Port Hacking, New South Wales; G.P. Whitley. Other Material Australia: Queensland: ex QM 1.20309, I, Lower Logan River. New South Wales: AMS 1.15854-006, I, Nullica River; AMS 1.22491-001, 2, Ballina; AMS IB.2304, I, Middle Harbour, Sydney Harbour; AMS 1.25038-001, 4, Brunswick Heads; AMS IB.999, 2, Tweed Heads; AMS 1.22960-001, I, Apple Tree Bay, Sydney; AMS 1.16957-002, I, Bull's Head Bay, Sydney Harbour; AMS 1.16956-001, I, Lane Cove River; AMS 1.30473-001, I, Bobbin Head; AMS 1.30080-001, I, Cugden Creek; AMS 1.29889-001, 3, Towra Point, Botany Bay; AMS IB.7273, 2; AMS 1.28505-005, I, Telegraph Creek, Jervis Bay; AMS 1.30077-001, I, Towra Point, Botany Bay; AMS IB.4746, I, Wolli Creek, Turrella; AMS 1.18065-010, 7, Tweed River; AMS 1.18061-006, I, Iluka; AMS IB.875, I, Port Hunter; AMS 1.25030-001, 2, Patonga; AMS 1.23429-001, I, Dee Why Lagoon, Sydney; AMS IB.876, I, Clarence River. Australia: Queensland: AMS 1.22795-001, 1(31.5), N of Tangalooma, Moreton Island, J. Dickie, 11 November 1975; AMS 1.25376-001, 4(29.5-38), Coomera Island, R. Morton, 22 October 1985; AMS 1.25375-001, Coomera Island, Queensland Fisheries, 22 October 1984.AMS lA.7344, 4(33-45.5), lagoon behind breakwater, Nambucca Heads, M. Ward, 20 September 1937; AMS IA.4762, 1(32.5), Flat Rock Creek, Jervis Bay, W.A. Rainbow, January 1931; AMS 1.21866-001, 1(32), Careel Bay, Pittwater, P. Weate and students, 17 October 1978; AMS 1.15857004, 1(26.5), Tuross River at Prince's Highway, D. Buckmaster, 10 September 1970; AMS 1.29891-002, Diagnosis A slender, rather flat-headed Mugilogobius with second dorsal and anal rays 1,8-10; pectoral rays 13-17; longitudinal scales 45-59; TRB 16-22; circumpeduncular scales 22-23; 10+17 vertebrae; predorsal naked or with 2-21 (mean 4) small scales on nape midline; scales on body mostly cycloid, with ctenoid scales restricted to caudal peduncle and isolated patch behind pectoral fin; no elongate spines in first dorsal fin, though tips usually free; unpaired fins low, rounded; body mottled brownish with midlateral row of darker spots and irregular Paratype of Waiteopsis paludis AMS IA.3918, 1(19), same data as holotype. Holotype of Ellogobius abascantus AMS IA.6850, 1(37), Bateman's Bay, New South Wales; G.P. Whitley, March 1936. Paratypes ofEllogobius abascantus New South Wales: AMS IA.6508, 1(28), Hen and Chicken Bay, Sydney Harbour, M. Gray, 4 September 1935; AMS 1.32053-001, 2(27.5-35.5), creek near Hanging Rock camp grounds, Bateman's Bay, G.P. Whitley, 22 March 1936. Table 25 Morphometries as percentages of SL or HL, as indieated, of Mugilogobius platynotus (Giinther, 1861). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. I. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital I. in HL Pectoral I. in SL Pelvic I. in SL Caudal I. in SL Longest Dl spine in SL h Other material examined (but not used in description) Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 28.5 61.3 75.7 19.7 27.9 42.9 51.7 13.9 7.6 20.0 10.7 22.6 19.2 37.7 20.8 16.8 12.9 22.2 9.4 33.4 55.3 75.5 19.7 14.3 25.6 15.1 31.8 26.4 58.1 33.7 22.1 19.1 26.3 15.7 30.9 49.9 66.9 16.7 11.1 23.1 13.1 28.0 22.5 48.6 29.0 19.6 15.9 23.8 13.2 22.4 46.1 61.8 13.7 8.3 19.2 10.5 22.3 21.5 36.1 23.7 15.8 12.1 19.2 11.1 29.7 61.3 75.7 24.0 14.8 26.8 14.3 29.0 28.9 43.9 34.1 22.6 18.8 26.2 14.6 28.2 51.9 67.4 17.9 11.8 23.6 13.1 25.6 24.9 40.3 28.4 19.9 16.3 24.1 12.6 22.8 14.1 27.0 21.6 40.5 25.2 20.5 16.4 22.6 _ .-·42-5.. . . Revision of MUgilogobius Figure 158 Jaws and suspensorium of Mugilogobius platynotus, female, ex AMS 1.25038-001, Brunswick Heads, New South Wales. Scale bar = 1 mm. markings; restricted to temperate Australia, from southern Queensland to Victoria. Description Based on 37 specimens, 17--45.5 mm SL. An asterisk indicates counts of holotype of Gobius platynotus. First dorsal V (in three), VI* (in 34); second dorsal 1,8*-10 (mean 1,9); anal 1,8-10 (mean 1,9*), pectoral rays 13-17 (mean 16*), segmented caudal rays 1517 (mean 16*); caudal ray pattern 9/7; branched caudal rays 7/6 to 9/7 (mean 8/7, 8/8 in holotype); unsegmented (procurrent) caudal rays 7/7 to 8/8; longitudinal scale count 45-59 (mean 53, 56 in holotype); TRB 16-22 (mean 19, 21 in holotype); predorsal scale count 0*-21 (mean 4); circumpeduncular scales 18-28* (mean 22). Gill rakers on outer face of first arch 2+5 to 4+6 (modally 3+6). Pterygiophore formula 3-12210 (in seven), 3-11210 (in one). Vertebrae 10+17 (in 11). Neural spine of second and third vertebra slender and pointed (in four) or slightly broadened at tip (in three). One (in one) or two epurals (in nine). Two (in seven) or three (in four) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid relatively low, forming long bridge across quadrate (Figure 158). Body compressed, usually rounded anteriorly. Head wider than deep, often especially so in males, HL 3.0--4.5 (mean 3.4) in SL. Depth at posterior preopercular margin 1.6-2.3 (mean 2.0) in HL. Width at posterior preopercular margin 1.3-1.9 (mean 1.5) in HL. Mouth large, subterminal, slightly oblique, forming angle of 15-25° with body axis; jaws reaching past posterior margin of eye in adult males and to below anterior half of eye in females (to posterior half of eye in holotype). Lips usually smooth, fleshy fimbriae occasionally present on inner edge of upper lip; lower lip free at sides, fused across front. Upper jaw 1.7-2.5 (mean 2.5 in 161 females, 2.1 in males) in HL. Eyes rather small, lateral, high on head, top usually forming part of dorsal profile, 3.5-5.2 (mean 4.3) in HL. Snout broad, rounded, 3.1-4.5 (mean 3.8) in HL. Interorbital broad, flat, 2.9-4.8 (mean 3.6) in HL. Interorbital, snout and sometimes side of head with sparsely scattered, fine, dark-pigmented villi (visible in holotype). Body depth at anal origin 4.27.3 (mean 5.9) in SL. Caudal peduncle compressed, length 3.7-5.2 (mean 4.3) in SL. Caudal peduncle depth 6.6-9.5 (mean 7.7) in SL. First dorsal fin rather low, generally rounded, with tips of first to fifth spines free, third and fourth spines longest or subequal and may be somewhat extended but not greatly so; spines slightly longer in males than females; all fin spines barely reaching second dorsal fin origin when depressed. First dorsal spine always shorter than next three. Second dorsal spine length 6.5-10.7 (mean 7.9) in SL. Third dorsal spine length 6.4-10.7 (mean 7.9) in SL. Fourth dorsal spine length 7.6-8.6 (mean 8.3) in SL. Second dorsal and anal fins low, rounded, posteriormost rays not much longer than anterior rays, fins falling well short of caudal fin base when depressed (usually with gap of at least five scales). Pectoral fin somewhat rounded, central rays longest, 4.4-6.3 (mean 5.1) in SL; rays usually all branched but for uppermost. Pelvic fins very short and round, reaching to less than half distance to anus, 5.2-8.3 (mean 6.3) in SL. Caudal fin rounded, 3.8-5.2 (mean 4.2) in SL. No mental fraenum, chin smooth. Anterior nostril in tube at preorbital edge, tube oriented down and forward, preorbital usually distinctly curved forward to accommodate nostril. Posterior nostril oval, placed closer to anterior centre margin of eye than to midpoint between eye and preorbital edge. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in 10), with low irregular fleshy ridge or raised bumpy flange (in nine), or with one or more fleshy knobs or flaps (in 16). Gill rakers on outer face of first arch mostly rudimentary, very short, smooth, longest one or two rakers near angle of arch; rakers on inner face of first arch few, quite stubby; outer rakers on other arches about same size as inner rakers of first arch; inner rakers on other arches nearly twice length of first arch inner rakers. Tongue tip blunt to rounded, sometimes quite short. Teeth in outermost row of upper jaw largest, stout and curved, largest teeth restricted to front of jaw; three to four rows of small sharp teeth behind this row; one or two rows at side of jaw; in females, teeth of outermost row not particularly enlarged. Lower jaw with three to five rows of small pointed teeth across front, teeth all tending to point posteriorly; no individual teeth particularly enlarged; usually only one row of teeth at side of jaw. H.K. Larson 162 A B Sensory papillae pattern longitudinal, as in Figure 159. Three s rows on snout, middle row often consisting of one papilla only; other rows of three to seven papillae each. Mental f papillae in two rows of three papillae each; row i extending in U -shape around chin. ,', \ .. ·t.·.. ... " \ ' \ 1 Figure 159 Mugilogobius platynotus, papillae pattern, AMS 1.15857-004, A, lateral view, scale bar = 1 mm; B, ventral view of chin (not to scale), Scalation omitted. Predorsal scales, if present, small, cycloid and evenly sized; nape midline naked or with 2-21 (mean four) scales on midline in 21 specimens (only one specimen with 21 scales among the Westernport Bay material, of which the others had none or up to 12 scales). Operculum with patch of small cycloid scales on upper third to half; occasionally naked. Cheek always naked. Pectoral base with cycloid scales, occasionally naked. Prepelvic area with patch of small cycloid scales before pelvics, occasionally naked. Belly with isolated patch of weakly ctenoid scales under pelvics (covering anterior % to 1/3 of belly), rest of scales cycloid. Most of body scales cycloid; ctenoid scales on side of body in two patches, one behind pectoral fin, with remaining ctenoid scales on caudal peduncle, extending forward up to behind rear of second dorsal fin. Genital papilla in male elongate, flattened, with blunt tip; in female, short, rounded and bulbous. Head pores absent as in all Mugilogobius. Figure 160 Coloration of fresh material Good colour photographs shown in Kuiter (1993: 358, two lower photographs) and Hoese and Larson (1994: 793, figure 700). Hoese and Larson (1994) illustrate an adult male with head and body dull brownish grey, scale margins darker brown, dorsal fins with broad submarginal yellow band, fin margins dark brown (difficult to distinguish from black background); lower half of first dorsal fin and lower three-quarters of second dorsal fin light greyish brown; first dorsal fin with two longitudinal dark brownish grey streaks crossing fin and intense black blotch at rear margin of fin; blotch partly extending anteriorly into uppermost dark brownish grey streak; second dorsal fin with two rows of vertically oriented dark brown spots; anal fin yellowish grey with narrow white margin; caudal, pectoral and pelvic fins plain pale yellowish grey. Male specimen in Kuiter (1993: 358, adult male, upper, and female, lowermost; upper picture almost identical to that in Hoese and Larson, 1994, probably same specimen) with some indistinct brown mottling on side of body and fine dark brownish vermiculation over top and side of head; on second dorsal fin, lowermost row of dark brown spots and blotches interspersed with light brownish grey patches; iris marbled brown and gold with distinct gold rim around pupil. Female specimen similar to male, but paler and with fins more translucent (during printing, colour apparently shifted toward greenish yellow); side of head with indistinct brownish bars and irregular streaks, with short bars from eye to upper jaw; dorsal surface of head and upper parts of opercle and preopercle with dark vermiculation; belly whitish; iris pale golden with light brown speckling. Coloration of preserved material Light brown to pale yellowish brown (depending on preservation), with indistinct scattered brown mottling and fine vermiculation on side of body and MlIgilogobills platynotus. Holotype of Ellogobills abscantlls Whitley, 39 mm SL, AMS 1.6850, Bateman's Bay, New South Wales. Revision of Mugilogobius Figure 161 163 Mugilogobius platynotus. In fresh water, Tooradin, Victoria. From colour slide by Rudie Kuiter. top of head; lower half of body usually without mottling; scale margins often narrowly outlined with brown, giving finely reticulate appearance in close view (Figures 160-161). Many specimens with midlateral brownish band formed by series of indistinct small dark blotches (nine or more) and mottling; brown band often discontinuous; small rounded brown spot always present at mid-eaudal base; series of broken-up narrow oblique brownish bars visible on upper anterior half of body, including brown oblique shoulder bar. Top of head and nape with fine brown vermiculation and spotting; side of head with brownish mottled, vermiculate or reticulate pattern; in some specimens two or three indistinct bars running from eye to upper jaw; usually dark brownish blotch on opercle close to rear preopercular margin. Pectoral base usually dusky with brownish blotch on upper half. Underside of head, branchiostegal rays and anterior part of breast plain dusky to brown. Belly pale; peritoneum dark brown dorsally and on sides, pale ventrally. Entire gill chamber pale. First dorsal fin spines and lower half of fin light to dark brown to greyish, fin with narrow dark brown margin and broad submarginal white to translucent whitish band; rear of fin posterior to fifth fin spine with black rounded spot; small specimens and females usually with pale first dorsal fin markings and black spot at rear of fin replaced by dusky brownish spot, lower half of fin translucent to pale dusky with narrow brownish longitudinal streak along centre of fin. Second dorsal fin translucent dusky to light brown to greyish with narrow submarginal whitish band and narrow brown to dusky margin; dusky lower portion of fin with two series of vertically oriented dark brown short streaks and blotches, proximal series usually of larger, more diffuse blotches. Anal fin plain dusky to brownish. Caudal fin plain dusky; large or heavily pigmented specimens sometimes with diffuse brownish spotting close to fin base (large male from Victoria with narrow brownish grey margin around entire fin), spotting sometimes coalescing with small brown spot at midbase of caudal (last spot of midlateral series). Pectoral fin with rays dusky to brownish; membrane dusky in some specimens. Pelvic fins pale, dusky or brownish; fraenum pigmented if rest of fin pigmented. Comparisons Mugilogobius platynotus is most similar to M. stigmaticus (both having similar high scale counts), and the two species overlap in range (New South Wales and southern Queensland). The two are most easily distinguished by predorsal scale number and extent (0-21, mean of 4, in M. platynotus, scales just reaching preopercular edge or either side of naked midline; versus 17-30, mean of 22, in M. stigmaticus, with scales extending halfway between preopercular border and eyes, and midline always scaled). Also, M. platynotus mostly has cycloid scales on the body, with ctenoid scales restricted to two areas, one patch behind the pectoral fin and an area on the caudal peduncle extending forward to behind rear of the second dorsal fin. Mugilogobius stigmaticus has a more extensive covering of ctenoid scales on the body, which usually extends forward to behind the pectoral fin. This species has a relatively low metapterygoid; the metapterygoid in M. stigmaticus is broader than in M. platynotus but not as broad as in other species of the genus. Distribution Specimens are known from Victoria to southern Queensland, south-eastern Australia (Figure 123). The record by Hoese and Larson (1994) from South Australia is dubious (Westernport Bay may be the western-most locality from which the species is known). Ecology Ivey (1951) gave a description of keeping this species in captivity for about a year, during which time she tried to acclimatise the fish to fresh water. Her description of them regularly leaping out of the H.K. Larson 164 water suggests that the change was not welcomed by the fish, which came from rock platform pools at Double Bay, Sydney, a marine habitat. The leaping behaviour described by Ivey is used by many gobioids seeking to escape from an unsuitable environment. I am quite familiar with the great leaps of which Mugilogobius is capable, as well as its ability to remain alive out of the water for considerable lengths of time (this latter ability shared by Calamiana sp. novo 3 and possibly others of that genus). Ivey's description of the fish blowing groups of bubbles indicates that the fish were probably kept in a very small container. This behaviour agrees with that reported by Gee and Gee (1991) for M. paludis (= M. platynotus), M. stigmaticus, Chlamydogobius sp. (it is uncertain which species they had), Mugilogobius sp. 9 (= Chlamydogobius ranunculus), Pseudogobius olorum and 15 other gobioids held in aquaria. More than half the fish would perform aquatic surface respiration when available oxygen fell to below 2.1 ppm. Bubble-gulping was observed in nine species (including all the gobionellines tested) and was often used by the fish in conjunction with other behaviours which increased access to oxygen. That Ivey's fish slowly ceased bubble-gulping may indicate that they were able to adapt to the low oxygen and low salinity environment. Bell et al. (1984) found this species to be a permanent resident of a mangrove site in Botany Bay, studied over a three-year period; they presented limited life history and biomass data. Morton et al. (1987) and Morton et al. (1988) described the abundance and diets of several fishes inhabiting a saltmarsh inlet at Coomera Island, in southern Moreton Bay, Queensland. The fishes studied included M. platynotus (reported as M. paludis), M. stigmaticus and Acentrogobius species. Unfortunately, neither Morton et al. (1987) nor Morton et al. (1988) separated the data for any of the gobiids sampled, so it is useful only in a very general way. The gobiids were mostly carnivorous, consuming algae, culicid larvae, other insects, arachnids, crustacea and detritus during autumn and winter, and ate exclusively culicid larvae and other insects in the summer. The habitat occupied by M. platynotus in these two studies was the main Figure 162 inlet to the saltmarsh (chiefly vegetated by the mangroves Avicennia and Rhizophora, with saltcouch grass Sporobolus and the succulent Salicornia towards the centre of the island). Salinity ranged from 22-37 %0 and temperature from 14-27SC. The specimens from Westemport Bay, Victoria, were recorded as being from an estuary with a soft mud and sand substrate in still, clear water with some "aquatic vegetation" (P. Unmack, in litt.). Remarks Whitley (1930) indicated that AMS IA.3917 was the type of his Waiteopsis paludis, but he neglected to state that there were actually two specimens in the bottle. The holotype (larger of the two) was illustrated in Whitley (1951: 405, figure 13), in which he clearly stated that the illustrated fish is the holotype. Koumans (1931) was puzzled by Whitley's characteristically brief generic diagnosis and asked "Does this genus belong to Gobiidae?". This species has recently been referred to in the literature as Mugilogobius paludis (Figure 162). From examination of the holotype of Gobius platynotus Giinther, 1861, (BMNH 1859.5.7.71), it would appear that it is conspecific with M. paludis. Although the type locality is not known, the naked nape, mostly cycloid scales, high number of circumpeduncular scales, vertebral number of 10+17, and large mouth (for a female) place Giinther's species as senior synonym to paludis. Mugilogobius platystomus (Giinther, 1872) Figures 162-168; Tables 5-8, 26 Gobius platystoma Giinther, 1872: 664, plate 63, figure B (Port Mackay, Queensland, Australia); McCulloch 1929: 372. Gobius mertoni Weber, 1911: 37-38, figures 5-6 (Panua Bori River on Sungi Manumbai, Wokam; forest creek at Dobo; forest creek at Udjir; Aru Islands, Indonesia) (in part); Nijssen et al., 1993: 232. Mugilogobius mertoni: Tan and Tan 1994: 356. Mugilogobius cf stigmaticus: Pusey and Kennard 1994: 27, 29, 80. Mugilogobius platynotus. Holotype of Waiteopsis paludis Whitley, 18 mm SL, AMS IA.3917-18, Gundamaian, Port Hacking, NSW. Revision of Mugilogobius Material Examined Holotype of Gobius platystoma BMNH 1871.9.13.179, 1(40), Port Mackay, Queensland; Godeffroy Collection. Paralectotype of Gobius mertoni Indonesia: ZMA 112.660, 1(18), forest creek, Udjir, Aru Islands, H. Merton, 15 April 1908; paralectotype of Gobius mertoni, SMF 6692, 1(20), Udjir, Aru Islands, H. Merton, 15 April, 1908; paralectotype of Gobius mertoni, SMF 6697-8, 2(13.514.5), Dobo, Pulau Wamar, Aru Islands, H. Merton, 6 March 1908; paralectotype of Gobius mertoni, SMF 6693-4, 1(12), Udjir, Aru Islands, H. Merton, 15 April, 1908. Other Material Singapore: CAS 36031, 4(23.5-34), Serangoon, March 1938. Belau: CAS 54690, 18(18-37.5), freshwater streamlet at Ngarbaged, Koror, Sumang et al., 2 October 1955; AMS 1.27217-001, 8(14-18), streamlet in taro swamp below Conservation Office, Koror, G. Bright, 22 February 1978. Australia: Northern Territory: NTM 5.14297-001, 2(31-32), drain under road by Darwin High School, D. Wilson, February, 1990; AMS 1.24691-002, 2(13.514.5),250 m from crossing, Leader's Creek, Darwin, D. Hoese and S. Reader, 14 September 1984; NTM 5.10694-018, 2(21-22.5), mangrove creek at Gunn Point, H. Larson and R. Williams, 20 September 1982; NTM 5.1867-001, 3(17.5-19), Marchinbar Island, Wessel Islands, P. Latz, 19 October 1972. Queensland: AMS 1.22959-002, 7(27-34), lagoon behind wharf at AIMS, Townsville, J. Gee, 16 July 1981; NTM 5.14204-001, 3(32-38), swamp adjacent to Tully River, B. Pusey, October 1993; NTM 5.14202-002, 2(28-33.5), Jarra Creek, Tully River, B. Pusey, 18 October 1993; NTM 5.14205-001, 2(30-40), swamp near mouth of Murray River, B. Pusey, 24 October 1993. Other material examined (but not used in description) Indonesia: ZRC 34197-8, 2, Pulau Bintan, Rhiau Archipelago; CMK 10640, 5, saltwater lake, Pulau Satonda, Sumbawa. Australia: Northern Territory: Figure 163 165 NTM 5.14290-010, 15, Leader's Creek; NTM 5.14289-008, 12, Leader's Creek. Queensland: AMS 1.21228-005, I, 10 km 5 of Bowen; AMS 1.21280-001, 2, between Mackay and Sarina. Diagnosis A moderately slender Mugilogobius with second dorsal and anal rays 1,7-9 modally 1,8; pectoral rays 14-17; longitudinal scales 39-57; TRB 12-18; circumpeduncular scales 14-19; predorsal scales small, 18-30, reaching at least to above preopercular margin; ctenoid scales on body in two patches, one extending forward in narrow wedge to below second dorsal fin origin and other patch behind pectoral fin; second or third spine of dorsal fin longest, spines often elongate but not filamentous; body with about seven oblique to nearly vertical oval brown blotches or bars along side and pair of dark spots at caudal base; known from Singapore, Belau, Indonesia and north-eastern Australia. Description Based on 44 specimens, 12-40 mm SL. An asterisk indicates counts of holotype of Gobius platystoma (Figure 163). First dorsal VI*; second dorsal 1,7-9 (mean 1,8*); anal 1,7-9 (mean 1,8*), pectoral rays 14-17 (mean 16*), segmented caudal rays 15-16 (mean 16*); caudal ray pattern 7/6 to 9/7 (modally 8/7); branched caudal rays 13-16 (mean 15); unsegmented (procurrent) caudal rays 6/6 to 7/7 (modally 6/7); longitudinal scale count 39-57* (mean 45); TRB 12-18 (mean 16*); predorsal scale count 18-30 (mean 22, 28 in holotype); circumpeduncular scales 14-19 (mean 16, 19 in holotype). Gill rakers on outer face of first arch 3+7 to 4+8 (modally 4+7). Pterygiophore formula 312210 (in 12), 3-11310* (in two). Vertebrae 10+16 (in 14), 11+15* (in one). Neural spine of first three vertebrae narrow, pointed (in seven), or second to third neural spine slightly broadened (in four). Two epurals (in 12). Two (in 12) or three (in three) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid moderately broad, broadest toward centre; forming bridge to quadrate. Six ossified gill rakers (in one 34 mm specimen). Mugilogobius platystomus. Holotype of Gobius platystoma Giinther, 1872, 41 mm SL, BMNH 1871.9.13.179, Port Mackay, Queensland. H.K. Larson 166 Body relatively slender, compressed, less so anteriorly. Head depressed, always wider than deep, HL 3.1-5.1 (mean 3.5) in SL; cheeks sometimes inflated. Depth at posterior preopercular margin 1.62.1 (mean 1.8) in HL. Width at posterior preopercular margin 1.2-1.5 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of about 25° with body axis; jaws enlarged in males, reaching past rear margin of eye in males, and to below front half of eye to mid-eye in females (as in holotype). Lips smooth, fleshy fimbriae on inner edge of upper lip and less often, across front of lower lip (fimbriae sometimes very close to outer edge); lower lip free at sides, fused across front. Upper jaw 1.8-3.1 (mean 2.6 in females, 2.2 in males) in HL. Eyes lateral, high on head, top usually forming part of dorsal profile, 3.2-5.1 (mean 4.2) in HL. Snout rounded, 3.2-4.8 (mean 3.8) in HL. Interorbital broad, flat, 2.4-5.1 (mean 3.3) in HL. Unscaled portion of top of head above preopercular margin up to snout tip covered with fine villi, sometimes extending into preorbital region. Body depth at anal origin 4.8-7.8 (mean 5.6) in SL. Caudal peduncle compressed, length 3.8-6.0 (mean 4.4) in SL. Caudal peduncle depth 6.5-10.5 (mean 7.5) in SL. First dorsal fin triangular, tips of second to fourth spines free and usually elongate but not greatly so (i.e. not filamentous), second or third spines longest or subequal; spines longer in males than females; spines often falling short of second dorsal fin origin when depressed. First dorsal spine always shorter than next three. Second dorsal spine length 5.1-8.6 (mean 6.7) in SL. Third dorsal spine length 4.4-9.6 (mean 6.5 in males, 7.2 in females) in SL. Second dorsal and anal fins low, pointed posteriorly, posteriormost rays longest; dorsal rays rarely reaching, and anal rays always falling short of, caudal fin base when depressed; however, in one syntype of Gobius mertoni, SMF 6692, second spine reaching back to second element in second dorsal fin. Pectoral fin oval to rounded, central rays longest, 4.1-5.6 (mean 4.8) in SL; rays usually all branched but for uppermost. Pelvic fins short, oval, reaching half to one-third distance to anus, 4.7-9.2 (mean 5.6) in SL. Caudal fin rounded to oval, 3.14.5 (mean 3.8) in SL. No mental fraenum, chin smooth. Anterior nostril in short tube placed just at edge of upper lip, tube oriented down and forward, preorbital produced forward to accommodate nostril. Posterior nostril oval to rounded, placed near anterior centre margin of eye. Gill opening extending forward to just under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in five), with low fleshy ridge or raised bumpy flange (in 21), or with distinct fleshy knobs or flaps (in 18). Gill rakers on outer face of first arch very short, stubby, sometimes finely papillose at tips, longest rakers near angle of arch; rakers on irmer face of first arch and outer face of second also stubby - Figure 164 Mugilogobius platystomus, papillae pattern. AMS 1.22959-002, Townsville, Queensland. Scale bar =1 mm. and finely papillose; inner rakers on other arches twice length of first arch inner rakers. Tongue tip blunt to concave; almost bilobed in five specimens from Queensland. Outer teeth in upper jaw largest, stout and curved, especially across front, three to four rows of very small sharp teeth behind this row; one or two rows at side of jaw; teeth larger in males than in females. Lower jaw with four or five rows of small pointed teeth across front, most teeth oriented upright, teeth in innermost row or two largest and stoutest, all pointing posteriorly; one or two rows of teeth at side of jaw. Predorsal scales small, evenly sized, reaching forward to above rear preopercular margin or slightly further, or halfway between preopercular margin and rear of eyes (as in holotype). Operculum with small cycloid scales; lower third to half unscaled. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area with small cycloid scales in patch before pelvic fins at least; if most of breast scaled, anteriormost region near isthmus, naked. Belly with all scales cycloid or with isolated patch of ctenoid scales under pelvic fins, covering anterior third of belly or less. Ctenoid scales on side of body in two patches, with narrow wedge extending forward up to about gap between dorsal fins and an isolated small patch behind pectoral fin (in holotype and 35 other specimens); sometimes all scales behind pectoral fin cycloid; seven specimens with narrow wedge of ctenoid scales extending along midside to behind pectoral fin (only one or two rows of ctenoid scales anterior to mid-dorsal gap). Genital papilla in male elongate, flattened, with pointed tip; in female, short, rounded and bulbous, with blunt to round tip. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 164. Three s papillae rows on snout, middle row of one to five papillae. Mental f papillae in single row or two short rows of either side of symphysis; 167 Revision of Mugilogobius Figure 165 Mugilogobius platystomus. Aquarium specimen from Darwin, NT. From colour slide by Neil Armstrong. mandibular i row sometimes extending right around chin. Coloration of fresh material Fresh specimens with colour pattern very like that of preserved specimens (Figure 165). Body light greyish brown, whitish grey ventrally, and body markings dark brown, with two black spots on caudal fin base; first dorsal fin with yellow area at front of fin, just below central blackish streak and black spot at rear. Coloration of preserved material Head and body greyish brown to pale yellowish brown, scale margins darkest; with dark brown oblique to vertical oval blotches and bars on sides, posteriormost bar just before caudal base with centre intensified as square blotch; two horizontal brown stripes crossing side of head (Figure 166). First three or four dark bars on side slanting forward, next three slanting posterioriy, with last bar or blotch vertical; considerable variation occurring in shape and orientation of body bars. Anteriormost body bar ("shoulder" bar) most oblique, slanting over pectoral base, ending on nape above opercle. Second body bar usually most conspicuous, forming dark brown, vertically oriented oval blotch behind pectoral fin. Small brown, square saddles, blotches and spots crossing mid-dorsal line, some coalescing with body bars; posteriormost saddle crossing upper base of caudal fin. Nape with brown spots and irregular to vermiculate streaks along either side of midline; interorbital and snout with brown vermiculation sometimes forming short streaks to upper half of eye. Figure 166 Side of head light brownish yellow with diffuse, short, oblique brown line from rear of eye extending above preopercle onto nape; short, dark brown, oblique line from lower anterior edge of eye to middle of upper jaw; distinct, dark brown, horizontal line running along lower edge of eye (often joining dark line from eye to jaw) across preopercle, ending on dusky brownish opercle; and dark brown horizontal line from near upper edge of upper lip extending across cheek to end on opercle. Rear edge of opercle often dark brown, in some specimens colour extending part-way onto branchiostegal membranes. Ventral margin of preopercle sometimes narrowly outlined with dark brown. Pectoral base dusky brown to yellowish, with horizontal dark bar crossing at midpoint. Breast and underside of head pale to dusky brown, chin often darkest. Belly plain whitish yellow. First dorsal fin dusky to translucent on proximal third, centre of fin with blackish streak widening posterioriy to black blotch, outer part of fin whitish to dusky with dusky to blackish margin; free tips of spines dusky to whitish. Second dorsal fin dusky proximally, with submarginal white to translucent band, fin margin dusky to dark grey; proximal twothirds of fin dusky with about two rows of vertically oriented blackish or brown streaks or oval blotches, two evenly spaced dark brown blotches at base of fin. Anal fin plain dusky to dark grey with whitish to translucent margin. Caudal fin plain dusky, membrane darker than rays, forming indistinct narrow streaks; two distinct dark brown to blackish oval spots at base, placed one above the other, usually oriented slightly obliquely, pointing Mugilogobius platystomus, 39 mm 5L, NTM 5.14204-001, Tully River, Queensland. .... a-. 00 ｾｬ｢ＧｾＢ <lO' ｾ＿Ａ I1JAPAN ;* . 30' # p. '.. , t . RYUKYU IS. • .... .' ". 20": 20' HAWAIIAN ISLANDS P . • ; MAIl1ANA IS. Ck.1LWf'lNES LACCADIVE IS. ': .. 10' Maldlv•• ｪｾｾ｜Ｎ｀Ｍｾ • :.' ", Ｎｾ .. 10' . ,': MARSHALlIS. ' CAROUNE IS: (,,-\-... Kosrae ｬｾｯＱ＼ KIRIMTI O' ､ＧＺｾｓｏｌｍｎ .} ":', SEYCHEllES ... 10' (J 20' o • ptafystoma rambaiae rivutus n. sp. *: stigmaticus tigrinus n, sp. witsoni n. sp, ＧｾＮ •... 30' ' .' <lOo V 110' .. ', SAMOA '." Ch, FIJI:' . '.' 20' 'LOYALlY IS. 30' *... 60' 0' 10' .. MADAGASCAR 40' ... ,.: , Maurrtius "'J' " IS. '.'...:, "c.' -. • . 100' 120' l<lO' ,_.. ｾｊ｣ 160' <lOo 0 180' 160' ::t ｾ r Figure 167 Distribution of Mugilogobius platystomus, M, rambaiae, M. rivulus n. sp., M. stigmaticus, M. tigrinus n. sp. and M. wilsoni n. sp. Ul o ::l Revision of Mugilogobius Table 26 169 Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius platystomus (Giinther, 1872). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic !. in SL Caudal!. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 28.8 59.1 74.8 20.8 19.7 47.2 65.2 12.9 9.0 16.6 9.5 22.4 19.6 36.4 24.6 18.9 10.8 22.1 10.4 31.9 62.2 86.4 20.8 14.9 26.2 15.5 30.9 31.0 55.3 41.1 24.7 22.2 31.9 22.6 29.5 55.8 73.2 17.7 12.2 23.0 13.5 27.7 24.3 46.2 33.2 21.6 17.7 26.5 15.9 27.1 53.1 66.0 15.6 9.3 19.4 12.3 20.8 21.3 32.4 19.7 17.9 17.1 22.6 12.3 30.9 61.8 80.2 20.0 14.0 26.2 14.8 28.9 30.2 42.7 38.4 24.2 21.3 30.3 18.8 28.7 56.8 73.4 18.5 12.2 23.1 13.4 26.0 24.3 39.1 30.1 21.0 18.5 26.4 14.6 22.5 15.5 27.0 22.6 52.2 37.4 20.0 15.8 13.8 toward last square body bar near caudal base. Pectoral fin with rays dusky to brown, membranes translucent. Pelvic fins dusky to dark brown, fraenum paler. Peritoneum brown on sides, dorsal midline and ventral area unpigmented. Comparisons This species is most similar to M. stigmaticus and M. littoralis sp. novo Mugilogobius littoralis sp. novo has two isolated sections of ctenoid scales on the body as does M. platystomus, but has a black peritoneum (versus brown dorsally, pale ventrally in M. platystomus), one small dark spot on the body near the caudal fin base and a plain caudal fin (versus two black spots on caudal fin and one on the body near base), and a diffuse colour pattern with narrow vertical bars and spots (versus oblique blotchy bars on the body). Mugilogobius platystomus is distinguished from M. stigmaticus under Comparisons for that species. Distribution Found from northern Australia to Belau, Indonesia and Singapore (Figure 167). Ecology Found in freshwater to marine habitats, usually from small brackish to freshwater streams running through mangrove or rain forest, or swamp. Darwin Figure 168 specimens inhabited water of 19-26%0 salinity and Tully River specimens were from habitats ranging from pure freshwater to 7%0. Remarks The holotype of Gobius platystoma Giinther is an aberrant specimen, having 11+15 vertebrae, dorsal pterygiophore formula of 3-11310 (shared only by one cleared and stained specimen from the Wessel Islands, NT), and a high number of lateral scales. Other characters are typical of the species. As indicated earlier, the paralectotypes of Gobius mertoni Weber consist of two species of Mugilogobius and a Pandaka. ZMA 112.660 (Figure 168), 1(18); SMF 6692,1(20) and SMF 6697-6698, 2(13.5-14.5) are M. platystomus. In SMF 6693-6696, one, 12.5 mm SL, is M. platystomus and three, 8-11 mm SL, are a species of Pandaka. The illustration of M. platystomus in Weber (1911: figure 5), is in error in showing the first dorsal fin spine longest. The artist has also exaggerated the triangular form of the dorsal fin; but the colour pattern more or less agrees with recent material. Pusey and Kennard (1994) recorded this species as M. stigmaticus from the wet tropics of Queensland (Tully and Murray Rivers). These specimens are very heavily pigmented, with blackish blotches and enlarged spots on the body (Figure 166); otherwise they resemble the Townsville specimens. Mugilogobius platystomus. Paralectotype of Gobius mertoni, 18 mm SL, ZMA 112.660, Udjir, Aru Islands. H.K. Larson 170 Mugilogobius rambaiae (Smith, 1945) Figures 9F, 16A, 167, 169-173; Tables 5-8,27 Vaimosa rambaiae Smith, 1945: 538, plate 9a (Bangkok, Thailand); Suvatti 1981: 204, plate 306, figure 128, plate 307. Vaimosa valigouva: Axelrod et al., 1990: 865 (misidentification). Mugilogobius rambaiae: Kottelat 1989a: 19; Tan and Tan 1994: 356. Mugilogobius sp. undet; Roberts 1989: 168. Guinea: WAM P.30975-007, 7(13.5-27), Buraratere Creek, Kikori River, G.R. Allen, 3 March 1995; WAM P.30966-002, 16(13.5-25.5), Kopi Chevron Camp, Kikori River, G.R. Allen, 27 February 1995. Other material eXilmined (but not used in description) Burma: SMNS 18851, 1, Arakan Province, Kyeintali Chaung River. Malaysia: ZRC unregistered (THH 9553), 1, near Kuching, Sarawak; NTM 5.14351-001, 1, same data as previous. Indonesia: ROM 68716, 1, Ajkwa River, near Timika, Irian Jaya. Material Examined Holotype ofVaimosa rambaiae USNM 119646, 1(41.5), caught from shallow slough of the klong [canal] behind Department of Fisheries, central Bangkok, Nai Pongse, Thailand; 28 May 1931. Paratypes ofVaimosa rambaiae USNM 119647, 9(24-33), same data as holotype. Other Material Thailand: CMK 5367, 2(12-17.5), estuary at Ao Kammala, Phuket, M. Kottelat, 23 April 1985. Singapore: CAS/SU 36032, 30(21.5-36), Serangoon, March 1938; ZRC 19886-95, 24(24-34.5), tidal stream near Woodlands, N. and K. Lim, 30 May 1990; ZRC 26972-3, 2(29.5-33.5), tidal stream 5 of Woodlands customs checkpoint, P. Ng et al., 30 January 1992; ZRC 29554, 1(25), Sungei Buloh mangroves, A. Wong et al., November 1992. Malaysia: NTM 5.14349-001, 7(12-25), coastal stream at Pantai Keracut, NW corner Penang, K. Urn, N. Sivasothi, 27 January 1994. Sri Lanka: USNM 316172, 1(23), mouth of Kelani River, Mattakuliya village near Colombo, T. Roberts, 12 March 1970; ZMH 7992, 1(31), lagoon at Panadhure, SW Sri Lanka, Duncker, 30 July 1909. Indonesia: ZRC 32864-6, 3(20-21), Pulau Bintan north, Rhiau Archipelago, T.H.T Tan, 12 May 1993; CAS 49460, 4(8-16.5), man-made canal entering Sungai Landak, 5 km E of Pontianak, Kapuas River basin, Borneo, T. Roberts and S. Woerjaotmodjo, 10 July 1976; BMNH 1937.6.14.2223, 2(38.5-41), East Indies; BMNH 1935.5.27.27, 1(27.5), Celebes or Borneo, Arnold. Papua New Figure 169 Diagnosis A moderately slender Mugilogobius with second dorsal rays 1,7-8; anal rays 1,7-9; pectoral rays 1417; longitudinal scales 28-38; TRB 8-12; circumpeduncular scales 11-13; predorsal scales 1120, small, reaching up to behind eyes, one to several scales sometimes larger than others; scales on body mostly ctenoid; third to fourth dorsal fin spines longest; body greyish, scales with short vertical or curved dark lines, distinct brown oblique shoulder bar present; known from fresh to brackish water in Thailand and Burma throughout the Malaysian Peninsula, Sri Lanka and Papua New Guinea. Description Based on 48 specimens, 12-41.5 mm SL. Counts of holotype of Vaimosa rambaiae (Figure 169) indicated by asterisk. First dorsal VI*; second dorsal 1,7-8* (mean 1,8); anal 1,7*-9 (mean 1,8), pectoral rays 14-17 (mean 15*), segmented caudal rays 14-16 (mean 16*); caudal ray pattern 6/4 to 9/7 (modally 8/7, 7/7 in holotype); branched caudal rays 10-16 (modally 16, 14 in holotype); unsegmented (procurrent) caudal rays 7/7 to 7/8 (modally 7/7); longitudinal scale count 28-38 (35 in holotype, mean 33); TRB 8-12 (mean 11*); predorsal scale count 11-20 (mean 16, 11 in holotype); circumpeduncular scales 11-13 (mean 12*). Gill rakers on outer face of first arch 2+6 to 4+7 (modally 2+6). Pterygiophore formula 312210 (in 20). Vertebrae 10+16 (in 33), 11+15 (in one). Neural spines of first three vertebrae bifid or expanded at tip (in 14) or slender and pointed (in four). Two epurals (in 30); one epural in one Mugilogobius rambaiae. Holotype of Vaimosa rambaiae Smith, 32.5 mm SL, USNM 119646, Bangkok. Revision of Mugilogobius specimen (two epurals partly fused at base). Two (in 18) or three (in 15) anal pterygiophores before haemal spine of first caudal vertebra. Body compressed, more rounded anteriorly, HL 2.6-4.8 (mean 3.5) in SL. Head wider than deep, somewhat flattened in adults; cheeks sometimes inflated. Depth at posterior preopercular margin 1.52.0 (mean 1.7) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.3) in HL. Mouth subterminal, slightly oblique, forming angle of about 20-25° with body axis; jaws generally reaching to below posterior half of eye in males and to below mid-eye to anterior half of eye in females (to below mid-eye in holotype). Lips usually smooth, fleshy fimbriae present mostly on inner edges of upper lip; lower lip free at sides, fused across front. Upper jaw 1.9-3.7 (mean 2.8 in females, 2.4 in males) in HL. Eyes lateral, high on head, forming part of dorsal profile, 3.2-4.3 (mean 3.7) in HL. Snout rounded, 3.2-4.4 (mean 3.8) in HL. Interorbital broad, flat, 1.2-3.9 (mean 3.0) in HL. Top of head, from in front of scaled nape forward to snout tip, usually covered with fine villi, sometimes relatively sparse (dense in holotype). Body depth at anal origin 3.4-6.2 (mean 5.3) in SL. Caudal peduncle compressed, length 3.4-5.2 (mean 4.2) in SL. Caudal peduncle depth 4.8-8.8 (mean 7.4) in SL. First dorsal fin generally low, tips of spines free but not filamentous, third and fourth spines longest or subequal, fourth usually longest; spines slightly longer in males than females; spines just reaching (or falling short of) second dorsal fin origin when depressed. First dorsal spine always shorter than next three. Second dorsal spine length 7.8-9.6 (mean 8.6) in SL. Third dorsal spine length 5.7-9.8 (mean 8.0) in SL. Fourth dorsal spine length 5.5-13.4 (mean 8.2) in SL. Second dorsal and anal fins low, fins rounded anteriorly, elongate posteriorly, last dorsal rays sometimes reaching upper caudal rays when depressed (anal rays falling short of caudal when depressed). Pectoral fin rounded to oval, central rays longest, 3.8-5.6 (mean 4.9) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, may reach about halfway to anus, 3.4-8.0 (mean 5.3) in SL. Caudal fin rounded to rectangular, 3.0-4.1 (mean 3.8) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, placed just behind upper lip, tube short, oriented down and forward, preorbital slightly curved forward to accommodate nostril. Posterior nostril oval, placed closer to anterior centre margin of eye. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in two) or with low irregular fleshy ridge or raised bumpy flange (in 11), or with distinct fleshy knobs and flaps (in 28). Gill rakers on outer face of first arch very short knobs, longest raker below angle of arch; rakers on inner face of first arch very rounded, stubby, with fine spiny papillae at tip; 171 inner rakers on other arches not much longer than first arch inner rakers (three rakers ossified in cleared and stained specimen). Tongue tip usually blunt to rOlmded, sometimes concave. Outer teeth in upper jaw largest, stout and curved, about three rows of small sharp inward-pointing teeth behind this row; usually two rows at side of jaw; teeth generally rather smaller in females. Lower jaw with about five rows of small pointed teeth across front, outermost row often oriented upright, inner rows pointing inward; no teeth particularly enlarged; usually only one row of teeth at side of jaw. Predorsal scales small, usually evenly sized, reaching forward to behind eyes. Anterior one to three scales sometimes somewhat larger than rest of predorsal scales, but no scale so large as to fit across interorbital space (as in chulae species-group). Operculum with small cycloid scales on upper twothirds at least. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1/4 to 1/3 of belly), rest of scales cycloid. Ctenoid scales on side of body extending forward in wedge up to behind pectoral fin; specimens from Papua New Guinea with ctenoid scales forming narrow wedge, sometimes broken into two patches below gap between dorsal fins. Genital papilla in male elongate, flattened, with blunt to rounded tip; in female, papilla short, bulbous, tip rounded to blunt. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 170. Cheek row c sometimes continuous below eye; papilla row e ending just after turning up onto rear edge of preopercle; three s rows on snout, with three to four papillae in each row; f row behind mandibular symphysis with four to six papillae in straight line. ＬＺｾｰＱＨＵｬＺ .. \Y . ｾＮＧ .. ,':" .... ｾ .:., :> .; . '" .... "c Figure 170 MlIgilogobills rambaiae, papillae pattern. Para type of Vaimosa rambaiae, USNM 119647. Scales omitted. Scale bar = 1 mm. 172 Table 27 H.K. Larson Morphometrics as percentages of SL or HL, as indicated, of Mugilogobius rambaiae (Smith, 1945). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. 1. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital 1. in HL Pectoral 1. in SL Pelvic 1. in SL Caudal 1. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 21.0 69.0 74.7 16.1 13.3 19.0 11.3 23.0 26.4 41.4 31.0 29.9 59.4 75.3 19.8 13.5 24.7 14.2 27.0 26.3 43.6 33.6 20.5 18.8 27.1 14.5 38.5 66.3 85.3 29.8 16.4 29.5 20.7 31.1 30.0 52.9 40.0 26.5 21.2 33.1 18.3 27.5 52.0 62.7 17.4 10.9 22.6 12.6 22.8 23.3 35.1 25.5 17.9 16.8 23.3 11.3 21.0 51.3 65.1 16.1 9.6 19.0 11.3 23.0 26.4 30.0 26.7 18.8 12.5 24.1 7.5 31.0 69.0 84.3 22.0 16.1 26.9 15.2 29.2 31.2 41.4 39.7 23.6 21.5 29.6 17.6 27.7 60.9 76.1 18.7 13.2 24.1 13.3 26.5 28.7 36.4 31.6 20.9 18.8 26.7 12.6 12.5 7.5 Coloration of fresh material Smith (1945) gave a detailed description of living specimens, as follows: Entire body and head except abdomen soft heliotrope gray; back with about 5 indistinct darker bands, which extend obliquely downward and forward to middle of side, the first band under first dorsal fin; each scale of back and side with a narrow reddish brown lunate spot; abdomen yellow-green; underside of head with 7 dark brown, strongly curved, parallel cross lines, the first one immediately behind and following curvature of the lower lip; first dorsal grayishgreen on basal half, heliotrope gray on free edge; with a submarginal transverse band of white involving the first 5 membranes and basally thereto a dark purplish gray band, which merges into the green shade at the base of the fin; second dorsal similar to first, but with a row of elliptical dark purplish gray spots in the paler purplish median band, a spot on each membrane; entire caudal fin uniform pale apple green, with about 7 curved, wavy, cross stripes of heliotrope gray; anal pale grayish-blue at base, pale lavender distally, all rays heliotrope gray; ventral and pectoral fins pale green to greenish yellow. Suvatti (1981: plate 307) included a copy of Luang Masya's painting of this species, reproduced as a Figure 171 black and white photograph in Smith (1945). The reproduction of the colour plate is rather shifted toward the red, but shows most of the colours indicated by Smith. A similar painting, of several specimens, was presented to the former Queen of Siam, Rambai Bami (Smith 1945), and presumably is still held by the Thai royal family. Axelrod et al. (1990) show a captive female which is light brownish grey, with a pale yellowish abdomen. Each body scale on the side and back has a dark brown vertical line, the spotting and streaks on the dorsals, anal and caudal fin is black, the shoulder bar is dark brown to blackish and there are two indistinct dusky saddles across the upper sides. The head has a network of reddish brown lines, forming rounded interspaces, and the iris is reddish. Coloration of preserved material Head and body pale whitish yellow to pale greyish yellow (depending on state of preservation); with most distinctive marking being broad dark brown oblique bar extending from behind pectoral base up onto predorsal area (bar falling short of meeting its counterpart on nape midline); four to six less distinct oblique brown saddles crossing back Mugilogobius rambaiae. Paratype of Vaimosa rambaiae Smith, 34.5 mm SL, USNM 119647, Bangkok. Revision of Mugilogobius Figure 172 173 Mugilogobius rambaiae, 34.5 mm SL, ZRC 19886-19895, Woodlands, Singapore. (Figures 171-172). Each scale of back and sides with short vertical to gently curved brown line near centre of scale; vertical lines more pronounced and placed closer to scale margin on scales on posterior half of body; scales also often narrowly outlined with grey to brown. Brown dorsal saddles or bars often indistinct but discernible, always variably developed among individuals. Top of head with indistinct blotches or marbling, usually several irregular lines joining across interorbital space or extending onto nape from upper rear margin of eye. Side of head with dark brown lines forming reticulate network of pale interspaces, especially prominent on cheek; dark lines often forming oblique stripes across opercle; two or three brown lines extending from front of eye onto preorbital and snout, uppermost line sometimes joining its counterpart across snout tip, forming broad Ushape. Underside of head whitish to dusky, with five to eight narrow brown lines across branchiostegal membranes and chin, lines curving anteriorly over ventral midline. Breast dark brown, brown extending up to pectoral base. Pectoral base dark brown, usually intensified in centre, forming dark oval spot. Belly and lower sides pale whitish to whitish yellow. First dorsal fin with lower two-thirds dusky, with blackish area posteriorly; outer third of fin transparent to translucent whitish with narrow dusky brown margin. Dusky area of fin often with two rows of dark brown streaks and blotches, blotches alternating with fin rays; medium-sized black spot posteriorly. Second dorsal fin similar, fin mostly dusky with transparent to translucent whitish submarginal band and narrow brownish margin; dusky area of fin often with blotches and vertical streaks on membranes between rays; three to four dark brown blotches sometimes present along base of fin. Anal fin plain dusky brownish. Caudal fin with many rows of thin irregular dark brown to blackish lines and/ or rows of fine spots, none particularly enlarged at base of fin; on upper and lowermost edge, fin plain dusky or with blackish streaks on membranes aligned with fin rays. Pectoral fin translucent, dusky at bases of rays. Pelvic fins and fraenum plain dusky to brown. Comparisons Mugilogobius rivulus sp. novo is very similar in colour pattern to this species, but has a pair of dark spots at the caudal fin base, with dark spotting clustered near these spots on the caudal fin; it also has the first dorsal fin spine elongate and white (in both sexes), while M. rambaiae has a low first dorsal fin with no elongate spines, and the first spine is never the longest. In general appearance, M. rambaiae has a reticulate pattern with narrow dorsal saddles sometimes present, while M. rivulus sp. novo looks mottled and banded, with the reticulate scale margin pattern less obvious. The anteriormost nape scales are slightly larger than those posterior to them in many specimens of M. rambaiae, but these scales are all small and evenly sized in M. rivulus sp. novo Other species pairs (such as the allopatric chulae/wilsoni sp. novo and sympatric fasciatus sp. nov./tigrinus sp. nov.) recognised in this paper differ from each other in an similar manner: elongation of different dorsal fin spines and differences in coloration. Distribution Specimens are known only from Thailand, Burma, Malaysia, Singapore, Indonesia, Sri Lanka and Papua New Guinea (Figure 167). Ecology Axelrod et al. (1990: 865) indicate that this species can be kept in a freshwater aquarium, and they refer to it as the "Queen of Siam Goby". Remarks The paratypes (USNM 119647) and an unknown number of other specimens ("many") were kept in a glass jar from the 28th May, 1931, to 2 December, 1931, according to the label, with the type specimens (and see Smith 1945). Two female specimens of these paratypes are not Mugilogobius, but are Calamiana kabilia. The Papua New Guinea specimens (Figure 173) represent a considerable distribution gap, as the Kikori River flows into the Gulf of Papua. The species identified by Axelrod et al. (1990: 864) as Vaimosa rambaiae is a species of Rhinogobius. 174 H.K. Larson Figure 173 Mugilogobiu5 rambaiae, 27 mm SL, WAM P.3097S-Q07, Kikori River, Papua New Guinea. Mugilogobiu5 rexi sp. novo Figures 87, 174-176; Tables 5-8,28 "Gelbe Grundel": Kottelat 1989b: 683, figure 7. Material Examined Holotype Indonesia, Sulawesi: MZB 5949, 29.5 mm SL male, mouth of Batuopa River, about 3 km S of Timampu, Lake Towuti, M. Kottelat, 22 June 1988. 9; anal rays 1,7-9; pectoral rays 12-14; longitudinal scales 27-30, TRB 8-10; circumpeduncular scales 11-13; predorsal scales 11-19, scales on nape reaching nearly up to eyes or past preopercular edge; vertebrae modally 11+16; often with about six short rows of transverse papillae below the eye, transverse rows sometimes developed as part of longitudinal rows c and cp; yellow (male) or yellowish grey (female) colour when live; restricted to the freshwater Lakes Towuti and Mahalona, central Sulawesi, Indonesia. Paratypes Indonesia, Sulawesi: NTM 5.12706-003, 16(1531), rice padi at Watidi, about 5 km 5E of Timampu, Lake Towuti, H. Larson and R. Williams, 15 September 1989; NTM 5.12707-002, 32(13-31), Tominaga, Lake Towuti, H. Larson and R. Williams, 1989; CMK 6500, 9(11-24.5), south-west coast of Lake Mahalona, M. Kottelat, A. Werner, 16 March 1989; CMK 6490, 6(18-26), 4-7 km E of Timampu, Watidi, Lake Towuti, M. Kottelat, A. Werner, 15 March 1989; MZB 5940, 1O(15-22.5),same data as holotype; NTM 5.14702-001, 15(14.5-33), same data as holotype; CMK 6205, 18(13-32.5), same data as holotype. Other material examined (but not used in description) Indonesia, Sulawesi, Lake Towuti: CMK 6218, 1, 5 of Timampu; CMK 6267, 4, same locality as preceding; CMK 6250, 1, same as preceding; CMK 6476, 1, Telok Balaote; CMK 6471, 1, 5ungei Batuopa; CMK 9751,7, Tandjung 5ubalaote. Diagnosis A small Mugilogobius with second dorsal rays 1,8- Figure 174 Description Based on 45 specimens, 14.5-33.0 mm SL. An asterisk indicates counts of holotype (Figure 174). First dorsal VI*; second dorsal 1,8*-9 (mean 1,8*); anal 1,7-9 (mean 1,8*); pectoral rays 12-14 (mean 13*); segmented caudal rays 15-16* (mean 16); caudal ray pattern modally 9/7; branched caudal rays 6/4 to 8/6 (modally 6/5); unsegmented (procurrent) caudal rays 6/6 to 6/7; longitudinal scale count 27-30 (mean 29, 28 in holotype); TRB 8*-10 (mean 9); predorsal scale count 11-19 (mean 15*); circumpeduncular scales 11-13 (mean 12*). Gill rakers on outer face of first arch 2+6 to 3+7 (modally 3+6). Upper and lower few pectoral rays unbranched, often only central four to eight rays branched. Dorsal pterygiophore formula 3-12210 (in seven), 3-11310 (in one). Vertebrae 11+16 (in 10), 11+17 (in one). Neural spines of first vertebra narrow and pointed (in two), or first two spines slightly broader than remainder (in one). Metapterygoid low and narrow, with slender process forming distinct bridge to quadrate. Fifth ceratobranchial rather slender; moderate flange Mugilogobiu5 rexi n. sp., holotype, 29.5 mm SL, MZB 5949, mouth of Batuopa River, Lake Towuti, Sulawesi. Revision of Mugilogobius Table 28 175 Morphometries as percentages of SL or HL, as indieated, of Mugilogobius rexi sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. l. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbitall. in HL Pectoral l. in SL Pelvic l. in SL Caudal!. in SL Longest D1 spine in SL Holotype 31.5 57.0 55.9 no 13.6 23.7 12.5 30.1 26.9 36.6 28.0 22.0 22.0 26.8 11.8 Males Males Minimum Maximum 30.0 50.0 45.8 17.9 10.3 22.6 9.0 25.0 24.2 31.3 16.7 19.4 19.7 22.9 9.2 34.1 58.9 55.9 23.5 14.4 28.8 13.2 31.9 33.3 38.0 28.0 24.5 25.1 30.5 14.6 present ventrally. Two epurals (in 10). Three (in 11) anal pterygiophores before haemal spine of first caudal vertebra. Head and body compressed, more so posteriorly. Head compressed or rather square in cross-section, head depth often greater than width but sometimes equal to width; HL 2.9-3.3 (mean 3.1) in SL; profile pointed. Depth at posterior preopercular margin 1.6-1.9 (mean 1.8) in HL. Width at posterior preopercular margin 1.6-2.3 (mean 2.0) in HL. Mouth relatively small, usually terminal, oblique, forming angle of about 25-30° with body axis; jaws reaching from below anterior margin of eye to anterior half of eye. Upper jaw length 2.6-3.4 (mean 3.0 in females, 2.9 in males) in HL. Lips fleshy, narrow, smooth, usually with fine fimbriae on inner edges of both lips; lower lip mostly free, fused across narrow tip of jaw. Eyes not particularly large, placed high on head dorsolaterally, often forming part of dorsal profile; eye width 3.0-4.1 (mean 3.6) in HL. Snout oblique, bluntly rounded in dorsal view; snout profile usually quite pointed, 3.1-4.3 (mean 3.6) in HL. Interorbital moderate to relatively narrow, flat or slightly convex, 3.6-{).0 (mean 4.4) in HL. Usually without any fine fleshy villi on naked areas of head. Body slender, depth at anal origin 4.1-5.6 (mean 4.7) in SL. Caudal peduncle long, compressed, length 3.5-4.5 (mean 3.9) in SL. Caudal peduncle depth 7.6-11.2 (mean 8.7) in SL. First dorsal fin low and rounded, often not reaching second dorsal fin when depressed. Third, sometimes fourth, dorsal fin spine usually longest. Third dorsal spine 7.5-10.8 (mean 8.6) in SL in 7.8-10.0 (mean 8.8) in females; fourth dorsal spine maximum length 6.9-10.0 (mean 8.0) in males, 7.8-9.0 (mean 8.4) in females. Second dorsal fin low, not much higher anteriorly than posteriorly. Anal fin low, with posteriormost rays usually longer than Males Mean Females Minimum Females Maximum Females Mean 32.8 54.9 50.0 21.7 12.8 26.0 11.8 28.0 27.4 34.4 23.1 21.8 21.7 21.7 12.6 30.0 51.6 43.3 19.5 10.7 22.5 10.2 23.3 24.2 29.4 20.3 19.1 18.5 24.2 10.0 34.1 62.6 61.5 24.2 14.8 28.2 12.6 31.5 32.0 37.0 26.1 23.8 22.2 29.4 12.9 32.0 56.8 52.0 21.3 12.9 25.9 11.3 27.8 28.4 33.0 23.2 21.6 20.6 26.5 11.9 anterior rays. Depressed second dorsal and anal fin rays only reaching halfway along caudal peduncle. Pectoral fins narrow, roughly rectangular, 4.1-5.2 (mean 4.6) in SL. Pelvic fins long and oval, reaching nearly to anus in adults; fins reaching anus in small specimens, fin origin below pectoral base, length 4.0-5.1 (mean 4.6) in SL in males, 4.5-5.4 (mean 4.9) in females. Caudal fin rounded to slightly truncate, 3.3-4.4 (mean 3.7) in SL. No mental fold or fraenum. Anterior nostril in short tube at edge of preorbital, directed forward and downward over upper lip. Posterior nostril rounded, with low rim, midway between eye and tip of snout, but usually closer to eye. Gill opening extending forward nearly to below rear preopercular margin. Gill rakers on outer face of first arch short, stubby, with very fine spiny papillae sometimes present; rakers on inner face of first arch similar in size to those of outer face, with very fine papillae at tips. Outer rakers on second arch short, stout; rakers on inner face of second arch slightly longer, with fine spiny papillae. Pectoral girdle smooth in all specimens, with no knobs or flange. Tongue blunt to concave. Teeth in outer row of upper jaw enlarged, caniniform, widely spaced, followed by four rows of smaller, sharp conical teeth across front; two rows at sides. Outermost row of teeth in lower jaw largest, but not particularly enlarged, outer row widely spaced, followed by three rows smaller teeth across front; one or two rows at sides. Teeth similar in males and females. Predorsal scales cycloid, small and even, with none enlarged, scales reaching forward to above preopercular margin or further, sometimes to close behind eyes. Operculum with cycloid scales, lower third to half of opercle unscaled. Cheek naked. Pectoral base scales cycloid, sometimes scattered over base. Prepelvic area scales cycloid, breast H.K. Larson 176 .... " '" I, B .. " a , Ｚｾ ..... ,. .. : -:-:-:- ｾＺ ....... . ＬＮｾ /::: ". -)' ｾＮ r ..... \ .' J/ " ••• ｾＮＬ : '/" -•.•.:..: "1.:.." '. ｾＬＯＮｾ "\ ) I " 1\ '. '. '----' c Figure 175 MugiIogobius rexi n. sp., papillae pattern. Paratypes. A, CMK 6205, lateral view; scales omitted; B, CMK 6500, dorsal view of interorbital area; C, CMK 6500, ventral view of chin area. Band C not drawn to scale. Scale bar = 1 mm. usually fully scaled. Belly with ctenoid scales anteriorly at least, with anterior half and up to entire belly with ctenoid scales; scales next to anus always cycloid, Genital papilla in male slender, elongate and flattened, narrowing toward tip; papilla in female short, rounded and stout, with no lobes at tip. Head pores absent as in all Mugilogobius species. Sensory papillae on head variable, could be referred to either as an abbreviate transverse pattern or a proliferated longitudinal pattern (Figure 175). Five to six short rows of transverse papillae under eye comprising rows c and cp; rows sometimes present as partly developed longitudinal rows. Row p around eye sometimes including several short transverse rows. One s row on each side of snout just above lip; longitudinal r row at level of posterior nostril. Two f rows on chin oriented longitudinally, on either side of chin. Coloration of fresh material From colour slides. Entire fish light yellow, with scale margins narrowly outlined in light yellowish brown. Nape, top of head and area behind eye with scattered mottling of light yellowish brown; dusky brown mark present between eye and upper lip. Unpaired fins translucent with dusky fin rays; first dorsal fin with narrow greyish brown submarginal line, second dorsal fin with greyish brown stripe along centre; other fins plain, pectorals and pelvics translucent. Iris golden brown above, gold below. From my field notes. Live fish translucent pale yellow, often with vivid yellow "faces" and vivid yellow on underside of head and on branchiostegal rays. Dorsal midline with groups of very fine golden speckles arranged around faintly dusky blotches. Scale margins on upper back outlined with dusky greyish brown. Specimens from NTM S.12707-002 with lower two-thirds of first dorsal fin plain yellow with dark line dividing yellow portion from transparent outer third of fin; second dorsal fin plain yellow with outer third transparent, narrow dusky line separating transparent area from rest of fin, second narrow dusky line running just above bases of rays; anal fin yellow with pinkish tinge and outer third of fin transparent with narrow dusky line at base of transparent portion; caudal fin translucent pale yellow. Most fish dull translucent yellowish or very light brown mottled with brown, and dark scale margins. Kottelat (1989b) said that these fish" ... exhibit a very conspicuous sexual dimorphism: females are grey or brown, while sexually active males are intense yellow". Coloration of preserved material Head and body dusky yellowish to yellowish grey, paler ventrally (Figures 174, 176). Most scale margins on head and body narrowly outlined with light brown; broader light brown sub-marginal line usually present as vertical line or rounded spot in centre of each scale, latter pattern usually present on posterior half of body. Top of head, nape and upper part of body with scattered brown spots and irregular mottling; mottling often forming short zigzag lines. In some specimens, about seven brown blotches or saddles crossing dorsal midline. Interorbital and snout with light brown diffuse vermiculation and short streaks; oblique, short brown streak at upper rear edge of eye and one or two short brown streaks or blotches on preorbital. Side of head with one or more short diffuse lines or Revision of Mugilogobius Figure 176 177 Mugilogobius rexi sp. nov., paratypes, CMK 6490, 32 and 28.5 mm SL, Watidi, Sulawesi. From colour slide by Maurice Kottelat. blotches across cheek, broken-up vermiculation and spotting on opercle. Lips and underside of head plain light brown. Breast pale yellowish, often with light brown blotch just before pelvics. Peritoneum brown, pale ventrally. First dorsal fin translucent with light brown fin rays and diffuse light brown margin, and series of narrow brown horizontal streaks along proximal third of fin. 5econd dorsal fin translucent with narrow light brown margin and series of oblique to horizontal streaks along centre of fin; fin rays brownish, darker at bases. Anal fin plain dusky to light brown, darker posteriorly. Caudal fin translucent with light brown fin rays. Pectoral fin translucent. Pelvics translucent to brown, fraenum unpigmented. Comparisons This species most closely resembles Mugilogobius lepidotus sp. nov., also from Lake Towuti, in that both species are small and slender, and have low scale counts in comparison with M. amadi or M. sarasinorum (also 5ulawesi lake endemics). Mugilogobius rexi sp. novo and M. lepidotus sp. novo both usually have narrow, pointed neural spines, with very little broadening of the tips found in M. amadi. Distribution Restricted to Lakes Mahalona and Towuti, central 5ulawesi (Figure 87). of at least a metre's depth, over silty substrate, particularly near clumps of aquatic plants and the freshwater mangrove Barringtonia racemosa. Kottelat (1989b; his translation) gives some information about this species: ... 'yellow goby' exhibit a very conspicuous sexual dimorphism: females are grey or brown, while sexually active males are intense yellow ... Females are also slightly smaller than males. They were observed almost everywhere in the lake, along the shores, but were much more common among submerged vegetation, especially flooded Pandanus bushes. Etymology Named for Rex Williams, technical officer at the Museum and Art Gallery of the Northern Territory, Darwin, in recognition of his careful work and commitment to the NTM fish collection, and who deserves to have a goby named after him. Mugilogobius rivulus sp. novo Figures 167, 177-181; Tables 5-8, 29 Material Examined Holotype Australia: Northern Territory: NTM 5.14065-001, 27.5 mm 5L male, drain at Leanyer 5wamp, C. Jones, 30 September 1994. Paratypes Remarks These gobies hover above the substrate in a manner reminiscent of the marine gobiine genus Arnblygobius Bleeker, in a slightly diagonal position, with the caudal fin curved to one side (personal observation). In Lake Towuti, they inhabited water Australia: Northern Territory: NTM 5.12740-001, 2(28-30.5), Darwin River, D. Wilson, February 1990; NTM 5.12741-001, 1(33), Berry 5prings, K. Martin, R. Wells, J. Kum Jew, 5 November 1977; NTM 5.14305-001, 4(18.5-28.5), drain under Gilruth Avenue, Darwin, D. Wilson, 22 January 1990; NTM --------------------- ._- 178 H.K. Larson 5.14306-001, 1(45), Leanyer Swamp, C. Jones, October 1991; NTM 5.14304-001, 1(41), Leanyer Swamp, C. Jones, 23 August 1994; NTM 5.11483001, 10(13-21), Leader's Creek, Gunn Point, D. Hoese, 5. Reader, D. Beechey, 14 September 1984; AM5 1.24690-001, 25(11.5-30.5), same data as preceding; AM5 1.24691-002, 14(10-17), pool at Leader's Creek, Gunn Point, D. Hoese, 5. Reader, 14 September 1984; NTM 5.13744-022, 35(11-21), Brooking's Creek, Howard Springs, R. Williams, 30 June 1993. Other material eXilmined (but not used in description) Australia: Northern Territory: NTM 5.14291-002, 6, Leader's Creek, Gunn Point; NTM 5.14289-002, 19, Leader's Creek, Gunn Point; NTM 5.14290-003, 138, Leader's Creek, Gunn Point; AM5 1.23927-009, 1, Blackmore River. Diagnosis A robust Mugilogobius with second dorsal rays 1,7-8; anal rays 1,7-9; pectoral rays 14-16; longitudinal scales 29-33; TRB 9-12; circumpeduncular scales 12-13; predorsal scales 1219, small, reaching to above preopercular margin; scales on body mostly ctenoid; first spine of dorsal fin white and always longest, often filamentous; body with distinct dark oblique shoulder bar, scale margins rimmed with narrow dark line forming reticulate pattern usually overlaid by narrow dark bars and dorsal saddles, two dark spots at caudal base; known only from the Northern Territory, Australia. Description Based on 30 specimens, 15-45 mm 5L. An asterisk indicates counts of holotype (Figure 177). First dorsal V (in two), VI* (in 28); second dorsal 1,7-8 (mean 1,8*); anal 1,7-9 (mean 1,8*), pectoral rays 14-16 (mean 15*), segmented caudal rays always 16*; caudal ray pattern 7/5 to 8/7* (modally 8/6); branched caudal rays 12-15* (mean 14); unsegmented (procurrent) caudal rays 8/8 to 9/8; longitudinal scale count 29-33 (mean 31, 32 in holotype); TRB 9-12 (mean 10, 11 in holotype); Figure 177 predorsal scale count 12-19 (mean 17, 16 in holotype); circumpeduncular scales 12-13* (mean 12). Gill rakers on outer face of first arch 3+6* to 4+8 (modally 3+7). Pterygiophore formula 3-12210 (in two). Vertebrae 10+16* (in two). Neural spine of second vertebra expanded or bifid at tip (in two). Two epurals (in two). Two (in one) or three (in one) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid broad, but not greatly expanded dorsally. Body compressed, more rounded anteriorly, HL 3.1-3.7 (mean 3.3) in 5L. Head wider than deep, somewhat flattened in adults. Depth at posterior preopercular margin 1.5-1.9 (mean 1.7) in HL. Width at posterior preopercular margin 1.2-1.6 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of about 20-25° with body axis; jaws generally reaching to below posterior half of eye in males and to below mid- to anterior half of eye in females (not quite reaching mid-eye in holotype). Lips smooth or with fleshy fimbriae present mostly on inner edge of upper lip; lower lip free at sides, fused across front. Upper jaw 2.0-3.2 (mean 2.9 in females, 2.6 in males) in HL. Eyes lateral, high on head, forming part of dorsal profile, 3.1-4.6 (mean 3.6) in HL. Snout rounded, 3.1-4.4 (mean 3.7) in HL. Interorbital broad, flat, 2.5-4.8 (mean 3.3) in HL. Top of head from in front of scaled nape forward to snout tip often covered with fine villi (sparse in holotype). Body depth at anal origin 4.4--6.2 (mean 5.4) in 5L. Caudal peduncle compressed, length 3.5-4.5 (mean 4.1) in 5L. Caudal peduncle depth 6.4-8.3 (mean 7.3) in 5L. First dorsal fin generally low, tips of spines other than first free but not filamentous, second to fifth spines nearly always falling short of second dorsal fin origin when depressed. First dorsal spine always longest, often filamentous, 3.1-7.9 (mean 5.6 in females, 4.8 in males) in 5L. Second dorsal and anal fins low, fins somewhat rounded anteriorly, pointed posteriorly, with posterior rays not much longer than anteriormost; last dorsal and anal rays falling well short of upper caudal rays when depressed (except in 45 mm 5L specimen). Pectoral fin rounded to oval, central rays longest, 3.7-5.1 (mean Mugilogobius rivulus sp. nov., holotype, 28 mm SL, NTM 5.10465-001, Leanyer Swamp, Northern Territory. Revision of Mugilogobius Table 29 179 Morphometries as percentages of SL or HL, as indicated, of Mugilogobius rivulus sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. 1. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital 1. in HL Pectoral 1. in SL Pelvic 1. in SL Caudal 1. in SL Longest D1 spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 29.5 60.5 76.5 17.1 11.6 24.4 13.5 27.2 24.7 39.5 30.9 22.9 19.6 27.3 22.5 29.5 55.4 69.2 16.2 9.5 22.1 12.9 24.2 23.3 32.0 25.4 20.0 17.1 26.4 16.5 32.6 60.9 80.5 22.9 15.1 26.7 15.6 32.3 32.0 50.4 40.6 27.4 21.8 31.4 32.4 30.5 58.6 72.7 18.7 12.8 24.7 13.9 27.7 27.4 38.7 31.8 23.4 20.4 29.0 21.9 27.2 52.1 62.5 16.2 10.8 23.2 12.1 22.9 21.7 31.1 20.8 19.5 18.9 27.2 12.7 32.0 65.4 78.7 21.4 17.1 28.6 14.5 28.9 31.4 41.7 39.7 24.9 23.3 30.9 25.9 30.5 58.5 71.9 18.8 13.3 25.3 13.6 26.3 28.6 34.4 29.9 23.0 20.9 29.2 18.2 4.4) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, reaching at least halfway to anus, 4.3-5.8 (mean 4.9) in SL. Caudal fin rounded to oval, 3.2-3.8 (mean 3.5) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, placed just behind upper lip, tube short, oriented down and forward, preorbital curved forward to accommodate nostril. Posterior nostril round to oval, placed near anterior centre margin of eye. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in two) or with irregular fleshy ridge or raised knobby flange (in 16), or with distinct fleshy knobs and flaps (in seven). Gill rakers on outer face of first arch very short knobs, longest raker at angle of arch; rakers on inner face of first arch rounded, stubby, with fine spiny papillae at tip; inner rakers on other arches not much longer than first arch inner rakers (four rakers ossified in cleared and stained specimen). Tongue tip blunt to concave. Outer teeth in upper jaw largest, stout and curved, behind this row, three to four rows of small sharp inward-pointing teeth; one or two rows at side of jaw, teeth generally rather smaller in females. Lower jaw with three to four rows of small pointed teeth across front, outermost row often oriented upright, inner rows pointing inward; no teeth particularly enlarged; one to two rows of teeth at side of jaw. Predorsal scales small, usually evenly sized, reaching forward to about halfway between preopercular margin and rear of eyes. Operculum with small cycloid scales on upper half to twothirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1/4 to 1/3 of belly), rest of scales cycloid, sometimes entire belly with cycloid scales. Ctenoid scales on side of body extending forward in wedge up to behind pectoral fin. Genital papilla in male elongate, flattened, tip rounded to slightly expanded; in female, papilla short and rounded. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 178. Papilla row e ending just after turning up onto rear edge of preopercle; three s rows on snout (sometimes central row with one or two papillae); each f row behind mandibular symphysis with four to five papillae arrayed in straight line; some specimens with dorsalmost end of row a bending slightly posteriorly. Coloration of fresh material From notes based on live specimen. Body light brownish grey with margins of scales on sides of body outlined in dark greyish brown. Narrow dark greyish brown saddles crossing back and dark '. P.@/-,...CJ ,. :, , C> .. : '. '- . .. . " c cp' . " Figure 178 "", Mugilogobius rivulus, papillae pattern. NTM S.14304-001, Leanyer Swamp, Darwin, NT. Scalation suggested only. Scale bar = 1 mm. 180 H.K. Larson greyish brown markings at base of tail fin. Lower half of head brown with rounded light brownish grey spots on cheeks and opercles; spots on cheek dull greyish yellow with lower half of head plain brownish grey. Two dully iridescent pale yellow spots on anterior part of opercle and dark brown to blackish oblique line through eye, extending from rear of mouth up to nape. Underside of head with dark brown narrow lines. First dorsal fin dusky brown with brown yellow margin, and pinkish white submarginal band; first spine (elongate) pale whitish to dull yellow (colour can be changed at will). Second dorsal fin dusky brownish with pinkish white submarginal band. Base of caudal fin dull yellowish with brown spots. Fin rays all brownish, membranes dusky. Coloration of preserved material Head and body pale whitish yellow to pale greyish yellow (depending on state of preservation), most distinctive marking being broad dark brown oblique shoulder bar or oval blotch extending from behind pectoral base toward predorsal area (bar not extending on nape) (Figures 179-180). Four to eight indistinct oblique brown saddles crossing back; six to eight vertical blotches or bars on upper side of body, variably developed in intensity among individuals, first blotch often oval, dark brown and similar in appearance to oblique shoulder bar; and two oval to triangular brown spots on base of caudal fin. Each scale of back and sides with short vertical to gently curved brown line near centre of scale; curved lines more pronounced on scales on posterior half of body and closer to scale margin; scales usually narrowly edged with grey to brown. Small brown blotches and X-shaped markings often present along midside of body, especially posteriorly. Top of head with indistinct blotches or marbling, usually several irregular lines joining across interorbital space, and one brown line extending onto nape from upper rear margin of eye; interorbital and snout plain dusky or with short vermiculate brown lines. Side of head with dark brown lines forming reticulate network of pale interspaces, especially prominent on cheek and opercle; two or three brown lines extending from front of eye onto preorbital and snout. Underside of head whitish to dusky, with seven narrow brown lines crossing branchiostegal membranes and chin, lines curving anteriorly over ventral midline (Figure 181); occasionally, indistinct eighth line crossing breast just behind branchiostegal membrane insertion. Breast usually dark brown, brown extending up to pectoral base. Pectoral base dark brown, with pale line crossing fin ray bases. Belly and lower sides pale whitish to whitish yellow. First dorsal fin with lower two-thirds dusky, with blackish area posteriorly, sometimes forming distinct black spot; outer third of fin transparent to translucent whitish with narrow dusky brown margin, fin spines often whitish, first dorsal spine nearly always white to translucent whitish. Second dorsal fin mostly dusky with transparent to translucent whitish submarginal band and narrow Figure 179 Mugilogobius rivullls sp. nov., female paratype, 28 mm 5L; NTM 5.12740-001, Darwin River, NT. Figure 180 MlIgilogobius rivullls sp. nov., AM5 1.24690-001, Leader's Creek, Gunn Point, NT. From colour slide by Doug Hoese. Revision of Mugilogobius Figure 181 181 Mugilogobius rivulus sp. nov., underside of head. AMS 1.24690-001, Leader's Creek, Gunn Point, NT. From colour slide by Doug Hoese. brownish margin; dusky area of fin often with blotches and vertical streaks on membranes between rays. Anal fin plain dusky brownish, margin whitish or translucent. Caudal fin plain dusky with two oval to triangular spots at base (spots usually small, but sometimes as large as pupil), scattered brown spots on centre of fin near base, brownish streak often present over procurrent rays. Pectoral fin dusky, translucent to white band at fin ray bases. Pelvic fins plain dusky to brown, fraenum usually translucent. Comparisons This species is very similar to M. rambaiae, and differs mainly in coloration and in always having the first dorsal fin spine longest. Mugilogobius rambaiae has a low first dorsal fin with no elongate spines (the first spine is always shorter than the third or fourth). In many specimens of M. rambaiae, the anteriormost nape scales are slightly larger than those posterior to them but these scales are all small and evenly sized in M. rivulus sp. novo The two species are very similar in coloration, but show some differences. Mugilogobius rivulus sp. novo has two dark spots at the caudal fin base, with dark spotting clustered near these spots on the fin, with most of the fin plain dusky. In M. rambaiae the caudal fin has many vertically aligned rows of narrow irregular dark brown to blackish lines and/ or rows of fine spots and streaks, and the lower fin margin is sometimes plain. Distribution Specimens are known so far only from the Northern Territory, Australia (Figure 167). Ecology This species is usually found in shallow, relatively still, brackish to freshwater creek habitats. It also appears be fairly tolerant of what looks, to an observer, to be sub-optimal habitat (storm drains). This species has been collected with M. platystomus in Leader's Creek, Gunn Point, NT, in a non-flowing (late dry season), brackish (19%0 salinity) Melaleuca forest creek, among dense leaf litter. Recently collected specimens from fresh water creeks on Melville Island, NT (specimens not examined for this paper), were nearly always found associated with drifts of leaf litter. Their dark live colour matched the dark brown dead leaves, as did an atyid shrimp species also associated with these leaf drifts. Etymology From the Latin rivulus, meaning rill or small brook, in reference to the habitats in which this species may be found. Mugilogobius sarasinorum (Boulenger, 1897) Figures 16F, 87,182-185; Tables 5-8, 30 Gobius sarasinorum Boulenger, 1897: 427, plate 28, figure 1 (Lake Posso); Weber 1913: 212; Kottelat and Sutter 1988: 56. Tamanka sarasinorum: Koumans 1953: 158-159; Whitten et al., 1988: 295; Larson and Kottelat 1992: 233; Kottelat et al., 1993: 154, plate 73. Mugilogobius sarasinorum: Kottelat et al., 1993: 154 (as synonym). H.K. Larson 182 Material Examined Lectotype Indoniesia, Sulawesi: NMBA 1844, 1(62), Lake Posso. Paralectotypes Indonesia: Sulawesi: NMBA 1843, 1(51), NMBA 1845, 1(56), NMBA 2731, 1(57); BMNH 1897.3.8.1-4, 4(42-63), Lake Posso, Sarasin. RMNH 14364, 1(49.5); NTM S.12698-003, 42(16-47), Lake Poso, 7 September 1989, H. Larson and R. Williams; ZMA 113.628, 2(39-43), Poso Lake, E.C. Abendanon, May 1910; NTM S.12700-002, 4(24-46), Tentena, Lake Poso, H. Larson and R. Williams, 10 September 1989; ZSI/CMK 6237, 23(17-33.5), E shore of Lake Poso between Tentena and Peura, M. Kottelat, 25 June 1988. Other material examined (but not used in description) All from Lake Poso. Paralectotype, NMBA 1846, 1. NMBA 2710, 1; NMBA 2732, 1; NMBA 2730, 1; NMBA 2733, 1. Diagnosis A rather deep-bodied Mugilogobius with somewhat compressed head, pointed profile, second dorsal and anal rays 1,8-9; pectoral rays 1518; predorsal scales 17-25; lateral scales 38-51; TRB 13-17; circumpeduncular scales 16-22; predorsal scales 17-25, small, and extending forward almost to behind eyes; sensory papillae on head occasionally forming multiple rows on cheek, males dark brown to greyish brown, females lighter with narrow bars and moUling on body; endemic to freshwater Lake Poso, central Sulawesi, Indonesia. Description Based on 30 specimens, 17-63 mm SL. An asterisk indicates counts of lectotype (Figure 182). First dorsal VI*; second dorsal 1,8*-9 (mean 1,8); anal 1,8*-9 (mean 1,8); pectoral rays 15-18 (mean 17, 16 in lectotype); segmented caudal rays 15-17 (mean 16*); caudal ray pattern modally 9/7; branched caudal rays 6/6 to 8/7 (mean 7/6); Figure 182 unsegmented (procurrent) caudal rays 8/8 to 8/9; longitudinal scale count 38-51 (43 in lectotype, mean 46); TRB 13-17* (mean 15); predorsal scales 17-25 (20 in lectotype, mean 21); circumpeduncular scales 16-22 (17 in lectotype, mean 18). Gill rakers on outer face of first arch 0+4 to 3+9 (modally 2+8). Dorsal pterygiophore formula 3-12210 (in five). Vertebrae 11+15 (in six). Neural spines of first vertebra stout, pointed (in four) and may be bifid at tip (in one). Metapterygoid low and narrow, with slender process forming distinct bridge to quadrate. Fifth ceratobranchial rather slender; tall, moderate flange present ventrally. Two epurals (in five). Three (in six) anal pterygiophores before haemal spine of first caudal vertebra. Head and body rather compressed, more so posteriorly. Head cylindrical to nearly triangular in cross-section (apex dorsally), head width usually greater than depth, although depth sometimes equal to width, HL 2.7-3.2 (mean 3.0) in SL; profile rounded to rather pointed. Depth at posterior preopercular margin 1.5-2.0 (mean 1.8) in HL. Width at posterior preopercular margin 1.5-1.8 (mean 1.6) in HL. Mouth relatively small, terminal to slightly subterminal (upper jaw anterior to lower), very slightly oblique, forming angle of about 10-20° with body axis; jaws reaching from below anterior margin of eye to anterior half of eye. Upper jaw 2.8-3.4 (mean 3.1 in females, 3.0 in males) in HL. Lips fleshy, narrow, smooth, with fine fimbriae on inner edges of both lips; lower lip mostly free, fused across tip of jaw. Eyes not particularly large, placed high on head dorsolaterally, sometimes forming part of dorsal profile; eye width 3.1-5.1 (mean 3.9) in HL. Snout oblique, rounded in dorsal view; snout profile usually rather pointed, sometimes rounded; lectotype with snout with slight bump in centre formed by ascending premaxillary process; length 3.1-4.1 (mean 3.6) in HL. Interorbital moderate to relatively narrow, flat, 3.6-6.5 (mean 5.3) in HL. Fine scattered villi sometimes present on naked areas on top of head (interorbital, upper part and side of snout). Body depth at anal origin 4.0-7.7 (mean 5.1) in SL. Caudal peduncle compressed, Mugilogobius sarasinorum. Lectotype of Gobius sarasinorum Boulenger, 64 mm SL, NMBA 1844, Lake Poso, Sulawesi. Revision of Mugilogobius Table 30 183 Morphometries as percentages of SL or HL, as indicated, of Mugilogobius sarasinorum (Boulenger, 1897). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. \. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital\. in HL Pectoral\. in SL Pelvic \. in SL Caudal\. in SL Longest 01 spine in SL Lectotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 33.5 67.3 66.3 25.0 15.3 24.4 31.2 50.6 57.6 17.4 9.8 23.1 10.0 24.5 19.4 31.1 15.4 21.4 17.7 23.3 12.6 37.0 62.9 68.7 24.0 16.2 28.0 13.5 31.9 32.5 35.9 27.5 25.3 23.0 29.7 16.7 33.2 56.0 62.1 20.0 12.3 25.8 11.0 28.7 26.4 33.1 19.7 22.8 21.3 26.5 15.6 32.6 50.0 55.2 13.0 10.0 22.1 10.0 25.9 19.7 29.5 15.5 19.4 18.8 23.5 11.3 34.4 67.3 66.3 25.0 15.9 28.4 13.2 29.6 32.8 33.8 23.1 25.3 23.7 29.2 16.9 33.5 56.9 61.3 20.3 12.8 25.6 11.0 27.6 26.1 32.1 18.9 22.1 20.9 25.9 13.8 27.4 19.7 31.7 19.2 25.3 21.9 23.7 16.9 length 3.5-4.5 (mean 3.9) in SL. Caudal peduncle depth 7.4-10.0 (mean 9.2) in SL. First dorsal fin rounded, low, second to fourth spine longest, usually third in females, third or fourth in males; fin usually reaching second dorsal origin when depressed but often falling short of fin origin in small females. Second dorsal spine length 7.3-8.2 (mean 7.6). Third dorsal spine length 5.9-8.9 (mean 7.3) in SL. Fourth dorsal spine length 6.0-6.8 (mean 6.4) in SL. Second dorsal fin similar in height to first dorsal fin; posteriormost rays not much longer than anteriormost; rays falling well short of caudal fin base when depressed. Anal fin similar to second dorsal fin, posteriormost rays often longest, falling short of caudal fin base when depressed. Pectoral fin oval, central rays longest, 3.9-5.1 (mean 4.5) in SL; all rays branched but for uppermost. Pelvic fins large, oval, rays usually reaching anus, 4.2-5.6 (mean 4.8) in SL. Caudal fin moderate in size, truncate to rounded posteriorly, 3.4-4.3 (mean 3.8) in SL. Chin usually smooth, without mental fraenum; sometimes row i running along behind lower lip anterior to row I, with flesh between rows inflated (Figure 183). Anterior nostril in short tube, placed at edge of preorbital, usually pointing down and forward over upper lip, preorbital slightly curved outward near nostril. Posterior nostril oval, sometimes with slightly raised rim, placed nearly halfway between preorbital edge and eye, usually closer to eye. Gill opening slightly oblique, usually extendin.g forward to under opercle. Inner edge of pectoral girdle with small rounded knobs or bumps (in 21), fleshy irregular flange (in seven) or smooth (in one). Gill rakers sometimes reduced to low bumps or absent. Outer gill rakers short; outer rakers on third and fourth arches slightly longer; inner rakers usually rather longer than outer rakers. Tongue tip blunt or concave. Outermost teeth in upper jaw largest, stout, curved and sharp, largest teeth across front; behind this row, two to five rows of small sharp curved teeth; usually only two to three rows of teeth at sides of jaw. Lower jaw with three to six rows of sharp curved teeth across front, two to three rows at sides; outermost row teeth often largest and stoutest but all teeth generally fairly evenly sized, oriented nearly upright; Figure 183 Mugilogobius sarasinorum, papillae pattern. A, RMNH 14364, lateral view; B, CMK 6237, ventral view of chin area (not to scale). Scales omitted. Scale bar = 1 mm. r--------------------------------------------- 184 H.K. Larson innermost teeth evenly sized, curved, often pointing posteriorly. Teeth similar in males and females. Predorsal scales cycloid, evenly sized, usually reaching forward to close behind or nearly to eyes. Operculum covered with small cycloid scales on upper two-thirds. Cheek always naked. Pectoral base covered with small cycloid scales. Prepelvic area covered with small cycloid scales. Belly scales cycloid, with patch of ctenoid scales anteriorly, close to base of pelvics; ctenoid scales extending at least halfway down belly, but not reaching anus. Ctenoid scales on side of body extending forward up to behind pectoral base. Genital papilla in male slender, quite elongate and flattened, narrowing toward tip; papilla in female rounded, bulbous and stout, shallow groove sometimes present along ventral surface. Head pores absent, as in all Mugilogobius species. Sensory papillae on head in longitudinal pattern, as in Figure 183. Rows b, d and e sometimes doubled or proliferated by extra papillae arranged in alternating pattern on either side of row (doubled or proliferated rows usually visible in larger specimens). Three s rows present, each consisting of three or more papillae. Mental f rows each of four or five papillae, or row continuous across chin; rows parallel to row i, sometimes extending anterior to rowf Coloration of fresh material From colour prints and slides of captive specimen. Head and body dull yellow to yellowish grey, with slightly brighter yellow patches dorsally, most prominent markings being five yellowish brown rounded to oblong blotches evenly spaced along midside of body (last spot at caudal base), blotches about equal to eye in size; remainder of upper half of head and body with scattered yellowish brown irregular blotches and moUling, about nine variably shaped narrow saddles or blotches crossing dorsal midline. Abdominal area plain pale pinkish brown; elsewhere plain greyish brown, lips yellowish. Opercle with bluish white iridescent patch in centre, few irregular brown lines and vermiculations over patch. Top of head and snout with light yellowish brown vermiculation; one short, oblique, relatively Figure 184 dark brown mark just behind upper rear corner of eye. Iris dull gold, with brown shading dorsally. Unpaired fins translucent pale bluish white, with dull yellow mottling and shading toward bases. Second dorsal fin with narrow, faint whitish margin and scattered brown spots along bases of fin rays. Anal fin with basal halves of fin rays brownish. Caudal fin with yellowish brown, vertically oriented oblong blotch across bases of fin rays, and smaller, round yellowish brown spot just above blotch; blotch dull whitish colour distally; vertically oriented rows of small translucent yellowish or yellowish brown spots on proximal two-thirds of fin, fin rays same colour or slightly darker. Pectoral fin translucent with fin rays very pale brownish. Pelvic fins translucent dull yellowish. My field notes on M. sarasinorum caught at night indicate that they were translucent with: ...about 6 oval 'clear brown' blotches along sides surrounded by whitish speckling. Fin margins pale blue, to whitish. Pelvics of some are dusky, with blue edges. Iris pale gold. Back and upper sides marbled with 'clear brown' and whitish speckles in between, and narrow bar from above pectoral base crossing nape to above other pectoral base. Kottelat et al. (1993) refer to males being almost black and females grey with dark narrow bars. Boulenger (1897) described his (live or freshly dead?) specimens as "Yellowish brown to dark brown above, without markings; fins brown to blackish; ventrals sometimes whitish". Coloration of preserved material Head and body plain light brown to dark brown, darker specimens always males, pale specimens may be either sex; darker dorsally and lighter brown ventrally (Figures 184-185). Dorsal surface of head, nape and upper part of body with indistinct greyish brown to brown small splotches and mottling; mottling sometimes forming irregular line along midside of body, or consisting of two staggered series of indistinct brown rounded spots. Some specimens with about five to seven indistinct brown narrow bars or rounded spots crossing dorsal midline; some females with four or five thin irregular bars on anterior half of body (extensions of mid-dorsal blotches). Head Mugilogobius sarasinorum, 50 mm SL, RMNH 14364, Lake Poso, Sulawesi. Revision of Mugilogobius Figure 185 185 Mugilogobius sarasinorum, CMK 6237, 28 and 34 mm SL, Lake Poso, Sulawesi. From colour slide by Maurice Kottelat. with indistinct scattered small brown spots and speckles sometimes forming two faint stripes on cheek below eye. Opercle and top of head with slightly more distinct brown spotting and vermiculation. All fins plain translucent light brown to dark brown, almost black in dark males, pectoral fins usually paler than others. Both dorsal fins with very narrow marginal white or translucent band; no black spot on rear part of first dorsal fin. Anal fin plain dark brown. Caudal with translucent to whitish rear margin, entire fin plain brown or with vertically aligned rows of brown spots, spots most pronounced on proximal half of fin. Pelvics translucent to dusky brown, rear margin may be translucent. Peritoneum speckled dusky brown, fading to pale belly. A colour illustration of recently dead specimens is in Kottelat et al. (1993: plate 73). Comparisons This species is closest in appearance to a very small M. amadi, which also lives in Lake Poso (see Comparisons under that species). Mugilogobius amadi has a protruding lower jaw, slender head and body, and plain dark brown colour in both sexes. gravel and large limestone rocks. The local people called them bungu or gabus. They also used the latter name for Ophicephalus and anything vaguely gobioid in appearance. One specimen was examined for gut morphology (the usual short-looped form was present); the gut contained the remains of a small fish and insect larvae (possibly chironomids). Five specimens examined for meristics were parasitised by one or more copepods. Remarks Boulenger (1897) stated he had 14 specimens, of which eight have been examined for this study. The whereabouts of the remaining six is unknown, and they are not referred to in Kottelat and Sutter (1988). The proliferated papillae in this species resemble the arrangement found in gobiines such as Acentrogobius, Glossogobius, Favonigobius and Oplopomus (Hoese, 1983; Gill, 1994). Mugilogobius stigmaticus (De Vis, 1884) Figures 167, 186-188; Tables 5-8, 31 Gobius stigmaticus De Vis, 1884: 686 (Moreton Bay, Queensland). Distribution Restricted to Lake Poso, central Sulawesi, Indonesia (Figure 87). Ecology This species could be observed moving around, at night, in silty sand shallows in front of the Pamona Indah Hotel at Tentena, at the north end of Lake Poso; they were not found there during the day. Usually the species was caught in open areas on a mud and gravel substrate, with some aquatic plants present, but were also found on banks of steep Mugilogobius devisi McCulloch and Ogilby, 1919: 223-224, figure 2 (unnecessary replacement name for Gobius stigmaticus De Vis, 1884); McCulloch 1922: 83; McCulloch 1929: 376; Koumans 1940: 160. Ellogobius stigmaticus: Whitley, 1933: 92-93; Whitley 1937: 18. Mugilogobius stigmaticus: Whitley 1930: 123; Koumans 1940: 206-207; Bell et al., 1984: 37; Morton et al., 1987: 222; Gee and Gee 1991: 19, 21-24. H.K. Larson 186 Material Examined Lectotype of Gobius stigmaticus Australia: Queensland: AMS 1.358, 1(37), Moreton Bay, Queensland Museum. Paralectotype of Gobius stigmaticus Australia: Queensland: AMS 1.40439-001, 1(39), same data as lectotype. Other Material Australia: Queensland: QM 1.14069, 21(14-39), William Parade, Yeronga, R.J. MacKay, J. Johnson and R. Leggett, 30 March 1977; QM 1.5021, 1(30), Pinkenbah, Brisbane; AMS 1.22904-001, 1(30.5), Calliope River, Gladstone, 1 February 1978; AMS 1.17157-003, 1(18.5), Ross River; QM 1.7002, 3(25.532), Chelmer, Brisbane River; QM 1.9651, 1(39), Clontarf, Moreton Bay, D. Rayment, 3 September 1968; QM 1.25238, 1(29), Boggy Creek, Brisbane River mouth, J. Johnson, J. Short, P. Lawless, 12 July 1988; AMS 1.31627-002, 5(17-26), Boggy Creek, Brisbane River; AMS 1.6213, 1(27), Caloundra; QM 1.25225, 11(16.5-29.5), Aquarium Passage, Doboy Creek, Brisbane River, J. Johnson, 22 June 1988; WAM P.28814-001, 6(13-42), Moreton Bay, V. Wadley, P. Young, 1972. New South Wales: AMS 1.14888, 1(32), Southwest Rocks, J. Wright; AMS 1.29891-001, 9(25-29.5), Towra Point, Botany Bay, NSW Fisheries, 9 February 1979. Other material examined (but not used in description) Australia: Queensland: Probable paralectotypes of Gobius stigmaticus, AMS 1.361, 2, Moreton Bay. QM 1.8048-9, 2, Brisbane River; QM 1.3948, 31, Breakfast Creek, Brisbane; QM 1.20309, I, Lower Logan River; QM 1.4337, I, Brisbane River; QM 1.28573, 7, Pine River mouth; QM 1.31196, 13, Saltwater Creek, Rothwell; ANSP 122177, 2, Rockhampton; QM 1.4070, I, Bulimba Creek, Tingalpa; QM 1.11031, I, Norman Creek; AMS IA.1577, 2, Breakfast Creek, Brisbane; QM 1.13390, 2, Deception Bay. New South Wales: AMS lA.4150, I, Trial Bay; AMS 1.16961-001, I, Parramatta River; AMS lA.5463, I, Woy Woy. Diagnosis A robust Mugilogobius with second dorsal rays 1,7- Figure 186 9 (modally 1,8); anal rays 1,7-8 (modally 1,8); pectoral rays 14-18; longitudinal scales 35-51; TRB 12-18; circumpeduncular scales 12-21; predorsal scales 1725, mostly small, anterior few sometimes larger, extending to between rear preopercular margin and rear of eyes; ctenoid scales on body extending forward in wedge to behind pectoral fin; third spine of dorsal fin modally longest, but no spines elongate; body with about seven brown blotches along midside, three oval to round dark spots at caudal base; known only from eastern Australia. Description Based on 31 specimens, 18.5-42 mm SL. An asterisk indicates counts of lectotype (Figure 186). First dorsal VI*; second dorsal 1,7-9 (mean 1,8*); anal 1,7-8* (mean 1,8), pectoral rays 14-18 (mean 17, 16 in lectotype), segmented caudal rays 16-18 (mean 16*); caudal ray pattern 7/7 to 10/8 (modally 9/7); branched caudal rays 14-18 (mean 16*); unsegmented (procurrent) caudal rays 8/8 (in one); longitudinal scale count 35-51 (mean 42, 45 in lectotype); TRB 12-18 (mean 15, 16 in lectotype); predorsal scale count 17-25 (mean 20, 19 in lectotype); circumpeduncular scales 12-21 (mean 17, 18 in lectotype). Gill rakers on outer face of first arch 3+6 to 5+7 (modally 4+7). Pterygiophore formula 3-12210* (in nine). Vertebrae 10+16* (in 17). Neural spine of first three vertebrae narrow, pointed (in six), or second neural spine slightly broadened (in two). Two epurals (in 11). Two (in 13) or three (in two) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid moderately broad, not expanded dorsally; forming bridge to quadrate. Body robust, compressed posteriorly, more rounded anteriorly. Head wider than deep, HL 3.03.5 (mean 3.4) in SL; cheeks sometimes inflated in large specimens. Depth at posterior preopercular margin 1.4-1.9 (mean 1.7) in HL. Width at posterior preopercular margin 1.1-1.5 (mean 1.4) in HL. Mouth subterminal to barely terminal, slightly oblique, forming angle of about 25° with body axis; jaws reaching to below mid-eye or to posterior half of eye, jaws not much enlarged in males (barely to below posterior half of eye in lectotype). Lips smooth, fleshy fimbriae may be present on inner edge of upper lip; lower lip free at sides, narrowly Mugilogobius stigmaticus, lectotype of Gobius stigmaticus De Vis, AMS 1.358, 37 mm female. 187 Revision of Mugilogobius Table 31 Morphometries as percentages of SL or HL, as indicated, of MlIgilogobills stigmaticlIs (De Vis, 1884). Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. I. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital \. in HL Pectoral I. in SL Pelvic \. in SL Caudal\. in SL Longest Dl spine in SL Lectotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 30.3 58.0 68.8 22.4 28.7 57.3 68.8 18.6 11.2 23.5 12.7 25.7 21.6 35.4 24.7 19.6 15.0 19.6 11.3 33.1 70.9 87.2 21.7 16.0 28.8 15.0 30.2 27.7 50.4 37.2 22.2 20.6 26.2 14.8 30.7 60.4 76.3 20.2 13.3 25.3 14.1 27.4 24.3 41.2 31.1 20.9 18.4 23.8 13.5 28.2 53.7 65.5 17.3 9.7 24.1 12.4 24.1 23.0 35.2 25.3 16.7 14.1 20.0 10.6 31.2 65.9 85.8 22.4 15.1 30.2 16.7 28.3 28.8 43.4 35.4 22.2 '9.6 25.6 15.1 29.7 59.2 74.3 19.6 12.6 26.6 14.2 26.2 25.4 38.4 30.1 19.9 18.0 23.5 14.1 25.7 15.1 25.9 24.1 42.0 26.8 18.6 15.1 fused across front. Upper jaw 2.0-2.8 (mean 2.6 in females, 2.4 in males) in HL. Eyes dorsolateral, high on head, top usually forming part of dorsal profile, 3.5-4.6 (mean 4.1) in HL. Snout rounded, 3.3-4.2 (mean 3.8) in HL. Interorbital broad, flat to slightly concave, 2.7-4.1 (mean 3.3) in HL. Top of head above preopercular margin up to snout tip usually covered with fine villi (most easily seen in specimens with intact mucous coat). Body depth at anal origin 4.5-5.8 (mean 5.1) in SL. Caudal peduncle compressed, length 3.3-4.2 (mean 3.9) in SL. Caudal peduncle depth 6.0-8.0 (mean 7.1) in SL. First dorsal fin low, triangular to rounded, tips of second to fifth spines often free, sometimes lengthened but not elongate, second to fourth spines longest or subequal, third spine modally longest; spines slightly longer in males than females; spines usually falling well short of second dorsal fin origin when depressed, sometimes free tips of spines reaching origin. First dorsal spine always shorter than next three. Second dorsal spine length 6.6-8.1 (mean 7.5 in males, 7.1 in females) in SL. Third dorsal spine length 6.6-9.4 (mean 7.7 in males, 8.2 in females) in SL. Fourth dorsal spine length 6.8-8.7 (mean 7.5 in males, 8.2 in females) in SL. Second dorsal and anal fins quite low, rounded anteriorly, posteriormost rays longest and somewhat pointed, rays falling well short of caudal fin base when depressed. Pectoral fin oval to rounded, central rays longest, 4.5-6.0 (mean 4.9) in SL; rays all branched but for uppem10st. Pelvic fins short, rounded to oval, reaching halfway to twothirds distance to anus, 4.8-7.1 5.6) in SL. Caudal fin short, rounded posteriorly, 3.8-5.1 (mean 4.3) in SL. No mental fraenum, chin smooth. Anterior nostril in short tube, placed at edge of upper lip, tube directed down and forward, preorbital curved forward to accommodate nostril. Posterior nostril oval to round, placed close to anterior centre margin of eye. Gill opening extending forward to under opercle, sometimes only just so. Inner edge of shoulder girdle smooth (in eight), with low fleshy or bony ridge or raised bumpy flange (in 15), or with distinct fleshy knobs or flaps, often placed on fleshy flange (in nine). Gill rakers on outer face of first arch very low knobs, longer near angle of arch; rakers on inner face of first arch about same size as outer rakers at angle of arch; inner and outer rakers on other arches same length as first arch inner rakers. Tongue tip blunt to concave. Outer teeth in upper jaw largest, stout and curved, three to four rows of evenly sized, small sharp teeth behind this row; usually two rows at side of jaw; teeth in males slightly larger than in females. Lower jaw with four or five rows of small pointed teeth across front, outermost row oriented upright, inner rows all pointing posteriorly; teeth in innermost row largest and stoutest toward midside of jaw; one or two rows of teeth at rear of jaw. Predorsal scales small, evenly sized, reaching forward to preopercular margin or further, extending up to behind eyes; in nine specimens (including paralectotype), anteriormost scale or two larger than others, but not enlarged or partly entering interorbital space; predorsal scales in lectotype small, reaching past preopercular margin. Operculum with small cycloid scales on upper half to two-thirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1/3 to 1/2 of belly), rest of scales cycloid. Ctenoid scales on side of body extending forward in wedge to behind pectoral fin. Genital papilla in male moderate to elongate, flattened, with blunt tip; in female, short, rounded, often bulbous. --------------------------- ---- H.K. Larson 188 ) '.' ) '\ 1 ' ' " '\' ) . .> . ｾ .ｾＧＮ.ＺＢＵ｣ｦＬＩﾧＨ .. . . . .' "\ ［ｾＩ \ , "- :J ...J./) )1 :J.. ...J .... "))'\ ..........................: J j'!'" ) .... ' . .' "c' . :. ｊＮｾｊ "cp' ...... J -,:: )I -.../) : V : ..... ＧＭＺｾ .. :' I ..' Figure 187 Mugilogobius stigmaticus, papillae pattern. QM 1.5021, Brisbane. Scale bar = 1 mm. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 187. Rear portion of broken c row consisting of three or more papillae. Three s rows on snout, central row shortest. Mental f row arranged in single or double row, or in small cluster. Coloration of fresh material Living captive specimens from Tin Can Bay, Queensland, not differing greatly from preserved specimens. Head and body greyish yellow to greyish brown with grey-brown to yellow-brown markings, scale margins narrowly edged with dark grey; lower half of body plain pale greyish yellow, belly pinkish yellow. Brown oblique shoulder bar and square brown blotch on side behind pectoral fin sometimes visible. Head darker, almost dark khaki colour, with lower half of head plain grey-brown; dorsal surface of head with brown vermiculation. First dorsal fin grey with narrow dark grey margin and two broad translucent pink to whitish pink bands above and below narrow dark grey central band which widens posteriorly into dark grey to black blotch at rear of fin; narrow grey triangular patch along base of anterior half of fin; fin rays light grey to blackish. Second dorsal fin Figure 188 light to dark grey, with submarginal pink to whitish pink band below grey fin margin; other markings grey to brown, pattern as for preserved specimens. Anal fin pale greyish to light pinkish grey, lighter proximally. Pectoral and pelvic rays hyaline to light greyish. Caudal fin plain grey to light yellowish grey, fin rays slightly darker, with small grey spots scattered near base of fin. Iris dark golden, with broad dark brown margin. Coloration of preserved material McCulloch and Ogilby (1919) illustrate a type specimen, showing typical markings. Head and body pale brownish yellow to light brown with darker brown markings, plain whitish to yellowish ventrally. Seven or eight square or irregularly-shaped cross-hatched brown blotches along midside of body and an offset row of slightly smaller, similar blotches on upper side of body, forming chequered or obliquely blotched pattern; posteriormost three or four blotches most distinct (Figure 188). Brown blotches sometimes coalescing on anterior half of body, forming oblique bars, including bar extending from behind pectoral base and ending on nape over opercle. Scaly sheath covering base of caudal fin with two oval, dark brown spots oriented toward posteriormost midlateral brown blotch Gust at hypural crease); these three spots forming blotchy "V"; mid-lateral blotch and lower caudal spot sometimes joining to form bar; upper caudal spot sometimes darker than lower. Side of head with variably intense dark lines over pale background; if network diffuse, two almost vertical brown lines from lower edge of eye to rear and mid-point of jaw usually remaining distinct. Opercle with similar network of brown lines, rear of opercle edged with dark brown, narrow to moderately broad line. Underside of head plain whitish to dusky; broad, diffuse brown streak extending from rear margin of opercle onto branchiostegal membranes; streak ending at membrane insertion on isthmus. Predorsal plain to mottled with diffuse brown spots and patches, oblique brown streak from upper rear margin of eye often running obliquely onto nape; interorbital Mugilogobius stigmaticus, 40 mm SL, QM 1.14069, Yeronga Creek, Queensland. Revision of MlIgilogobills and snout with brown vermiculation and narrow blotches. Pectoral base pale with horizontal brown bar crossing just above mid-point; upper half of fin often darker than lower. First dorsal fin translucent to dusky, with central broad brown band which intensifies posteriorly, forming black spot; transparent to whitish submarginal band immediately above; narrow fin margin and any long fin spines dusky to dark brown. Second dorsal fin whitish to dusky, with one to three rows of vertically oriented elongate brown to blackish streaks on proximal two-thirds of fin; two dark brown blotches usually present at base of fin; submarginal whitish band present; outer margin of fin dusky to dark brown. Anal fin brown to dark grey with white margin. Caudal fin plain dusky to translucent, membrane sometimes darker than rays, forming diffuse dark streaks; only distinct markings are two dark brown oval spots at base described above; brown triangular blotch at base of upper procurrent rays often present. Pectoral fins translucent to dusky. Pelvic fins pale to dusky, fraenum unpigmented. Peritoneum dark brown dorsally, fading at lower sides to pale ventrally. Comparisons Most similar to M. platynotus and M. platystomus, two species whose ranges overlap with M. stigmaticus; see under M. platynotus for comparison with that species. Mugilogobius stigmaticus can be distinguished from M. platystomus by the colour pattern of the head, having bars and a reticulate pattern (versus stripes in M. platystomus), three paired spots at caudal fin base (versus two big spots); the continuous extent of ctenoid scales along the sides (versus two patches of ctenoid scales, in a few specimens only, the two areas joined by one or two rows of ctenoid scales); and the relatively low, nonelongate first dorsal fin spines (versus first four spine tips always free of membrane and often elongate). Distribution This species is restricted to the east coast of Australia. Specimens are known from TownsvilIe, Queensland, to New South Wales (as far south as Botany Bay) (Figure 167). Ecology Morton et al. (1987) described the abundance and diets of several fishes inhabiting a saltmarsh inlet at Coomera Island, in southern Moreton Bay, Queensland; the fishes studied included M, stigmaticus (see Ecology under M. platyllotus). Gee and Gee (1991) described this species' reaction to hypoxia (see Ecology under M. platynotus). 189 Remarks In De Vis' original description (1884), it is clear that he was using more than one specimen, as he described colour variations, but he gave no indication of the number of specimens he had. Of the two specimens labelled syntypes in AMS 1.358, the smaller (37 mm) is here designated as lectotype of Gobius stigmaticus De Vis, as it is somewhat less dehydrated, shows the colour pattern better and has the usual predorsal scale arrangement (scales small and evenly sized) (Figure 186). McCulloch and Ogilby (1919), in their redescription of this species (as Mugilogobius devisi), commented on the differences in nape scale size. They made it clear that De Vis' "cotypes" of Gobius stigmaticus consisted of two specimens only, of yellow-brown colour, with the darker patterning well-preserved. The two specimens (36-39 mm SL) labelled syntypes in AMS 1.361 have all nape scales small and evenly sized, and are stained dark brown and appear to have had a somewhat different preservation history than AMS 1.358. McCulloch and Ogilby (1919) did not refer to these specimens and may not have considered them to be types. These specimens would have arrived and been catalogued at AMS at the same time as 1.358 (May 1886; similar handwriting for both these lots), and in the original AMS register book, 1.361 is labelled as "Gobius stigmaticus = type of Gobius brunneus De Vis" (S. Reader, personal communication). De Vis did not describe a Gobius brunneus; this is a "museum name". It is possible that McCulloch and Ogilby overlooked the specimens in 1.361 and the specimens are genuine types of Gobius stigmaticus. The specimens are therefore indicated as being "probable paralectotypes", until more information is found regarding these specimens. Mugilogobius devisi McCulloch and Ogilby (1919: 223) was an unnecessary replacement name for Gobius stigmaticus De Vis, 1884; they thought it was preoccupied by Smaragdus stigmaticus Poey (= Gobiollellus). McCulloch and Ogilby expressed some doubt about this, in a footnote, but created the name anyway. Koumans (1940) pointed out their mistake, and accepted that M. stigmaticus was a good species. It seems that McCulloch was still uncertain in 1929 (1929: 376), as he stated that stigmaticus was "regarded as preoccupied", under M. devisi in his checklist of Australian fishes. Mugilogobius tigrinus sp. novo Figures 167, 189-190; Tables 5-8, 32 Material Examined I-lolotype ZRC 40283, 16.5 mm SL female, mangrove creek, Sungei Pandan, Singapore, H. Larson, P. Ng, D. Murphy, 29 December 1993. H.K. Larson 190 Table 32 Morphometries as percentages of SL or HL, as indicated, of Mugilogobius tigrinus sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. 1. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL lnterorbitall. in HL Pectoral 1. in SL Pelvic 1. in SL Caudal 1. in SL Longest Dl spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 28.5 55.3 70.2 18.2 11.5 27.3 11.5 21.3 34.0 36.2 25.5 20.6 20.0 24.8 18.2 28.3 55.3 63.2 17.1 10.0 25.9 12.1 19.6 27.9 26.3 20.9 19.0 19.4 23.5 15.9 34.5 65.5 74.1 20.8 13.8 30.6 13.8 27.5 32.7 41.0 40.8 24.8 25.0 28.8 28.6 30.7 59.7 69.2 19.4 12.6 28.7 12.9 24.0 30.7 34.4 27.7 22.3 21.9 26.2 21.5 26.2 55.3 64.1 17.9 10.0 25.8 11.5 19.6 26.9 26.7 19.6 19.4 18.2 23.0 16.0 32.5 64.6 77.1 20.8 13.1 31.5 13.3 25.6 35.4 36.2 30.8 24.5 24.6 27.3 31.1 29.6 60.4 71.6 19.5 11.9 28.4 12.7 22.9 31.6 32.2 26.3 21.2 21.2 25.2 22.4 Paratypes Thailand: U5NM 316184, 1(17), brackish Nypa palm swamp along N bank of Maenam Bang Pakong, about 14 km N of Chonburi Chachoengsao Province, 5ta. T-27, 22 December 1970; URM P.12664, 10(12.5-16), mangrove swamp, Phuket, H. 5enou and C. Vidthayanon; NTM 5.13953-017, 6(9.5-15.5), small mangrove creek at Klong Bang 5ai, Phuket, H. Larson, D. Hoese and PMBC staff, 8 December 1993; NTM 5.14318-001, 1(18), Chantaburi, Mahidolia project, T. Wongratana; NTM 5.14288-003,3(13-16), ponds within research station at Ta-Chalaab, Chantaburi Province, Chulalongkorn University and NIFI, 2 June 1990. Malaysia: ZRC 7641-57, 17(11.3-22.5), probably Malaysia, probably D.5. Johnson in 195!Hi8. Singapore: NTM 5.13957007, 3 (9.5-17.5), same data as holotype; AM5 1.37570-002, 1(19.5), 5ungei Pandan, D. Hoese and K. Urn, 22 December 1993. Diagnosis A rather small, slender Mugilogobius with relatively compressed body and second dorsal and anal rays 1,6-7; pectoral rays 14-16; longitudinal scales 24-28; TRB 8-10; circumpeduncular scales 11-12; predorsal scales 9-12, beginning close behind eyes, anteriormost scale greatly enlarged; scales on body mostly ctenoid; first or second spine of dorsal fin longest, sometimes elongate; head with large eyes and short pointed snout; body with four complete black bands, two half bands and one rounded to elongate black spot on caudal base; known only from mangrove habitats in peninsular Thailand, Malaysia and 5ingapore. Description Based on 35 specimens, 9.5-22.5 mm 5L. Counts of holotype (Figure 189) indicated with asterisk. First dorsal VI*; second dorsal 1,6-7* (mean 1,7); anal 1,6-7* (mean 1,7), pectoral rays 14-16 (mean 15*), segmented caudal rays 16*; caudal ray pattern modally 9/7*; branched caudal rays 6/6 to 9/6 (modally 7/6, 6/5 in holotype); unsegmented (procurrent) caudal rays 6/8 to 7/7; longitudinal scale count 24-28 (mean 27*); TRB 7*-10 (mean 8); predorsal scale count 9-12 (mean 10*); circumpeduncular scales 11-12* (mean 12). Gill rakers on outer face of first arch 3+7 to 5+7 (modally 4+8). Pterygiophore formula 3-12210 (in four). Vertebrae 10+16 (in eight). Neural spine of second vertebra expanded at tip (in one), or pointed (in four). Two epurals (in four). Two (in five) or three (in one) anal pterygiophores before haemal spine of first caudal vertebra. Metapterygoid relatively low and narrow, reaching across to quadrate. Fifth ceratobranchials slender, with shortbased flange on ventral surface. Cleithrum with distinct flange on inner medial face. Body slender, quite compressed (less so anteriorly). Head wider than deep, but not greatly so, HL 2.9-3.8 (mean 3.4) in 5L; somewhat rectangular in cross-section. Depth at posterior preopercular margin 1.5-1.8 (mean 1.7) in HL. Width at posterior preopercular margin 1.3-1.6 (mean 1.4) in HL. Mouth subterminal, oblique, Figure 189 Mugilogobius tigrinus sp. nov., holotype, 17 mm SL female, ZRC 40283, Sungei Pandan, Singapore. Revision of Mugilogobius forming angle of about 30-35° with body axis; jaws reaching to below mid-eye in males and to below anterior half of eye in females (nearly to below mideye in holotype). Lips usually smooth, small fleshy fimbriae often present on inner edges of upper lip and front of lower lip; lower lip free at sides, fused across front. Upper jaw length not much different between males and females, 1.4-3.8 in HL (mean 3.0 in females, 2.9 in males). Eyes large, lateral, high on head, top usually forming part of dorsal profile, 2.8-3.9 (mean 3.3) in HL; eye width greater than snout length. Snout short, profile pointed to slightly rounded (usually pointed), 2.8-5.1 (mean 4.3) in HL. Interorbital broad, flat, 1.4-5.1 (mean 3.7) in HL. Top of head from anterior portion of interorbital forward to snout tip usually with sparse scattering of fine villi. Body depth at anal origin 4.8-5.9 (mean 5.2) in SL. Caudal peduncle long, compressed, length 3.2-3.9 (mean 3.5) in SL. Caudal peduncle depth 7.3-8.7 (mean 7.9) in SL. First dorsal fin low, triangular, tips of first to fourth spines may be free, first or second spines longest and sometimes elongate; spines slightly longer in males than females; spines barely reaching second dorsal fin origin when depressed; elongate spine, if present, reaching up to third element of second dorsal fin. First dorsal spine usually longest (on average, first spine longest in females, second spine longest in males), ＳＮＲｾＳ (mean 4.7) in SL. Second dorsal spine length ＳＮＵｾＳ (mean 5.6) in SL. Second dorsal and anal fins short-based, low, posteriormost rays longest, rays falling well short of caudal fin when depressed. Pectoral fin oval to somewhat pointed, central rays longest, 4.0-5.3 (mean 4.6) in SL; rays usually all branched but for uppermost. Pelvic fins short, oval, reaching past halfway to two-thirds of distance to anus, 4.0-5.5 (mean 4.7) in SL (fin nearly reaches anus in one 19.5 mm SL male). Caudal fin oval, rounded posteriorly, 3.5-4.4 (mean 3.9) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, short, placed on edge of upper lip, tube oriented down and forward, preorbital usually produced forward to accommodate nostril. Posterior nostril rounded, placed close to anterior centre margin of eye. Gill opening extending forward to under opercle. Inner edge of pectoral girdle with several to many small rounded knobs or flattened lobes (triangular to square in shape). Gill rakers on outer face of first arch slender but very short, with tiny spiny papillae at tip, longest rakers near angle of arch; rakers on inner face of first arch usually longer than those of outer face, often equal to longest raker; outer rakers on second and remaining arches short, with tiny papillae at tip, and inner face rakers relatively long and slender, as with inner face rakers on third and fourth arches. Tongue tip usually blunt, sometimes concave. Outer teeth in upper jaw largest, but not greatly enlarged, 191 Figure 190 MlIgilogobills tigrinlls n. sp., papillae pattern. Paratype, URM P.12664. Scales omitted. Scale bar = 1 mm. curved and pointed, three or four rows of very small sharp teeth behind this row; one or two rows of small teeth at side of jaw; outer teeth in upper jaw larger in males than in females. Females with lower jaw with five or six rows of evenly sized, small pointed teeth across front, usually no teeth particularly enlarged; one or two rows of teeth at sides. In males, about five rows of curved, pointed teeth across front, teeth increasing in size posteriorly with innermost row teeth largest; usually only one row of teeth at side of jaw. Predorsal scales about same size as body scales but for anteriormost, scale considerably enlarged, close behind eyes and partly entering interorbital space. Operculum with cycloid scales on upper half to two-thirds. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area completely covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior % to Vz of belly), rest of scales cycloid; some specimens with only few ctenoid belly scales. Ctenoid scales on side of body extending well forward to behind pectoral fin. Genital papilla in male elongate, flattened, narrowing to pointed or blunt tip; in female, papilla large, rounded, often bulbous, tip blunt. Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 190. Papillae in rows c and i somewhat more widely spaced than those of rows b and d, but papillae not enlarged. Usually three s rows, of one to three papillae each; some specimens only with two s rows. Papilla row f on chin of two papillae. Coloration of fresh material I noted live fish in Singapore as being whitish with black bands and caudal spot, with large dark eyes and short, pointed, dark snout. First dorsal fin whitish with large black spot and elongate white first spine. H.K. Larson 192 Coloration of preserved material Head and body whitish to very light brown (depending on preservation), with dark brown markings (Figure 189). Six narrow dark brown bars on body and one crossing head; four bars encircling (or nearly encircling) body; all bars (but for fourth) two or three scales wide. First brown bar crossing head just behind first predorsal scale, extending down onto opercle and intensifying as blackish blotch. Second brown bar crossing back directly above pectoral base, and ending just behind upper part of base. Third brown bar beginning at first dorsal fin origin or just behind it and extending to belly, stopping short of crossing midline. Fourth brown bar diffuse, sometimes indistinct, three to four scales wide and very short, ending well before midside of body; bar occupying interdorsal space. Fifth brown bar commencing approximately below middle of second dorsal fin and forming continuous band around body; bar slightly oblique (slanting anteriorly) in some specimens. Sixth brown bar forming continuous band around caudal peduncle, placed between midpoint of peduncle and rear of soft dorsal fin. Seventh brown bar encircling caudal peduncle just at beginning of caudal procurrent rays; bar sometimes slightly oblique (slanting posteriorly). On caudal fin, just above mid-point of fin base, dense dark brown to black rounded spot or short oblong bar; round spot usual. Top of head and snout diffuse to dark brown; anteriormost (enlarged) scale usually brown; brown bar sometimes joining eyes across interorbital space. Dark brown streak or broad line extending from anterior lower edge of eye forward to upper lip, reaching point halfway between anterior nostril and rictus. From centre rear of eye, diffuse dark brown streak or line extending obliquely up to first dark brown bar (on predorsal region); second dark brown streak or broad line running from lower margin of eye horizontally across to preopercular margin. Pectoral base pale with diffuse round brown spot on upper half. Dusky to dark brown oval patch covering most of breast; belly pale. Branchiostegals and lips dusky to brown; preopercular and opercular edges sometimes narrowly outlined in dark brown. First dorsal fin dusky anteroventrally, with small to very large round black spot or blotch occupying most of fin; broad white submarginal band present; fin margin dark brown to black; elongate (first or second) dorsal spine white, or occasionally, dusky. Second dorsal fin dusky with submarginal white band and brown margin; centre of fin with distinct brown to blackish blotch (continuation of fifth body bar) usually present. Anal fin dusky with whitish margin. Caudal fin plain dusky apart from black spot at base already described, with dark brown to blackish upper and lower edges near fin base. Pectoral fin translucent to dusky, fin rays outlined with dusky pigment. Pelvic fins dusky to brownish, fraenum whitish, unpigmented. Comparisons This species looks very similar to M. fasciatus and is syntopic with it in Thailand and Singapore. The two species can be most easily distinguished by colour pattern and longest first dorsal spines. The bands are wide with five encircling the body in M. fasciatus, and narrow with four encircling the body in M. tigrinus. The first brown band on the body begins at a level above the pectoral base in M. tigrinus and at the first dorsal fin origin in M. fasciatus; M. fasciatus has a black-bordered white stripe from the eye to the mouth, while M. tigrinus has a single blackish stripe from the eye to the middle of the upper jaw. Mugilogobius fasciatus has the second to fourth first dorsal spines free and elongate, while M. tigrinus has the first or second spine elongate. When alive, M. tigrinus has a very distinctive physiognomy with its pointed snout and large dark eyes distinguishing it immediately from M. fasciatus. Distribution Specimens are so far known only from peninsular Thailand, Malaysia and Singapore (Figure 167). Ecology Mugilogobius tigrinus has been found in shallow small mangrove creeks and pools. Remarks Mugilogobius tigrinus and M. fasciatus have the lowest lateral scale counts of all Mugilogobius species, with a mean of 26 (other species have mean scale counts of 29 or higher, usually 30-32). The two species also share similar colouring and habitat preference. Etymology From the Latin tigrinus, pertaining to tigers, in reference to the narrow bands or stripes across the body of this species. Mugilogobius wilsoni sp. novo Figures 167, 191-193; Tables 5-8, 33 ?Mugilogobius duospilus: Davis 1988: 164. Material Examined Holotype NTM 5.13993-001, 21 mm SL male, creek by Darwin High School, D. Wilson, February 1990. Para types Australia: Northern Territory: USNM 316174, Revision of Mugilogobius 193 28(13.5-26.5), desiccating mud pool cut off from creek at Yirrkala, Cape Arnhem, RR Miller, 30 July 1948; NTM 5.13993-002, 3(19-25), same data as holotype; NTM 5.10439-001, 3(9-17), Danger Point, hand, H. Larson, 30 April 1982; AM5 1.23940-001, 7(12-30), creek off Casuarina Beach, Darwin, 0-1 m, rotenone, D. Rennis, 7 August 1983; NTM 5.13178001, 20(10-14), tidal creek behind Lifesaving Club, Casuarina Beach, G. Dobson, June 1989; NTM 5.12433-008,4(12-15), creek at Base Bay, Vanderlin Island, 0-1 m, dipnet, H. Larson and W. Houston, 27 July 1988; NTM 5.13518-012, 14(15-22), small mangrove creek, E arm Bing Bong Creek, Gulf of Carpentaria, H. Larson, R Lau and K. Akerman, 5 September 1992. Queensland: AM5 IA.6560, 6(13.527.5), Plantation Beach, Lindeman Island, G. Whitley, 2 September 1935; AM5 IA.6638, 2(1118.5), Bowen, G. Whitley, 16 August 1935; AM5 1.20785-038, 1(15.5), Hartley's Creek, 1-2 m, rotenone, colI. D. Hoese, 15 December 1978; AM5 1.21259-004, 19(11.5-23), creek 200 m 5 of Cape Tribulation, D.F. Hoese, 11 August 1979. Other material examined (no data taken) Australia: Western Australia: ex QM 1.22552, 1, Pago Ruins Creek. Northern Territory: NTM 5.13876-001, 3, Marchinbar Island, Wessel Islands; NTM 5.12990-003, 5, Vestey's Beach, Darwin; NTM 5.13885-003, 1, West Arm, Darwin Harbour; NTM 5.14347-001, 8, Ludmilla Creek, Darwin; NTM 5.10694-017, 9, Gunn Point; NTM 5.10420-003, 1, Elizabeth River; NTM 5.14292-001, 21, golf links canal, Darwin; U5NM 308905, 2, Oenpelli; AM5 1.23939-014, 1, Darwin. Queensland: AM5 1.26849001, 3, Point Lookout; QM 1.13293, 1, Hinchinbrook Island; AM5 IA.6666, 18, Lindeman Island. Diagnosis A moderately slender Mugilogobius with second Table 33 dorsal rays 1,7-8; anal rays 1,6-8; pectoral rays 1316; longitudinal scales 26-31; TRB 8-13; circumpeduncular scales 12-13; predorsal scales llB, anteriormost enlarged, placed close behind eyes; usually 12 circumpeduncular scales; scales on body mostly ctenoid; dorsal fin rounded, without elongate or filamentous spines, black spot usually on anterior half of fin; body grey with five to seven oblique black bands across sides, bands interspersed with black spots and chevron-shaped black markings, two oval spots on base of caudal fin; known only from mangrove creeks of northern Australia. Description Based on 42 specimens, 13-30 mm 5L. An asterisk indicates of holotype (Figure 191). First dorsal V (in one), VI" (in 41); second dorsal 1,7"-8 (mean 1,7); anal 1,6-8 (mean 1,7"); pectoral rays 13-16 (mean 14, 15 in holotype), segmented caudal rays 15-16 (mean 16"); caudal ray pattern 9/ 7; branched caudal rays 7/6 to 9/7 (modally 8/7"); unsegmented (procurrent) caudal rays 6/7 to 7/8 (mostly indiscernible in available X-rays); longitudinal scale count 26-31 (mean 29, 28 in holotype); TRB 8-11 (mean 10"); predorsal scale Figure 191 Mugilogobius wilsoni sp. nov., holotype, 21 mm SL, NTM S.13993-001, Darwin, Northern Territory. Morphometries as percentages of SL or HL, as indicated, of Mugilogobius wilsoni sp. novo Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. !. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital!. in HL Pectoral!. in SL Pelvic !. in SL Caudal!. in SL Longest Dl spine in SL Holotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 31.9 53.7 68.7 20.5 14.8 26.2 13.3 29.9 31.3 43.3 43.3 21.0 20.5 27.1 14.8 21.5 53.0 64.4 14.0 9.8 19.5 9.5 24.4 25.3 34.0 31.4 15.5 13.5 20.0 10.5 32.9 63.0 79.1 21.2 15.5 29.7 15.2 31.6 36.6 45.8 44.6 23.0 21.5 30.4 17.0 29.7 56.8 71.6 19.0 12.3 26.8 13.5 28.2 29.7 39.9 37.4 21.3 19.3 26.1 15.1 25.9 54.5 58.4 18.4 10.5 24.0 12.6 23.6 27.8 33.3 31.7 17.8 16.8 22.8 10.2 31.5 63.2 74.5 21.7 14.9 30.8 14.5 29.3 34.9 38.2 43.1 22.6 20.0 27.7 16.2 28.4 58.8 70.1 19.9 13.0 27.2 13.6 26.6 30.4 25.8 35.8 20.7 18.7 25.1 14.1 --- __ _----------_._- H.K. Larson 194 count 11*-13 (mean 12); circumpeduncular scales 12-13 (mean 12"). Gill rakers on outer face of first arch 3+7 to 7+10 (modally 4+9). Pterygiophore formula 3-12210 (in two). Vertebrae 10+16 (in 17). Neural spine of second and third vertebra broad, expanded at tip (in two). One (in one) or two (in one) epurals. Two (in four) anal pterygiophores before haemal spine of first caudal vertebra. Body compressed, usually somewhat rounded anteriorly. Head wider than deep, HL 3.0-4.7 (mean 3.5) in SL; head slightly depressed anteriorly in large males; cheeks sometimes inflated in large males. Depth at posterior preopercular margin 1.61.9 (mean 1.8) in HL. Width at posterior preopercular margin 1.3-1.7 (mean 1.4) in HL. Mouth subterminal, slightly oblique, forming angle of about 20-25° with body axis; jaws generally reaching to below mid-eye to posterior half of eye in males and to below anterior half of eye in females (to below mid-eye in holotype). Lips usually smooth, fleshy fimbriae often present on inner edges of upper lip and front of lower lip; lower lip free at sides, broadly fused across front. Upper jaw 2.2-3.0 (mean 2.8 in females, 2.5 in males) in HL. Eyes lateral, high on head, top usually forming part of dorsal profile, 2.2-3.9 (mean 3.3) in HL. Snout slightly rounded, 3.2-4.2 (mean 3.7) in HL. Interorbital broad, flat, sometimes slightly concave in adult males, 1.4-3.2 (mean 2.8) in HL. Top of head from just behind eyes to snout tip usually covered with fine villi (density variable, usually visible only in specimens with well-preserved mucous coat). Body depth at anal origin 4.6-7.1 (mean 5.2) in SL. Caudal peduncle compressed, length 3.3-5.1 (mean 3.8) in SL. Caudal peduncle depth 6.6-10.5 (mean 7.4) in SL. First dorsal fin somewhat rounded to triangular, tips of spines usually not free, second or third spines longest or subequal; spines slightly longer in males than females; spines barely reaching second dorsal fin origin when depressed, and often falling short of origin. First dorsal spine always shorter than next two. Second dorsal spine length 5.9-9.5 (mean 6.7 in males, 7.2 in females) in SL. Third dorsal spine length 6.4-9.8 (mean 6.9 in males, 7.5 in females) in SL. Second dorsal and anal fins about equal in height to first dorsal, posteriormost rays longest; second dorsal rays just reaching caudal base when depressed (males) or falling well short (females), anal rays not reaching caudal fin when depressed. Pectoral fin rounded, central rays longest, 4.3-6.5 (mean 4.9) in SL; rays usually all branched but for uppermost. Pelvic fins short, rounded to oval, reaching half to two-thirds of distance to anus, 3.8-7.4 (mean 5.3) in SL. Caudal fin rounded, 3.3-5.0 (mean 4.0) in SL. No mental fraenum, chin smooth. Anterior nostril tubular, placed at edge of upper lip, tube short, oriented down and forward, preorbital curved forward slightly to accommodate nostril. Posterior nostril rounded to oval, placed close to anterior centre margin of eye. Gill opening usually extending forward to under opercle. Inner edge of pectoral girdle smooth with no ridge or flange (in two), with smooth bony flange (in three), with low irregular fleshy ridge (in four), most specimens with distinct fleshy knobs or bumps (in 32); knobs rounded and fleshy or rather flattened flaps. Gill rakers on outer face of first arch very short, with small spiny papillae on outer face only, longest few rakers near angle of arch; rakers on inner face of first arch twice as long, tips with fine spiny papillae; outer rakers on remaining arches low, stubby, with papillae on tips; inner rakers on remaining arches with spiny papillae, equal in length to first arch inner rakers. Tongue tip usually blunt, concave in three specimens. In males, outermost teeth across front of upper jaw largest, pointed and widely spaced; teeth at sides of jaw smaller; jaw with two to four rows of small curved sharp teeth, two rows at sides. Upper jaw teeth in females similar, but all teeth much smaller. In males, lower jaw teeth in three or four rows across front, teeth stout, sharp, all curving inward, usually only two rows at sides of jaw. Females similar in arrangement, but teeth much smaller and finer. Anteriormost predorsal scale enlarged, placed at rear of interorbital space and often intruding partway; remainder of nape scales smaller, all cycloid. Operculum with small cycloid scales on upper third to half or more. Cheek always naked. Pectoral base covered with cycloid scales. Prepelvic area covered with small cycloid scales. Belly with isolated patch of ctenoid scales under pelvics (covering anterior 1,4 to 1;2 of belly); rest of scales cycloid. Ctenoid scales on side of body extending forward in wedge to behind pectoral fin. Genital papilla in male elongate, flattened, narrowing to rounded tip; in female, short, rounded and blunt-tipped. Figure 192 Mugilogobius wilsoni n. sp., papillae pattern. Paratype, USNM 316174. Scales omitted. Scale bar = 1 mm. Revision of Mugilogobius Head pores absent as in all Mugilogobius. Sensory papillae pattern longitudinal, as in Figure 192. Papilla row p in characteristic "eyebrow" shape; row often set in groove. Cheek papilla row c broken, with wide gap under eye; rear portion of row often of only one papilla. Preopercular margin papilla row e not extending far up vertical limb of preopercle. Three 5 rows on snout, usually of three or more papillae each; middle row sometimes of only two papillae. Two f row papillae behind mandibular symphysis. Coloration of fresh material From my field notes. Body grey, whitish below, with scale margins narrowly outlined in black, with black to dark grey oblique bars and spots; fins clear with grey and black markings, with no blue or yellow present. Colour slide of captive specimen (Figure 193). Body pale yellowish grey with grey to black markings; outer third of first dorsal fin yellow above rounded black spot, outer edge of fin with very narrow brown margin; second dorsal fin and upper and lower edges of caudal fin with broad white margin. Coloration of preserved material Head and body pale yellowish to greyish white (depending on preservation), with scale margins usually outlined with dark brown to blackish, giving overall reticulate appearance; side of body with six or seven dark brown to blackish oblique bars (usually joining across dorsal midline) (Figure 191). No obvious sexual dichromatism. Anteriormost bar angling forward behind pectoral base up to above opercle; bar extending onto nape and sometimes meeting its counterpart Figure 193 Mugilogobius wilsoni sp. novo Captive specimen from Dunk Island, Queensland. From colour slide by K. Uhlenhunt. 195 on nape midline; bars usually separated by two or three scales, or coalescing (as in holotype). Second bar beginning at midbase of second dorsal fin and extending down and forward obliquely, ending midlaterally. Remaining four to five bars equally spaced along sides of body, tending to be oriented vertically; lower half of bars often sharply angled back posteriorly, forming chevrons. Short upright brown streaks and/or variably shaped blotches sometimes interspersed with body bars; margins of scales between bars outlined with dark brown, giving overall reticulate background pattern including chevrons and Xs and Ws. At caudal base, dark brown chevron or vertical bar present just before hypural crease; (apex of chevron facing anteriorly. On base of caudal fin on fin-base scales, vertically oriented pair of distinct black round to oval spots; area immediately surrounding spots usually pale, giving ocellate appearance. Side of head with two horizontal stripes. Uppermost (broader) stripe running from midupper lip around lower edge of eye and across upper preopercle and ending on opercle as series of indistinct blotches or streaks. Lowermost head stripe very narrow and slightly wavy, sometimes partly broken up; stripe running from corner of mouth and reaching, or ending just before, edge of preopercle. Some indistinct dusky vermiculation and blotches sometimes present on cheek between head stripes, or forming indistinct wavy stripe extending from just above corner of mouth nearly to rear of preopercle. Opercle with two irregular oblique streaks or blotches on upper half (remainder of upper preopercular stripe); lower half plain dusky. Snout tip with dark brown vermiculation and spotting. Lips plain dusky, or with pencil-thin darker lines along edges. In males, interorbital space, snout and lips often distinctly dark brownish grey, contrasting with rest of body. Underside of head plain dusky; usually thin dark lines visible following lower lip margin, preopercular margin and skin folds across isthmus; several lines often crossing branchiostegal membranes; diffuse or dense dark grey to blackish wide band running from lower half of each opercle, crossing branchiostegal membranes and joining its counterpart. Usually, diffuse brown band crossing anterior part of breast and extending up to below pectoral base. Pectoral base light brown, with distinct rounded brown spot on centre of upper half. Abdomen and ventral edge of caudal peduncle pale yellowish to whitish. First dorsal fin with dark grey margin and large black spot or band. Black spot present from second to fourth spines; spot rounded or forming broad blotch or band. Area in front of black spot dusky grey; area above translucent to whitish. Second dorsal fin dusky grey with grey margin, translucent to whitish submarginal band and short vertical H.K. Larson 196 blackish streaks on membranes between spines on lower half of fin; some streaks and spots very dark, forming rows of black blotches. Anal fin evenly dusky brown to grey; sometimes indistinct vertical blackish streaks on membranes visible; margin translucent. Pectoral fins evenly dusky grey. Pelvics translucent to dusky, centre of fins often dusky. Caudal fin usually dusky with two black spots described above on base; fine blackish streaks radiating out from fin base, becoming diffuse and vanishing about halfway along fin. Comparisons This species looks very much like M. chulae, especially in basic colour pattern. However, the first dorsal spines are never filamentous as in M. chulae, and the black spot on the first dorsal fin is positioned differently (in M. wilsoni the spot is placed between the second and fourth spines, while in M. chulae it occupies the space between the third and sixth spines). Mugilogobius wilsoni appears to be a smaller species, with the largest known specimen observed being 30 mm SL (that of M. chulae being 38.5 mm SL). Distribution Specimens are known only from northern Australia (Western Australia, the Northern Territory and Queensland) (Figure 167). Ecology This species has mostly been collected from small muddy mangrove creeks and may not travel very far up into more brackish habitats. Specimens from the collection which included the holotype were kept in captivity for some weeks in seawater and fed on live brine shrimp. They were quite active and made attractive aquarium fish. David Wilson (Territory Wildlife Park) was able to induce spawning in this species after his specimens had spent about two weeks in captivity. The fish were from a tidal stream which was fresh at the time of collection, and the adults were maintained in fresh water. Eggs hatched after six to seven days at about 27°C. Larvae were noted as being very small, freeswimming, but unfortunately required far smaller food items than were available and did not survive. recognition of his continuing help and enthusiasm in collecting gobies and for promoting the keeping and appreciation of native Australian freshwater fishes. Nomina Dubia There are seven names which remain here as nomina dubia but are possibly species of Mugilogobius. Most type specimens for these names are apparently lost or destroyed. Herre's types have suffered particularly, many of which were kept at the Bureau of Science, Manila, which was destroyed in World War H. In the following cases, the original descriptions have not given sufficient information to positively identify the fishes without a specimen, even when an illustration accompanied the description. Gobius magniloquus Day, 1876: 296 (Madras, India). The possible syntype, ZSIC 159, is lost or destroyed. It was reported as lost according to the ZSI type register, in a letter to me from Dr R.P. Barman (19 August 1991). The original description (Day 1876: 296) indicates that the predorsal is fully scaled, with a large scale just behind the eyes (a large scale can be found in Stigmatogobius, Redigobius, Pseudogobius and the chulae-group of Mugilogobius) and the maxilla reaches to below the rear edge of the eye (similar to Mugilogobius and Redigobius). The high number of predorsal scales given by Day (15) is similar to counts for some species of Mugilogobius. Gobius poeyi Steindachner, 1867: 350 (Barbados). The holotype (NMW 30608) was examined at Vienna in 1988. The specimen is considerably bleached, with the only discernible colour pattern a brownish spot on the first dorsal fin (Figure 194). In the original description, Steindachner reported two parallel rows of black-brown spots on the second dorsal fin; a blackish spot between the last two first dorsal fin spines; the body brown with dark spots in two rows along the upper half of the body; two poorly-defined spots on scales at the base of the tail fin, with these spots the same size as those on the body; and small dark spots on the caudal rays. Four species of Mugilogobius share similar counts to this specimen: notospilus, stigmaticus, littoralis and myxodermus. Steindachner's name is older than any Remarks The specimens identified as M. duospilus by Davis (1988) were not available for study, but they were probably of this species. Two specimens in poor condition from the Aru Islands (SMF 6737), may possibly be this species. Etymology This species is named for David Wilson, of the Territory Wildlife Park, Berry Springs, NT, in Figure 194 Holotype of Gobius poeyi Steindachner, 31 mm SL, NMW 30608, Barbados. Photograph by A. Schumacher. Revision of MlIgilogobills of these, but it is not possible to determine to which species poeyi belongs, without further information as to where the specimen actually came from, and how it came to be labelled as being from Barbados. No other specimens of Mugilogobius have been reported from the Barbados (or elsewhere in the Atlantic), and it is highly probable that the reported locality for this species is a mistake. The original catalogue entry for this species was checked for this study by Harald Ahnelt (formerly of NMW), and all data agrees with the locality recorded on the specimen label. Correspondence on this problem is in progress with Or B. Herzig of NMW. Rhinogobius schultzei Herre, 1927: 185 (Fabrica, Occidental Negros, Philippines). The syntypes (BSM 12407 and BSM 26833) were destroyed in WWIl. The original description was based on two specimens (28-37 mm in length) from a river. Koumans (1940: 186) regarded the species as probably belonging to Stigmatogobius (the genus in which he placed species now usually assigned to Pseudogobius or Redigobius). He also stated that the first scale behind the eyes was enlarged. From Herre's description of preopercular pores and protruding chin, it is unlikely that the specimens belonged to a species of Mugilogobius, but possibly were Eugnathogobius or Stigmatogobius, based on these characters and Koumans' observation of a large scale behind the eyes. Tamanka arguellesi Roxas and Ablan, 1940: 308, plate 7 (Dagupan, Pangasinan Province, Philippines). Judging from the original description and figure, this species appears to belong to the abei-group of Mugilogobius. Roxas and Ablan's description of "a pair of terminal pores" on the upper lip probably refers to the anterior nostril and the "prominent tubular pore" on the "upper margin of snout, very far advanced from each eye" refers to the posterior nostril (shown in Roxas and Ablan 1940: plate 7). The holotype (47 mm SL; BSM 31951) and 13 BSM paratypes were destroyed in WWIl. 197 description". The Eugnathogobius. species is possibly Vaimosa villa Herre, 1927: 154, plate 12, figure 2 (Molo, Iloilo Province, Philippines). Described from the holotype (BSM 13195, 36 mm SL) and one paratype (BSM 13228, 35 mm SL) destroyed in WWII. Koumans listed catalogue number of types and considered it to be "A good species of Mugilogobius" (1940: 186). I have been unable to determine with what species it agrees. Pandaka Herre, 1927 Pandaka Herre, 1927 (Pandaka pusilla Herre, 1927: 197, plate 15, figures 1-2, Sitankai, Philippines, by original designation). Berowra Whitley, 1928 (Cobius lidwilli McCulloch, 1917: 185, plate 31, figure 2, Cowan Creek, Hawkesbury River, New South Wales, Australia, by original designation and monotypy). Berowara Takagi, 1966 (lapsus for Berowra). Diagnosis Very small gobies distinguished by following combination of characters. Second dorsal and anal fin rays 6-8, all segmented; 14-16 pectoral rays; segmented caudal rays 17, in 9/8 pattern; 20-24 scales in lateral series; predorsal scales absent; gill opening restricted to just under opercle; pectoral girdle smooth or with narrow bony ridge; distinct lobe or fleshy spur present on rear edge of branchiostegal membrane; headpores absent; sensory papillae sparse, arranged in reduced longitudinal or transverse pattern (Figure 195); no fine villi on head; jaws short, mouth terminal with lower jaw anteriormost and no sexual dimorphism in jaw length; teeth small, sharp and curved, in several rows; genital papilla in female short, rounded, often bulbous, and short and pointed in male; gut short, with one simple loop; body small Tamanka maculata Aurich, 1938: 154 (Timpuk-See, Jolo, Sulu Province, Philippines). The syntypes were not found at ZMH during my visit in 1988, and are possibly lost. The ZMH collection was stored in subways during World War 11, and a number of bottles were stolen for the alcohol they contained (Or H. Wilkens, personal communication). The species is probably a Mugilogobius or Tamanka, and may be a sisterspecies to T. siitensis, based on Aurich's description. Tamanka umbra Herre, 1927: 223 (Palawan, Philippines ). The holotype (BSM 10600) and six paratypes (4260 mm SL, BSM 26893-98) were destroyed in WWII. Koumans (1940: 188) listed catalogue numbers of types and stated that they "agreed with the a Figure 195 Pandaka pygmaea, papillae pattern. CAS 47916, Malabon, Philippines. Scale bar = 1 mm. H.K. Larson 198 Figure 196 Pandaka pygmaea, 11 mm SL, CAS 47916, Dagat-Dagatan Lagoon, Malabon, Philippines. and slender with small, short-based fins; body with one to several dark bands and spots, with row of partly subcutaneous spots along lower mid-line of caudal peduncle (Figure 196). Pterygiophore formula 3-12210; modally two epurals, often partly fused together dorsally; vertebrae 10+14-15, modally 10+15; two to four preanal pterygiophores, usually two; neural spines of first few vertebrae slender, pointed; palatine and pterygoid relatively stout, almost equal in length, palatine not reaching quadrate; metapterygoid very low, slender, short, never reaching toward quadrate (Figure 197); quadrate very low; fifth ceratobranchial relatively slender, with high flange on ventral surface; scapula unossified; occasionally one ossified gill-raker; first epineural inserting just behind parapophysis. Remarks Birdsong et al. (1988) placed Pandaka and Brachygobius in their "Gobiopterus Group" of gobies, based on their sharing a low vertebral count (10+15=25), two epurals and a dorsal pterygiophore formula of 3-12210. The only other genera in their "Gobiopterus Group" were Gobiopterus and Mistichthys, two "neotenous" genera united by other characters. Pandaka and Brachygobius, despite their low vertebral count, belong with Birdsong et aI.'s "Gobionellus Group". The genus is distributed from the Indian Ocean and western tropical Pacific, in estuarine and freshwater habitats (Figure 198), free-swimming in small groups or dense schools over sand and mud bottoms, typically in mangroves. This genus will be formally revised in a separate paper (in preparation). PRELIMINARY LIST OF VALID SPECIES Pandaka lidwilli (McCulloch, 1917) Figure 198 Gobius Iidwilli McCulloch, 1917: 185, plate 21, figure 3 (Cowan Creek, New South Wales, Australia). Pandaka pusilla Herre, 1927 Figures 197-198 Pandaka pusilla Herre, 1927: 197, plate IS, figures 12 (Sitankai, Philippines). Pandaka pygmaea Herre, 1927 Figures 195-196, 198 Pandaka pygmaea Herre, 1927: 198, plate IS, figure 3 (Philippines [possibly from Malabon]). Pandaka rouxi (Weber, 1911) Figure 198 Gobius rouxi Weber, 1911: 40, figure 9 (Panua Bori River near Sungi Manurnbai, Wokam; Waskai River near Sungi Manumbai, Wokam; Sungi, Waskai, Wokam; Dungi Kololobo, Kobroor; near Seltutti, Kobroor, Aru Island, Indonesia). Pandaka silvana (Bamard, 1943) Figure 198 Gobius silvanus Barnard, 1943: 258, figure 33 (Knysna Lagoon, South Africa). Pandaka minuta Smith, 1959: 205, figure 23 (Ibo and Warnizi Islands, Mozambique). Figure 197 Jaws and suspensorium of Pandaka ptlsilla, male, ex CAS 38588, Coron, Philippines. Scale bar = 1 mm. Pandaka trimaculata Akihito and Meguro, 1975 Figure 198 Pandaka trinwcuIata Akihito and Meguro, 1975b: 63, ｾ (') <: (ji' O· ｾ ::l p.t? Cf)?J. (0' :"* 30' o ..,., 1.w' ::: I1JAPAN ｾＮＬ C§, ｾ C" it . '" , ..:- 20· (l HAWAIIAN ISLANDS ｾＧｌｬｐｦＱｎｅｓ } oｾ . : MAR1ANA IS Ｇｖ｛ｪｾｃａｾｏｬｎｅ " 10' ｍａｾｈｬｉｓ :,' IS: Ｈｾ Kosroe ;' .. <"<1'«-1< KIRIMTJ "s 0' 0' J'::l:"SOlOMON IS. ..•. "\ "o.V 10' 10' •...: ＧｾＮ 20' AUSTRALIA ..... SAMOA ｃｊｾＧ FIJI ; . • 20' 'LOYALTY IS. 30' 30' v .w' 1.w' Figure 198 Type localities of species of Pandaka. TASMAN5fA 160' ell! o .w' 180' 160' .... '.t:l '.t:l 200 H.K. Larson figures 1-3A (mouth of Miyara River, Ishigakijima, Okinawa Prefecture, Japan). Pseudogobius Popta, 1922 Pseudogobius Popta, 1922 (Gobius javanicus Bleeker, 1856: 88, Java, by subsequent designation, by Aurich 1938: 158). Lizagobius Whitley, 1933 (Mugilogobius galwayi McCulloch and Waite, 1918: 50, plate 3, figure 1, South Australia, by monotypy). Diagnosis Distinguished by following combination of characters. Second dorsal fin rays 1,6-9; anal fin rays 1,6-9; second dorsal and anal fin rays modally equal in number; pectoral rays 14-17; segmented caudal rays 16, in 9/7 pattern; 24-30 scales in lateral series; predorsal scales 6-11, usually with large scale Figure 199 Figure 200 Pseudogobius javanicus, papillae pattern, WAM P.30806-003, male, Maratua Island, Kalimantan Timur. Scale bar = 1 mm. Pseudogobius o[on/m, papillae pattern, male, AMS 1.20190-003, Chap man River, Kangaroo Island, South Australia. Scale bar = 1 mm. Figure 201 Jaws and suspensorium of Pseudogobius javanicus, female, NTM 5.11125-029, Sanur Beach, Bali. Scale bar = 1 mm. Figure 202 Jaws and suspensorium of Pseudogobius melanostictus, male, USNM 268186, Negros Oriental, Philippines. Jaws artificially gaped for clarity. Scale bar = 1 mm. present close behind eyes; no fine villi on head; rear portion of oculoscapular canal absent, no preopercular pores; papillae on head in longitudinal pattern, cheek papillae in rows a, C and cp larger and more widely spaced than those in rows b and d (Figures 199-200); gill opening restricted; pectoral girdle smooth or with low flange, without distinct fleshy lobes or flaps; tongue tip blunt to rounded; long intestine and stomach coiled in corkscrew manner about each other; mouth often inferior, with inflated snout overhanging upper lip; jaws usually short with thin to almost absent lips; teeth shape and arrangement sometimes differing between sexes; genital papilla elongate and flattened in male, and short and rounded in female. Pterygiophore formula 3-12210; vertebrae 10+1517; two epurals; two to three (modally two) pre-anal pterygiophores; first few neural spines usually slender, pointed, tips sometimes expanded or bifid; metapterygoid short and wide, often expanded " III :So '"o' ::l eo- o A ;'" Ｚｾ .lO' ｾＮ 0[ . tvUKYU IS. ｾ 30'. ::r. ;:: '" . l: .... ｾ 20' ｾＩ LACCJJ)/\/E IS. ': .. Makftv., ｓｾｉ • : ......IIANA IS. ｾＩｪＧ :: : ':, ＬＺｾＬｾｒｓｈａｕ l5. 10" ':"KIlII8A n 0' ... : CAROUNE IS. !COStCMI 0' ;;Ji\ ｾ .... SlYCHEUIS .; ·t 10' .. ';-1 u 20' ... :." '. SOlOMON l5. .. Ｂｾｑ｜ 10' " .. , , Mourllivs ......DAGASCAR avicennia javanicus ... melanosticta A oIorum • poicilosoma * • ｾＺＧ ,..-" \j .lO' Figure 203 .lO' 60' ｦＱｾＹＮＧＺ LOYALTY fS. ,', :: .. 20". 30'. 30' 20" SAWOA Ｎｾ • .... AO': IlO" Distribution of Pselldogobills species examined (in press n.b.: P. masago inadvertently omitted). N o .... 202 H.K. Larson dorsally but not forming bridge to quadrate; palatine short and broad, only reaching partly down pterygoid; pterygoid always much shorter than palatine (Figures 201-202); fifth ceratobranchial slender to broad, flattened and porous with thin fine teeth, low ridge or flange on ventral surface; none or up to five gill rakers ossified; top of scapula above foramen occasionally ossified. Remarks Popta (1922) first created the name Pseudogobius, including within it Gobius javanicus Bleeker and Pseudogobius penango Popta, but did not specifically designate a type species for the genus. Gobius javanicus is a valid species of Pseudogobius and P. penango is a species of Redigobius as currently understood. Koumans (1931) used the name Pseudogobius, based on a "museum name" of Bleeker's, in synonymy with Stigmatogobius. He included under Stigmatogobius two other museum names: G. isognathus Bleeker and Pseudogobius neglectus Bleeker. Koumans did not refer to Popta. Aurich (1938) referenced the genus as: "Pseudogobius (Bleeker, Museumsname) Aurich (= Stigmatogobius Blkr. partim.). Typ: javanicus Blkr." In his description of species of Vaimosa, the term "Typ" is used as being typical of a group, rather than as type species and the word Type is used for type species; however, elsewhere in his paper, "Typ." also can refer to type species. It is uncertain, therefore, if Aurich's "Typ: javanicus Blkr" is a type species designation. Aurich attributed the genus to himself, despite apparently being aware of Popta's work (he refers to penango Popta, but Popta does not appear in his bibliography). It could be argued that Pseudogobius Aurich is a homonym of Pseudogobius Popta and that no type species has been designated. Herre (1953b) referred to Koumans' discussion of Stigmatogobius and again included Pseudogobius as a synonym of Stigmatogobius. Akihito and Meguro (1975a), in their discussion of Pseudogobius in Japan, concluded that Pseudogobius Popta was the correct generic name associated with the holotype of Gobius javanicus: "As stated by Aurich (1938: 160), the generic name Figure 204 Pseudogobius Popta should be adopted for this species...". Chatterjee (1980), in a footnote, agreed with their statements. Eschmeyer and Bailey (1990: 339) and Eschmeyer (1990: 648) list all three Pseudogobius names and their authors and indicated that if javanicus was a valid species, it should be designated as type of Popta's genus. In the interests of stability Gobius javanicus Bleeker is so designated here as the type species of Pseudogobius Popta. The genus has some resemblance in appearance to Redigobius, which differs by having a simple uncoiled gut, greater development of the oculoscapular canal, often having a compressed head and body, in the larger lips and mouth, the jaws often greatly exaggerated in males, and it exhibits mostly free-swimming behaviour (hovering). Pseudogobius is widely distributed in the tropical to warm temperate Indo-west Pacific, (Figure 203), in freshwater and estuarine habitats over muddy substrates, seagrass beds or in mangroves. This genus will be revised in full in a separate paper (in prep.); a preliminary list of probably valid species of the genus are given below. PRELIMINARY LIST OF VALID SPECIES Pseudogobius avicennia (Herre, 1940) Figure 203 Vaimosa avicennia Herre, 1940a: 17, plate 12 (Kranji River, Singapore). Mugilogobius avicennia: Tan and Tan 1994: 356. Material Examined Holotype CAS 33006,1(31.5), Kranji River, Singapore, A.W. Herre, 10 March 1937. Pseudogobius javanicus (Bleeker, 1856) Figures 12£, 20B, 21B, 199,201,203-204 Gobius javanicus Bleeker, 1856: 88 (Patjitan, central Java). Vaimosa piapensis Herre, 1927: 147, plate 10, figure 3 Pseudogobius javanicus. 28 mm SL, CMK 6083, Tanjung Pasir, Java. Revision of Mugilogobius (Piapi Creek, Dumaguete, Oriental Negros, Philippines); Herre 1953b: 768. Stigmatogobius javanicus: Herre 1953b: 764-768. Pseudogobius javanicus: Bleeker 1983: plate 438, figure 14; Tan and Tan 1994: 356. Material Examined Holotype of Gobius javanicus RMNH 4549, 1(34), in river, Patjitan [= Tanjung Pacinan], Java, Indonesia; 1879. Other Material Indonesia: NTM S.11125-016, 111 (18-33), mangroves at Send Sanur beach, Ball, H. Larson, T. Gloerfeldt-Tarp, P. Kailola, 10 June 1983; ZSM/CMK 6083,4(26-29.5), S of Tanjung Pasir, Tanjerang District, West Java, M. Kottelat and F. Yuwono, 29 May 1988. Singapore: CMK 6012, 24(12-31), canal into sea, W coast, M. Kottelat and P. Ng, 21 May 1988. Pseudogobius masago (Tomiyama, 1936) Gobius ornatus masago Tomiyama, 1936: 73-74, figure 26 (coast of Tiba-Ken, Japan). Material Examined Paratype, ZUMT 34087, Chiba, Japan. Pseudogobius melanostictus (Day, 1876) Figures lIE, 160,202-203) Gobius melanosticta Day, 1876: 290, plate 63, figure 2 (Madras, India). Vaimosa serangoonensis Herre, in Herre and Myers 1937: 40, plate 2 (creek at Serangoon, Singapore). Vaimosa adyari Herre, 1945e: 402 (Adyar River, India). Material Examined Syntypes of Gobius melanosticta India: BMNH 1889.2.1.3388-97, 11(18-33), Madras, F. Day; NMW 84081.1-2, 2(22.5-36), Madras; MNHN A18, 1(32), Madras, F. Day. Holotype ofVaimosa serangoonensis CAS 30984, 1(28.5), Serangoon, Singapore, AW. Herre, 18 March 1934. Holotype ofVaimosa adyari CAS 39864, 1(21), Adyar River, Madras, India, AW. Herre, 4 January 1941. Other Material Sri Lanka: ZMH 19309, 4(20-32), lagoon, Panadhure, Duncker, 30 July 1909. Singapore: SMF 18199, 1(40), H. Berkenkamp, July 1983. 203 Philippines: USNM 268186, 11(27-32.5), Siquijor Island, Negros Oriental, L. Knapp and party, 14 May 1979. Indonesia: CMK 6286, 3(31.5-35.5), Pangandaren, Java, Vivaria Indonesia, 8 July 1988; CMK 4546, 10(14-31), Bungus Bay, near Padang, Sumatra, P. Bianco and M. Kottelat, 29 November 1984. Papua New Guinea: USNM 316051, 2(27-35), stream behind Trobriand Hotel, Kiriwinna, Trobriand Islands, B. Collette, 6 June 1970. Pseudogobius olorum (Sauvage, 1880) Figures 200, 203 Gobius olorum Sauvage, 1880: 43 (Swan River, Western Australia). Mugilogobius galwayi McCulloch and Waite, 1918: 50, plate 3, figure 1 (Patawalunga Creek, near Adelaide, South Australia). Material Examined Syntypes of Gobius olorum MNHN A.1913, 3(38-41), river in Australia, Castelnau. Holotype ofMugilogobius galwayi SAMA F.583, 1(50.5), Patawalonga Creek, near Adelaide, South Australia. Other Material Australia: Western Australia: NTM unregistered, 13(35-40), Swan River, H. Gill, 27 November 1984; NTM unregistered, 5(36-52), Swan River, H. Gill, 30 January 1985. South Australia: SAMA F.5123, 7(2744.5), near Longpoint, Coorong, Strathalbyn Naturalist's Club, 11-12 February 1984. Pseudogobius poicilosomus (Bleeker, 1849) Figure 203 Gobius poicilosoma Bleeker, 1849: 31 (Pasuruan, Java). Material Examined Syntypes of Gobius poicilosoma RMNH 4488, 7(22-31), Pasuruan, Java, Indonesia, 1847. Other Material Papua New Guinea: USNM 316208, 3(30.5-32.5), Dogura Creek, SE of Port Moresby, B. Collette, 21 June 1970; BPBM 15836, 2(32.5-33), Meiro River, Madang, G. Alien, J. Randall and R. Steene, 15 August 1973. Northern Territory: NTM 5.12429015, 63(15-25), creek on E side Vanderlin Island, H. Larson and W. Houston, 22 July 1988. Queensland: ROM 38801, 24(14-32), 5aunder's Beach, N of Townsville, R. Winterbottom, 4 October 1981. 204 H.K. Larson Redigobius Herre, 1927 Redigobius Herre, 1927 (Gobius sternbergi Smith, 1902b: 169, fig., Lake Buhi, Philippines, by monotypy). Ostreogobius Whitley, 1930 (GiIlichthys australis Ogilby, 1894: 367, New South Wales, Australia, by original designation and monotypy). Parvigobius Whitley, 1930 (Parvigobius immeritus Whitley, 1930: 122, replacement name for Gobius flavescens De Vis, 1884: 689, Moreton Bay, Queensland, by original designation and monotypy). Microgobius Koumans, 1931 (listed as synonym of Stigmatogobius; Gobius hoevenii Bleeker, 1851b: 426, Sambes River, Indonesia, and Gobius tambujon Bleeker, 1854: 319, Indonesia). Preoccupied by Microgobius Poey. Cyprinogobius Koumans, 1938 (Lophogobius chrysosoma Bleeker, 1875: 114, Borneo, by original designation). Diagnosis Distinguished by following combination of characters. Dorsal fin rays 1,6-8, anal fin rays 1,6-7; modally with one more ray in second dorsal fin than in anal fin; pectoral fin rays 15-20; 17 segmented caudal rays, in 9/8 pattern; lateral scales 22-30; TRB 7-11; predorsal scales often large, 6-16, extending close up to behind eyes; sensory canals on head with complete oculoscapular and preopercular canals and pores; sensory papillae arrangement variable, modally arranged in longitudinal pattern (Figure 205), one species with partly transverse pattern (Figure 206); rows a and c consisting of large, widely spaced papillae in some species, usually all papillae small and closely Figure 205 Redigobius dispar. Headpores and papillae pattern, male, USNM 263330, Lake Buhi, Luzon, Philippines. Scale bar = 1 mm. Figure 206 Redigobius chrysosomus. Headpores and papillae pattern, male, AMS 1.24683-003, Blackmore River, NT. Scale bar = 1 mm. spaced, as in row p if present, usually replaced by headpores; rows b and d always consisting of small close-set papillae; row c broken under eye, with rear portion of row modally consisting of one papilla; no fine villi on dorsal surface of head; pectoral girdle smooth, with few fleshy knobs or lobes, or with low flange on anterior edge of cleithrum; jaws terminal, males with (sometimes greatly) enlarged mouths; anterior nostril in tube, close to edge of upper lip or distinctly overhanging lip; genital papilla variable in male, flattened and square, triangular or elongate in shape, and in female, short, rounded and bulbous; simple short gut with one loop present. Dorsal pterygiophore formula 3-12210; 25-27 vertebrae, 11-12 precaudal and 14-16 caudal; two epurals; three or four, rarely two, anal fin pterygiophores before first caudal haemal spine; neural spines of first few vertebrae modally slender and pointed, sometimes bifid or slightly broadened at tip; palatine and pterygoid usually slender, palatine sometimes quite short, palatine not quite reaching quadrate; metapterygoid short, occasionally expanded dorsalIy, not forming bridge to quadrate (Figure 207); rear edge of preopercle with posteriorly facing groove or ridge; fifth ceratobranchial stout, triangular, with stout teeth, short high flange on ventral surface; upper corner of scapula modally ossified above foramen; one to three gill-rakers ossified; first epineural inserting on or behind parapophysis of first vertebra. Remarks Miller (1987) erroneously included Sphenentogobius Fowler as a synonym of Redigobius (Sphenentogobius is a junior synonym of Hemigobius). A full revision of this genus is in progress as a separate project. There are 44 nominal species, of Revision of Mugilogobius 205 Other Material Sri Lanka: USNM 164012, 3(28-29), 1955; CMK 7184, 8(26.5-36), Bolagod Lake, R. Pethiyagoda, 14 December 1989. Queensland: AMS 1.22041-013, 15(18-27.5), Mowbray River, D.F. Hoese, H.K. Larson and G.R. Allen, 13 September 1980; AMS 1.22721-005, 15(19.5-26.5), mouth of Daintree River, G. Hardy and A Ayling, 26 June 1980. Redigobius bikolanus (Herre, 1927) Figure 208 Vaimosa bikolana Herre, 1927: 151, plate 11, figure 2 (Puru, Legaspi, Albay Province, Philippines). Gobius flavescens De Vis, 1884: 689 (Moreton Bay, Queensland, Australia) (primary homonym of Gobius flavescens Fabricius). Parvigobius immeritus Whitley, 1930: 122 (replacement name for Gobius flavescens De Vis). Vaimosa horiae Herre, 1936b: 280 (Bab-el-Thuap, Pelew Group) (in part). Vaimosa osgoodi Herre, 1935c: 420 (Viti Levu Island, Suva Harbour, Fiji). Vaimosa montalbani Herre, 1936c: 359, plate 1, figure 3 (Lake Naujan, Mindoro, Philippines). B Figure 207 Jaws and suspensoria of: A, Redigobius dispar, male, ex USNM 263330; B, Redigobius macrostomus, female, ex AMS I.19341-002, Clarence River, New South Wales. Scale bar = 1 mm. which about 11 may be valid. A preliminary list of probably valid species is provided. Some of the descriptions of the seven nomina dubia listed under Mugilogobius could equally apply to species of Eugnathogobius or Pseudogobius as well as Redigobius, so caution is required when dealing with these names. Redigobius species occur in Indo-west Pacific estuarine and fresh waters; if in fresh water, often found close to tidal influence (Figure 208). PRELIMINARY LIST OF VALID SPECIES Redigobius balteatus (Herre, 1935) Figure 208 Vaimosa balteata Herre, 1935a: 419 (Majalibit Inlet, Waigiu Island). ?Acentrogobius balteatops Smith, 1959: 200, plate 9G (Inhaca, Mozambique). Material Examined Holotype of Vaimosa balteata FMNH 17386, 1(21), Majalibit Inlet, Waigiu [Irian Jaya], Indonesia, AW. Herre, 7 June 1929. Pseudogobius bikolanus: Aurich 1938: 161. Gobius johnstoniensis Koumans, 1940: 166 (Johnstone River, Innisfail, Queensland, Australia). Mahidolia pagoensis Schultz, 1943: 229, 240, figure 20 (Pago Pago, Tutuila Island, Samoa). Vaimosa novaehebudorum Fowler, 1944: 180, figures 27-28 (New Hebrides). Stigmatogobius minutus Takagi, 1957: 114, figure 5, plate 6E (Tomari estuary, Satsuma Peninsula, Kagoshima Prefecture, Japan). Stigmatogobius versicolor Smith, 1959: 197, figure 12 (Mahe, Seychelles). Redigobius bikolanus: Tan and Tan 1994: 357. Material Examined Holotype ofVaimosa osgoodi FMNH 17387, 1(16), river entering Suva Harbour, Viti Levu, Fiji, AW. Herre, 13 March 1929. Holotype ofVaimosa montalbani CAS 30967, 1(20), Lake Naujan, Philippines, AW. Herre, 28 November 1933. Holotype of Mahidolia pagoensis USNM 116113, 1(15), stream at village of Pago Pago, Tutuila Island, Samoa, L.P. Schultz, 2 June 1939. ｾ Cl' 180' 4lr. 30' ｾ 20' ｾ LACCADIVE IS, .' • '. 10' Maldlve. ｲｾＧｾｍ｜ .... 20' '. MARlANAIS, ,.:.,., .: . ::":....:i ... " CAROUNE IS.' Kosrae .: IORIIlATI .' 0' * balteatus [) '. 20' , MADAGASCAR 30' . . . :;gs::I::l" •:; SEYCHEUES ': 10' Mauritius Cl ' \' .1 , '. ') .,: I er. MARSHAlL IS. t ";":''. • bikolanus • chrysosoma • dewaalii dispar o ieptochilus o macrostoma l::" penango 'Y roemeri '* 0' ＢＺＧｾＮ SOLOMON IS• c:\ . ". ,.,, ｾＮ .' ｙｬａｏｾ 1er. SAMOA 4 ＬＢｾｉｊｆ If:l ,., '. ." .: 20' IS, 30' o 40' 120' 140' .• D TAW.AN!E.4 = ｾ r Figure 208 Distribution of Redigobius species, based on material examined, U1 o ::: Revision of Mugilogobius Holotype of Vaimosa novaehebudorum ANSP 71392, 1(21), New Hebrides, E. Jackson, 1943. 207 Mugilogobius pongolensis Kok and Blaber, 1977: 163, figures 1-2 (Nsimbu River, Pongolo River floodplain, 2ululand, South Africa). Other Material Material Examined Philippines: CAS 26377, 76(10.5-21), Lake Bombon, Luzon, A.W. Herre, May 1931; CAS 37648, 7(20.5-27.5), Old Ayukitan, Dumaguete, A.W. Herre, 7, 13 August 1940. Belau: FMNH 47241, 2(22-26), Babelthuap Island, A.W. Herre, 14 October 1933. New Caledonia: MNHN 1992.428, 14(22.529), River Pirogues, Southern Province, PEDCAL 62,2 October 1991. Lectotype of Gobius dewaalii Redigobius chrysosomus (Bleeker, 1875) Figures 206, 208 Lophogobius chrysosoma Bleeker, 1875: 114 (Bandjermasin, Borneo and Amboina). Cyprinogobius chrysosoma: Koumans 1938: 12. Material Examined RMNH 4489, 4(35.5-37), Bandjermasin, Borneo, Indonesia. Other Material South Africa: AMS 1.27220-001, 25(12-56), Nhololo Dam, Pongola River, H. Kok, 17 July 1977; AMS 1.27101-001, 4(23-26.5), Potter's Channel, St Lucia system, Natal, H. Kok, 11 September 1976; SAM 2.47173, 3(24.5-32), east shore SLake, St Lucia, M. Wapenaar, June 1964. Redigobius dispar (Peters, 1868) Figures 14, 18C, 205, 207A, 208-209 Gobius dispar Peters, 1868: 264 (Luzon, Philippines). Buhi, Philippines). Pseudogobius dispar: Aurich 1938: 164. Cyprinogobius dispar: Koumans 1938: 13. Vaimosa dispar: Herre 1953b: 767. Australia: Northern Territory: AMS 1.24683-003, 139(15.5-26), Blackmore River, D. Hoese, S. Reader and D. Beechey, 8 September 1984. Queensland: AMS 1.22044-003, 5(14-29), upstream from mouth, Daintree River, D.F. Hoese, H.K. Larson, G.R. Allen and W. Starck, 14 September 1980. Redigobius dewaalii (Weber, 1897) Figure 208 Gobius dewaalii Weber, 1897: 144 (Umgeni River, Natal, South Africa). Gobius maxillaris Davies, 1949: 375 (Knysna River, Figure 209 Other Material Gobius sternbergi Smith, 1902b: 169, figure (Lake Syntypes of Lophogobius chrysosoma South Africa). ZMH 103.238, 1(28), Umgeni River, Natal, South Africa, M. Weber, 1894. Material Examined Syntypes of Gobius dispar 2MB 6703, 4(25-36), Luzon, Jagor; 2MB 6702, 7(29-42), same data as preceding; 2MB 6700, 11(3140), Batu Lake, Jagor, Philippines. Syntypes of Gobius stembergi USNM 50536, 3(17.5-24), Lake Buhi, Luzon, Philippines, F.W. Richardson, 5 July 1901. Other Material Philippines: USNM 263330, 29(14.5-32), Lake Buhi, Camarines Province, T. Roberts, 27 April 1976. Redigobius dispar, CMK 11334, 38 mm SL, Halmahera Island, Maluku, Indonesia. H.K. Larson 208 Redigobius leptochilus (Bleeker, 1875) Figure 208 Acentrogobius leptochilus Bleeker, 1875: 131 (Amboina). ?Gobius oyensi Beaufort, 1913: 137, figure 4 (Buru and Cerarn). ?Lophogobius wera Popta, 1922: 27 (Wera River, Sumbawa, Sunda Archipelago, Indonesia). Material Examined Syntypes of Pseudogobius penango SMF 6580-1, 2(26-30), Rumbia, J. Elbert, 1909; SMF 6579, 1(29.5), Mengkoka, Penango village, Indonesia, J. Elbert, October 1909; RMNH 10666,1(29.5), Rumbia, J. Elbert, Sunda Expedition 1909-1910. Paralectotype of Gobius latifrons ex NMBA 1847-52, 19.5 mm SL female, Lake Matanna and Kalaena River, Indonesia. Material Examined Holotype of Acentrogobius leptochilus RMNH 4663, 1(38.5), Amboina. Syntype of Gobius oyensi ZMA 113.263, 1(44), upper course of Toebah River, western Cerarn, Indonesia, L.F. de Beaufort, 27 February 1910. Holotype of Lophogobius wera SMF 6551, 1(40), Wera River, Sumbawa, Indonesia, J. Elbert, 12 December 1909. Other Material Indonesia: NTM 5.14154-001, 1(26), Tongoloka River, Sumbawa, K. Martin, 1 April 1995. Other Material Indonesia: Sulawesi: CMK 6143, 53(12.5-39.5), on road from Palopo to Rantepao, Tandung River, Kabupaten Luwu, M. Kottelat, 12 June 1988; CMK 6444, 8(15-30), Sungei Torak, 19 km on road from Enrekang to Pare Pare, M. Kottelat and A. Werner, 9 March 1989; CMK 11395, 8(1334.5), un-named tributary entering Salo Larona from the south about 300 m down-river of bridge in road to power plant, M. Kottelat, 12 February 1995; RMNH 12076, 2(41-42), unspecified locality. Redigobius roemeri (Weber, 1911) Figures 9C, 22C, 208 Redigobius macrostomus (Giinther, 1861) Figures 2078,208 Gobius macrostoma Giinther, 1861: 44 (Australia). Secondary homonym of Gobiopsis macrostomus Steindachner, 1861. Gobius microphthalmus Giinther, 1861: 550 (replacement name for Gobius macrostoma Giinther,1861). Gillichthys australis Ogilby, 1894: 367 (New South Gobius roemeri Weber, 1911: 39, figure 8 (Panua Bori River near Sungi Manumbai, Wokam and Matora River near Sungi Manumbai, Wokarn, Aru Is, Indonesia). Gobius reticularis Weber, 1911: 39, figure 7 (Walde near Wokamar, Wokam; Matora River near Sungi Manumbai, Wokam; near Papakula, Kobror; near Seltutti, Kobroor, Aru Island, Indonesia) (in part). Cyprinogobius micropthalmus: Koumans 1938: 13. ? Vaimosa macrognatha Herre, 1927: 145, plate 10, figure 2 (Lake Taal, Batangas Province, Philippines). Material Examined Vaimosa horiae Herre, 1936b: 280 (Bab-el-Thuap, Holotype of Gobius macrostoma Pseudogobiopsis roemeri: Herre 1953b: 761 (Bungau, Wales, Australia). Pelew Group) (in part). BMNH 1953.12.31.6, 1(33), Australia, J. Gould. Sulu Province). Other Material Material Examined Australia: New South Wales: AMS I.16954-018, 10(18.5-30), Cowan Creek, Hawkesbury River, J. Paxton and H. Recher, 22 November 1972. Syntype of Gobius roemeri Redigobius penango Popta, 1922 Figure 208 Pseudogobius penango Popta, 1922: 36 (Rumbia and Penango, Great Sunda Island, Celebes). SMF 6703, 1(34), Sungei Manumbai, Panura Boribei, River, Wokam, Aru Islands, Indonesia, H. Merton, 14 March 1908. Syntype of Gobius reticularis ZMA 112.661, 17 mm SL female, forest creek near Woakmar, Aru Island, Indonesia, H. Merton, 17 March 1908. Revision of Mugilogobius Syntypes ofVaimosa horiae CAS 29070, 2(26.5-29.5), Bab-€l-thuap, Y. Hori, 14 October 1933; CAS 29071, 9(21.5-28), same data as preceding. Other Material Philippines: USNM 31635, 4(15-33), Casagatan River, Zambales Province, Luzon, T. Roberts, 14 March 1976; FMNH 50948, 12(12.5-37), Lapulapu River, Lapulapu, Iwahig, Palawan, Werner, 4 March 1947. Belau: CAS 76087, 21(22.5-40), Arakitaoch Stream, Airai, Babelthuap Island, Sumang and Marbon, 27 November 1956. Stigmatogobius Bleeker, 1874 Stigmatogobius Bleeker 1874: 323 (type species Gobius pleurostigma Bleeker 1849: 28, Surabaya, Borneo, by original designation and monotypy). Diagnosis Distinguished by following combination of characters. Always with distinctive transverse sensory papillae pattern (Figure 2); second dorsal and anal rays 1,7-8, usually one more anal fin ray than in second dorsal fin; 15-17 segmented caudal rays, modally 17, in 9/8 pattern; pectoral rays 1321; first dorsal fin relatively tall, pointed; 12-14 circumpeduncular scales; one enlarged predorsal scale close behind eyes, other scales much smaller; headpores absent or reduced, infraorbital pore absent, rear portion of oculoscapular canal usually absent, interorbital canals separate, not joined posteriorly, anterior interorbital pores and preopercular pores present or absent; no fine villi on head; tongue blunt to bilobed; anterior nostril in tube, not always extending downward over upper lip; shoulder girdle usually smooth, rarely with bony flange or small fleshy flaps present; genital papilla slender, elongate and flattened in male, short and bulbous in female; gut short, with one simple loop; body colouring striking, pale with black spots or vertical bar. 209 Pterygiophore formula 3-12210; one or two epurals (usually two); vertebrae 10-12+15-16; usually two (sometimes one) anal pterygiophores before first haemal spine and first haemal spine curving around the second anal pterygiophore in all but one species; first few neural spines narrow, pointed; high number of procurrent caudal rays (9/8 to 10/10); metapterygoid broad, expanded dorsally, often partly overlapping quadrate (Figure 210); palatine and pterygoid relatively slender, palatine nearly reaching quadrate; upper part of scapula ossified; low ridge or groove along rear edge of preopercle; fifth ceratobranchial slender, stout teeth present, high flange on ventral surface. Remarks The first haemal spine is curved around the second anal pterygiophore in three species (Figure 19), reminiscent of the arrangement seen in the ptereleotrine microdesmids (Hoese, 1984; Birdsong, et al., 1988). The absence of an infraorbital pore may appear to place Stigmatogobius closer to the Awaous/ Stenogobius group of gobionellines than to the Mugilogobius group, but it falls out with the latter in phylogenetic analyses. The species of this genus will be formally revised in a separate paper (in prep.). Stigmatogobius are robust-bodied gobies which hover above the substrate in fresh to estuarine waters of the Indo-Malayan region (Figure 211). Miller et al. (1980) discussed phenetic clustering relationships of Stigmatogobius, and used two species for their haemoglobin electrophoresis analyses of the "knight goby" of the aquarium trade. One they identified as S. sadanundio while the other, unidentified, was likely to have been S. pleurostigma; both species are often confused in the aquarium trade and fish-keeping magazines. PRELIMINARY LIST OF VALID SPECIES Stigmatogobius borneensis (Bleeker, 1851) Figure 211 Gobius borneensis Bleeker, 1851a: 10 (Banjermassing, Borneo). Stigmatogobius singapurensis Bleeker, 1878: 204 (Singapore). ?Gobius beccarii Perugia, 1892: 1010 (Sarawak). Material Examined Syntypes of Gobius borneensis RMNH 6175, 8(36-45), Banjermassing, Borneo, Indonesia, 1849. Figure 210 Jaws and suspensorium of Stigmatogobius sadanundio, male, ex CMK 6278, Medan, Sumatra. Scale bar = 1 mm. Holotype of Stigmatogobius singapurensis RMNH 4660, 1(36.5). 80' 50' ARABIAN SEA o 0, t: LACCADIVE ';', ISLANDS • ' 10' o . o ", '<;) ,.. Ｎｾ I', Maldives . .' '0 I' .:; 0" 0" ',' , ••• * SEYCHELLES . ... , .,' borneensis • pleurostigma • sadanundio ' .: ....- . ｾ 10j ｾ＼ｩＧｏｾ C::::::;;:;"'2r:J;? ｾ pO''' '""""" o o ,08 0 0" Ｂｾ '. o ｾＬ 60' 80' 100' :x ｾ Figure 211 Distribution of Stigmatogobius species, based on material examined. rU1 o ::I Revision of Mugilogobius Figure 212 211 Stigmatogobius pleurostigma, ZMA 120.459, 48 mm, 5L Tambak 5umur, Java. Remarks The placement of Gobius beccarii Perugia here, as a possible synonym of S. borneensis, follows Tortonese (1963). Stigmatogobius sadanundio (Hamilton, 1822) Figures 2, 19,211,213 Gobius sadanundio Hamilton, 1822: 52 (estuaries near Calcutta). Gobius apogonius Cantor, 1850: 1164 (Pinang). Stigmatogobius pleurostigma (Bleeker, 1849) Figures 19, 211-212 Gobius pleurostigma Bleeker, 1849: 28-29 (Surabaya, Java). Vaimosa spilopleura Smith, 1933: 66-68 (Chantabun River, southeastern Thailand). Material Examined Material Examined Pakistan: ZMA 100.117, 1(38}, Piali River, Ganges Delta, L.P. de Beaufort, 6 January 1938. Singapore: ANSP 77797, 4(34-51}, mullet pond, Pulo Ubin, Singapore Fisheries Department; CMK 8315, 4(4262}, mangrove near Sungei Buloh, M. Kottelat and D. Murphy, 8 April 1992. Indonesia: ZSM/CMK 6278, 4(37-48}, Medan, Sumatra, 8 July 1988. Syntypes of Gobius pleurostigma RMNH 6173, 33(28-64}, Surabaya, Java, Indonesia, 1848. Stigmatogobius sella (Steindachner, 1881) Gobius sella Steindachner, 1881: 212 (Borneo). Holotype ofVaimosa spilopleura ?Gobius beccarii Perugia, 1892: 1010 (Sarawak). KUMF 182, 1(45.5}, steamer landing, Chantabun estuary, Indonesia, H.M. Smith, 25 June 1931. Vaimosa brocki Herre, 1936a: 9 (Singapore Harbour). Material Examined Paratypes ofVaimosa spilopleura, Syntypes of Gobius sella Thailand: KUMF 1894, 4(34-49}, steamship landing, Chantabun estuary, July 1931; KUMF 1891, 1(37.6}, Tachin River, 16 December 1927. NMW 30107-30108, 2(33.5-40}, Indonesia, E. Pfeiffer. Other Material Holotype ofVaimosa brocki Indonesia: ZMA 120.459, 5(45-49}, fishpond, Tambak Sumur, Java, P.N. van Kampen, December 1906; CMK 7275, 11(10.5-30}, Sungei Lukit, Pandang Island, Riau Province, Sumatra, M. Kottelat, 12 February 1991. Figure 213 CAS 30965, 1(28.5}, Singapore harbour, A.W. Herre, March 1934. Other Material Indonesia: CAS 49464, 7(13-31}, Sungai Durian, Stigmatogobius sadanundio, 40 mm 5L, NTM 5.13954, Ao Nam Bor, Phuket, Thailand. 212 H.K. Larson Kapuas River basin, Borneo, T. Roberts and S. Woerjoatmodjo, 13 July 1976. Sumatra: CMK 7336, 17(8-26), estuary, Padang Island, Riau Province, M. Kottelat, 14 February 1991. Remarks The syntypes (MNSG C.E.12656) of Gobius beccarii Perugia have not yet been examined. Tamanka Herre, 1927 Tamanka Herre, 1927: 220, Jolo, Philippines (Tamanka siitensis Herre, 1927, by original designation). Diagnosis Relatively large, with second dorsal rays 1,6-9; anal rays 1,7-8, pectoral rays 15-28, always 17 segmented caudal rays, in 9/7 pattern, lateral scales 44-56; body with ctenoid scales posteriorly, usually extending up to behind pectoral fin in narrow wedge; predorsal scales small, cycloid, 11-27, all evenly sized and reaching forward past rear edge of preopercle; TRB 14-19; circumpeduncular scales 18-24; headpores always absent. Head papillae pattern longitudinal, rows composed of small, evenly sized papillae, papillae in rows a, c, cp and p small and closely spaced; long p row on top of snout and around eye forming "raised eyebrow" shape; on side of head, papilla row c broken under eye, rear portion consisting of two or more small papillae; three s rows present on snout, modally of three or more papillae, anteriormost row placed just above upper lip. Occasionally, dorsal surface of head with fine villi on interorbital space and top of snout. Gill opening oblique, pectoral girdle smooth, without lobes or fleshy flange. Mouth terminal; jaws slightly longer in males than in females. Snout broad, rounded, profile rounded to somewhat pointed. Anterior nostril in tube and oriented down and forward over upper lip, preorbital usually curving outward slightly around base of nostril. Genital papilla in male slender, elongate and flattened, broad at base and narrowing toward tip; papilla in female short, bulbous and stout, narrowing toward tip and broad at base, groove present along ventral surface. Simple short gut with one loop present. Body and fins very dark brown and almost no pattern discernible. Pterygiophore formula 3-12210; modally two epurals; two pre-anal pterygiophores. Vertebrae 26, 10 precaudal and 16 caudal. Neural spines of first three vertebrae stout, and often broadly expanded at tip. Palatine and pterygoid relatively slender, nearly equal in length, with palatine slightly longer, palatine reaching quadrate; quadrate somewhat forked, dorsal arm broad; metapterygoid broad and expanded dorsally (greatest expansion in mature males), upper limb overlapping quadrate. Fifth ceratobranchial stout, narrowly triangular, with curved sharp teeth and narrow triangular to pointed flange ventrally. Scapula unossified, fused to cleithrum. No gill rakers ossified. Glossohyal broadly spatulate. Remarks Herre (1927) created the genus Tamanka, describing four new species for the genus. He separated Tamanka from Vaimosa Jordan and Seale, 1906, due to the" much smaller and more numerous scales (38 to 54 in a longitudinal series), and by having many small cycloid scales on the opercles instead of a few large ctenoid ones" (Herre 1927). The species originally included in Tamanka by Herre (1927) were T. bivittata, T. siitensis, T. tagala and T. umbra. The types of these were housed in the Bureau of Science, Manila, and all holotypes were destroyed in World War 11. Paratypes of T. siitensis survived, however, as the USNM was sent a batch of 12 specimens. Herre also sent large numbers of non-type specimens of T. siitensis to CAS (which exchanged material with USNM, FMNH and AMS). From Herre's descriptions of the three "lost" species, T. bivittata can be identified as Mugilogobius abei Jordan and Snyder, 1901 (based on the description and figure); T. tagala is possibly a Mugilogobius, and T. umbra could be either Mugilogobius or Eugnathogobius. Koumans (1953) noted that Tamanka was very close to Mugilogobius, but provisionally retained the two as separate genera, because "According to Herre ... Tamanka differs from Mugilogobius by more numerous scales in the longitudinal line, 38-54 Table 34 Frequency distribution of second dorsal and anal fin ray counts in Tamanka siitensis Herre, 1927. Species Second dorsal rays 6 7 8 9 siitensis 1 Table 35 Frequency distribution of pectoral ray counts in Tamanka siitensis Herre, 1927. 1 35 Anal rays 7 8 2 2 Species 15 16 17 18 siitensis 1 13 22 2 Table 36 37 Frequency distribution of transverse backward scale counts in Tamanka siitensis Herre, 1927. Species 14 15 16 17 18 19 siitensis 1 7 10 16 4 1 Revision of Mugilogobius Figure 214 213 Tamanka siitensis. Neotype of Tamanka siitensis Herre, 64 mm SL male, USNM 87128, Lake Siit, Jolo, Sulu Archipelago, Philippines. instead of 37, and in having many small scales on the opercle instead of a few large scales" (Koumans 1953: 155). He did consider that "these characters are not so exact", and added Tamanka ubinensis Herre, 1937 (a junior synonym of Calamiana variegata), to the genus. The genus Tamanka was considered to be valid by Larson and Kottelat (1992). Material Examined Philippine Islands Sulu Archipelago, Jolo Island: Neotype USNM 87128, 1(64), Lake Siit, A.W. Herre, 11 June 1921. Paratypes Tamanka siitensis (Herre, 1927) Figures 6C, 90, llB, 16E, 92, 214-216; Tables 34-38 USNM 355555, 11(35-64), same data as neotype. Tamanka siitensis Herre, 1927: 220-222, plate 17, figure 3 (Lake Siit, north coast Jolo Island); Koumans 1931: 117; Aurich 1938: 152-154; Herre 1953b: 766; Koumans 1953: 160; Birdsong et al., 1988: 213. ?Tamanka maculata Aurich, 1938: 154 (Lake Tirnpuk, Other Material CASjSU 26368, 82(28-65), Lake Siit, colI. A.W. Herre, 18 July 1929; FMNH 40794-9, 6(34-41), Lake Panamao, A.W. Herre, 12 September 1940; CASjSU 38624, 64(22-52), same data as preceding; AMS 1.19194-001,15(34-40), same data as preceding. Jolo, Sulu Province). Table 37 Frequency distribution of lateral scale counts in Tamanka siitensis Herre, 1927. Species 44 45 siitensis 1 1 Table 38 46 47 48 49 50 51 52 53 54 55 56 3 3 6 4 3 4 2 8 2 1 Morphometries as percentages of SL or HL, as indicated, of Tamanka siitensis Herre, 1927. Character Head length in SL Head depth in HL Head width in HL Body depth in SL Body width in SL Caud. ped. \. in SL Caud. ped. d. in SL Snout length in HL Eye width in HL Jaw length in HL Interorbital\. in HL Pectoral \. in SL Pelvic \. in SL Caudal\. in SL Longest D1 spine in SL Neotype Males Minimum Males Maximum Males Mean Females Minimum Females Maximum Females Mean 36.9 58.1 66.1 21.7 33.1 50.0 49.5 17.5 10.0 21.5 11.4 24.0 18.4 34.4 22.9 19.1 17.2 21.7 11.0 39.4 62.2 66.1 23.2 14.1 26.7 14.5 28.7 28.1 47.5 34.3 26.2 22.0 26.6 22.0 35.9 53.1 54.8 19.9 12.3 24.0 12.5 25.8 22.4 39.7 27.3 21.7 19.4 24.5 13.8 33.2 55.9 57.2 22.4 14.2 28.2 13.3 27.4 28.0 44.9 30.1 24.4 21.4 26.8 13.8 37.5 55.9 57.2 22.4 14.2 28.2 13.3 27.4 28.0 44.9 30.1 24.4 21.4 26.8 13.8 35.1 52.5 53.8 19.5 12.2 25.4 12.2 24.0 24.9 36.8 25.4 21.3 19.0 24.6 13.0 22.7 13.0 28.0 23.3 47.5 34.3 23.0 19.5 25.3 22.0 214 Diagnosis As for genus. Description Based on 39 specimens, 28-65 mm SL. An asterisk indicates counts of neotype (Figure 214). First dorsal VI*; second dorsal 1,6-9 (mean 1,8*); anal 1,7-8 (mean 1,8*), pectoral rays 15-18 (mean 17*); segmented caudal rays always 17*; caudal ray pattern 10/7*; branched caudal rays 8/6 to 10/6 (modally 9/7); unsegmented (procurrent) caudal rays 8/9 or 9/9; longitudinal scale count 44-56 (54 in neotype, mean 51); TRB 14-19 (18 in neotype, mean 16); predorsal scales 11-27 (25 in neotype, mean 23); circumpeduncular scales 18-24 (20 in neotype, mean 21). Gill rakers on outer face of first arch 3+9 to 6+11 (modally 4+10). Dorsal pterygiophore formula 3-12210* (in four). Vertebrae 10+16 (in four). Neural spines of first three vertebra (at least second spine) broad, blunt-tipped and expanded dorsally. Metapterygoid relatively low, slightly expanded dorsally, in two small (41-41 mm SL) from Lake Panamao, expanded dorsally in two larger (51 and 73 mm SL) specimens from Lake Siit; distinct bridge to quadrate. Fifth ceratobranchial long and very slender; very short-based, tall flange present ventrally. Two epurals (in three); one in one specimen. Two (in four) anal pterygiophores before haemal spine of first caudal vertebra. Body relatively compressed, more so posteriorly. Head depressed anteriorly, always wider than deep, HL 2.5-3.0 (mean 2.9) in SL; profile bluntly rounded or pointed. Depth at posterior preopercular margin 1.6-2.1 (mean 1.9) in HL. Width at posterior preopercular margin 1.5-2.0 (mean 1.9) in HL, preopercular area sometimes slightly inflated in males. Mouth moderately large, terminal, slightly oblique, forming angle of about 25-30° with body axis; jaws reaching to below middle of eye. Upper jaw 2.1-2.9 (mean 2.7 in females, 2.5 in males) in HL. Lips fleshy, narrow, smooth, with fine fimbriae on inner edges of both lips; lower lip mostly free, fused near tip of jaw. Eyes large, placed dorsolaterally, sometimes forming part of dorsal profile; eye width 3.4-5.4 (mean 4.3) in HL. Snout broad, rounded in dorsal view; snout profile rounded to somewhat pointed; some specimens from Lake Siit with concave interorbital and snout with raised bump in centre formed by ascending premaxillary process; snout 3.5-4.6 (mean 4.1) in HL. Interorbital broad, flat to concave, 2.9-4.7 (mean 3.9) in HL. Occasionally fine villi on interorbital and snout. Body depth at anal origin 4.3-6.0 (mean 5.2) in SL. Caudal peduncle compressed, length 3.5-4.2 (mean 4.1) in SL. Caudal peduncle depth 6.9-9.2 (mean 8.1) in SL. First dorsal fin rounded, low, usually third or fourth spine longest; fin not reaching second dorsal origin when depressed and often falling well short H.K. Larson of fin origin. Longest dorsal spine length 6.9-9.1 (mean 8.0) of SL. Third dorsal spine length 4.5-9.1 (mean 7.9) in SL. Fourth dorsal spine length 4.5-8.6 (mean 8.0) in SL. Second dorsal fin usually taller than first dorsal fin; posteriormost rays usually longer than anteriormost; fin falling well short of caudal fin base when depressed. Anal fin similar to second dorsal fin, posteriormost rays longest, rays falling short of caudal fin base when depressed. Pectoral fin oval, central rays longest, 3.8-5.6 (mean 4.7) in SL; all rays branched but for uppermost. Pelvic fins oval, rays not reaching anus, 4.5-{).0 (mean 5.3) in SL. Caudal fin moderate in size, oval, rounded posteriorly, 3.7-4.6 (mean 4.1) in SL. Chin smooth, without mental fraenum. Anterior nostril in very low tube, placed at edge of upper lip, preorbital slightly curved outward near nostril. Posterior nostril oval, sometimes with slightly raised rim, placed nearly halfway between preorbital edge and eye, slightly closer to eye. Gill opening somewhat oblique, usually extending forward to under opercle, sometimes nearly to preopercular edge. Inner edge of pectoral girdle smooth without flanges or lobes; one specimen with low fleshy flange on one side. Gill rakers on outer face of first and second arch rather short, with fine spiny papillae on inner faces of rakers on lower limb, rakers longer toward angle of arch; rakers on inner face of first arch short, similar in size to those of first arch, with fine papillae at tips; inner rakers on second arch, outer and inner rakers of third and fourth arches longer, spiny papillae at tips. Tongue concave to blunt or irregularly shaped at tip, nearly trilobed in neotype; tongue sometimes "folded" along longitudinal axis. Outer teeth in upper jaw largest, relatively small, straight and sharp, evenly spaced along length of jaw, largest teeth across front; behind this row, two to four rows of small sharp curved teeth. Lower jaw with four to six rows of sharp curved teeth across front, two to three rows at sides; outermost row teeth largest and stoutest, oriented nearly upright; innermost teeth pointing posteriorly. Teeth similar in males and females. Predorsal scales cycloid, mostly evenly sized, reaching forward to past preopercular margin, extending forward to about halfway between rear of eyes and preopercular margin. Operculum covered with small cycloid scales on upper third to three-quarters. Cheek always naked. Pectoral base covered with small cycloid scales. Prepelvic area covered with small cycloid scales. Belly scales cycloid, with patch of weakly ctenoid scales anteriorly, close to base of pelvics; ctenoid scales sometimes extending nearly halfway down belly. Ctenoid scales on side of body posteriorly, extending forward in narrow wedge up to behind pectoral base or at least to below first dorsal fin; occasionally wedge broken, with small ctenoid Revision of Mugilogobius Figure 215 Tamanka siitensis, papillae pattern. CAS/SU 38624. Scales omitted. Scale bar = 1 mm. patch behind pectoral fin and ctenoid wedge reaching to below second dorsal fin origin. Genital papilla in male slender, elongate and flattened, broad at base and narrowing toward tip; papilla in female short, bulbous and stout, narrowing toward tip and broad at base, groove present along ventral surface. Head pores absent. Sensory papillae on head in longitudinal pattern, as in Figure 215. All papillae small, close-set, equal in size. Three s rows present, each of three or more papillae. Coloration of fresh material The only available information is that given by Herre in the original description. He described the colour of large males as black to umber, with smaller ("younger") ones deep brown, and "... fins black, with a series of white spots forming a crossband near margin of first dorsal, and a white crossband near margin of second dorsal; sometimes the ventrals have a narrow pale or whitish margin along sides" (Herre 1927). Coloration of preserved material Head and body plain light brown to dark brown (darker specimens usually, but not always, mature males), darker dorsally and lighter brown ventrally (Figure 216). Dorsal surface of head, nape and Figure 216 215 upper part of body sometimes with indistinct brown mottling; often small brown blotch visible just behind upper rear margin of eye, and indistinct brown blotches on opercle. All fins brown to dark brown to almost black, pectoral fins usually paler than others. Some small specimens (30-40 mm SL) with very light dusky fins. First dorsal fin with narrow submarginal white band, tips of fin spines dusky to dark brown. Second dorsal fin with similar narrow submarginal white band; band commences at about second soft ray and widens slightly posteriorly. Anal fin plain dark brown. Caudal and pelvic fins with translucent to whitish rear margins, or entire fin plain brown. Peritoneum brown, ventral midline pale. Comparisons This species is most similar in appearance to M. cagayanensis (see under Remarks for that species). Distribution Known only from two crater lakes on Jolo Island, Sulu Archipelago, Philippines (Figure 92). Ecology Lake Siit and Lake Panamao are land-locked, apparently freshwater lakes close to the coast. Herre (1927) described Lake Siit as a" ... small, deep, fresh-water crater lake beside the sea ... ". Aurich (1938) stated that Lake Seit (== Siit) and Lake Timpuk (from which he described Tamanka maculata) were occasionally joined together by encroachment of the sea. Remarks Herre (1927) described T. siitensis from a series of males ＨＴｾＵ mm in length) taken from a sample of 136 specimens, ＱＹｾＵ length, from Lake Siit, but he did not give a length for the type, BSM 11452, which was destroyed in Manila during WWII. Herre stated that the Bureau of Science also had six "typical" specimens 31-54 mm in length, from the Titunod River, Lanao Province, Mindanao (these specimens Tamanka siitensis. Paratype of Tamanka siitensis Herre, 53 mm SL, CAS 38624, Lake Panamao, Jolo, Sulu Archipelago, Philippines. 216 have not been found). The species does not appear to have not been collected in recent times; the very real presence of pirates operating in the Sulu Sea may have some influence in this. A neotype for Tamanka siitensis Herre (USNM 87128) is designated here, to ensure stability for the genus Tamanka, as two of the four species originally placed in the genus are unidentifiable at present, and only the type species, T. siitensis, is placed in the genus in this revision. The specimen was chosen in that it was a large male in good condition and resembled the illustration given by Herre (1927). The remaining paratypes are in USNM 355555. Aurich (1938) described Tamanka maculata from Lake Timpuk, Jolo Island, Sulu Archipelago; this lake is neighbour to Lake Slit, in which T. siitensis is found. It is possible that Aurich's Tamanka maculata may be closely related to T. siitensis. Aurich (1938: 154) seemed convinced that the two species, despite being found in neighbouring lakes occasionally joined by the action of the sea, were indeed separate species. The syntypes were not found at ZMH, where Aurich's type material was deposited, in 1988, and are possibly lost. The ZMH collection was stored in subways during World War 11, and a number of bottles were stolen for the alcohol they contained (Or H. Wilkens, personal communication). The fin ray and scale counts given in Aurich's description of Tamanka maculata do not differ from the range for T. siitensis, although the lateral scale count (40-45) is rather low. Of the 82 specimens in CAS/SU 26368, only six are female. A number of specimens, from both lakes, have encysted parasites scattered over their bodies and fins. BIOGEOGRAPHY The present knowledge of Mugilogobius and its relatives, as is true for many gobioids, is limited by collecting effort. It is not by accident that the Mugilogobius-group fauna of northern Australia is better known than that of Papua New Guinea, for example, as this is largely due to sampling in the former region since 1974 by myself and Ooug Hoese (AMS). Within the Indo-Malaysian Archipelago, recent collecting effort by Maurice Kottelat (CMK) and Peter Ng and staff at the National University of Singapore (ZRC) has revealed much of this fauna. However, there are still many gaps, which may prove to be due to low collecting effort rather than being real gaps in distribution Therefore any discussion of distribution and biogeography is speculative, and the following must be considered as preliminary. The centre of diversity of the Mugilogobius-group is the Indo-Malayan Peninsula, with Chlamydogobius being the only genus found outside this region, i.e. restricted to Australia. The most plesiomorphic H.K. Larson genus, Redigobius, shares a similar distribution to Mugilogobius and Pseudogobius. Mugilogobius is striking in its ability to occupy a broad range of habitats and its wide distribution (Figures 83, 87, 92, 123, 167). However, it does not occur in the Red Sea, the west coast of Africa or the New World. Its distribution is similar to, but more restricted than, that of Periophthalmus, a mangrove associate which has one species on the west African coast but none in the New World, a distribution unique among gobioids (Murdy, 1989). However, Mugilogobius is not restricted to mangroves per se, but to shallow estuarine habitats which by definition, often incorporate mangroves. The most widespread, and apparently the most abundant, species of Mugilogobius is M. mertoni, known from South Africa and the Seychelles, the Gulf of Aden, throughout South-east Asia to Japan, northern Australia and the western Pacific. Mugilogobius cavifrons is the next most widespread species; it is abundant, reaches the Pacific Plate via Caroline Islands conduit and has two forms (see Remarks under this species' account). In the initial work for this paper, these two forms were considered as separate species (parvus from Taiwan, Japan and the Philippines; cavifrons from Indonesia and the New Guinea region). The distribution of these two appears to be separated by Wallace's Line, and closely resembles that observed for two sibling species of the coral reef fish Siganus (Woodland, 1986: figure 2). Within the chulae and abei species-groups there are several other species which are reasonably Widespread: M. chulae and M. fusca from the first group, and M. platystoma and M. notospilus from the second. Among the less widely-distributed species, Mugilogobius abei is restricted to the east Asian region: Japan, southern China, Korea and Taiwan. Two species are restricted to peninsular Thailand, Malaysia and Singapore (M. fasciatus and M. tigrinus). Five species are restricted to the Australian continent (M. littoralis, M. platynotus, M. rivulus, M. stigmaticus and M. wilsoni). Mugilogobius fusculus is known only from Papua New Guinea, while M. filifer is known from northern Australia and Papua New Guinea. The greatest diversity of Mugilogobius species appears to be in Australia, with nine species (jilifer, littoralis, mertoni, notospilus, platynotus, platystomus, rivulus, stigmaticus, wilsoni). The next most speciose area is the Indo-Malayan Archipelago, which has eight species (cavifrons, chulae, fuscus, fasciatus, mertoni, platystomus, rambaiae, tigrinus). Further collecting in the latter region may well tip the balance, however. For example, only three species are known from the island of Borneo (chulae, mertoni and rambaiae), and it is probable that additional species may be found there. Revision of Mugilogobius Within Australia, there are some apparent barriers to distribution for the Mugilogobius-group. The known ranges of Mugilogobius filifer and M. littoralis extend across north-western Australia but both stop at the top of Cape York and do not extend down the eastern side. These disjunctions may possibly be explained by the Cape York Peninsula barrier (Springer and Williams, 1994; Springer and Larson, 1996). This is the transition from welldeveloped coral reef habitat to shallower water and less-qeveloped reefs as one traverses Torres Strait from east to west and enters the Sahul Shelf. It is not clear how this transition influences coastal estuarine fishes, but a faunal shift does exist. In the Torres Strait, only one species of Mugilogobius, M. filifer, has been recorded. Mugilogobius platystoma and M. mertoni are known from the Aru Islands, the Northern Territory and north Queensland (Daintree River and Tully), M. wilsoni is known from northwestern Australia to the bottom of the Gulf of Carpentaria, and from Cape Tribulation (east side of Cape York), while M. notospilus is known from northern New Guinea and the Daintree River, north Queensland. None of these species is known from the Torres Strait. However, as is the case for most species of this group, limited collecting effort may be disguising the true distributions. Similarly, there is very little information about the distribution of Mugilogobius in the Indian Ocean region, with only three species (Juscus, mertoni, rambaiae) recorded so far. This low number may be due to collecting effort, but is suspected to be real. The Sulawesi freshwater Mugilogobius and the five desert-dwelling species of Chlamydogobius (sistergroup to Mugilogobius) all belong to isolated populations. The Sulawesi species have a precaudal/caudal vertebrae pattern of 11+15-16, somewhat similar to the pattern in Chlamydogobius, which has a 10-11+16-18 vertebral pattern. The similarity in vertebral pattern is probably due to convergence. The temperate species M. platynotus has 10+17 vertebrae, unlike its probable sisterspecies, M. stigmaticus (both species restricted to the east coast of Australia). One possibility may be that Mugilogobius platynotus is closer to the ancestor of Chlamydogobius than any other species of Mugilogobius. This species also resembles Chlamydogobius in morphology and colouring (compare Kuiter, 1993: 358 and Larson, 1995: plates 1-2). The ancestor of Chlamydogobius may have entered the interior of Australia about 15-20 million years ago (early to mid-Miocene) (Larson, 1995). However, rather than assume a relationship based on vertebral number and arrangement between M. platynotus, the Sulawesi Mugilogobius species, and Chlamydogobius, it is considered more likely that the increase in vertebrae is a response to the environments in which the fish are found (e.g. lower temperature, isolation). Chlamydogobius 217 ranunculus, probably the least-derived species of its genus, is estuarine, and probably resembles the ancestral population. The Sulawesi Mugilogobius species are landlocked, found in the tectonic lakes of central Sulawesi. Lake Matano and Lake Poso are very deep, flooded rift lakes of central Sulawesi. Lake Matano is the deepest South-east Asian lake known; 590 m deep in its western half, and is believed to have been under sea water during late Tertiary and early Quaternary (1.5-5 million years ago) (Whitten et al., 1988). Lake Poso is also deep (450 m), but does not share its drainage system with the rest of the central tectonic lakes (Mahalona, Masapi, Matano, Towuti, Wawantoa). These five are collectively called the Malili Lakes and are all partly isolated from each other (Kottelat, 1989b, 1991). All of the indigenous freshwater fishes of Sulawesi's rivers and lakes are of marine derivation, unlike the rest of Indonesia (Kottelat et al., 1993). Kottelat (1991) described a possible scenario for colonisation of the Sulawesi tectonic lakes by the marine ancestor of the present-day freshwater fish family Telmatherinidae. These fishes only inhabit Sulawesi and the Misool and Batanta Islands, western Irian Jaya (Saeed and Ivantsoff, 1991). Eastern Sulawesi collided with western Sulawesi about 15 million years ago but remained submerged until the late Miocene (10 million years ago) (Audley-Charles, 1987). According to Kottelat, Eastern Sulawesi may thus have included brackish lakes available as habitat for the marine ancestors of the gobiids (and telmatherinids and adrianichthyids), which then colonised fresh water habitats as they developed. This scenario may equally apply to the Sulawesi lake-dwelling gobiids. The Sulawesi Mugilogobius do not seem to have diverged to the same extent as have the telmatherinids and adrianichthyids which inhabit the same lakes (Kottelat, 1990a-c, 1991). It is probable that these latter two families of freeswimming, plankton-picking species were able to take better advantage of the resources provided by the lakes. All the present Sulawesi lake-dwelling gobiids belong to only two genera: six Mugilogobius and six Glossogobius species (the taxonomy and relationships of the Glossogobius are being studied by Doug Hoese, AMS, and Jerry Alien, WAM). Closely related ancestral populations may have given rise both to the lake-dwelling species groups in Sulawesi and the isolated Jolo Archipelago of the Philippines, as there are some shared similarities. For example, species from both these localities have terminal mouths and are relatively plain in colour, and often become large. Alternately, similarity between these species could be due to similar evolutionary pressure within the lakes. The palaeogeography of the Philippines "remains uncertain" (Audley-Charles, 1981). A few primary 218 freshwater fishes have used the Sulu Archipelago as stepping stones from the Malaysian Archipelago to Mindanao (Earl of Cranbrook, 1981), and estuarine fishes could have also. It is possible that the ancestors of Mugilogobius cagayanensis and Tamanka siitensis island-hopped along the estuaries of these islands during a time of lower sea levels and have since become isolated as sea levels rose. J010 Island is one of the string of islands extending from the north-east tip of Borneo (Sabah) to Mindanao (Figure 92), while Cagayan Sulu (called Lupa Pula on some Indonesian maps), though isolated off northern Sabah, sits on the island shelf near the 100 m depth mark. When the distribution for each genus of the Mugilogobius-group is examined, it becomes apparent that a few genera are "successful" in that many species of these genera are Widespread and found across a variety of habitats. Four genera, Mugilogobius, Redigobius, Pandaka and Pseudogobius, are distributed across the Indo-west Pacific (see Figures 83, 87, 92, 123, 167, 198, 203, 208). These genera include many species which are estuarine or salt-tolerant. Some species of Mugilogobius and Pandaka have adapted to being totally freshwater dwelling and breeding. The genus Pseudogobius, present on the east coast of India, is apparently absent from Sri Lanka, the western Indian coast and eastern Africa, unlike Mugilogobius, Pandaka and Redigobius (Figure 203). Pseudogobius is restricted to estuarine habitats, with most species inhabiting mangroves, never penetrating very far into freshwater or into more marine environments, while its relatives Mugilogobius, Pandaka and Redigobius include estuarine and freshwater-tolerant species. The distribution of Pseudogobius might reflect the distribution of mangroves. This was indicated to be the limiting factor for' Periophthalmus distribution by Murdy (1989), who stated that the" ... west coast of India south of Bombay and the southern end of the Arabian Peninsula are largely devoid of mangrove ... ". Another key to the limited distribution of Pseudogobius might be its diet, as Pseudogobius is a herbivore (Gill and Potter 1993), while the other three genera mentioned are carnivorous. The temperate species Pseudogobius olorum has been shown to prefer seagrass habitat in the laboratory and the field (Gill and Humphries 1995), and other species may have specific habitat requirements not available on the Pacific Plate. The probable sister-group to Pseudogobius, Hemigobius, has one species in the Indo-Malay Archipelago and one more widely distributed species which extends to northern Australia and southern New Guinea (Figure 74). The species of this genus are restricted to mangrove habitats, which may have prevented them from extending any further eastward (onto the Pacific Plate, for H.K. Larson instance). It is possible that the genus has not extended further west along the east Indian coastline; however, low collecting effort may be the reason. Additionally, Hemigobius may have a dietary requirement which is not available elsewhere, as both species in the genus have very long, tightly coiled guts. Four genera of the Mugilogobius-group (Brachygobius, Caecogobius, Eugnathogobius and Stigmatogobius) are all generally restricted to fresh or brackish waters of the Indo-Malayan Archipelago (Figures 43, 59, 211). These "freshwater" genera appear to be limited by Wallace's Line, which is well-known in the context of primary freshwater fishes, and this boundary is also associated with a number of inshore marine fish faunas (Woodland, 1986). The Pacific Plate is a significant biogeographic unit for marine fishes (Springer, 1982: figure 2), and the distributions of many groups (families and genera) of Indo-Pacific shallow-water fishes do not extend onto the Pacific Plate. Among the Gobiidae, many of the genera that drop out as one goes east are composed of freshwater-dependent species. This is true for the Mugilogobius-group. Species of this group occur on the Philippine Plate or the Indian-Australian Plate. Species of Mugilogobius and ｒ･､ｩｧｯ｢ｾｵｳＬ for example, appear to have arrived on the Pacific Plate using the Caroline Islands Conduit (of Springer 1982) which connects onto the Pacific Plate via the estuaries of Micronesia (Palau to Kosrae). The only known Hawaiian Mugilogobius (in the centre of the Pacific Plate) is an introduced population, which probably arrived in ballast water. The fresh and brackish waters on the islands of the Pacific Plate have been colonised by relatively few genera belonging to three groups Of gobiids: the eleotridines (Eleotris, Ophiocara, Oxyeleotris), gobionellines (Awaous, Stenogobius, Mugilogobius, Pandaka, Pseudogobius, Redigobius) and sicydiines (Lentipes, Sicyopterus, Sicyopus, Stiphodon) (Springer, 1982; Ryan, 1991). Some of these genera (Ophiocara, Oxyeleotris, Mugilogobius, Pandaka, Pseudogobius, Redigobius) are limited in their distribution to the margins or eastward along the Caroline Islands conduit to Ponape and Kosrae, while others (Eleotris, Awaous, Stenogobius, Lentipes, Sicyopterus, Sicyopus, Stiphodon) extend further out onto the Pacific Plate proper. In Eleotris, the sicydiines and Awaous, the adults breed in freshwater with the larvae being swept down to the sea, whereupon post-larvae and juveniles migrate upstream some months later (e.g. Ryan, 1991; Smith and Hobson, 1996). No such migration has been observed for species of the Mugilogobius-clade; for example, adults and tiny juveniles of Mugilogobius, Hemigobius and Pseudogobius are regularly collected together. Gosline (1971) and Miller (1973) indicated that as 219 Revision of Mugilogobius a general rule that less derived groups tend to survive in eryhaline or freshwater habitats while the sea is occupied by their more derived descendants. Ancestral gobiids were probably estuarine species, although while the butines (currently regarded as plesiomorphic to gobionellines and gobiines)· are mostly estuarine, the Rhyacichthyidae and Odontobutidae, currently regarded as the most primitive gobioids, are all freshwater species. It is possible that the rhyacichthyids and odontobutids were derived from estuarine forms which have not survived. Most Mugilogobius species are found in very shallow, muddy to sandy habitats where there is some freshwater input. Gnatholepis may be the only circumtropical, fully marine genus among the gobionellines (the Indo-Pacific gobionelline Oxyurichthys includes several estuarine species, although most species are coastal to offshore marine in habitat preference). ACKNOWLEDGEMENTS My very many thanks to curators and staff at the institutions who loaned specimens or sent me wonderful mystery gobies, especially Jerry AlIen, WAM; His Majesty, Emperor Akihito, BLIH; Eric Anderson, RUSli Genie Bohlke, ANSP; Dave Catania, CASi Barry Chernoff, FMNHi G. Duhamel, MNHN; Bill Eschmeyer, CAS; Karsten Hartel, MCZi Masayoshi Hayashi, YCM; Barbara Herzig, NMW; S.H. Hoda, University of Karachii Doug Hoese, AMS; P. Hulley, SAMi J.-c. Hureau, MNHNi Akihisa Iwata, formerly of BLIHi Susan Jewett, USNM; Jeff Johnson, QM; Maurice Kottelat, Switzerlandi Friedhelm Krupp, SMFi Kelvin Lim, ZRCi Mark McGrouther, AMS; Katsusuke Meguro, BLIH; Nigel Merrett, BMNHi Gary Nelson, AMNHi J0rgen Nielsen, ZUMCi Han Nijssen, ZMA; Peter Ng, University of Singapore; Martien van Oijen, RMNHi H.-J. Paepke, 2MB; Brad Pusey, Griffith University, Nathani Jack Randall, BPBM; K. Sakamoto, ZUMTi Hiroshi Senou, Tokyo; Terry Sim, SAMA; Bill Smith-Vaniz, formerly of ANSPi Arnold Suzumoto, BPBMi Chavalit Vidthayanon, NIFI; H. Wilkens, ZMH; Rick Winterbottom, ROM; Kai-Erik Witte, University of Konstanzi Prachit Wongrat, KUMF; Thosaporn Wongratana, Chulalongkorn University, Bangkoki Tetsuo Yoshino, URM. Many thanks to those people who helped provide fresh specimens, information, and/or photographs: Mark Adams, SAMA; Neil Armstrong, ANGFA; Rick Bawden, NTMi I-Shiung Chen, Bristol Universityi Satish Choy, formerly of University of Bruneii Karen Coombes, NTM; Graeme Dobson, formerly of NT Health, Darwin; Ross Felixi the late John Glover, SAMAi Bruce Grose, ANGFA, Victoriai Russell Hanley, formerly at NTMi Bruce Hansen, ANGFA; Paul Horner, NTMi Hans Horsthemke, GermanYi Chris Jones, Northern Territory Universityi Richard Kimber, Alice Springs; Keith Martin, Dames and Moore, Darwini Dirk Megirian, NTM; N. Oseko, Japani Ukkrit Satapoomin, Biological Research Centre, Phuketi M. Selway, Darwin; T. Suzuki, Amagasakikita Senior High School, Japani K. Uhlenhunt; Peter Unmack, Arizona State Universityi Rob Wager, Queensland Department of Primary Industries, Brisbanei ANGFA, Victoria; Graham White, NT Fisheries, Darwini Rex Williams, NTMi Dave Wilson, Territory Wildlife Park, Darwin; Dr Wu Han-ling, Shanghai University. Many thanks to Tony Gill, Doug Hoese and Charles Webb, for their helpful comments and advice on various versions of this paper. Thanks also to Mella Parshen-Kempfer and Judy Marsland for patiently translating from the German for me, and to Neil Smit for translating from the Dutch. This paper formed part of a PhD thesis submitted to the University of Queensland, Brisbane. My thanks to the Museum and Art Gallery of the Northern Territory for its continuing support during this project. 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Nominal species Taxonomic allocation (this paper) Ellogobius abascantus Whitley, 1937 Ctenogobius abei Jordan and Snyder, 1901 Mugilogobius adeia Larson and Kottelat, 1993 Vaimosa adyari Berre, 1945 Brachygobius aggregatus Berre, 1940 Gobius alcocki Annandale, 1906 Gnathogobius aliceae Smith, 1945 Gobius amadi Weber, 1913 Stigmatogobius amblyrhynchus Bleeker, 1878 Stigmatogobius amblystoma Zander, 1972 Gobius apogonius Cantor, 1850 Tamanka arguellesi Roxas and Ablan, 1940 Gillichthys australis Ogilby, 1894 Vaimosa avicennia Berre, 1940 Vaimosa balteata Berre, 1935 Acentrogobius balteatops Smith, 1959 Gobius beccari Perugia, 1892 Tamanka bivittata Berre, 1927 Vaimosa bikolana Berre, 1927 Hemigobius bleekeri Koumans, 1953 Gobius borneensis Bleeker, 1851 Vaimosa brocki Berre, 1936 Vaimosa cagayanensis Aurich, 1938 Glossogobius campbellianus Jordan and Seale, 1908 Gobius cavifrons Weber, 1909 Vaimosa cardonensis Berre, 1940 Gobius chilkensis Jenkins, 1910 Vaimosa chinensis Berre, 1935 Lophogobius chrysosoma Bleeker, 1875 Vaimosa chulae Smith, 1932 Vaimosa crassa Berre, 1945 Caecogobius cryptophthalmus Berti and Ercolini, 1991 Mugilogobius devisi McCulloch and Ogilby, 1919 Gobius dewaalii Weber, 1897 Gobius dispar Peters, 1868 Gobius doriae Giinther, 1868 Stigmatogobius duospilus Fowler, 1953 Gobius durbanensis Barnard, 1927 Gobius eremius Zietz, 1896 Gobius flavescens De Vis, 1884 Vaimosa jontinalis Jordan and Seale, 1906 Vaimosafusca Berre, 1940 Tamanka fusculus Nichols, 1951 Mugilogobius galwayi McCulloch and Waite, 1918 Gobius gastrospilos Bleeker, 1853 Chlamydogobius gloveri Larson, 1995 Gobius hoevenii Bleeker, 1851 Vaimosa horiae Berre, 1936 Parvigobius immeritus Whitley, 1930 Stigmatogobius inhacae Smith, 1959 Stigmatogobius isognathus Bleeker, 1878 Chlamydogobius japalpa Larson, 1995 Gobius javanicus Bleeker, 1856 Gobius johnstoniensis Koumans, 1940 Vaimosa jurongensis Herre, 1940 Vaimosa kabilia Berre, 1940 Brachygobius kabiliensis Inger, 1958 Vaimosa karatunensis Aurich, 1938 Vaimosa koumansi Mukerji, 1935 Gobius latifrons Boulenger, 1897 Vaimosa layia Berre, 1953 Acentrogobius leptochilus Bleeker, 1875 Mugilogobius platynotus (Giinther, 1861) Mugilogobius abei Gordan and Snyder, 1901) Mugilogobius adeia (Larson and Kottelat, 1993) Pseudogobius melanostictus (Day, 1876) Brachygobius aggregatus Berre, 1940 Brachygobius nunus (Bamilton, 1822) Calamiana kabilia (Berre, 1940) Mugilogobius amadi (Weber, 1913) Redigobius? Redigobius Stigmatogobius sadanundio (Bamilton, 1822) Mugilogobius ? Redigobius macrostomus (Giinther, 1861) Pseudogobius avicennia (Berre, 1940) Redigobius balteatus (Berre, 1935) Redigobius balteatus (Berre, 1935) Stigmatogobius sella (Steindachner, 1881)? Mugilogobius abei Gordan and Snyder, 1901) Redigobius bikolanus (Berre, 1927) Hemigobius mingi (Berre, 1936) Stigmatogobius borneensis (Bleeker, 1851) Stigmatogobius sella (Steindachner, 1881) Mugilogobius cagayanensis (Aurich, 1938) Eugnathogobius oligactis (Bleeker, 1875) Mugilogobius cavifrons (Weber, 1909) Redigobius ?Pseudogobius Redigobius Redigobius chrysosomus (Bleeker, 1875) Mugilogobius chulae (Smith, 1932) Hemigobius hoevenii (Bleeker, 1851) Caecogobius cryptophthalmus Berti and Ercolini, 1991 Mugilogobius stigmaticus (De Vis, 1884) Redigobius dewaalii (Weber, 1897) Redigobius dispar (Peters, 1868) Brachygobius doriae (Giinther, 1868) Mugilogobius notospilus (Giinther, 1877) Mugilogobius mertoni (Weber, 1911) Chlamydogobius eremius (Zietz, 1896) Redigobius bikolanus (Berre, 1927) Mugilogobius notospilus (Giinther, 1877) Mugilogobius fuscus (Berre, 1940) Mugilogobius fusculus (Nichols, 1951) Pseudogobius olorum (Sauvage, 1880) Pseudogobius Chlamydogobius gloveri Larson, 1995 Hemigobius hoevenii (Bleeker, 1851b) Redigobius bikolanus (Berre, 1927) Redigobius bikolanus (Berre, 1927) Mugilogobius mertoni (Weber, 1911) Redigobius Chlamydogobius japalpa Larson, 1995 Pseudogobius javanicus (Bleeker, 1856) Redigobius bikolanus (Berre, 1927) Eugnathogobius siamensis (Fowler, 1934) Calamiana kabilia (Berre, 1940) Brachygobius kabiliensis Inger, 1958 Mugilogobius cavifrons (Weber, 1909) Redigobius? Mugilogobius latifrons (Boulenger, 1897) Mugilogobius mertoni (Weber, 1911) Redigobius leptochilus (Bleeker, 1875) H.K. Larson 232 Nominal species Taxonomic allocation (this paper) Vaimosa leveri Fowler, 1943 Gobius lidwilli McCulloch, 1917 Mugilogobius luzonensis Roxas and Ablan, 1940 Vaimosa macrognathos Herre, 1927 Gobius macrostoma Giinther, 1861 Tamanka maculata Aurich, 1938 Gobius magniloquus Day, 1876 Calamiana magnoris Herre, 1945 Glossogobius mas Hora, 1923 Vaimosa mawaia Herre, 1936 Gobius maxillaris Davies, 1949 Gobius melanosticta Day, 1876 Gobius melanurus Bleeker, 1849 Gobius mertoni Weber, 1911 Stigmatogobius micrognathus Rao, 1971 Gobius microphthalmus Giinther, 1861 Eugnathogobius microps Smith, 1931 Chlamydogobius micropterus Larson, 1995 Vaimosa microstomia Seale, 1910 Tamanka mindora Herre, 1945 Vaimosa mindora Herre, 1945 Gnatholepis mingi Herre, 1936 Ctenogobius minima Hora, 1923 Pandaka minuta Smith, 1959 Stigmatogobius minutus Takagi, 1957 Vaimosa montalbani Herre, 1936 Ctenogobius myxodermus Herre, 1935 Stigmatogobius neglectus Koumans, 1932 Gobius notospilus Giinther, 1877 Vaimosa novaehebudorum Fowler, 1944 Gobius nunus Hamilton, 1822 Mugilogobius obliquifasciata Wu and Ni, 1985 Gobiopsis oligactis Bleeker, 1875 Gobius olorum Sauvage, 1880 Vaimosa oratai Herre, 1940 Gobius ornatus masago Tomiyama, 1936 Vaimosa osgoodi Herre, 1935 Gobius oyensi Beaufort, 1913 Mahidolia pagoensis Schultz, 1943 Ctenogobius paludosus Herre 1940 Waiteopsis paludis Whitley, 1930 Glossogobius parous Oshirna, 1919 Pseudogobius penango Popta, 1922 Vaimosa perakensis Herre, 1940 Tamanka philippina Herre, 1945 Vaimosa piapensis Herre, 1927 Gobius platynotus Giinther, 1861 Gobius platystoma Giinther, 1872 Gobius pleurostigma Bleeker, 1849 Gobius poeyi Steindachner, 1867 Gobius poicilosoma Bleeker, 1849 Mugilogobius polylepis Wu and Ni, 1985 Mugilogobius pongolensis Kok and Blaber, 1977 Pandaka pusilla Herre, 1927 Pandaka pygmaea Herre, 1927 Chlamydogobius ranunculus Larson, 1995 ,Gobius reticularis Weber, 1911 Vaimosa rambaiae Smith, 1945 Vaimosa rivalis Herre, 1927 Gobius roemeri Weber, 1911 Gobius rouxi Weber, 1911 Brachygobius sabanus Inger, 1958 Gobius sadanundio Hamilton, 1822 Gobius samberanoensis Bleeker, 1867 Redigobius Pandaka lidwilli (McCulloch, 1917) Mugilogobius or Pseudogobius Redigobius Redigobius macrostomus (Giinther, 1861) Mugilogobius or Tamanka Pseudogobius? Mugilogobius? Redigobius? Calamiana kabilia (Herre, 1940) Eugnathogobius? Eugnathogobius siamensis (Fowler, 1934) Redigobius dewaalii (Weber, 1897) Pseudogobius melanostictus (Day, 1876) Hemigobius mingi (Herre, 1936) Mugilogobius mertoni (Weber, 1911) Redigobius? or Pseudogobius? Redigobius macrostomus (Giinther, 1861) Eugnathogobius microps Smith, 1931 Chlamydogobius micropterus Larson, 1995 Pseudogobius? or Redigobius? Mugilogobius mertoni (Weber, 1911) Calamiana mindora (Herre, 1945) Hemigobius mingi (Herre, 1936) Pseudogobius Pandaka silvana (Barnard, 1943) Redigobius bikolanus (Herre, 1927) Redigobius bilrolanus (Herre, 1927) Mugilogobius myxodermus (Herre, 1935) Eugnathogobius oligactis (Bleeker, 1876) Mugilogobius notospilus (Giinther, 1877) Redigobius bikolanus (Herre, 1927) Brachygobius nunus (Hamilton, 1822) Hemigobius hoevenii (Bleeker, 1851b) Eugnathogobius oligactis (Bleeker, 1875) Pseudogobius olorum (Sauvage, 1880) Eugnathogobius siamensis (Fowler, 1934) Pseudogobius masago (Tomiyama, 1936) Redigobius bikolanus (Herre, 1927) Redigobius Redigobius bilrolanus (Herre, 1927) Eugnathogobius paludosus (Herre, 1940) Mugilogobius platynotus (Giinther, 1861) Mugilogobius cavifrons (Weber, 1909) Redigobius penango (Popta, 1922) Eugnathogobius oligactis (Bleeker, 1875) Mugilogobius cavifrons (Weber, 1909) Pseudogobius javanicus (Bleeker, 1856) Mugilogobius platynotus (Giinther, 1861) Mugilogobius platystomus (Giinther, 1872) Stigmatogobius pleurostigma (Bleeker, 1849) Mugilogobius Pseudogobius poicilosomus (Bleeker, 1849) ?Calamiana polylepis Wu and Ni, 1985 Redigobius dewaalii (Weber, 1897) Pandaka pusilla Herre, 1927 Pandaka pygmaea Herre, 1927 Chlamydogobius ranunculus Larson, 1995 Redigobius roemeri (Weber, 1911) Mugilogobius rambaiae (Smith, 1945) Redigobius? or Eugnathogobius? Redigobius roemeri (Weber, 1911) Pandaka rouxi (Weber, 1911) Brachygobius sabanus Inger, 1958 Stigmatogobius sadanundio (Hamilton, 1822) Redigobius? Revision of Mugilogobius 233 Nominal species Taxonomic allocation (this paper) Vaimosa sapanga Herre, 1927 Gobius sarasinorum Boulenger, 1897 Rhinogobius schultzei Herre, 1927 Gobius sella Steindachner, 1881 Vaimosa serangoonensis Herre, 1937 Vaimosa siamensis Fowler, 1934 Tamanka siitensis Herre, 1927 Gobius silvanus Bamard, 1943 Tamanka sinensis Herre, 1935 Stigmatogobius singapurensis Bleeker, 1878 Vaimosa spilopleura Smith, 1933 Chlamydogobius squamigenus Larson, 1995 Gobius sternbergi Smith, 1902 Gobius stigmaticus De Vis, 1884 Thaigobiella sua Smith, 1931 Tamanka tagala Herre, 1927 Tamanka talavera Herre, 1945 Gobius tambujon Bleeker, 1854 Vaimosa tessellata Herre, 1927 Gobius tigrellus Nichols, 1951 Pandaka trimaculata Akihito and Meguro, 1975 Tamanka ubinensis Herre in Herre and Myers, 1937 Tamanka umbra Herre, 1927 Sphenentogobius vanderbilti Fowler, 1940 Vaimosa valigouva Deraniyagala, 1936 Apocryptes variegatus Peters, 1868 Gobius vergeri Bleeker, 1867 Stigmatogobius versicolor Smith, 1959 Vaimosa villa Herre, 1927 Lophogobius wera Popta, 1922 Brachygobius xanthomelas Herre, 1937 Gobius xanthozona Bleeker, 1849 Stigmatogobius yanamensis Rao, 1971 Vaimosa zebra Aurich, 1938 Vaimosa zebrina Herre, 1950 Pseudogobiopsis wuhanlini Zhang and Chu, 1997 Redigobius Mugilogobius sarasinorum (Boulenger, 1897) Mugilogobius? or Pseudogobius? Stigmatogobius sella (Steindachner, 1881) Pseudogobius melanostictus (Day, 1876) Eugnathogobius siamensis (Fowler, 1934) Tamanka siitensis Herre, 1927 Pandaka silvana (Bamard, 1943) Mugilogobius chulae (Smith, 1932) Stigmatogobius bomeensis (Bleeker, 1851) Stigmatogobius pleurostigma (Bleeker, 1849) Chlamydogobius squamigenus Larson, 1995 Redigobius dispar (Peters, 1868) Mugilogobius stigmaticus (De Vis, 1884) Brachygobius xanthozonus (Bleeker, 1849) Eugnathogobius? Mugilogobius? Mugilogobius cavifrons (Weber, 1909) Redigobius Redigobius Eugnathogobius? Pandaka trimaculata Akihito and Meguro, 1975 Calamiana variegata (Peters, 1868) Mugilogobius? or Eugnathogobius? Hemigobius mingi (Herre, 1936) Mugilogobius chulae (Smith, 1932) Calamiana variegata (Peters, 1868) Redigobius. Redigobius bikolanus (Herre, 1927) Mugilogobius? Redigobius Brachygobius xanthomelas Herre, 1937 Brachygobius xanthozonus (Bleeker, 1849) Pseudogobius Mugilogobius chulae (Smith, 1932) Calamiana mindora (Herre, 1945) Engnathogobius siamensis (Fowler, 1934) Guide to Authors Subject Matter: Reviews, observations and results of research into all branches of natural science and human studies will be considered for publication. However, emphasis is placed on studies pertaining to Western Australia. Longer papers will be considered for publication as a Supplement to the Records of the Western Australian Museum. Short communications should not normally exceed three typed pages and this category of paper is intended to accommodate observations, results or new records of significance, that otherwise might not get into the literature, or for which there is a particular urgency for publication. All material must be original and not have been published elsewhere. 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A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placement

A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placement

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