A review of the diversity and impact of invasive non-native species in tropical marine ecosystems

Tropical marine ecosystems are biologically diverse and economically invaluable. However, they are severely threatened from impacts associated with climate change coupled with localized and regional stressors, such as pollution and overfishing. Non-native species (sometimes referred to as ‘alien’ species) are another major threat facing these ecosystems, although rarely discussed and overshadowed by the other stressors mentioned above. NNS can be introduced accidentally (for example via shipping activities) and/or sometimes intentionally (for aquaculture or by hobbyists). Understanding the extent of the impacts NNS have on native flora and fauna often remains challenging, along with ascertaining when the species in question actually became ‘invasive’. Here we review the status of this threat across key tropical marine ecosystems such as coral reefs, algae meadows, mangroves, and seagrass beds. We aim to provide a baseline of where invasive NNS can be found, when they are thought to have been introduced and what impact they are thought to be having on the native ecosystems they now inhabit. In the appended material we provide a comprehensive list of NNS covering key groups such as macroalgae, sponges, seagrasses and mangroves, anthozoans, bryozoans, ascidians, fishes, and crustaceans.N

Interactions between marine megafauna and plastic pollution in Southeast Asia

Southeast (SE) Asia is a highly biodiverse region, yet it is also estimated to cumulatively contribute a third of the total global marine plastic pollution. This threat is known to have adverse impacts on marine megafauna, however, understanding of its impacts has recently been highlighted as a priority for research in the region. To address this knowledge gap, a structured literature review was conducted for species of cartilaginous fishes, marine mammals, marine reptiles, and seabirds present in SE Asia, collating cases on a global scale to allow for comparison, coupled with a regional expert elicitation to gather additional published and grey literature cases which would have been omitted during the structured literature review. Of the 380 marine megafauna species present in SE Asia, but also studied elsewhere, we found that 9.1 % and 4.5 % of all publications documenting plastic entanglement (n = 55) and ingestion (n = 291) were conducted in SE Asian countries. At the species level, published cases of entanglement from SE Asian countries were available for 10 % or less of species within each taxonomic group. Additionally, published ingestion cases were available primarily for marine mammals and were lacking entirely for seabirds in the region. The regional expert elicitation led to entanglement and ingestion cases from SE Asian countries being documented in 10 and 15 additional species respectively, highlighting the utility of a broader approach to data synthesis. While the scale of the plastic pollution in SE Asia is of particular concern for marine ecosystems, knowledge of its interactions and impacts on marine megafauna lags behind other areas of the world, even after the inclusion of a regional expert elicitation. Additional funding to help collate baseline data are critically needed to inform policy and solutions towards limiting the interactions of marine megafauna and plastic pollution in SE Asia

Denial of long-term issues with agriculture on tropical peatlands will have devastating consequences

Non peer reviewe

Taxonomy based on science is necessary for global conservation

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Preliminary checklist of fishes obtained from South Java Deep-Sea(SJADES) Biodiversity Expedition 2018

10.26107/RBZ-2021-0051RAFFLES BULLETIN OF ZOOLOGYSupplement No. 36496-526Singapor

FIGURE 14. Aulopareia vadosa n in A review of the gobiid fish genus Aulopareia (Gobiidae: Gobiinae) with description of a new species from Kuwait and discussion of the status of Gobius cyanomos Bleeker

FIGURE 14. Aulopareia vadosa n. sp., 32mm SL holotype, ZRC 54321, Sulaibikhat Bay, Kuwait. Photograph by Heok Hui Tan.Published as part of <i>Larson, Helen K. & Jaafar, Zeehan, 2022, A review of the gobiid fish genus Aulopareia (Gobiidae: Gobiinae) with description of a new species from Kuwait and discussion of the status of Gobius cyanomos Bleeker, pp. 493-516 in Zootaxa 5155 (4)</i> on page 513, DOI: 10.11646/zootaxa.5155.4.2, <a href="http://zenodo.org/record/6691350">http://zenodo.org/record/6691350</a&gt

Geosesarma Aedituens, A New Terrestrial Crab (Crustacea: Decapoda: Brachyura: Sesarmidae) From Bali, Indonesia

Naruse, Tohru, Jaafar, Zeehan (2009): Geosesarma Aedituens, A New Terrestrial Crab (Crustacea: Decapoda: Brachyura: Sesarmidae) From Bali, Indonesia. Raffles Bulletin of Zoology 57 (1): 183-187, DOI: 10.5281/zenodo.534186

Oxuderces NEXIPINNIS (CANTOR 1849

OXUDERCES NEXIPINNIS (CANTOR, 1849) FIGURES 2, 3B, 4A, 5A, 6, 7A, 8A, 9A, 10B, 14; TABLE 1 Apocryptes nexipinnis Cantor, 1849: 1170 (type locality, Sea of Penang, Malaysia) Apocryptes cantoris not Day, 1871 (in part Day, 1876, specimens from Madras, India) Apocryptichthys livingstoni Fowler, 1935: 162 (type locality, Paknam, Thailand) Syntypes examined: BMNH 1860.3.19.568-69, 2 undet., 65.0 – 67.8 mm SL, Sea of Penang, Malaysia. Other type material examined: Apocryptichthys livingstoni holotype: ANSP 63091, male, 73.2 mm SL, mouth of Me Nam Chao Praya, Paknam, Thailand. Other material examined: Sixty-four specimens, 19.9 – 80.6 mm SL. RMNH.PISC 12091, male (66.8 mm), 2 females (65.9 – 73.0 mm), east coast Java Island, Indonesia; RMNH.PISC 12092, 5 males (63.0 – 76.4 mm), 14 females (60.0 – 79.9 mm), east coast Java Island, Indonesia; RMNH.PISC 12433, female (55.6 mm), Java Island, Indonesia; RMNH.PISC 12750, female (79.4 mm), Surabaya, Indonesia; RMNH.PISC 33844, female (57.6 mm), Indonesia; RMNH.PISC 17382, female (72.0 mm), Pulau Weh, Sumatra, Indonesia; USNM 119547, female (64.0 mm), off Tachalon, Gulf of Thailand; USNM 278447, female (43.6 mm), southern side of river mouth, Muar river, Johore, Malaysia; USNM 279359, 12 males (19.9 – 50.0 mm; 1 C&S, 35.7 mm), 14 females (22.1 – 31.5 mm; 1 C&S, 47.9 mm); USNM 288662, 1 male (C&S, 39.7 mm), 2 females (C&S, 21.6 – 26.8 mm), southern side of river mouth, Muar river, Johore, Malaysia; ZRC 39777, male (80.6 mm), Sungai Selising, Matang mangroves, Perak, Malaysia; ZRC 53919, 3 males (50.3 – 53.2 mm), 3 females (33.4 – 53.3 mm), Matla mudflats, Jharkali, South 24 Parganas, West Bengal, India. Differential diagnosis: Oxuderces nexipinnis is differentiated from O. dentatus by the following characters: first hemal spine terminating above midlength of first anal-fin pterygiophore (vs. extending ventrally below mid-length of first anal-fin pterygiophore; Fig. 10); mouth superior (vs. terminal); conspicuous dermal invagination posterior to the point of attachment of pelvic-fin base (vs. dermal invagination absent; Fig. 6); head relatively long (27.3 – 30.3%SL vs. 25.1 – 27.7%SL). 1 mm Description: Head markedly depressed, wider than deep, head length 27.3 – 30.3%SL, head width 32.6 – 44.4%HL, head depth 30.4 – 42.7%HL. Eye diameter 8.7 – 16.3%HL; interorbital distance 3.0 – 4.8%HL, eye without dermal cup. Snout profile pointed, snout length 12.0 – 15.5%HL. Mouth superior, lower jaw terminating just anterior to upper jaw, gape wide; jaw length 41.3 – 59.7%HL; distinct notch in middle of upper lip between two medial teeth. Teeth in both jaws caninoid, unevenly spaced, in single row; one or two prominent and elongate canines on each side of premaxillary symphysis, canine teeth extending slightly anteroventrally and reaching beyond lower jaw when mouth closed, teeth decreasing in length posteriorly; teeth in lower jaw more uniform in size, teeth all shorter than medial canine teeth of upper jaw, teeth absent posteriorly, no canine tooth on each side of symphysis internal to anterior margin of lower jaw. Body compressed and gracile; body depth at anus 8.6 – 12.9%SL, body width at anus 4.5 – 7.3% SL. Predorsal long, predorsal length 28.2 – 31.6%SL. D1 and D2 connected by membrane for entire height, base of D1 and D2 54.9 – 59.4%SL; base of anal fin 35.9 – 40.4%SL. No membrane connecting D2 or A to caudal fin. Pelvic fin short and rounded, not reaching genital papilla, pelvic-fin length 14.0 – 18.2% SL. Pectoral-fin length moderate, 14.0 – 19.2%SL. Caudal fin long, lanceolate, 20.8 – 24.8%SL. D1 with six spinous dorsal rays (VI); all elements of D2 and A fins segmented; D2 with 25 – 26 elements, A fin with 24 – 26 elements; pectoral rays 21 – 24; caudal fin segmented rays 17; dorsal procurrent rays 5, ventral procurrent rays 4. Two epural bones. Lateral longitudinal scale count 50 – 54, scales on body small and embedded in skin anteriorly, and increasing in size posteriorly. All specimens without scales on dorsal region anterior to D1, ventral region of the head, cheek, operculum and pectoral-fin base except for largest specimen examined (ZRC 39777, 80.6 mm) which has cheek scales and 14 predorsal midline scales; all scales small and embedded. Raised and distinct free-neuromast rows on head. Conspicuous dermal invagination posterior to point A First dorsal-fin anterior elements of attachment of pelvic-fin base, invagination deep and appearing as a ‘pit’ (Fig. 6). Coloration: Preserved specimens uniformly light to dark brown with black spots on dorsal region of head and nape. Darkened brown to black upper lip and pectoral-fin base. Dark brown spot between 20 th and penultimate rays, positioned at mid-height of D2. Murdy (1989: 20) based his description of live coloration on material from Peninsular Malaysia that he had identified as O. dentatus (= O. nexipinnis here): ‘Head and trunk greyish blue; 7 greyish-blue spots along lateral midline starting at a point equal to origin of first dorsal fin and ending at a point equal to about 75% of the length of D2; a suffusion of shiny, bluish scales on posterior half of trunk; 6 dusky, saddle-like blotches on dorsum starting at terminus of D1 with 4 across the base of D2 and 1 on caudal peduncle; D1 translucent; D2 translucent except for a thin, faint dusky stripe through the median part of fin and a large black ocellus near distal tips of last 4 rays; caudal, anal and pelvic fins translucent; pectoral fin translucent but with a large, dusky blotch on upper base; nape with dusky reticulations; anterior nostril with a black anterior margin; venter shiny white.’ In the Malacca Straits, along the western coast of Peninsular Malaysia, Takita, Agusnimar & Ali (1999: 132) identified O. nexipinnis (reported as O. dentatus) by the characteristic ‘...bright green eyes and depressed head, and a dark spot posteriorly on the dorsal fin’. They reported that O. nexipinnis occurs on the mudflats of Bengkalis and Penang islands in Peninsular Malaysia, at sites facing the sea and not influenced by freshwater inflow (Takita et al., 1999). At Teluk Bahang in Penang Island, this species was found low on the intertidal zone, an area which was exposed for only several days per lunar cycle, prompting the hypothesis that O. nexipinnis is probably active underwater (Takita et al., 1999). This hypothesis is corroborated by Polgar & Bartolino (2010) who found larger individuals of O. nexipinnis (reported as O. dentatus) in the lower intertidal zones (more aquatic) and smaller individuals in higher intertidal zones (more terrestrial) at their study site in Tanjung Piai, Johor, along the western coast of Peninsular Malaysia. Etymology: Latin, ‘ nexus’ meaning ‘tied together’ and ‘ pinnis ’ meaning ‘fins’, referring to the continuous dorsal fins. Distribution and ecological note: Oxuderces nexipinnis is distributed in the Indo-West Pacific, here reported from the southern part of the South 1 mm China Sea, Gulf of Thailand, Java Sea, Andaman Sea and Bay of Bengal (Fig. 13). This species lives on intertidal soft-bottom habitats, predominantly mudflats. Adults have been observed to swim through the muddy substrate with only a thin film of water above them, through which they would occasionally poke their heads (Murdy, 1989, reported as O. dentatus). This species has been reported to store air in its burrows, especially in the spawning chambers (Ishimatsu et al., 1998, reported as O. dentatus). In Malaysia, it is a common prey of homalopsine snakes such as Bitia hydroides (see Jayne, Ward & Voris, 1995, reported as O. dentatus) and Cerberus rynchops (see Jayne, Voris & Heang, 1988, reported as O. dentatus). Remarks: Apocryptes nexipinnis, described by Cantor (1849), was classified in the genus Oxuderces, and synonymized with O. dentatus by Murdy (1989). We resurrect this name from synonymy as Oxuderces nexipinnis. The BMNH syntypes are the left sides (heads and bodies) of two individuals that have been dried, filleted and stuffed with cotton wool (Fig. 14). Cantor (1849) did not indicate the number of specimens he had on hand and reported the total length of the specimen (or specimens) on which he based his description as 3 3/ 8 inches, or 85.7 mm TL. He further indicated that the caudal fin comprised 1/5 of the TL, which would mean that the specimen (or specimens) was approximately 68 mm SL. The BMNH syntypes measure approximately 67.8 mm SL (possibly a male) and 65 mm SL (possibly a female). Although the larger specimen is closer in size to that given by Cantor (1849), we do not designate it as a lectotype because both specimens are extremely distorted, as well as incomplete, and we cannot identify either with certainty as the primary specimen on which the description was based. DISCUSSION Examination of type specimens and a review of the literature confirmed the hypothesis that Oxuderces comprises two distinct species. Specimens from coastal areas of southern China agree with the detailed description of the type specimen of Oxuderces dentatus (see Springer, 1978). Contra Springer (1978) and Murdy (1989), we consider Apocryptes cantoris Day, 1871 (in part) and Apocryptichthys livingstoni Fowler, 1935 to be synonyms of Apocryptes nexipinnis Cantor, 1849, rather than of Oxuderces dentatus Eydoux & Souleyet, 1848. Material from Peninsular Malaysia, Thailand, Indonesia and India agrees with the type specimens and original description of Apocryptes nexipinnis which we classify here as the second valid species of Oxuderces. Contra Murdy (1989), we do not classify Apocryptodon wirzi Koumans, 1937 in the genus Oxuderces. After examination of type and non-type specimens of Apocryptodon wirzi, we conclude that this species has characters diagnostic of the inferred sister genus Apocryptodon (Murdy, 1989; Agorreta et al., 2013), and does not conform to our revised concept of Oxuderces. Oxuderces nexipinnis uniquely has a conspicuous dermal invagination of unknown function just posterior to the point of attachment of the pelvic-fin base. The epidermal surfaces of all oxudercine gobies are barriers to the external environment and aid with respiration when individuals are near the surface or out of the water. Consequently, the skin of amphibious oxudercine gobies, such as Boleophthalmus, Periophthalmodon, Periophthalmus and Scartelaos, has been examined in detail to hypothesize adaptations to terrestriality (see Park et al., 2003; Zhang et al., 2003). In contrast, little is known of the form and function of the skin of the oxudercine gobies that do not readily emerge out of water, such as species of the genera Apocryptodon, Oxuderces and Parapocryptes. Here we report for the first time some of the modifications of the skin on the head of Oxuderces, in which the dorsal part of the head and the eyes are covered with a thick epidermis that gradually thins ventrally (Figs 4A, 5A). This differs from the epidermal layer in the dorsal region of the head in Apocryptodon, which is relatively thin (Figs 4B, 5B). In addition, O. nexipinnis uniquely has a conspicuous dermal invagination of unknown function just posterior to the point of attachment of the pelvic-fin base. Swollen cells in the middle epidermal layer, hypothesized to inhibit desiccation while on land (Yokoya & Tamura, 1992; Suzuki & Hagiwara, 1994), were reported in Boleophthalmus, Periophthalmodon, Periophthalmus and Scartelaos (Suzuki, 1992; Zhang et al., 2000, 2003; Park, 2002). We identify these cells in Oxuderces, but not in Apocryptodon. Dense networks of capillaries close to the epidermal surface have been reported in Boleophthalmus, Periophthalmodon, Periophthalmus and Scartelaos (Zhang et al., 2000, 2003), assumed to aid in cutaneous respiration while out of water. We observed these capillaries in both Apocryptodon and Oxuderces. In many fish, a thickened epidermis and associated modifications to the epidermis are correlated with the ability to breathe atmospheric air (see Park, Kim & Kim, 2014: 206). Periophthalmus, for example, can undergo cutaneous respiration facilitated by the intraepithelial capillaries that characterize the thickened epidermis (Park, Kim & Kim, 2000). Likewise, the thickened epidermis in Oxuderces may function as an accessory respiratory organ for gaseous exchange, possibly aiding in respiration in hypoxic habitats. Both species of Oxuderces further share a unique morphology: a fleshy, interorbital trough (Fig. 3A, B). The function of this trough is unknown, and it has not been reported in any other gobioid fishes. The presence and position of a single, median sensory pore C in the anterior part of the trough suggests that the function of the trough may be sensoryrelated. We are encouraged by the details revealed by histological preparation of the heads of oxudercine gobies, and plan to continue that examination as part of our larger study of oxudercine goby phylogeny. ACKNOWLEDGEMENTS We are grateful to the following collections managers and curators who allowed us access to material under their care: Mark Sabaj Perez (ANSP), Mark McGrouther (AMS), Dave Catania (CAS), Gavin Dally (NTM), Jeffrey Williams (USNM), Gento Shinohara (NSMT-P), James Maclaine (BMNH), Tracy Heath (BMNH), Ronald de Ruiter (RMNH), Denis Vallan (NMBA), Urs W uest € (NMBA), Kelvin Lim (ZRC) and Kazuo Sakamoto (ZUMT). We are thankful for the help extended by Greg Erickson, Jeffrey Clayton, Kris Murphy, Andrew Rao, Nalani Schnell and Harry Taylor. Victor Springer provided constructive comments on a draft version of the manuscript. Helen Larson and two other anonymous reviewers improved the manuscript with their suggested edits. Harry Taylor photographed the specimens in Figure 14. Sandra Raredon photographed the specimens in Figures 2, 3 and 12B. Helen Wimer prepared the histological slides. Lisa Palmer, Laura Tancredi, Amanda Lawrence and Erin Bilyeu, through the office of Paul Kimberly, assisted with photography of the histological slides. Colleen Lodge prepared Figures 3 – 5 and 7 – 10. Erin Clements Rushing and Leslie Overstreet, Smithsonian Institution Libraries, provided the image in Figure 1. This study was supported by grants from the Systematic Research Fund (2012/13) and the Leonard P. Schultz Fund, Smithsonian Institution, awarded to the first author. The Herbert R. and Evelyn Axelrod Chair in Systematic Ichthyology in the Division of Fishes (USNM) supported the preparation and publication of this paper. REFERENCES Agorreta A, San Mauro D, Schliewen U, Van Tassell JL, Kova � ci c � M, Zardoya R, R uber € L. 2013. Molecular phylogenetics of Gobioidei and the phylogenetic placement of European gobies. Molecular Phylogenetics and Evolution 69: 619 – 633. Bauchot ML, Whitehead PJP, Monod T. 1982. Date of publication and authorship of the fish names in Eydoux & Souleyet’s zoology of La Bonite, 1841 – 1852. Cybium 3e serie. Bulletin de la Société Francaise D’Ichtyologie 6: 59 – 73. Beon MS, Oh MK, Lee YJ, Kim CH, Park JY. 2013. A comparative study on vascularization and the structure of the epidermis of an amphibious mudskipper fish, Scartelaos gigas (Gobiidae, Teleostei), on different parts of the body and the appendages. Journal of Applied Ichthyology 29: 410 – 415. Berg LS. 1940. Classification of fishes both recent and fossil. Travaux de L’Institut Zoologique de L’Académie des Sciences de l’U.R.S.S 5: 87 – 345. Birdsong RS. 1975. The osteology of Microgobius signatus Poey (Pisces: Gobiidae), with comments on other gobiid fishes. Bulletin of the Florida State Museum Biological Sciences 19: 1 – 187. Bleeker P. 1849. Bijdrage tot de kennis der Blennioiden en Gobioiden van den Soenda-Molukschen Archipel, met beschrijving van 42 nieuwe soorten. Verhandelingen van het Bataviaasch Genootschap van Kunsten en Wetenschappen 22: 1 – 40. Bleeker P. 1874. Esquisse d’un systeme naturel des Gobioides. Archives Neerlandais de Sciences Naturelles, Haarlem 9: 289 – 331. Bloch ME, Schneider JG. 1801. Systema Ichthyologiae. Iconibus ex Illustratum. Post Obitum Auctoris Opus Inchoatum Absolvit. Correxit, Interpolavit Jo. Gottlob Schneider. Berlin: Facsimile, Cramer & Swann: Codicote. Boulenger GA. 1904. Fishes (Systematic Account of Teleostei). The Cambridge Natural History, London 7: 541 – 727. Cantor T. 1849. Catalog of Malayan fishes. Journal of the Asiatic Society of Bengal 18: 983 – 1443. Chu YT. 1931. Index Piscium Sinensium. Biological Bulletin of the St John’s University 1: iv+1 – iv291. Clayton DA. 1993. Mudskippers. In: Ansell AD, Gibson RN, Barnes M, eds. Oceanography and Marine Biology: An Annual Review 31: 507 – 577. Day F. 1871. On the fishes of the Andaman Islands. 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A synopsis of the fishes of China Part VII. Quarterly Journal of the Taiwan Museum 9: 161 – 354. Genten F, Terwindhe E, Danguy A. 2009. Atlas of fish histology. Enfield, NH: Science Publishers. Gill T. 1872. Arrangement of the families of fishes. Smithsonian Miscellaneous Collections 247: xlvi+1 – xlvi 49. Gill T. 1893. Families and subfamilies of fishes. Memoirs of the National Academy of Science 6: 127 – 138. Golvan YJ. 1965. Catalogue systématique des noms de genres de poissons actuels. Paris: Masson et Cie. Gosline WA. 1968. The suborders of Perciform fishes. Proceedings of the United States National Museum 124: 1 – 78. G unther € A. 1861. Catalogue of the acanthopterygian fishes in the collection of the British Museum, Vol. 3. London: Taylor and Francis. G unther € A. 1880. An introduction to the study of fishes. Edinburgh: Adam and Charles Black. Herre AW. 1927. Gobies of the Philippines and the China Sea. Monographs of the Bureau of Science, Manila, Philippine Islands 23: 1 – 352 + 26 pls. Hoese DF. 1984. Gobioidei: relationships. In: Moser HG, ed. Ontogeny and systematics of fishes. Special Publication of the American Society of Ichthyologists and Herpetologists No 1. Lawrence, KS: Allen Press, 588 – 591. Hoese DF, Gill AC. 1993. Phylogenetic relationship of eleotridid fishes (Perciformes: Gobioidei). Bulletin of Marine Science 52: 415 – 440. Ishimatsu A, Gonzales TT. 2011. Mudskippers: front runners in the modern invasion of land. In: Patzner RA, Van Tassell JL, Kova�ci�c M, Kapoor BG, eds. The biology of gobies. Enfield, NH: Science Publishers, CRC Press, 609 – 638. Ishimatsu A, Hishida Y, Takita T, Kanda T, Oikawa S, Takeda T, Khoo KH. 1998. Mudskippers store air in their burrows. Nature 391: 237 – 238. Jayne BC, Voris HK, Heang KB. 1988. Diet, feeding behavior, growth and numbers of a population of Cerberus rynchops (Serpentes: Homalopsinae) in Malaysia. Fieldiana 50: 1 – 15. Jayne BC, Ward TJ, Voris HK. 1995. Morphology, reproduction and diet of the marine homalopsine snake Bitia hydroides in Peninsula Malaysia. Copeia 1995: 800 – 808. Jordan DS. 1905. A guide to the study of fishes, Vol. 2. New York: Henry Holt and Company. Koumans FP. 1937. Notes on Gobioid fishes. 10. On the collection of the museum of Basle, with description of a new species of Apocryptodon. Zoologische Mededeelingen 20: 24 – 26. Larson HK. 2001. A revision of the gobiid fish genus Mugilogobius (Teleostei: Gobioidei), and its systematic placement. Records of the Western Australian Museum Supplement 62: 1 – 233. Lindberg GU. 1971. Opredelitel’i kharakteristika semeistv ryb mirovoi fauny. Russia: Leningrad. Murdy EO. 1989. A taxonomic revision and cladistic analysis of the oxudercine gobies (Gobiidae: Oxudercinae). Records of the Australian Museum 11 (Supplement): 1 – 93. Murdy EO. 2006. A revision of the gobiid fish genus Trypauchen. Zootaxa 1343: 55 – 68. Park JY. 2002. Structure of the skin of an air-breathing mudskipper, Periophthalmus magnuspinnatus. Journal of Fish Biology 60: 1543 – 1550. Park JY, Kim IS, Kim SY. 2000. Histology of skin of the amphibious fish, Periophthalmus modestus. Korean Journal of Biological Sciences 4: 315 – 318. Park JY, Lee YJ, Kim IS, Kim SY. 2003. A comparative study of the regional epidermis of an amphibious mudskipper fish Boleophthalmus pectinirostris (Gobiidae, Pisces). Folia Zoologica 52: 431 – 440. Park JY, Kim IS, Lee YJ. 2006. A study on the vascularization and structure of the epidermis of the air-breathing mudskipper, Periophthalmus magnuspinnatus (Gobiidae, Teleostei), along different parts of the body. Journal of Applied Ichthyology 22: 62 – 67. Park JY, Kim IS, Kim SY. 2014. Morphology and histochemistry of the skin of the mud loach, Misgurnus mizolepis, in relation to cutaneous respiration. Korean Journal of Biological Sciences 5: 303 – 308. Pezold F. 1993. Evidence for a monophyletic Gobiinae. Copeia 3: 634 – 643. Pezold F. 2004. Phylogenetic analysis of the genus Gobionellus (Te

Oxuderces DENTATUS EYDOUX & SOULEYET 1848

GENUS OXUDERCES EYDOUX & SOULEYET, 1848 Oxuderces Eydoux & Souleyet, 1848: 181 (type species Oxuderces dentatus Eydoux & Souleyet, 1848, type by monotypy) Composition: Two allopatric species: O. dentatus Eydoux & Souleyet, 1848 (Figs 3A, 11, 12) and O. nexipinnis (Cantor, 1849) (Figs 2, 3B, 6, 14). The distribution of each species is shown in Figure 13. Differential diagnosis: Oxuderces is differentiated from all other oxudercine gobies by five putative synapomorphies: (1) single, anterior interorbital pore C that sits in a fleshy, interorbital trough supported internally by expanded, curved medial margins of the frontal bones and lined externally with a thick epidermis and thin dermis (vs. no fleshy, interorbital trough, medial margins of the frontal bones expanded but not curved, and epidermis extremely thin and dermis thick); (2) highly thickened epidermis over the eye (vs. a thickened dermis); (3) neural spine of the fourth vertebra broad and spatulate (vs. narrow and pointed); (4) anterior ceratohyal elongate and notched posterior to insertion of fourth branchiostegal (vs. not elongate, or elongate but not notched); and (5) acute angle between metapterygoid – symplectic – quadrate strut and the anguloarticular (vs. angle obtuse). Description: Anterior portion of head markedly depressed; snout profile broadly pointed; gape wide; distinct notch in middle of upper lip between two medial premaxillary teeth; lips thick, posterior lower lip protruding distally. Roof of mouth with fleshy palp that is elliptical and with pointed tips, palp studded with papillae. Teeth in both premaxilla and dentary in single row; one or two prominent, elongate canine teeth on each side of premaxillary symphysis, canine teeth extending slightly anteroventrally and projecting beyond lower jaw when mouth closed, all teeth on premaxilla caninoid; dentary with discontinuous caninoid teeth; no canine tooth on each side of symphysis internal to anterior margin of dentary; dentary with small dorsally directed flange on each side just posterior to posterior-most tooth. Maxillo-dentary ligament with finger-like projections on dorsoposterior sides of lower lip; dentary with ligament on each side attaching medially just posterior to posterior-most tooth and antero-medially to flange (Murdy, 1989: fig. 73). Gill opening restricted, beginning from the region anterior to midpoint of pectoral-fin base, coursing anteroventrally, and ending just dorsal to pelvic-fin origin. Eyes positioned anterodorsally but not meeting medially, no dermal cup or membrane covering ventral portion of eye. Thickened dermal layer (or secondary cornea) over dorsal portion of eye, extending, and gradually thinning out ventrally and posteriorly (Figs 4, 5). Single anterior interorbital sensory pore (pore C) at anterior edge of fleshy trough or groove; trough covered externally by thickened dermal layer extending from dorsal area of eyes; a pair of anterior oculoscapular canal pores present. Posterior nostril large, anteroventral to eye; anterior nostril opening ventrally at tip of pendulous short tube overlapping upper jaw. Sphenotic bones small, not contacting eyes. Frontal bone elongate; lateral processes on medial portion of frontal distinct; frontal bone fused, forming interorbital bridge, slightly curved, not overlapping ethmoid anteriorly (Fig. 5A). Maxilla and premaxilla terminating posterior to eye, maxilla not reaching retroarticular. Palatine terminating before mid- If a specimen was damaged or abnormal (e.g. it has obvious parasites), we did not record data; this is noted in the table by the abbreviation DNT. Data for the holotype of O. dentatus are not included in the range reported for the species. ectopterygoid. Ectopterygoid thin and elongate, meeting with quadrate posteriorly (Fig. 7A). Metapterygoid terminating at hyomandibular junction; angle between metapterygoid – symplectic – quadrate strut and anguloarticular acute (vs. angle obtuse). Preopercle subcrescent-shaped, thin, meeting hyomandibula dorsally. Basihyal bone triangular with continuous cartilaginous anterior margin. Branchiostegal rays 5, rays thin. No laminar processes on parapophyses of fourth vertebrae; neural spine of fourth vertebra broad and spatulate. D1 and D2 connected by membrane for entire height, shallow indentation separating D1 and D2 due to varying height of fin elements. No membrane connecting D2 or A to caudal fin. D1 with six spinous elements, each spine with associated pterygiophore; spaces between first five D1 spines subequal, base of sixth spine positioned approximately midway between base of fifth D1 spine and first element of D2; all elements in D2 and A fins are segmented rays; penultimate and ultimate rays of D2 and A fins supported by single pterygiophore; caudal fin lanceolate, with 17 segmented rays; pelvic fin short and rounded, not reaching genital papilla. Caudal peduncle cartilage elongate, reaching anterior extent of penultimate vertebra. Vertebrae slightly elongate, vertebrae number 10 + 16, two epural bones present. Dense network of capillaries close to epidermal surfaces, especially on head. Males with triangular, conical genital papilla, with posterior tip pointed; females with bulbous, rectangular genital papilla. No other observed external sexual dimorphism. Etymology: Greek in origin, derived from the word ‘ oxyderkes ’ meaning ‘sharp-sighted’ (Murdy, 1989). Gender masculine. Remarks: The genus Oxuderces, and type species O. dentatus, was first described by Eydoux and Souleyet based on a single specimen collected from Macao along with some 200 other fish species aboard the corvette La Bonite between 1836 and 1837 (Bauchot, Whitehead & Monod, 1982; Figs 1, 11). The unique character of the holotype that prompted its description as a new genus and species (Eydoux & Souleyet, 1848:181) was a peculiar gill opening interpreted by Springer (1978:1) as ‘... continuous across the isthmus and restricted to the ventral surface of the head’. The holotype and, until 1978, the sole scientific specimen identified as O. dentatus also lacks pelvic fins (Fig. 12A; Springer, 1978: fig. 4b). Springer (1978) concluded that the absence of pelvic fins in the holotype was abnormal, although it was not considered so by Eydoux & Souleyet (1848) who compared Oxuderces to the ‘anarrhiques’, the French vernacular for wolffishes, currently classified as the family Anarhichadidae, which typically lack pelvic fins. Although Eydoux & Souleyet (1848) placed their new genus in the family Gobioides, congruent with the modern Gobiidae, classification of Oxuderces has been unstable (Springer, 1978). It has been subsequently placed in its own family, the Oxudercidae (G unther€, 1861; Gill, 1872, 1893; Reeves, 1927; Chu, 1931), which was classified alternatively as a trachinoid (Berg, 1940; Fowler, 1956; Schultz, 1960; Golvan, 1965; Lindberg, 1971), blennioid (Gosline, 1968) or gobioid fish (G unther€, 1880; Boulenger, 1904; Jordan, 1905). Many of these ichthyologists who reclassified O. dentatus did not examine the holotype or, even if they did, were probably diverted to a search for close relatives among other fishes that also lacked pelvic fins, such as the wolffishes. No other specimen of Oxuderces has been identified without pelvic fins. A After examining the holotype, Springer (1978) returned Oxuderces dentatus to the family Gobiidae, a decision followed by all subsequent workers (e.g. Hoese, 1984; Murdy, 1989; Pezold, 1993). The monotypic Apocryptichthys Day, 1876, allied historically with Oxuderces, was described for Apocryptes cantoris Day, 1871. Confusion surrounding the identity of Apocryptichthys cantoris stemmed from the inconsistent descriptions in the literature and figures by Day (1871, 1876, 1889; and see Springer, 1978: 10 for a tabulated summary). These descriptions varied from publication to publication. The original description of Apocryptichthys and the figure included of the type specimen Apocryptes cantoris (Day, 1876) conform to Oxuderces (probably referring to AMS B.8336 from Madras), whereas the original description of Apocryptes cantoris (probably referring to BMNH 1870.5.18.23 from the Andaman Islands) conforms to Scartelaos histophorus (Valenciennes, 1837). Springer (1978) designated a lectotype for Apocryptes cantoris Day, 1871 (BMNH 1870.5.18.23) from an unspecified number of syntypes and classified the species in the genus Boleophthalmus Valenciennes, 1837. Murdy (1989) reclassified this species in Scartelaos Swainson, 1839, a decision with which we concur, as does V. G. Springer (pers. comm.). As the original description of Apocryptichthys may be interpreted to include Oxuderces, we have included Apocryptichthys, in part, in our treatment of Oxuderces. Ecological note: Published information on the biology and natural history of Oxuderces may refer to what we now recognize as three allopatric species classified in two genera. We interpret information on specimens from eastern China to refer to O. dentatus, that on specimens from the Indo-West Pacific, excluding China, to refer to O. nexipinnis, and that on specimens from northern Australia and New Guinea to refer to Apocryptodon wirzi. General information on O. dentatus may refer to either one of the species of Oxuderces.Published as part of Jaafar, Zeehan & Parenti, Lynne R., 2017, Systematics of the mudskipper genus Oxuderces Eydoux & Souleyet 1848 (Teleostei: Gobiidae: Oxudercinae) with resurrection from synonymy of O. nexipinnis (Cantor 1849), pp. 195-215 in Zoological Journal of the Linnean Society 180 (5) on pages 201-206, DOI: 10.1111/zoj.12482, http://zenodo.org/record/571094