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1 aqua International Journal of Ichthyology Vol. 18 (2), 15 April 2012 Aquapress ISSN

2 aqua - International Journal of Ichthyology Managing Editor: Heiko Bleher Via G. Falcone 11, Miradolo Terme (PV), Italy Tel./Fax: heiko@aquapress-bleher.it Scientific Editor: Frank Pezold College of Science & Engineering Texas A&M University Corpus Christi 6300 Ocean Drive Corpus Christi, TX Tel frank.pezold@tamucc.edu Editorial Board: Gerald R. Allen Department of Aquatic Zoology, Western Australian Museum, Perth, Australia Nina G. Bogutskaya Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia Friedhelm Krupp Curator of Fishes, Senckenberg Research Institute and Natural History Museum, Frankfurt am Main, Germany Flávio C. T. Lima Museu de Zoologia da Universidade de São Paulo São Paulo, Brasil Axel Meyer Lehrstuhl für Zoologie und Evolutions biologie, Universität Konstanz, Germany Paolo Parenti Department of Enviromental Sciences, University of Milano-Bicocca, Milan, Italy Mário de Pinna Museu de Zoologia da USP, São Paulo, Brazil John E. Randall Bishop Museum, Honolulu, Hawaii, U.S.A. Richard Winterbottom Centre of Biodiversity & Conservation Biology, Royal Ontario Museum, Toronto, Canada Helen K. Larson Curator Emeritus, Fishes - Museum and Art Gallery of the Northern Territory. Darwin, Australia Scope aqua is an international journal which publishes original scientific articles in the fields of systematics, taxonomy, bio geography, ethology, ecology, and general biology of fishes. Papers on freshwater, brackish, and marine fishes will be considered. aqua is fully refereed and aims at publishing manuscripts within 2-4 months of acceptance. In view of the importance of color patterns in species identi - fication and animal ethology, authors are encouraged to submit color illustrations in addition to descriptions of coloration. It is our aim to provide the international scientific community with an efficiently published journal meeting high scientific and technical standards. Call for papers The editors welcome the submission of original manuscripts which should be sent in digital format to the scientific editor. Full length research papers and short notes will be considered for publication. There are no page charges and color illustrations will be published free of charge. Authors will receive one free copy of the issue in which their paper is published and an e-print in PDF format. Subscription Notice At least one volume (4 issues) of aqua is being published per year, each issue comprising pages (incl. cover). The subscription rate (for one volume = 4 issues): Personal subscription: Euro 75,00 (online edition) - Euro 100,00 (online edition + print edition incl. priority mail); Institutional subscription: Euro 800,00 (online edition) - Euro 1000,00 (online edition + print edition incl. priority mail). Subscription enquires should be sent to the publisher at the address given below or by to: aqua@aquapress-bleher.it - aqua@aquapress-bleher.com Special Publication Since 2003 Aquapress publishes a series of Special Publications, which are produced at irregular intervals. All Special Publications have about 100 or more pages and are available separately from regular issues of aqua. Enquiries about subscriptions and prices should be sent to the publisher at the address given here above or by to: aqua@aquapress-bleher.it - aqua@aquapress-bleher.com ISSN Publisher: Aquapress, Redazione aqua, I Miradolo Terme (Pavia), Italy Printer: Pronto Stampa Srl Bergamo Italy Copyediting and layout: Rossella Bulla 2012 aqua, International Journal of Ichthyology

3 aqua, International Journal of Ichthyology Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species James L. Van Tassell 1*, Luke Tornabene 2, Patrick L. Colin 3 1) American Museum of Natural History, Central Park West at 79th Street, New York, NY , U.S.A. *Corresponding author: jvantassell@gobiidae.com 2) Texas A&M University Corpus Christi, 6300 Ocean Drive, Corpus Christi, TX 78412, U.S.A. 3) Coral Reef Research Foundation P.O. Box 1765 Koror, Palau Received: 20 May 2011 Accepted: 30 September 2011 Abstract Bollmannia Jordan is a poorly studied group of American seven-spined gobies with representatives in the tropical and subtropical western Atlantic and tropical eastern Pa - cific oceans. We review the taxonomy of the western Atlantic species and provide redescriptions for the four valid species: B. boqueronensis, B. communis, B. eigenmanni and B. litura. Bollmannia jeannae is considered to be a junior synonym of B. boqueronensis. We also describe a new genus and species of deep-water goby and discuss its affinities to Bollmannia and other genera of the Microgobius group of the Gobiosomatini. An identification key is provided for all western Atlantic members of the Microgobius group. Results of this study highlight the need for a combined morphological and molecular phylogenetic analysis to resolve the relationships among the genera of the Microgobius group. Zusammenfassung Bollmannia Jordan ist eine kaum erforschte Gruppe amerikanischer Grundeln mit sieben Stacheln mit Vertretern im tropischen und subtropischen Westatlantik und im tropischen Ostpazifik. Wir überprüfen hier die Systematik der westatlantischen Arten und geben Neubeschreibungen der vier gültigen Arten: B. boqueronensis, B. communis, B. eigenmanni und B. litura. Bollmannia jeannae wird als jüngeres Synonym zu B. boqueronensis aufgefasst. Wir be - schrei ben außerdem eine neue Gattung und Art der Tief - wasser-grundeln und diskutieren ihre verwandtschaftlichen Bezüge zu Bollmannia und anderen Gattungen der Microgobius-Gruppe der Gobiosomatini. Schließlich wird ein Bestimmungsschlüssel für alle westatlantischen Angehörigen der Microgobius-Gruppe wiedergegeben. Das Ergebnis der vorliegenden Untersuchung unterstreicht die Not wen - digkeit einer kombiniert morphologischen und molekularbiologischen phylogenetischen Analyse, um die Verwandt - schaftsbeziehungen zwischen den Gattungen der Microgobius-Gruppe zu klären. Résumé Bollmannia Jordan est un groupe peu étudié de gobies américains à sept épines avec des représentants dans l Atlantique ouest tropical et subtropical et dans le Pacifique est tropical. Nous faisons une révision de espèces de l Atlantique ouest et donnons la redescription des quatre espèces reconnues : B. boqueronensis, B. communis, B. eigenmanni et B. litura. Bollmannia jeannae est considéré comme un synonyme plus récent de B. boqueronensis. Nous décrivons aussi un nouveau genre et une nouvelle espèce de gobie des eaux profondes et en discutons les affinités avec Bollmannia et d autres genres du groupe des Microgobius appartenant aux Gobiosomatini. Une clé de détermination est fournie pour tous les membres de l Atlantique ouest du groupe des Microgobius. Les résultats de cette étude mettent en évidence la nécessité d une analyse phylogénétique combinée, à la fois morphologique et moléculaire, pour tirer au clair les relations à l intérieur des genres du groupe Microgobius. Sommario Il genere Bollmannia Jordan comprende un gruppo di ghiozzi a sette spine poco studiati che abitano le aree tropi - ca li e subtropicali dell Atlantico occidentale e quelle tropicali del Pacifico orientale. Si presenta una revisione della tas sonomia delle specie atlantiche accompagnata da una nuova descrizione di quattro specie valide: B. boqueronensis, B. communis, B. eigenmanni e B. litura. Bollmannia jeannae è da considerarsi sinonimo di B. boqueronensis. Inoltre, viene descritto un nuovo genere e una nuova specie di ghiozzo di acque profonde discutendone le affinità con Bollmannia e altri generi del gruppo Microgobius della tribù Gobiosomatini. Per tutti i membri del gruppo Microgobius dell Atlantico occidentale è fornita una chiave dicotomica. I risultati di questo studio sottolineano la necessità di un analisi filogenetica combinata con dati morfologici e molecolari per risolvere le relazioni di parentela tra i generi del gruppo Microgobius. 61 aqua vol. 18 no April 2012

4 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species INTRODUCTION Bollmannia Jordan, 1890, is one of five genera that make up the Microgobius group of the tribe Gobiosomatini (Birdsong et al. 1988; Van Tassell & Baldwin 2004). The Gobiosomatini is a monophyletic group within the subfamily Gobiinae (sensu Pezold 1993; Thacker 2003; Thacker & Hardman 2005) and the tribe is often referred to as the American seven-spined gobies. Bollmannia currently comprises 14 nominal species, five of which were described from the Atlantic Ocean: Bollmannia eigenmanni (Garman, 1896), B. boqueronensis Evermann & Marsh, 1899, B. jeannae Fowler, 1941, B. communis Ginsburg, 1942, and B. litura Ginsburg, While attempting to identify specimens of Bollmannia collected from Panama in 2004, several problems were noted in the descriptions of the Atlantic species. The characters described for each of the species are not uniform across each of the descriptions. Several of the characters described as diagnostic are of little value as these characters are commonly destroyed when Bollmannia are trawled or dredged from considerable depths. In most cases the descriptions are based on a small number of specimens and do not adequately summarize the intraspecific variation seen among populations. Furthermore, there is a discrepancy between the description of B. litura and the corresponding holotype. As a result of these issues, the validity of several species is questioned. Bollmannia jeannae is not listed by Nelson et al. (2004), McEachern & Fechhelm (2005) or Murdy & Hoese (2002). Both B. jeannae and B. eigenmanni are omitted by Smith (1997). Eschmeyer (2011) lists all five nominal species as valid. This paper attempts to clarify these discrepancies and resolve the status of the Atlantic members of the genus. No formal review of western Atlantic Bollmannia has been published to this point. In an unpublished master s thesis, Bedenbough (1988) reviewed the Pacific members of the genus. Ginsburg addressed the genus in two incomplete, unpublished manuscripts, the drafts and notes of which have been found at the Smithsonian Institution s National Museum of Natural History. Work on these manuscripts likely began between 1939 and 1942, as some of his notes included the eastern Pacific Bollmannia species described in Ginsburg (1939) but not B. communis, which he described in Subsequent drafts by Ginsburg contained B. communis and additional specimens of each of the species discussed in prior drafts. In his unpublished works Ginsburg described and illustrated sensory papillae and head pore patterns for several species of Bollmannia. We describe these characters for each of the valid species in the western Atlantic. In addition to the five nominal species of western Atlantic Bollmannia, we also address a previously undescribed deepwater goby from the Caribbean. This species was originally documented as the filamentous goby by Colin (1974) and subsequently referenced by Dennis et al. (2004), where it was referred to as the sabre goby. The molecular phylogenies of Rüber et al. (2003) and Rüber & Van Tassell (unpublished, presented in 2006) suggested that this species belongs within the monophyletic Microgobius group of the tribe Gobiosomatini, specifically as the sister group to a monophyletic clade containing four species of Bollmannia. Despite the apparent close relationship between this species and Bollmannia, there are several prominent differences between the two in terms of morphology and habitat. Therefore, we describe this species and place it in a new genus, Antilligobius, within the Microgobius group of the Gobiosomatini. The affinities between Antilligobius n. gen. and the allied genera Bollmannia, Akko, Parrella, Palatogobius and Microgobius are discussed. Identification keys are provided for the Atlantic species of the Microgobius group of the Gobiosomatini. METHODS Methods for counts and measurements follow Böhlke & Robins (1968) in most regards. The last two elements in the second dorsal and anal fin stem from a single pterygiophore and are counted as a single ray. The upper and lower segmented caudal rays are frequently short with only one or two striae but are stouter than adjacent procurrent rays and are included in the counts of segmented caudal rays. Counts of branched caudal rays include rays with only slight terminal branching. Lateral scale counts are the number of diagonal rows beginning with the scale above and behind the posterodorsal corner of the operculum and ending at the posterior edge of the hypural plate. Transverse scale counts are the number of horizontal scale rows counted from the origin of the second dorsal fin diagonally to the origin of the anal fin. Caudal peduncle scales are counted beginning at the first full sized scale on top of the caudal peduncle immediately in front of the caudal fin, and following the scale rows down and forward to aqua vol. 18 no April

5 James L. Van Tassell, Luke Tornabene, Patrick L. Colin the ventral edge of the peduncle, then around and back to the original scale. Gill raker counts are total counts for the outer edge of the first gill arch. The following measurements are straight-line point to point measurements taken with digital calipers: standard length, snout tip to posterior edge of hypural plate; head length, snout tip to posterior edge of operculum, not including fleshy membrane; predorsal length, distance from the insertion point of the first dorsal spine to the tip of the snout; snout length,snout tip to the anterior rim of eye; upper jaw length, anterior tip of upper jaw to posterior margin of maxillary; eye diameter, horizontal diameter of orbit; pupil diameter, horizontal diameter of pupil; postorbital length, posterior rim of eye to posterior edge of operculum, not including fleshy membrane; body depth at first dorsal fin origin,vertical distance from origin of spinous dorsal fin to ventral profile, not including pelvic fin; least caudal peduncle depth, vertical distance at shallowest point between base of last anal ray to posterior edge of hypural plate; and caudal fin length, distance from origin of ray on hypural plate to tip of longest ray. Dorsal pterygiophore formula is that of Birdsong et al. (1988), patterns of sensory papillae are described according to Sanzo (1911) and cephalic canal pore terminology follows Akihito et al. (1988). In some cases papillae and pore patterns were more easily observed by staining with cyanine blue (Saruwatari et al. 2006). Osteological observations were made from radiographs and specimens cleared with trypsin and counter stained for cartilage and bone (Dingerkus & Uhler 1977). Institutional acronyms follow Sabaj Pérez (2010). In many cases scale counts of Bollmannia were determined by counting scale pockets or remnants thereof, as scales in this genus are highly deciduous and specimens are typically in poor condition as a result of collection via trawl or dredge. In many other specimens, including some holotypes, scale pockets were missing entirely. For this reason, the scale count ranges presented here are trends from a composite of specimens and should be regarded as approximations. Similarly, the fins are heavily damaged on most specimens of Bollmannia. The relative lengths of rays and spines described here should also be regarded as approximations. Papillae and pore patterns shown here are composite drawings from several specimens. In nearly every specimen of Bollmannia, portions of the lateralis canals are destroyed and in many cases entire rows of papillae are missing as a result of damage or poor preservation. In particular, the predorsal region, the posterior half of the preoperculum and the operculum are typically more heavily damaged than other regions. Thus the absence of extensive papillae or cephalic canals in these regions (specifically the canal between K and L ) on our illustrations may be subject to correction if specimens are examined in which papillae and canals are more perfectly preserved. Despite these issues there is some taxonomic value in the aforementioned characters. However, this value, and consequently the ability to confidently identify specimens of Bollmannia, is largely determined by the overall physical condition of the specimen. SYSTEMATICS Antilligobius, n. gen. Van Tassell & Tornabene Diagnosis: Antilligobius can be distinguished from other Gobiosomatini by the following combination of characters, not listed in order of taxonomic importance: first dorsal fin very elongate in both sexes, extending posteriorly at least to base of last anal fin ray when depressed, sometimes reaching middle of caudal fin; caudal fin elongate but with upper rays 3,4,5 shorter then corresponding lower rays, forming notch in upper margin of fin; pelvic fins united to form disk with well-developed frenum, frenum with smooth posterior margin, not supported with collagenous thickenings; trunk scales ctenoid, lateral scale rows 21-28; predorsal region, cheek, pectoral fin base and breast with cycloid scales; predorsal scales 8-10; first caudal vertebra with expanded haemal arch and tiny haemal spine; haemal arch with elongate parapophyses, riblike in appearance in lateral view; abdominal cavity elongated and extending through first haemal arch. Description: Body slightly elongate and laterally compressed; first dorsal spines VII; pterygiophore formula of first dorsal fin 3(221110); second dorsal fin I,11-12; first and second dorsal fins separate from each other and from caudal fin; pectoral rays 18-21; anal fin I,11-13, first two anal pterygiophores inserted anterior to first haemal spine, remaining pterygiophores in approximately 1:1 ratio with caudal vertebrae; total vertebrae 27, 11 precaudal and 16 caudal; pelvic fins with one spine and five rays; vomerine teeth absent; mouth large, % SL and inclined 30 from horizontal; 63 aqua vol. 18 no April 2012

6 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species eye large, diameter % SL; gill opening extending from upper edge of pectoral fin to ventral body midline; gill rakers on lower limb of first arch thin and elongate, rakers on upper arm short, stubby; urogenital papilla thin and conical in males, thick and bulbous in females; cephalic lateralis canal and pore system on head well-developed; sensory papillae in transverse pattern; ventral postcleithrum present; no obvious sexual dimorphism, except for urogenital papilla. Comparison: Antilligobius has a number of characters not present in other American seven-spined genera that warrant its placement in a new genus. These include: a very narrow, splinter-like metapterygoid; short slender gill rakers on the lower limb of the first arch; a short papillae row b, never extending to posterior of the pupil; a modified first haemal arch; the caudal fin elongate but with the upper rays 3,4,5 shorter than the corresponding rays on the ventral half of the fin, forming a notch in the upper margin of the fin. Antilligobius is most similar morphologically to Bollmannia and Parrella, sharing with those genera the presence of scales on the breast, belly, pectoral fin base and predominately ctenoid scales on the trunk. Bollmannia and Antilligobius also both possess scales on the cheek and a spatulate basihyal. The bright yellow stripe and blue colors of Antilligobius easily distinguish it from Bollmannia and Parrella which are drab brown with darker brown markings. Etymology: The generic epithet Antilligobius is formed from the Dutch Antillen, which refers to the region now known as the Antilles or Caribbean Sea, and the Latin gobius meaning small fish or gudgeon. To be treated as masculine. The name is given in reference to its distribution within the ancient Sea of the Antilles. Antilligobius nikkiae, n. sp. Van Tassell & Colin Sabre goby (Figs 1-2, 4-11) Fig. 1A-B. A. Antilligobius nikkiae, holotype, 30.2 mm SL, female, Curaçao, AMNH , preserved; B. Antilligobius nikkiae, paratype, mm SL, male, Curaçao, AMNH Photos by J. Van Tassell. Image editing by L. Tornabene. aqua vol. 18 no April

7 James L. Van Tassell, Luke Tornabene, Patrick L. Colin Table I. Morphometrics for type specimens of Antilligobius nikkiae (excluding cleared and stained specimen). Measurments other than SL are in % SL. AMNH AMNH AMNH AMNH AMNH ANSP ANSP USNM (holotype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) (paratype) Sex female female male male female male female female SL (mm) Eye diameter Upper jaw length Head length Postorbital length Depth at DI origin Least caudal peduncle depth Snout length Holotype: AMNH , 30.2 mm SL, female, Curaçao Sea Aquarium slope, Curaçao, N, W, quinaldine collection, collected by Adriaan Dutch Schrier, July 2004, depth 90 m, loose rock slope (Fig. 3). Paratypes: AMNH , 2 males, mm SL, 1 female, 26.7 mm SL, collected with holotype; ANSP , 29.7 mm SL, female, cleared and stained specimen, shelf edge of reef at La Parguera, Puerto Rico, 28 August 1979, Pat Colin; ANSP , 1 male 21.7 mm SL, 1 female 25.7 mm SL, shelf edge of reef at La Paruera, N, W, m, 45 slope, collected among small indentations in rocky structure of slope face, Puerto Rico, 22 November 1978, Pat Colin; AMNH , 1 male, mm SL, 90 m, JVT , Genbank AF491066, Curaçao Sea Aquarium slope, N, W, Curaçao, January 2001, Adriaan Dutch Schrier; USNM , 1 female, 31.6 mm SL, m, west coast of Isla de la Juventud, N, W, Cuba, 27 December 1997, R. Grant Gilmore. Description: Morphometric data given in Table I. Counts of the holotype (Fig. 1) indicated by an asterisk, followed by number of specimens with each count in parentheses. Median and paired fins: first dorsal VII*(9); first four dorsal spines very elongate, extending to base of last anal ray when depressed, extending to middle of caudal fin in some specimens; second dorsal I,11(4), I,12*(5); anal fin I,11(1), I,12*(1), I,13(7); pectoral rays 18(2), 20*(6), 21(1); pectoral fin elongate, reaching vertical through anterior end of anus; caudal fin elongate but with upper rays 3,4,5 shorter than corresponding lower rays, forming notch in upper margin of fin; segmented caudal rays 16(3), 17* (6), branched rays 14*(6), 15(3); pelvic fin I,5*(9); pelvic fins united to form disk, disk extending 2/3 to ¾ distance from insertion of spine to anterior of anus; anterior margin of frenum extending halfway along pelvic spine, posterior margin of frenum smooth, without fimbriae and lacking collagenous thickenings. Scales: body completely covered with ctenoid Fig. 2. Antilligobius nikkiae in natural habitat, south-west Puerto Rico, approximately 90 m. Photos by P. Colin. 65 aqua vol. 18 no April 2012

8 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with cycloid scales; lateral scale rows 21(1), 25(4), 26*(2), 28(1); transverse scale rows 6(2), 7*(4); predorsal scales 7(1), 8(4), 9(1), 10*(2), scales extending anteriorly to behind eye; caudal peduncle scales 9(1), 10(3), 11*(2), 12(1); scales on cheek 3(4), 4*(2), 5(2); operculum scales 0*(4), 3(4); no modified basicaudal scales present. Head: head length % SL; mouth angled upwards 30 from horizontal, reaching posteriorly to vertical through pupil; upper jaw length % SL; teeth in upper jaw in 2-3 rows anteriorly, becoming two rows about midway along premaxilla, ending in single row at posterior end of premaxilla; all teeth conical with slightly recurved tips; teeth in outer row longest; teeth in lower jaw in two rows anteriorly, becoming single row about half way along dentary; teeth conical with slightly recurved tips; single recurved canine located at mid-dentary; tongue slightly emarginate; eye large, % SL; interorbital width approximately equal to pupil diameter; snout short, % SL; Fig. 3. Habitat at the type locality of Antilligobius nikkiae, Curaçao Sea Aquarium slope, approximately 90 m. Arrows indicate individual gobies. Photos by A. Schrier and C. Baldwin. Fig. 4. Sensory papillae and head canal pores of Antilligobius nikkiae. Drawing by J. Van Tassell. aqua vol. 18 no April

9 James L. Van Tassell, Luke Tornabene, Patrick L. Colin gill opening extending from upper edge of pectoral fin to ventral midline; gill rakers on lower arm of first arch 12-13, upper arm 5; gill rakers on lower arm of first arch thin and elongate, rakers on subsequent arches shorter and thicker; epibranchials present; anterior nostril a short erect tube; posterior nostril with raised rim along anterior edge; no barbels present. Genitalia: papilla elongate and conical in males, with distinct line of melanophores along dorsal edge present in one specimen, all others with no pigment; papilla short, bulbous, and without melanophores in females. Sensory papillae and head pores (Fig. 4): oculoscapular pores B, C(s), D(s), E, F, G, H present; posterior canal (typically between pores K and L ) absent; preopercular pores M, N, O present; single interorbital canal; papillae in transverse pattern, with 5 short vertical rows below eye, none extending below level of row d; row d long, beginning below anterior margin of eye, ending before transverse row 4; row b short, beginning below eye and posterior to 5th transverse row, ending well short of posterior margin of preopercle; row n continuous across dorsal midline; rows g and m reduced, each represented by 2-3 papillae on each side of dorsal midline. Pigment: body translucent pearlescent in life, tan in preservation; body with bright yellow stripe (white after preservation) along lateral midline beginning at eye and extending onto caudal fin; width of stripe about ¾ of eye diameter at widest point, becoming thinner on caudal fin, ending at or near tip of caudal fin; thin iridescent blue stripe along ventral surface of yellow stripe anteriorly, absent in preservation; dorsal midline of body with narrow iridescent blue stripe, absent in preservation; abdomen and gular regions more or less pale; lateral edge of upper and lower jaw reddish-orange; iris of eye iridescent blue along dorsal and ventral margins, with iridescent patches of yellow at the anterior and posterior margins, iris uniformly dark in preservative; first dorsal fin translucent and tinged with yellow on interspinal membranes, uniformly clear in preservation; membranes between spines 4-6 of first dorsal fin with diffuse dark streak, streak more prominent in preservation; second dorsal fin yellow at base, translucent distally, uniformly clear with dusky distal margin in preservation; anal fin translucent with scattered Fig. 5. Suspensorium and jaw osteology of Antilligobius nikkiae. Abbreviations ART, anguloarticular; D, dentary; MPT, metapterygoid; MX, maxilla; PMX, premaxilla; PT, ectopterygoid; QU, quadrate; SYM, symplectic; Photo by J. Van Tassell, editing by L. Tornabene. Fig. 6A-B. A. Cranial osteology (orbit) of Antilligobius nikkiae, lateral view; B. Cranial osteology (roof) of Antilligobius nikkiae, dorsal view. Abbreviations - F, frontal; ME, median ethmoid; SOC, supraoccipital; SPH, sphenotic. Photos by J. Van Tassell, editing by L. Tornabene. 67 aqua vol. 18 no April 2012

10 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Fig. 7. A. Antilligobius nikkiae left pectoral fin osteology; B. Antilligobius nikkiae pectoral girdle, ventral view; C. Antilligobius nikkiae right pectoral fin girdle, with reduced ossification. Abbreviations CL, cleithrum; COR, coracoid; PTM, posttemporal; RAD, radial; SCA, scapula; SCL, supracleithrum; VPCL, ventral postcleithrum. Photos by J. Van Tassell, editing by L. Tornabene. Fig. 8A-C. A. hypural plate of Antilligobius nikkiae; B. Ribs and haemal arch of Antilligobius nikkiae; C. precaudal vertebrae of Antilligobius nikkiae, note the large vertebral foramina. Abbreviations EPU, epural; HYP, hypural; HA, haemal arch; HS, haemal spine; NS, neural spine, PC, procurrent cartilage; PHYP, parahypural; PR, pleural rib; US, urostyle; VF, vertebral foramen. Photos by J. Van Tassell, editing by L. Tornabene. aqua vol. 18 no April

11 James L. Van Tassell, Luke Tornabene, Patrick L. Colin melanophores along dorsal and ventral edges and with horizontal yellow stripe along midline of fin in life, stripe pale in preservation; pelvic fin translucent with yellow dorsal margin in life, pale in preservation; caudal fin translucent with yellow stripes in life, dusky in preservation; two caudal fin stripes continuations of midlateral yellow body stripe and yellow stripe along base of anal fin; two additional yellow stripes on caudal fin, one thin one along dorsal fin margin and another along ventral margin; stripes pale in preservation; upper half of pectoral fin base yellow (white in preservation), lower half unpigmented; pectoral rays translucent. Osteology: based on ANSP , female, cleared and stained specimen. Suspensorium and jaws (Fig. 5): anterior arm of suspensorium consisting of palatine, relatively broad dorsally, tapering to point about midway along ectopterygoid; ectopterygoid articulating with quadrate along its anterodorsal edge; metapterygoid very thin, needle-like, not articulating with quadrate or its small cartilaginous posterior edge; no symplectic process on preopercle; ascending process of premaxilla extending dorsally to anterodorsal edge of rostral cartilage; maxilla narrow, lacking process along shaft, with ventral tip curved anteriorly; dentary reduced, lacking large pocket typically receiving anguloarticular; anguloarticular articulating with dentary along ventral groove in dentary and in small anterior pocket near symphysis of dentary. Cranial (Fig. 6): median ethmoid forming septum between eyes, anterior and posterior edges ossified; anterior of frontals overlapping median ethmoid; sagittal crest present along dorsal midline where left and right frontal bones fuse; sagittal crest confluent with supraoccipital crest; anterolateral edge of frontals possessing well-developed transverse process; this process forming open tube supporting cephalic sensory canal; sphenotic short, lacking well-developed anterior process. Pectoral girdle (Fig. 7): left side of specimen with four partly ossified radials and cartilaginous scapula; radials on right side not ossified, appearing fused into single cartilaginous plate along with scapula; posttemporal possessing single reduced lateral shelf; ventral postcleithrum present. Vertebral column and caudal skeleton (Fig. 8): precaudal vertebrae 11; caudal vertebrae (including terminal element) 16; vertebrae 3-11 with pleural rib articulating with respective parapophyses; twelve pairs of epineurals, articulating with vertebrae as follows: epineurals 1, 2 with corresponding parapophyses, 3-10 with corresponding pleural ribs, epineural 11 reduced and not articulating with pleural rib and 12 highly reduced and embedded in tissue; first caudal vertebra with expanded haemal arch and tiny haemal spine; haemal arch comprising elongate parapophyses, rib-like in appearance in lateral view; where parapophyses join ventrally to form haemal arch, a nubbin of bone (haemal spine) projecting posteriorly (Fig. 8B); rib-like lateral side of haemal arch easily confused with pleural ribs on radiographs; dorsal fin pterygiophore formula 3(221110); 2 pterygiophores of anal fin preceding first haemal arch; hypurals 1-2 not fused with hypurals 3-4; foramen on neural arches 6-22 singular and very large; neural arches 1, 24 through 27 with no foramen; neural arches 2, 3, and 4 with single small foramen; neural arch 5 with two foramina. Hyoid series (Fig. 9): branchiostegals 5, first branchiostegal small, thin, and attached to ventral surface of narrow portion of ceratohyal, following three branchiostegals blade-like in appearance and attached at ventrolateral surface of broad portion of ceratohyal; posteriormost branchiostegal broadest, articulating with ventral surface of epihyal; Fig. 9. Basihyal (BH) of Antilligobius nikkiae, spatulate in shape with only posterior one-third ossified. Photo by J. Van Tassell, editing L. Tornabene. 69 aqua vol. 18 no April 2012

12 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species basihyal broad, spatulate, slightly emarginate, with only posterior third ossified (Fig. 9); ventral hypohyals, dorsal hypohyals and urohyal ossified; urohyal extremely thin; basibranchials 2, 3 ossified; basibranchial 4 either absent, unstained or too small to discern; hyobranchials 1, 2, and 3 cartilaginous; ceratobranchials 1, 2, 3, 4 cartilaginous at Fig. 10. Cheek myology of Antilligobius nikkiae. Drawing by J. Van Tassell. Fig. 11. Illustration of Antilligobius nikkiae from Cayos los Indios, off the west coast of the Isla de Juventud, Cuba. Illustration by R. G. Gilmore. tips; ceratobranchial 5 completely cartilaginous and fused with lower pharyngeal plate; lower pharyngeal plate ossified, possessing large conical teeth with slightly recurved tips. Myology (Fig. 10): superficial jaw musculature composed of adductor mandibulae 1, adductor mandibulae 1 and adductor mandibulae 2; adducaqua vol. 18 no April

13 James L. Van Tassell, Luke Tornabene, Patrick L. Colin tor mandibulae 1 and 1 combined along most of their length and originating on pterotic and ventrally along anterior edge of preopercle; adductor mandibulae 1 separated anteriorly from adductor mandibulae 1 and inserting on primordial ligament near its anterior attachment to maxilla, then running ventrally along primordial ligament and combining with fibers of adductor mandibulae 1 ; adductor mandibulae 1 with two insertions, along ventral section of primordial ligament and separately by tendon attached to coronoid process; adductor mandibulae 1 and 2 differentiated by presence of ramus mandibularis V, visible between muscle masses; adductor mandibulae 2 originating along preopercle, ventral to adductor mandibulae 1 and continuing ventrally to end of preopercle; adductor mandibulae 2 inserting via tendon onto coronoid process of dentary, joining with tendon of adductor mandibulae 1 before inserting on coronoid process. Habitat: Antilligobius nikkiae is known from collections or observations from Curaçao, Bahamas, Cuba, Belize, Chinchorro Bank off eastern Yucatan (Quintana Roo) and Puerto Rico. It is a calcareous wall associate most often found in large numbers on the glacial sea level plateau at around 100 m (Grant Gilmore pers. comm.). Records of the habitat and behavior of this species are described below. Belize: Colin (1974) recorded what is probably A. nikkiae, which he listed as filamentous goby. His observations were made from a submersible on deep reef escarpments at Tobacco Reef and Glovers Reef, Belize, between 90 and 170 m (Colin 1974). The fish occurred in schools of hundreds of individuals and may have been the most common fish observed between 120 and 150 m in these areas. The gobies were reported as being slender, about 5 cm in length, with a silver body and blue eyes (consistent with the appearance of A. nikkiae in the field) and the first dorsal fin was extremely long and held erect. No specimens were collected or photographed in Belize. Bahamas: The first series was collected by SCUBA diving at Acklins Island at 73 m in 1973 by one of us (PC). Small groups were found hovering along a steep drop-off in small indentations in the reef face. Specimens were returned to the surface alive, and some were maintained for several months in aquaria. Dennis et al. (2004) reported the species to be common at m off Lee Stocking Island, Exuma chain, Bahamas. Cuba: Antilligobius nikkiae (Fig. 11) was captured using crystalline rotenone in DMSO from the Johnson-Sea-Link II submarine, Dive 3,069 on 27 December 1997 at a depth of 169 to 198 m. It was collected at Cayos los Indios, N, W, off the west coast of the Isla de Juventud and Golfo de Batabano (Grant Gilmore pers. comm.). Gilmore observed A. nikkiae in small schools typically on vertical calcareous walls, hovering amidst the gorgonians and sponges, hanging along the wall margin. He has also observed them Fig. 12. Pelvic frenum of Bollmannia, with collagenous supports and scalloped posterior margin. Photo by L. Tornabene. 71 aqua vol. 18 no April 2012

14 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species quite often over the sandy terrace that marks the last glacial sea level beach, typically at a depth of around 100 m. He and his colleagues have observed this species in San Salvador, Abaco, New Providence, Grand Bahama and Cat Island in the Bahama Islands; Chinchorro Bank off eastern Yucatan (Quintana Roo); and Terneffe, Glovers and Lighthouse reefs off Belize. Observations in Cuba were limited to the southern shore west of Cayo Largo to Cabo San Antonio. The Johnson-Sea-Link II observations were made from 1978 to Curaçao: Antilligobius nikkiae has been observed by technical divers, on talus slopes, starting at depths of 90 m and was quite common at 100 m (Fig. 3). They hovered a few inches to a foot above the substrate in small groups of 3-4, or in larger groups exceeding 24 individuals where they appeared to hunt for copepods or other zooplankton (Forrest Young, pers. comm.). Puerto Rico: Specimens of A. nikkiae were collected on two occasions at m depth on the shelf edge reef at La Parguera, Puerto Rico (Fig. 2). Two fish were taken on 22 November 1978 (ANSP ) while four specimens were collected on 28 August 1979 and later lost in the mail. Collection dates for these two fish from November 1978 are incorrectly listed in Dennis et al. (2004) as May Field notes record one deep dive on May 1978, but no fish were collected on that dive. Additional fish were seen on 21 August 1979 at another area along the shelf edge reef with similar geomorphology. Dennis et al. (2004) recorded the species from south-western Puerto Rico at 91 m. The gobies were found to hover slightly above the bottom, remaining close to one another and often facing in the same direction. The reef slope where the gobies were captured was steep,with an overall slope of about 45 from 45 to 90 m depth, then becoming near vertical below 90 m. The gobies were found among small indentations in the rocky structure of the reef face where the slope transitions between steep and vertical. They were found with other fish species typical of the deep vertical reef environment, including Ptereleotris helenae, Chromis enchrysura, Chromis scotti, Centropyge argi, Chromis insolata, Liopropoma mowbrayi, Liopropoma carmabi, Gramma linki and juvenile Lutjanus buccanella. The sabre gobies collected from Puerto Rico were captured alive using quinaldine-alcohol solution and returned directly to the surface with the SCUBA divers, where they were maintained in aquaria. Distribution: Known from Belize, Bahamas, Puerto Rico, Cuba, Mexico and Curaçao. The distribution is most likely widespread throughout the Caribbean wherever the glacial sea level plateau occurs. Etymology: The specific epithet nikkiae is for Nicole Laura Schrier, the daughter of Adriaan Dutch Schrier, owner of the Sea Aquarium in Curaçao, who collected many of the type specimens. The species is given the common name Sabre goby in reference to its long, filamentous first dorsal fin that resembles a sabre (a long sword with a curved blade). Bollmannia Jordan, 1890 Bollmannia Jordan, in Jordan & Bollman, 1890: 164 (type species Bollmannia chlamydes Jordan by monotypy). Diagnosis: Bollmannia can be distinguished from other Gobiosomatini genera by the following combination of characters, not listed in order of taxonomic importance: first dorsal of most species with black spot or blotch on posterior portion of fin; second dorsal I,11-15; anal I,10-15; pelvic fins with well-developed frenum supported with collagenous thickenings, frenum with scalloped posterior margin; caudal fin lanceolate; trunk scales ctenoid; predorsal region, cheek, pectoral fin base and breast with cycloid scales; predorsal scales Description: Body slightly elongate and laterally compressed; first dorsal VII; pterygiophore formula of first dorsal fin 3(221110); spines of dorsal fin frequently elongate in both sexes; second dorsal I,11-15; first and second dorsal fins separate from each other and from caudal fin; pectoral rays 19-26; anal I,10-15; trunk scales ctenoid; lateral scale rows 24-31; predorsal region, cheek, pectoral fin base and breast with cycloid scales; predorsal scales 6-10; scales highly deciduous; first pterygiophore of second dorsal fin inserted behind ninth neural spine, remaining pterygiophores arranged in 1:1 ratio with underlying vertebrae; first two anal pterygiophores inserted anterior to first haemal spine, remaining pterygiophores in 1:1 ratio with caudal vertebrae; total vertebrae 27, 11 precaudal, 16 caudal; caudal fin long, 30-50% SL, and lanceolate; pelvic fin with one spine and five rays, fins united, forming disk with well-developed frenum, frenum with thickened collagenous supports and scalloped margin (Fig. 12); teeth in both jaws arranged in multiple rows, teeth in inner and outer rows typically enlarged, evenly spaced, and more aqua vol. 18 no April

15 James L. Van Tassell, Luke Tornabene, Patrick L. Colin Fig. 13A-D. Sensory papillae and head canal pores of A. Bollmannia litura, B. B. boqueronensis, C. B. eigenmanni and D. B. communis. Abbreviations AN, anterior naris; PN, posterior naris. Drawings by L. Tornabene (A, D), or modified from Ginsburg (unpublished manuscript) by L. Tornabene (B, C). 73 aqua vol. 18 no April 2012

16 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species caninoid; teeth in midde rows smaller and more irregularly spaced; vomerine teeth absent; mouth large, 12-17% SL and inclined vertically; eye large, diameter % SL; gill opening extending entire length of pectoral fin base; gill rakers on lower arm of first arch elongate, rakers on upper arm short, stubby and poorly ossified; urogenital papilla thin and conical in males, thick and bulbous in females; cephalic lateralis canal and pore system on head well-developed (Fig. 13); sensory pa pillae pattern in transverse pattern (Fig. 13); ventral post-cleithrum present; nearly all species with dark conspicuous spot on posterior portion of first dorsal fin and sometimes on caudal peduncle; sexual di m or phism generally absent, with the exception of slightly larger dorsal fins in males of B. boqueronensis, and subtle pigmentation differences in B. communis. Bollmannia boqueronensis Evermann & Marsh, 1899 White-eye goby (Figs 13B, 14) Bollmannia boqueronensis Evermann & Marsh, in Evermann & Marsh 1899: 356 (type locality Puerto Real, Puerto Rico). Bollmannia jeannae Fowler, in Fowler 1941: 95, Figs7-9 (type locality Key West, Florida). Material examined: USNM 49366, holotype of B. boqueronensis, 15.5 m, Puerto Real, Puerto Rico; USNM , paratype, 3, 15.5 m, Puerto Real, Puerto Rico; UF , 2, m, Cordoba, Colombia, 12 July 1966; AMNH , 2 plus 1 cleared and stained, mm SL, 8.4 m, JVT , Bocas del Toro, Laguna de Chiriqui, Panama, 9 June 2004, Robertson et al.; AMNH , 4, mm SL, 14.3 m, JVT , Bocas del Toro, N, W, Panama, 12 June 2004, Robertson et al.; UF , 12, 36 m, Gulfo de Morrosquillo, Colombia, 13 July 1966; UF , 5, m, Caribbean Sea, N of Cabo Tiburon, Panama, 18 July 1966; USNM , 1, m, Key West, Florida, 13 February 1902; USNM , 2, 37 m, Gulf of Mexico, Silver Bay Station 54, N, W, Florida, 16 July 1957; USNM , 1, 49 m, Gulf of Mexico, N, W, Florida, 18 March 1885; USNM , 2, Tortugas Islands, Florida, 24 July 1924; USNM , 3, 73 m, Gulf of Venezuela, N, W, Venezuela, 10 November 1958; AMNH , 1, 47.7 mm SL, m, Spaanse Water, N, W, Curaçao, 11 February 2005, Robertson et al.; AMNH , 1, 49.4 mm SL, m, Spaanse Water, N, W, Curaçao, 11 February 2005, Robertson et al.; AMNH , 1, mm SL, 15 m, Spaanse Water, N, W, Curaçao, 10 February 2005, Robertson et al.; ANSP 69718, Bollmannia jeannae holotype, 58.8 mm SL, 34.7 m, Key West, Florida, 12 April 1940; ANSP 69719, B. jeannae paratype, 52.7 mm SL, 34.7 m, Key West, Florida, 12 April 1940; UPRM 3392, 1, 45.8 mm SL, 18 m, east end of Laurel Reef, Puerto Rico, 7 September 1975; UPRM 3788, 3, mm SL, no collection data (possibly collected with UPRM 3790); URPM 3791, 1, 51.8 mm SL, no collection data (possibly collected with UPRM 3790); UPRM 3790, mm SL, 18 m, Laurel Reef, La Parguera, Puerto Rico; UF , 3, mm SL, Golfo de Morrosquillo, N, W, Colombia, 13 July 1966; UF , m, field stations PLC-J19, Fig. 14. Bollmannia boqueronensis, 41.5 mm SL, Bocas del Toro, Panama, AMNH Photo by J. Van Tassell, editing by L. Tornabene. aqua vol. 18 no April

17 James L. Van Tassell, Luke Tornabene, Patrick L. Colin Caribbean Sea, Jamaica, 13 February 1971, P. L. Colin and N. Copland. Diagnosis: Second dorsal fin typically I,12; anal fin typically I,12; eye diameter % SL; spines of first dorsal fin not distinctly elongate; first and second dorsal fins with 3-5 brown or tan horizontal stripes; upper lip lightly pigmented to pale, no dark stripe above upper lip; pelvic disk lightly pigmented to pale; body with 4-5 dark brown blotches along lateral midline; blotches often connected by slightly lighter horizontal band of pigment along lateral midline; longitudinal papillae row b originating at vertical through anterior margin of pupil. Description: Body somewhat laterally compressed; depth at origin of first dorsal fin % SL; least depth of caudal peduncle % SL; size to 100 mm TL (Robins & Ray 1986). Median and paired fins: first dorsal VII, spines not notably elongate; second dorsal I,11(8), I,12*(35), I,13(2); anal I,11*(7), I,12 (36), I,13(2); pectoral rays 19(3), 20(2), 21*(7), 22(5), 23(1), longest rays reaching posteriorly to vertical through anus; pelvic I,5*(10), fifth ray longest, reaching posteriorly to or just falling short of anus; pelvic fin with well-developed frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(1), 17(7), 19*(1), branched rays 14(6), 15(1). Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; lateral scale rows 27-31; transverse scale rows 7; predorsal scales 9-10, scales extending anteriorly to vertical behind eye; caudal peduncle scales 11-13; no modified basicaudal scales present. Head: head length % SL; mouth large, angled upwards approximately degrees from horizontal; mouth reaching posteriorly to vertical through pupil; upper jaw length % SL; teeth in both jaws arranged in three rows; teeth in inner and outer rows enlarged, regularly spaced and caninoid; teeth in middle row smaller and irregularly spaced; eye diameter % SL; interorbital narrow, width less than diameter of pupil; snout short, % SL; postorbital length % SL; gill rakers on lower arm of first arch 10, upper arm 3; gill rakers on lower limb thin and elongate, rakers on subsequent arches shorter and thicker. Genitalia: papilla elongate and conical in males, no melanophores present; papilla short and bulbous in females. Sensory papillae and head pores (Fig. 13B): oculoscapular pores B, C(s), D(s), E, F, G, H, K and L present; in approximately half of specimens examined, anterior canal terminating with pore H, other half of specimens with anterior canal terminating with pore posterior to H, herein termed J, which may not be homologous with J sensu Akihito (1984); preopercular pores M, N, O present; papillae in transverse pattern with 4-5 rows present below eye; first transverse row beginning approximately below anterior margin of eye; last transverse row beginning below posterior margin of pupil; transverse rows often extending below longitudinal row b; row b beginning at vertical through anterior margin of pupil; row d beginning slightly posteroventral of first transverse papillae row, at vertical through anterior one-third of eye and continuing posteriorly to below posterior margin of eye, sometimes slightly further posteriorly; row n not continuous across dorsal midline, stopping slightly before pore D; row x beginning at ventral termination of row n and continuing posteriorly to just before posterior margin of operculum; two horizontal rows of papillae on operculum, one beginning slightly above and behind pore N, the other slightly below and behind pore O. Pigmentation: following description based on recently collected specimens or preserved specimens with well-preserved pigmentation. Overall body background color pale; body with 4-5 dark brown blotches along lateral midline; blotches typically connected by slightly lighter horizontal stripe of pigment along lateral midline; thin, light brown or tan horizontal stripe of pigment below dorsal midline and above lateral midline, beginning behind eye and continuing to base of caudal fin; this stripe often broken or incomplete and becoming less distinct posteriorly; abdomen and gular regions pale; in life, head often with bright iridescent orange vertical markings on preoperculum and operculum and orange stripes on upper and lower jaws; dorsal fins with 3-5 brown or tan horizontal stripes separated by pale or translucent stripes; posterior portion of first dorsal fin with distinct black spot ranging in size from as small as pupil to as large as eye; anal fin completely pale or lightly pigmented at base; pelvic fin lightly pigmented to pale; upper half of caudal fin often with 2-3 horizontal or slightly diagonal brown or tan 75 aqua vol. 18 no April 2012

18 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Table II. Comparison of Bollmannia litura holotype and original description. Measurements other than SL are in % SL. B. litura holotype Description of B. litura USNM (Ginsburg 1935) Sex male male SL 38.4 mm 39 mm Eye diamter Postorbital length Head length Maximum body depth (base of first dorsal fin) Least depth of caudal peduncle Head depth behind eye Head width behind eye Predorsal distance Caudal fin length broken 49 Second dorsal fin I,13 I,11 elements Anal fin elements I,13 I,11 Pectoral fin rays stripes similar to those on dorsal fins; midline of caudal fin often with brown horizontal stripe continuing from stripe on lateral midline of body; lower half of caudal fin pale; pectoral fin base and rays pale. Habitat: Specimens in this study were collected over sand and sandy-mud bottoms and flat areas with fine sediment adjacent to coral reefs at depths from meters. Distribution: Specimens examined in this study were collected from the eastern Gulf of Mexico, Florida Keys, Tortugas, Puerto Rico, Jamaica, Colombia, Panama, Curaçao and Venezuela. Additional museum lots exist, but have not been examined, from Guatemala (UF 15661), Honduras (UF ), Martinique (ANSP ), and Bahia, Brazil (MNHN ). Moreno- Mendoza et al. (2011) also report B. boqueronensis from Alacranes Reef Marine Park, Mexico. Remarks: Fowler s description of B. jeannae stated that B. jeannae differs from B. boqueronensis in coloration, pectoral fin length and cheek scale size (Fowler 1941). Ginsburg (1942) stated that the differences do not hold up when B. jeannae is compared to actual specimens of B. boqueronensis and that the illustrations of B. boqueronensis that accompany Evermann & Marsh s (1899) description are inaccurate. Ginsburg concluded that B. jeannae, which is known from only the holotype and one paratype, is almost certainly a synonym of B. boqueronensis. He offered no formal synonymy of the two species. Bollmannia jeannae falls within the range of B. boqueronensis in all measurements taken here. The coloration differences between the two species as described by Fowler (1941) do not hold when live or freshly preserved specimens of B. boqueronensis are compared, nor do the differences in scale size. The B. jeannae types differ from the B. boqueronsensis holotype in second dorsal and anal fin ray counts. Both type specimens of B. jeannae have I,11 in the second dorsal fin and I,12 in the anal fin, whereas B. boqueronensis was described as having I,12 in the second dorsal and I,13 in the anal fin. Based on our analysis, both I,11 and I,12 are counts that have been observed in both fins of B. boqueronensis. For these reasons, we agree with Ginsburg (1942) and consider B. jeannae to be a synonym of B. boqueronensis. Evermann & Marsh (1899) used the presence of a series of 3-5 large scales on the lower margin of the cheek as a unique and distinguishing characteristic for B. boqueronensis. Although this row of scales is rare or nonexistent in other species of Bollmannia, the deciduous nature of the scales on all species of Bollmannia diminishes the diagnostic utility of this character. Some specimens of B. boqueronensis examined in this study were missing some or all of these scales. Because scale pockets on the face are also frequently destroyed, it is difficult to determine if cheek scales were present at some point and lost during collection, or if scales were absent and thus this character is somewhat variable. Robins & Ray (1986), Humann & Deloach (2002) and McEachran & Fechhelm (2005) report B. boqueronensis as sometimes having orange and yellow spots on the dorsal fins, but these observations were almost certainly based on misidentified specimens of B. litura (which has been collected with B. boqueronensis), or B. communis. Ginsburg (unpublished manuscript) described B. boqueronensis as having an anterior oculoscapular lateralis canal ending with pore H (missing pore J ). Ginsburg used the absence of this character to divide Bollmannia into two distinct groups (proposed subgenera), with B. litura and B. communis both possessing pore J. Our study shows this charaqua vol. 18 no April

19 James L. Van Tassell, Luke Tornabene, Patrick L. Colin acter to be highly variable in B. boqueronensis, as approximately half of the specimens examined have pore J. The variation in this character does not appear to be correlated with geographic distribution, sexual dimorphism or size. Two specimens of Bollmannia from Alacranes Reef, Mexico were identified as B. boqueronensis by Moreno-Mendoza et al. (2011) despite possessing one spine and 13 rays in the anal and second dorsal fins. This combination of counts is more typical of B. communis, which is commonly reported from the western Gulf of Mexico; however, we agree with the identification of Moreno-Mendoza et al. (2011). Their specimens possess all of the diagnostic pigmentation features of B. boqueronensis described here and were also collected from sandymud sediment at the base of a coral reef, which is a common habitat for B. boqueronensis. Bollmannia communis is common over thick mud and is not typically associated with reef habitats. Comparisons: Bollmannia boqueronensis can be distinguished from B. communis in usually having fewer anal and second dorsal elements (typically I,12 in B. boqueronensis vs. typically I,13 in B. communis). It differs from B. litura and B. communis in having a lightly pigmented pelvic fin (vs. a dark pelvic fin), in having tan or brown stripes on the dorsal fin (vs. having yellow or orange stripes or spots) and in having dark spots and a stripe of pigment along the lateral midline (vs. dark spots that are rarely connected by a stripe). Bollmannia boqueronensis can be distinguished from B. eigenmanni in lacking a distinct dark stripe of pigment on or above the upper lip, having a slightly deeper caudal peduncle ( % SL in B. boqueronensis vs. less than 10% SL in B. eigenmanni), having three pores in the preopercular series (vs. two pores in the peropercular series) and in having longitudinal papillae row b originating at a vertical through the anterior margin of pupil (vs. originating more posterior in B. eigenmanni). Bollmannia communis Ginsburg, 1942 Ragged goby (Figs 13D, 15) Bollmannia communis Ginsburg, in Ginsburg 1942: 634 (type locality Gulf of Mexico, Louisiana). Materials examined: USNM , holotype, 1 male, 52.5 mm SL, 18 m, Gulf of Mexico, NW of Southwest Pass entrance, of Mississippi Delta, N, W, Louisiana, 7 September 2006; USNM , paratype, 40 m, Gulf of Mexico, N, W, Louisiana, 30 January 1939; USNM , paratype, 27 m, Gulf of Mexico, NW of Southwest Pass entrance, off Mississippi River Delta, N, W, Louisiana, 10 November 1938; USNM , paratype, 68 m, Gulf of Mexico, N, W, Louisiana, 23 January 1939; USNM , paratype,55 m, Gulf of Mexico, N, W, Louisiana, 13 May 1938; USNM , paratypes, 7, 77 m, Gulf of Mexico, N, W, Texas, 22 March 1938; USNM , paratype, 46 m, Gulf of Mexico, N, W, Texas, 4 February 1939; USNM , paratypes, 6, 64 m, Gulf of Mexico, N, W, Texas, 31 January 1939; USNM , paratypes, 3, 64 m, Gulf of Mexico, N, W, Louisiana, 13 May 1938; USNM Fig. 15. Bollmannia communis, Gulf of Mexico. Photo by Brandi Noble, NOAA Photo Library. 77 aqua vol. 18 no April 2012

20 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species , paratypes, 5, 64 m, Gulf of Mexico, east of Mustang Island, N, W, Texas, 23 January 1939; USNM , paratype, 31 m, Gulf of Mexico, east of Mustang Island, N, W, Texas, 22 April 1938; USNM , paratype, 82 m, Gulf of Mexico, N, W, Texas, 4 February 1939; USNM , paratypes, 7, mm SL, 60 m, Gulf of Mexico, N, W, Texas, 30 January 1939; USNM , paratypes, 50.3 m, Gulf of Mexico, east of Mustang Island, N, W, Texas, 22 April 1938; USNM , 20, Gulf of Mexico, N, W, Florida, 21 March 1962; USNM , 6, 44 m, Gulf of Mexico, Campeche Bank, 110 Miles of (SW) Campeche, Yucatan, N, W, Mexico, 19 August 1951; USNM , 11 of 12, 64 m, N, W, French Guiana, 30 June 1972; TCWC , 5, mm SL, 47.5 m, Gulf of Mexico, Mississippi/Alabama, 12 March 1988; AMNH , 16, Gulf of Mexico, NOAA Station 022, Texas, October 2006; AMNH , 12, Gulf of Mexico, Texas, NOAA Station 032, October 2006; AMNH , 12, Gulf of Mexico, Texas, NOAA Station 045, October 2006; UPRM 2196, 1, 40 mm SL, off Anasco, Puerto Rico, 7 August 1964, J. Randall; UPRM 3788, 2, mm SL, off Rio Anasco, Puerto Rico, 11 August 1987, B. Yoshioka; UPRM 1551, 42 mm SL, Anasco, Puerto Rico, 29 March 1963, Randall et al.; GCRL 17540, 58, mm SL, Gulf of Mexico, N, W, Louisiana, 26 August 1978; GCRL 28344, 2, mm SL, 35 m, Gulf of Mexico, N, W, Louisiana, 9 November 1997, W. E. Aguirre. Diagnosis: Second dorsal fin typically I,13; anal fin typically I,13; eye diameter % SL; middle spines of first dorsal slightly elongate; first and second dorsal fins with dark circles or ellipses with orange or yellow centers (centers pale in preservation); upper lip lightly pigmented to pale, no dark stripe above upper lip; pelvic fin heavily pigmented, black or nearly so; body with 4-5 dark brown blotches along lateral midline; blotches typically not connected by slightly lighter horizontal stripe of pigment along lateral midline; longitudinal papillae row b originating at vertical through anterior margin of pupil. Description: Body somewhat laterally compressed, depth at origin of first dorsal fin % SL; least depth of caudal peduncle % SL; size to 100 mm SL (Robins & Ray 1986). Median and paired fins: first dorsal VII, middle spines slightly elongate in some specimens; in males, fourth spine reaching posteriorly to base of fourth to sixth ray of second dorsal fin when depressed; spines slightly shorter in females; second dorsal I,12(23), 1,13*(82), I,14(3); anal I,11(1), I,12(15), I,13*(84), I,14(8); pectoral rays 19(3), 20(4), 21(4), 22(8), 23*(4), longest rays extending posteriorly to vertical through anus; pelvic I,5*(10), fifth ray longest, extending to anus or falling just short of anus; pelvic fin with well-developed frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(1), 17*(5), branched rays 14*(6). Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; scales in lateral series 24-29; transverse scale rows 6-7; predorsal scales 8-9, scales extending anteriorly to vertical behind eye; caudal peduncle scales 10-13; no modified basicaudal scales present. Head: head length % SL; mouth large, angled upwards approximately degrees from horizontal; mouth reaching posteriorly to vertical through pupil; upper jaw length % SL; teeth in both jaws arranged in three rows; teeth in inner and outer rows enlarged, regularly spaced and caninoid; teeth in middle row smaller and irregularly spaced; eye diameter % SL; interorbital narrow, width less than diameter of pupil; snout short, % SL; postorbital length % SL; gill rakers on lower arm of first arch 10, upper arm 3; gill rakers on lower limb thin and elongate, rakers on subsequent arches shorter and thicker. Genitalia: papilla elongate and conical in males, occasionally with melanophores on lateral edges; papillae short and bulbous in females. Sensory papillae and head pores (Fig. 13D): oculoscapular pores B, C(s), D(s), E, F, G, H, J and K, L present; anterior canal terminating in pore J ; preopercular pores M, N, O present; papillae in transverse pattern, with 4-5 transverse rows present below eye; first transverse row beginning approximately below anterior margin of eye; last transverse row beginning below posterior margin of pupil or slightly more posterior; middle transverse rows highly variable, frequently disjunct, sometimes aqua vol. 18 no April

21 James L. Van Tassell, Luke Tornabene, Patrick L. Colin extending below longitudinal row b; transverse rows on opposite sides of head often varying in a single individual; row b originating at vertical through anterior margin of pupil and continuing posteriorly to below posterior margin of eye, sometimes extending slightly further posteriorly; row d beginning slightly posteroventral of bottom of first transverse row, at vertical through anterior margin of pupil and continuing posteriorly to below posterior margin of eye, sometimes extending slightly further posteriorly; row n continuous across dorsal midline; row x beginning at ventral termination of row n and continuing posteriorly to above pore L. Pigmentation: following description based largely on recently preserved specimens. Comments on coloration in life based on color photographs of B. communis specimens from Gulf of Mexico (Fig. 15) which we have not examined. Overall body background color pale; body with 4-5 faint brown blotches along lateral midline; blotches rarely connected by slightly lighter horizontal stripe of pigment along lateral midline; abdomen and gular regions pale; top of head, snout, interorbital and area above jaw uniformly pigmented with scattered small dark melanophores; occasionally faint elongate patches of pigment present on preoperculum directly beneath and behind eye; base of first dorsal fin with two yellowish orange stripes in life (pale in preservation), stripes becoming broken distally and forming distinct ellipses with dark margins and bright yellow or orange centers in life; yellow and orange pigment on fins fading to pale in preservation; posterior portion of first dorsal fin with distinct black spot ranging in size from as small as pupil to as large as eye; distal margin of first dorsal fin with black stripe in males; second dorsal fin pigmented with ellipses similar to those of first dorsal fin; anal fin nearly pale at base, becoming more heavily pigmented distally; pelvic fin heavily pigmented, black or nearly so; upper half of caudal fin pigmented with ellipses similar to dorsal fins; lower half of caudal fin with two faint yellowish-orange diagonal stripes in life, fading in preservation; pectoral fin base pale; pectoral rays dusky to pale. Habitat: Specimens in this study were collected from mud bottoms at depths of m. Distribution. Known primarily from the Gulf of Mexico, with one confirmed record from French Guiana. Several lots of Bollmannia from Puerto Rico, (UPRM 1551, UPRM 2196, ANSP ) were tentatively listed as B. communis by Dennis et al. (2004); however, these specimens are in very poor condition and it is unclear as to whether these specimens are B. communis or B. litura. Comparisons: Bollmannia communis and B. litura are the most similar of all western Atlantic Bollmannia species. The two species both have similar sensory papillae and sensory pore/canal patterns, nearly identical median fin pigmentation and lack a black band on the upper lip. Bollmannia communis can be distinguished from B. litura in having a smaller eye (typically less than 10% SL in B. communis vs. typically more than 10% SL in B. litura) and from B. litura and all other species in having higher modal counts in the second dorsal and anal fins (typically I,13 in B. communis vs. typically I,10-12 in other western Atlantic Bollmannia species). Male B. communis have dark distal margins on the first dorsal fin whereas male B. litura do not. Bollmannia communis can be further distinguished from B. boqueronensis by pigmentation, as B. boqueronensis has tan or brown stripes on the dorsal fins, a pale pelvic disk and usually a brown stripe of pigment on the lateral midline and B. communis typically has yellow or orange (pale in preservation) stripes and ellipses on the dorsal fins, a dark pelvic disk and no stripe of pigment on the lateral portion of the body. Bollmannia communis can be distinguished from B. eigenmanni in having three pores on the preoperculum (M, N, O ) vs. having two (M, O ), having a dark pelvic fin (vs. a pale pelvic fin), lacking a distinct black stripe on or above the upper jaw (black stripe present in B. eigenmanni) and in having longitudinal papillae row b originating at a vertical through anterior margin of the pupil (vs. originating more posterior to this point in B. eigenmanni). Bollmannia eigenmanni (Garman, 1896) Shelf Goby (Figs 13C, 16) Gobius eigenmanni Garman, in Garman 1896:88, Pl. 3 (type locality Key West, Florida). Materials examined: USNM , holotype, 1 male, mm SL, 110 m, Key West, Florida, 1893; USNM , 1, 37.6 mm SL, m, Gulf of Mexico, N, W, Florida, 15 March 1885; UF , 1, 23.3 mm SL, m, Gulf of Mexico, South of Cailou Bay, N, W, Louisiana, 12 September 1962; UF , 1, m, north of Isla de Margarita, N, W, 79 aqua vol. 18 no April 2012

22 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Table III. Comparison of morphological characters across the Microgobius-group genera Antilligobius Bollmannia Parrella Microgobius Akko Palatogobius Type species Antilligobius nikkiae Bollmannia chlamydes Parrella maxillaris Microgobius signatus Akko dionaea Palatogobius paradoxus Van Tassell & Colin Jordan, 1890 Ginsburg, 1938 Poey, 1876 Birdsong & Robins, 1995 Gilbert, 1971 Precaudal vertebrae Caudal vertebrae Hypurals 1-2 fused to 3-4 No No No No No No and to urostyle DI DI pterygiophores 3(221110) 3(221110) 3(221110) 3(221110) 3(221110) 3(221110) DI elongate Yes, both sexes Yes, both sexes of some Yes, females of some Yes, males No No species species DII I,11-12 I,11-15 I,11-12 I,14-19 I,14-15 I,17-20 DII position Post 9th neural spine Post 9th neural spine Post 9th neural spine Post 9th neural spine Post 9th neural spine Post 9th neural spine A I,11-13 I,10-15 I,10-12 I,14-20 I,14-15 I,19-21 Anal fin position First two elements First two elements First two elements First two elements First two elements First two elements anterior to haemal spine anterior to haemal spine anterior to haemal spine anterior to haemal spine anterior to haemal spine anterior to haemal spine on 12th vertebrae on 12th vertebrae on 12th vertebrae on 12th vertebrae on 12th vertebrae on 12th vertebrae C segmented (Atlantic species only) C branched (Atlantic species only) C shape Lanceolate Lanceolate Lanceolate Lanceolate Lanceolate Lanceolate P V I,5 I,5 I,5 I,5 I,5 I,5 V shape Ovate Ovate Rounded Ovate Ovate Ovate Pelvic frenum Well developed, Well developed, Well developed, Well developed, Well developed, Absent smooth margin scalloped margin scalloped margin smooth margin smooth margin Cephalic lateral-line B' C(s) D(s) E F H' B' C(s) D(s) E F G H' B' D(s) F, H', B' D F G H' B' G' B' C(s) D(s) E F' canal pores (sometimes G) (or H, J') K' L' or B' D(s) F G H' (K'L' absent in or B' C(s) G H', B. eigenmanni) sometimes K' L' Opercular canal pores M' N O' M' N O' or M' O' Absent or M',N' M' N O' or M' N' Absent Absent Scale type Mainly ctenoid, Mainly ctenoid, Ctenoid with Cycloid + ctenoid Cycloid Cycloid some cycloid some cycloid reduced ctenii Predorsal scales Some 0 Scales cheek 2-3 scales Few large scales Absent Absent Absent Absent Scales opercular Absent Present Absent Absent Absent Absent Scales breast Present Present Present Absent Absent Absent Scales belly Present Present Present Absent Absent Absent Scales pectoral fin base Present Present Present Absent Absent Absent Scales transverse scale rows None Lateral scale rows Modified basicaudal scales Absent Absent Absent Absent Absent Absent Vomerine teeth Absent Absent Absent Absent Absent Present Angle of mouth 30 deg 30 deg 35 deg Highly variable 50 deg 35 deg between species and sex aqua vol. 18 no April

23 James L. Van Tassell, Luke Tornabene, Patrick L. Colin p and sex Extent of gill openings Upper pectoral fin Upper pectoral fin Upper pectoral fin Length of pectoral base Base of 4th pectoral Upper margin of orbit base to ventral base to ventral base to ventral to ray to pelvic to just below lower surface of body surface of body surface of body fin base margin of orbit Gill rakers shape Short slender Moderately long, Short stout Long slender Papilloid Moderately long, slender slender Gill rakers upper+lower Posterior nostrils Raised rim Slight raised rim No rim No rim No rim Raised rim Anterior nostrils Short tube Short tube Short tube Short tube Short tube Short tube Fleshy nape crest Absent Absent Absent Present - some species Absent Absent Male papillae Conical Conical Conical Conical Conical Short, bluntly pointed Female papillae Bulbous Bulbous Bulbous Bulbous Short, rounded Short blunt conical Color sexually dimorphic No Only No Some species No No B. communis Head sexually dimorphic No No Mouth and eye Some species No No larger in males Gas bladder Well developed Well developed Absent Well developed Small - thickened Absent Bony support of Canal supported by Portions of canal Canal supported by Canal supported by Canal supported by Canal supported by supraorbital sensory open trough enclosed in bony open trough open trough open trough open trough canal tube Sagittal crest on frontal Present Present Absent Present Absent Absent bone Basihyal shape Spatulate Spatulate Bifid Bifid Minute, reduced Narrow Process on metapterygoid Absent Absent Absent Absent Absent Absent overlapping quadrate Metapterygoid shape Very narrow, Broad, 2-3 times width Very broad, at least 3 Slightly broadened, less Very broad, at least 3 Very narrow splinter like of symplectic (male) times width of than 2 times width times width of symplectic symplectic of symplectic (sexually dimorphic) Symplectic process Absent Slightly developed Absent Slightly developed Absent Present on preopercle to absent to absent Lateral shelf on posttemp- Present (1 reduced) Present (2) Present (1) Absent Present (1) Absent oral (number of shelfs) Sphenotic length Moderate Short Short Short Long, extending to Short vertical through posterior margin of jaw Ventral postcleithrum Present Present Absent Absent Absent Present Shape of braincase Circular Circular Circular Circular Elongate Circular in dorsal view Sensory papillae row b Short, not forward Medium to long, Very long extending Short, anterior of Very long extending Long extending forward of last transverse row extending to anterior almost to 1st row pupil to posterior almost to posterior to under middle of eye but B. eigenmanni of eye in all but of eye or longer of pupil B. eigenmanni Sensory papillae row n Long, continuous across Long, continuous across Short, not continous Long, continuous Long, continuous Long, nearly continuous dorsal midline dorsal midline or nearly so across dorsal midline across dorsal midline across dorsal midline across dorsal midline Sensory papillae row x 1 Not continuous Almost continuous Almost continuous Continuous Continuous Almost continuous Sensory papillae row d Continuous Continuous Continuous Continuous Continuous Continuous Sensory papillae transverse None None None None Last transverse row None rows extending below d 81 aqua vol. 18 no April 2012

24 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Venezuela, 20 July 1968; AMNH 83384, 1, 20.1 mm SL, Gulf of Mexico, N, W, Florida, 29 August 1976; UF , 1, 36.2 mm SL, 41 m, Gulf of Mexico, N, W, United States, 5 June 1974; UF , 1, 23.8 mm SL, m, Golfo Triste, N of Puerto Cabello, N, W, Venezuela, 26 July 1968; UF , 1, 27.8 mm SL, m, W of Cartagena, N, W, Colombia, 1 August 1968; AMNH 85997, 1, mm SL, Gulf of Mexico, N, W, United States, 7 February 1978; AMNH 83254, 1, Gulf of Mexico, N, W, United States, 28 August 1976; YPM 3987, 1, 84 m, Gulf of Mexico, N, W, Louisiana, 10 April 1937, R/V Atlantis; YPM 3986, 4, 51 m, Gulf of Mexico, N, W, Louisiana, 25 March 1937, R/V Atlantis; YPM 3983, 4, 79 m, Gulf of Mexico, N, W, Louisiana, 10 April 1937, R/V Atlantis; YPM 3921, 3, 79 m, N, W, Louisiana, 10 April 1937; TCWC , 3, mm SL, 47.5 m, Gulf of Mexico, N, W, Mississippi and Alabama, 12 March 1988; TCWC , 1 cleared and stained, 28.3 mm SL, 47.5 m, Gulf of Mexico, N, W, Mississippi and Alabama, 12 March 1988; GCRL V69:3460, 1, 34 mm SL, N, W, Dominica, 5 March 1966, M/V Oregon. GCRL V81:17057, 1, 27.6 mm SL, 68.5 m, off Port O Conner, Gulf of Mexico, N, W, Texas, 8 September 1980, R. Mulcahy. Diagnosis: Second dorsal fin typically I,11; anal fin I,10-12; anterior spines of first dorsal fin elongate; eye diameter % SL; upper lip with distinct dark stripe of pigment on or directly above lip; pelvic disk lightly pigmented to pale; side of body with no distinct markings; longitudinal papillae row b originating at vertical through posterior margin of pupil. Description: Body somewhat laterally compressed, depth at origin of first dorsal fin % SL; body depth at caudal peduncle typically % SL (two specimens reaching 10.5% SL); size to 37 cm SL. Median and paired fins: first dorsal VII, first three spines elongate, third spine longest, sometimes reaching origin of eighth second dorsal ray when placed flat against back; second dorsal I,10(1), I,11*(12); anal I,10(6), I,11(3), I,12*(4); pectoral rays 19(2), 20(3), 21*(4), rays extending posteriorly to vertical through anus; pelvic I,5, fifth ray longest, reaching posteriorly to or just falling short of anus; pelvic fin with well-developed frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(1), 17(7), 18(1), branched rays 14(4). Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; lateral scales 27-29; transverse scale rows 7; predorsal scales 7-9, extending anteriorly to behind eye; caudal peduncle scales 10-11; no modified basicaudal scales present. Head: head length % SL; mouth large, angled upward approximately degrees from horizontal; mouth reaching posteriorly to vertical through pupil; upper jaw length % SL; teeth in both jaws arranged in three rows; teeth in inner and outer rows enlarged, regularly spaced and caninoid; middle row of teeth smaller and irregularly spaced; eye diameter % SL; interorbital narrow, width less than diameter of pupil; snout short, % SL; postorbital length Fig. 16. Bollmannia eigenmanni, 30.1 mm SL, Gulf of Mexico, preserved, TCWC Photo by L. Tornabene. aqua vol. 18 no April

25 James L. Van Tassell, Luke Tornabene, Patrick L. Colin % SL; gill rakers in upper arm 3-4, lower arm 9-10; gill rakers on lower limb thin and elongate, rakers on subsequent arches shorter and thicker. Genitalia: papilla elongate and conical in males, no melanophores present; papilla short and bulbous in females. Sensory papillae and head pores (Fig. 13C): oculoscapular pores B, C(s), D(s), E, F, G, H present; no enclosed posterior canal present over operculum; preopercular pores M, O present; papillae in transverse pattern, with 4-5 rows present below eye; first transverse row beginning at vertical just anterior to anterior margin of pupil; last transverse row beginning below posterior margin of eye; middle transverse rows somewhat variable, with anterior rows sometimes extending ventrally to row d; row b beginning at vertical through posterior margin of pupil and continuing posteriorly to posterior margin of preopercle; row d beginning slightly posteroventral of first transverse row, at vertical through anterior one-third of eye and continuing posteriorly to vertical through posterior margin of eye and sometimes slightly further posteriorly; row n nearly continuous across dorsal midline, stopping before pore D; row x sometimes broken into two segments, beginning over pore F and ultimately terminating over operculum. Pigmentation: no fresh or recently preserved specimens of B. eigenmanni were available for description. The museum specimens used in this study had no prominent pigmentation on the body and fins. For the most part the general color on the body and fins of these specimens is pale or dark brown, depending on the method and quality of preservation. The only pigmentation that is apparent after prolonged preservation is the distinct dark band of pigment on or directly above the upper jaw(diagnostic for this species) and a dark spot on the posterior portion of the first dorsal fin, and, in some specimens, a small patch of melanophores at the base of the caudal fin. Habitat: Some specimens in this study were taken over muddy bottoms. It has been collected at depths of m. Distribution: Bollmannia eigenmanni is fairly uncommon in collections and is primarily known from the Florida Keys (type locality) and throughout the Gulf of Mexico, where it has been collected in trawls with B. communis. Records also exist from Venezuela. A single specimen from Dominica was previously identified as B. eigenmanni by C. E. Dawson, however, this record is questionable (see Remarks below). Remarks: When describing B. litura, Ginsburg (1935) noted that specimens of B. eigenmanni were not available for comparison, thus he was forced to rely on the original description of B. eigenmanni for comparison of the two species. In an unpublished manuscript, Ginsburg later noted that the type of B. eigenmanni had been lost. The Gobius eigenmanni holotype (USNM ) has since been found at USNM. A single specimen from Dominica (GCRL 69:3460) was identified by C. E. Dawson as B. eigenmanni, likely due to the second dorsal and anal fin counts (I,11 and I,12 respectively), elongate first dorsal spines and a faint area of pigment above the upper jaw, which is restricted to the very tip of the snout on this specimen. There is no pigment on the remainder of the upper jaw. When compared with other B. eigenmanni this specimen has more pectoral rays (22) and gill rakers on the lower arm of the first arch (12) and also has a slightly shorter jaw, larger eye and shorter postorbital length. In addition, there are 3 pores in the preopercular canal series, whereas all other B. eigenmanni have two. For these reasons we do not consider this specimen to be B. eigenmanni. The identity of this specimen remains unclear and additional specimens are needed to clarify what species occurs in Dominica. Comparisons: Bollmannia eigenmanni is the most distinctive species of western Atlantic Bollmannia. It differs from all other genera in having 2 pores in the preopercular series (M, O ) versus having 3 (M, N, O ), a trait that Ginsburg (unpublished manuscript) used to separate B. eigenmanni into its own subgenus. Longitudinal papilla row b originates at a vertical through the posterior margin of the pupil in B. eigenmanni, whereas row b originates more anteriorly in the other Atlantic species. Bollmannia eigenmanni can also be distinguished from its Atlantic congeners by possessing a distinct dark stripe of pigment on or directly above the upper jaw and in having a shallower caudal peduncle (typically % SL in B. eigenmanni, with two specimens reaching 10.5% SL, vs. typically >10% SL in other species). It further differs from B. communis in averaging fewer rays in the second dorsal and anal fins (I,10-12, in B. eigenmanni vs. I,12-14 in B. communis) and from B. boqueronensis in having prolonged first dorsal spines. 83 aqua vol. 18 no April 2012

26 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Bollmannia litura Ginsburg, 1935 Citrine goby (Figs 13A, 17) Bollmannia litura Ginsburg, in Ginsburg 1939: 1, Pl. 1 (type locality Samaná Bay, Dominican Republic). Materials examined: USNM 93797, holotype, 1 male, 38.4 mm SL, m, Caroline station 59, Samana Bay, , , Dominican Republic, 16 February 1933; USNM , 2 females, mm SL, 31 m, Caroline station 59, Samana Bay, N, W, Dominican Republic, 16 February 1933; UF , 4 of 14, mm SL, m, Gulf of Venezuela, Venezuela, 27 July 1968; AMNH , 1, Portobello, N, W, Panama, 1 June 2007, Robertson et al.; AMNH , 6 females, 2 males, mm SL, 31.5 m, JVT , Bocas del Toro, N, W, Panama, 6 June 2004, Robertson et al.; AMNH , 3 females, 7 males plus 1 male cleared and stained, mm SL, 34.2 m, JVT , Bocas del Toro, N, W, Panama, 6 June 2004, Robertson et al.; UF , 1, m, Caribbean Sea, Colombia, 11 July 1966; USNM , 1 female, field number IU 8849, St. Lucia; UF , 15 of 16, m, Golfo de Venezuela, PIL 762, Venezuela, 27 July 1968; UF141434, 10, m, Cordoba, PIL 362, Colombia, 12 July 1966; UF , 75, mm SL, 12.8 m, NE of Cabo Res Puntas, N, W, Guatemala, 19 March 1968; UF , 4 (labeled as 5), mm SL, 18 m, Golfo de Morrosquillo, N, W, Colombia, 13 January 1966; UF , 31 (labeled as 30), mm SL, 35 m, Caribbean Sea, N, W, Venezuela, 27 July 1968; AMNH , 1, 32.6 mm SL, 25.2 m, JVT , Bocas del Toro, Panama, 2004, Robertson et al.; UF , 2, mm SL, 36 m, Golfo de Morrosquillo, N, W, Colombia, 13 July 1966; UF , 1, 41.3 mm SL, m, off Cartagene, N, W, Colombia, 1 August 1968; UF , 8 of 14 (2 cleared and stained removed under same number), mm SL, m, Caribbean Sea, station PIL 723, Venezuela, 21 July 1968; AMNH , JVT , 1, 35.5 mm SL, 20.7 m, JVT , Bocas del Toro, Laguna de Chiriquí, N, W, Panama, 6 June 2004, Robertson et al. Diagnosis: Second dorsal fin typically I,11 or I,12; anal fin typically I,11 or I,12; eye diameter % SL (9.5% in two specimens); middle spines of first dorsal fin elongate in some specimens; first and second dorsal fins with dark circles or ellipses with orange or yellow centers (centers pale in preservation); upper lip lightly pigmented to pale, no dark stripe above upper lip; pelvic disk heavily pigmented to black; body with 4-5 dark brown blotches along lateral midline; blotches very rarely connected to one another by narrow stripe of lighter pigment; longitudinal papilla row b beginning at vertical through anterior margin of pupil. Description: Body and head somewhat laterally compressed, depth at origin of first dorsal fin % SL; least depth of caudal peduncle % SL; size to 57.4 mm SL. Median and paired fins: first dorsal VII, middle spines slightly elongate in some specimens, fourth spine reaching posteriorly to base of fifth to seventh ray of second dorsal fin with placed flat along back, fifth and sixth spines somewhat shorter, typically not reaching beyond base of fourth ray of second dorsal fin when placed flat along back; Fig. 17. Bollmannia litura, mm SL, Bocas del Toro, Panama, AMNH Photo by J. Van Tassell, editing by L. Tornabene. aqua vol. 18 no April

27 James L. Van Tassell, Luke Tornabene, Patrick L. Colin Fig. 18A-E. Sensory papillae and head canal pores patterns for type species of Microgobius group genera; A. Akko brevis; B. Microgobius signatus; C. Bollmannia chlamydes; D. Parrella maxillaris; E. Palatogobius paradoxus. Abbreviations AN, anterior naris; PN, posterior naris. Drawings A,C-E by J. Van Tassell, B modified from Birdsong (1981). 85 aqua vol. 18 no April 2012

28 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species second dorsal I,11(17), I,12(36), I,13*(1); anal I,10(1), I,11(13), I,12(41), I,13*(1); pectoral rays 18(1), 19(1), 20(5), 21(4), 22*(2), 23(2), reaching posteriorly to vertical through anus; pelvic I,5*(10), fifth ray longest, extending posteriorly to or just falling short of anus; pelvic fin with welldeveloped frenum; pelvic frenum supported with collagenous rays (Fig. 12); caudal fin lanceolate, segmented rays 16(3),17*(10), branched rays 13*(3), 14(10). Scales: trunk completely covered with ctenoid scales, ctenii becoming larger posteriorly; predorsal region, cheek, operculum, pectoral fin base and pelvic fin base with scattered cycloid scales; lateral scale rows 24-28; transverse scale rows 6-7; predorsal scales 7-9, scales extending anteriorly to vertical behind eye; caudal peduncle scales 11-12; and no modified basicaudal scales present. Head: head length % SL; mouth large, angled upward approximately degrees from horizontal; mouth extending posteriorly to vertical through pupil; upper jaw length % SL; teeth in both jaws arranged in three rows; inner and outer rows of teeth enlarged and caninoid; middle row of teeth smaller than other rows and irregularly spaced; eye large, (9.5 in two specimens) % SL; interorbital narrow, width less than diameter of pupil; snout length % SL; postorbital length % SL; gill rakers on lower arm 10*(5), upper arm 3*(5); gill rakers on lower first arch thin and elongate, rakers on subsequent arches shorter and thicker. Genitalia: papilla elongate and conical in males, occasionally with melanophores on lateral edges; papilla short and bulbous in females. Sensory papillae and head pores (Fig. 13A): oculoscapular pores B, C(s), D(s), E, F, G, H, J and K, L present; anterior canal terminating posterior to H in a pore herein termed J, which may not be homologous with J sensu Akihito (1984); preopercular pores M, N, O present; papillae in transverse pattern, with 4-5 transverse rows present below eye; first transverse row beginning below anterior margin of eye; last transverse row beginning below and slightly behind posterior margin of pupil; middle transverse rows highly variable, often differing on each side of head of a single individual, frequently broken and sometimes extending below longitudinal row b; row b beginning at vertical through anterior margin of pupil, continuing posteriorly to vertical through posterior margin of eye, sometimes slightly further; row d beginning slightly posteroventral of termination of first transverse row, at vertical through anterior one-third of eye and continuing posteriorly to below posterior margin of eye, sometimes slightly further; row n continuous across dorsal midline; row x beginning at ventral termination of row n and continuing posteriorly to above pore L. Pigmentation: following description based on recently collected specimens or preserved specimens with well-preserved pigmentation. Overall body background color pale; body with 4-5 faint brown blotches along lateral midline, most conspicuous at base of caudal fin; blotches never connected by slightly lighter horizontal stripe of pigment along lateral midline; abdomen and gular regions pale; in life, head often with bright orange markings vertically on preoperculum and operculum and horizontal orange stripes on upper and lower jaws; dorsal fins with distinct circles or ellipses with dark margins and bright yellow or orange centers in life, with yellow and orange becoming pale in preservation; posterior portion of first dorsal fin with distinct black spot ranging in size from as small as pupil to as large as eye; anal fin nearly pale, in life with 1-2 yellowish orange stripes continuing onto lower half of caudal fin; pelvic fin heavily pigmented, considerably darker than other fins; upper half of caudal fin pigmented with ellipses similar to dorsal fins; lower half of caudal fin with two yellowish orange diagonal stripes in life, stripes fading in preservation; pectoral fin base pale; pectoral rays lightly pigmented, appearing dusky to pale. Habitat: Occurs at depths of m. Specimens in this study from Panama were collected over mud, mixed mud, shell and leaf litter bottoms. Distribution: This species has been recorded from Dominican Republic, Venezuela, Panama, Colombia, Guatemala, St. Lucia and also may occur off Puerto Rico (see Remarks on B. communis above). Remarks: The description of B. litura (Ginsburg 1935) states that B. litura differs from western Atlantic congeners primarily in numbers of second dorsal and anal fin elements. The species is described as having one spine and 11 rays in the second dorsal and anal fins, however, the specimen cataloged as the holotype (USNM 93797) has one spine and 13 rays in both fins. In addition to the differences in dorsal and anal fin elements, there are also discrepancies between the holotype and the description in eye diameter, body depth at the caudal peduncle, maximum body depth, least depth of aqua vol. 18 no April

29 James L. Van Tassell, Luke Tornabene, Patrick L. Colin caudal peduncle, depth and width of head at the posterior margin of the eye and in the length of the caudal fin, which, along with nearly every other fin, was in poor condition in the holotype when we examined it. The number of pectoral rays is also higher in the holotype than in the description (22 vs. 20), although the fins are heavily damaged and it was difficult to count the rays in the dorsalmost and ventralmost parts of the fin. The standard length, head length, predorsal distance and postorbital length were similar in the description and the holotype. Only one specimen of B. litura was known at the time of description; however, a second pair of Bollmannia litura (USNM ) was collected during the same expedition in which the holotype was collected, less than 5 miles from the type locality (Samana Bay, Domincan Republic) at similar depths (~20-30 m). These specimens were later identified as B. litura by Ginsburg and were included in notes from one of Ginsburg s unpublished manuscripts, making a total of three B. litura specimens analyzed by Ginsburg. One specimen (38.5 mm SL female) from USNM has counts of I,11 in both the second dorsal and anal fin and the second specimen (35.9 mm SL female) has counts of I,12 in both fins. Similar discrepancies in morphometics were noted in the two USNM specimens, as the standard lengths, head lengths, and post orbital lengths measured in this study were similar to those recorded in Ginsburg s notes, but other features measured in this study were smaller than those recorded in Ginsburg s notes. The differences in eye, depth and width measurements between Ginsburg s observations and ours may be due to specimen shrinkage over time, mainly in width and depth relative to length. But shrinkage cannot account for the discrepancies in counts of anal fin and second dorsal fin elements, the most notable inconsistency between the original description and the holotype. Ginsburg may have miscounted the fin and recorded the incorrect counts in the description as well as in his unpublished records. Alternatively, it is possible that at some point the holotype was switched with another specimen, with the original holotype now being lost. One reason for the former option being unlikely is Ginsburg s reputation of being meticulous and highly repetitive in his meristic counts. As noted by Myers (1976), Ginsburg excelled in the meticulous measuring and counting of meristic characters he demanded of himself the utmost accuracy. I have seen him spend the morning measuring and counting the same small sample two or three times over, and then return after lunch to repeat the process on the same specimens once or twice again. A second line of evidence supporting the scenario that the type was switched is that the description of B. litura, although inaccurate in its account of what now is catalogued as the holotype, is altogether quite accurate in its description of a morphologically distinctive species of Bollmannia. We have observed the species from multiple localities and note that several of its distinctive morphological characters were described by Ginsburg himself. We consider B. litura a valid species. If the holotype was indeed switched with another specimen, it is unlikely that it was one of the specimens from USNM , as these specimens are female, and the holotype was described as being male. The identification of the specimen currently cataloged as the B. litura holotype remains unclear. Counts and measurements from the cataloged holotype (USNM 93797) and the original description are compared in Table II. Comparisons: Bollmannia litura is most similar to B. communis but can be distinguished from that species in having a larger eye (typically greater than 10% SL in B. litura vs. less than 10% SL in B. communis) and in having lower modal counts in the second dorsal and anal fins (I,11-12 for B. litura vs. typically I,13 for B. communis). It can be distinguished from B. boqueronensis by lacking tan or brown stripes on the dorsal fins and brown blotches along the lateral midline. Bollmannia litura also typically has yellow or orange (pale in preservation) stripes and ellipses on the dorsal that are lacking in B. boqueronensis, and it has a much darker pelvic fin than B. boqueronensis. Bollmannia litura differs from B. eigenmanni in having three pores on the preoperculum (M, N, O ) vs. having two (M, O ), having a dark pelvic fin (vs. a pale pelvic fin), lacking a distinct black stripe on or above the upper jaw (stripe present in B. eigenmanni), having longitudinal papillae row b originating at a vertical through the anterior margin of the eye (originating more posteriorly in B. eigenmanni) and in having a deeper caudal peduncle ( % SL vs % SL). Etymology: The common name of citrine goby is given in reference to the bright yellowish-orange spots on the dorsal and caudal fins of this species, which are reminiscent of the yellow-orange colored gemstone citrine. 87 aqua vol. 18 no April 2012

30 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Key to genera and species of the Microgobius group of the Gobiosomatini (Atlantic species only) Below is an artificial identification key to the Atlantic species of the genera Akko, Bollmannia, Antilligobius, Microgobius, Palatagobius and Parrella. The characters used in the generic key do not necessarily apply to eastern Pacific members of the genus. The key to western Atlantic Microgobius is modified from Birdsong (1981) and the characters used in the key to Palatogobius are from Greenfield (2002). 1a) Second dorsal fin and anal fin confluent with caudal fin... Akko (A. dionea) 1b) Second dorsal fin and anal fin distinctly separate from caudal fin a) Vomerine teeth present; pelvic fin narrow, lacking a well-developed frenum; preopercular canal pores absent... Palatogobius 2b) Vomerine teeth absent; pelvic fin rounded or oval, with a well-developed frenum; preopercular canal pores present or absent a) Top of head naked; second dorsal typically I, Microgobius 3b) Top of head scaled; second dorsal typically I,11-13, rarely I, a) Preopercular canal pores absent, head dorsoventrally depressed, pelvic fin round Parrella (P. macropteryx) 4b) Preopercular canal pores present, head laterally compressed, pelvic fin oval a) Dorsal spines 1-4 greatly elongate, longest spine reaching to or extending beyond posterior margin of second dorsal fin when depressed; body with distinct broad yellow band along lateral midline (white in preservation); pelvic frenum with smooth posterior margin, frenum lacking thickened collagenous supports... Antilligobius (A. nikkiae) 5b) Dorsal spines 1-4 may or may not be elongate, but never extending to posterior margin of anal fin when depressed; body not pigmented as stated above; pelvic frenum with scalloped margin, frenum supported with collagenous thickenings (Fig. 12)... Bollmannia Key to Atlantic species of Palatogobius (characters from Greenfield 2002) 1a) Eye % SL; interorbital % SL; snout % SL; pectoral rays P. grandoculus 1b) Eye % SL; interorbital % SL; snout % SL; pectoral rays typically 19 (sometimes 18 or 20)... P. paradoxus Key to Atlantic species of Microgobius (modified from Birdsong 1981) 1a. Three pores in preopercular sensory canal; second dorsal fin elements typically I,17 or more; anal fin elements typically I,18 or more; great than 65 lateral scale rows b. Two pores in preopercular sensory canal; second dorsal fin elements typically I,16 or less; anal fin elements I,17 or less; fewer than 65 lateral scale rows a. Soft dorsal-fin elements I,19-20; anal fin elements I,20 (occasionally I,19); lateral scale rows about 77-90; scales mostly cycloid; females with pale bar edged in black on body above pectoral fin... M. signatus 2b. Soft dorsal fin elements I,17-18; anal fin elements I,18 (occasionally I,19); lateral scale rows 68-78; scales mostly ctenoid; no dark markings on body in either sex M. microlepis 3a. A fleshy median crest present on nape; a prominent dark spot on body below spinous dorsal fin origin; caudal fin usually greater than 40% of SL... M. meeki 3b. Fleshy median crest absent or poorly developed on nape; body with no dark spot below spinous dorsal origin or with many dark spots; caudal fin usually less than 40% of SL a. Scales mostly ctenoid; about 4 enlarged caninoid teeth in outer row of each dentary; interorbital width broad (about 4% SL); a broad yellow stripe on side with 2 narrow yellow stripes above... M. carri 4b. Scales mostly cycloid; about 8 enlarged caninoid teeth in outer row of each dentary; interorbital width narrow (less than 3% SL); no yellow stripe on body a. Three pores in lateral cephalic sensory canal; body with numerous dark blotches; mouth of males greatly enlarged (greater than 15% SL in males larger than 25 mm)... M. gulosus 5b. Two pores in lateral cephalic sensory canal; body without dark spots; mouth of males little enlarged (less than 15% SL in males) M. thalassinus aqua vol. 18 no April

31 James L. Van Tassell, Luke Tornabene, Patrick L. Colin Key to the Atlantic species of Bollmannia 1a) Upper jaw with distinct black stripe of pigment on or above upper lip; sensory lateralis canal on posterior margin of preopercle with 2 pores (M, O Fig. 13C); longitudinal papillae row b originating at vertical through posterior margin of pupil... B. eigenmanni 1b) Upper jaw dusky or lacking pigment completely; sensory lateralis canal on posterior margin of preopercle with 3 pores (M, N, O Fig. 13A, B, D); longitudinal papillae row b originating at vertical through anterior margin of pupil a) Pelvic fin lightly pigmented to pale; dorsal fins with 3-4 light brown to tan longitudinal stripes often continuing onto caudal fin; lateral midline of body with 4-5 brown blotches often connected with a slightly lighter horizontal stripe of pigment... B. boqueronensis 2b) Pelvic fin heavily pigmented to black; dorsal fins and upper half of caudal fin with dark ellipses with bright orange or yellow centers in life, center of ellipses pale in preservation; lateral midline of body with 4-5 faint brown blotches that are rarely connected by horizontal stripe of pigment a) Eye diameter greater than 10% SL; second dorsal and anal fin elements typically I,11-12 (rarely I,13)... B. litura 3b) Eye diameter less than 10% SL; second dorsal and anal fin elements typically I,13 (occasionally I,12 or I,14 in either the second dorsal or anal, but rarely in both)... B. communis Comments on the tribe Gobiosomatini (Materials examined listed in Appendix I) Birdsong (1975) first proposed the tribe Gobiosomatini (see Smith & Baldwin (1999)) for discussion on spelling of tribe name) as a monophyletic assemblage to include 19 genera of New World gobiids based on the presence of the following characters: (1) vertebral count of 11 precaudal, caudal; (2) first dorsal pterygiophore pattern of 3(221110); (3) scapula unossified; and (4) first 2 anal pterygiophores inserted anterior to the first haemal arch. Several of the sponge-dwelling genera do not possess those characters but were included based on previously suggested relationships (Böhlke & Robins 1969; Böhlke 1969) and the presence of hypural fusion in those genera and most genera within the Gobiosomatini a character rarely found in other genera of Gobiiformes. The tribe was later separated into the Gobiosoma and Microgobius groups (Birdsong et al. 1988). The Gobiosoma group is united based on: (1) fusion of hypurals 1-2 with 3-4 and the terminal vertebral element; (2) dorsal pterygiophore pattern of 3(221110); (3) vertebral counts of 11 precaudal and caudal vertebrae; (4) one epural; and (5) two anal pterygiophores anterior to the first haemal arch. The Microgobius group lacks hypural fusion and possesses the remainder of characters uniting the Gobiosoma group. The genera Robinsichthys Birdsong, 1988, Akko Birdsong & Robins, 1995 and Ophiogobius Gill, 1863, were later added to the tribe. Genera currently in the Gobiosoma group include Aboma, Aruma, Barbulifer, Elacatinus, Chriolepis, Eleotrica, Evermannichthys, Ginsburgellus, Gobiosoma, Gobulus, Gymneleotris, Nes, Ophiogobius, Pariah, Psilotris, Pycnomma, Risor, Robinsichthys, Tigrigobius and Varicus. Aboma, Elacatinus and Tigrigobius were included in Birdsong s genus Gobiosoma based on their subgeneric status as defined by either Böhlke & Robins (1969) for Elacatinus and Tigrigobius or Hoese (1971) for Aboma. The Microgobius group includes Akko, Bollmannia, Microgobius, Palatogobius and Parrella. The outgroup to the Gobiosomatini is currently unknown and may be composed of more than one species (Rüber et al. 2003). Several eastern Atlantic and Mediterranean genera, namely Odondebuenia, Corcyrogobius, Vanneaugobius and Gorogobius have been proposed as possible outgroups (Miller & Tortonese 1969; Miller 1972, 1978; Van Tassell et al. 1988) along with Egglestonichthys and Callogobius from the Indo-west Pacific (Miller & Wongrat 1979) based primarily on sensory papillae patterns. The presence of modified basicaudal scales in many of the Gobiosomatini along with similar pterygiophore insertion patterns suggest an eastern Atlantic/ Mediterranean origin for the outgroup (Van Tassell et al. 1988; Birdsong 1988; Birdsong et al. 1988). Molecular studies of the tribe have added additional support for the monophyly and proposed relationships of the tribe to eastern Atlantic and Mediterranean genera (Rüber et al. 2003; Rüber & Van Tassell, unpublished, presented in 2006). They indicate additional subgroups within the Gobiosoma group and areas in need of further investigation. Four subgroups can be recognized: Barbulifer group (Aruma, Barbulifer, Elacatinus, Enypnias, Evermannichthys, Ginsburgellus, Gobiosoma, Risor, 89 aqua vol. 18 no April 2012

32 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Tigrigobius); Nes group (Nes, Psilotris, Gobulus, Gymneleotris, Pycnomma, Chriolepis, Varicus); Aboma; and Robinsichthys. Aboma is sister to the Barbulifer Group and lacks fusion of hypural 1-2 with 3-4, adductor mandibulae 1 gamma and adductor mandibulae 2 gamma, characters present in the Barbulifer group (Van Tassell 1998). Most genera within the Nes group examined to date (Nes, Psilotris, Gobulus, Gymneleotris, Pycnomma, Chriolepis, Varicus) have pelvic fins that are separate to at least some extent (never with complete disks), lack adductor mandibulae 1, possess no head pores or canals, but have hypural fusion. Aboma and Robinsichthys lack hypural fusion, but this was unknown when the Gobiosoma Group was first proposed and their relationship to the remaining genera requires further investigation. These subgroups are used to facilitate comparisons among genera in the Microgobius group below. Comments on the Microgobius group The Microgobius group proposed by Birdsong et al. (1988) consisted of Microgobius, Bollmannia, Parrella and Palatogobius. This group was phenetically associated with the Gobiosoma group on the basis of a dorsal pterygiophore pattern of 3(221110), vertebral count of 11+16, one epural and two anal pterygiophores anterior to the first haemal arch. The Microgobius group was distinguished from the Gobiosoma group by lacking fusion of hypural 1-2 with 3-4 and the terminal caudal element (Birdsong et al. 1988). Birdsong & Robins (1995) added Akko to the Gobiosomatini, but did not comment further on relationships due to the large number of autapomorphies it possesses. Additional studies of mtdna (Rüber et al. 2003) and morphological characters (Van Tassell & Baldwin 2004) placed Akko in the Microgobius group. Additional characters that unite the group include dorsal papillae row n elongate, uniting across dorsum or nearly so, a feature present in all genera except Parrella; caudal fin lanceolate; body laterally compressed; metapterygoid process not overlapping the quadrate; and modified basicaudal scales absent. We place Antilligobius nikkiae within the Microgobius group of the Gobiosomatini based on the presence of the following characters: vertebrae ; dorsal pterygiophore formula 3(221110); anal pterygiophores anterior to the first haemal arch 2; hypurals 1-2 and 3-4 not fused; dorsal papillae row n elongate, uniting across dorsum; caudal fin lanceolate; body laterally compressed; metapterygoid process not overlapping the quadrate; and modified basicaudal scales absent. A total of 57 morphological characters, including papillae and head pore patterns (Fig. 18), are summarized for the 6 genera of the Microgobius group in Table III. DISCUSSION Antilligobius nikkiae has a number of characters not present in other genera of the Microgobius group that warrant its placement in a new genus. These include a very narrow, splinter-like metapterygoid; short slender gill rakers on the lower limb of the first arch; a short papillae row b, never extending posterior of the pupil, and the modified first haemal arch. The modified haemal arch and its elongate parapophyses, rib-like in appearance in lateral view, where parapophyses join ventrally to form the haemal arch, is easily confused with pleural ribs in radiographs. The modified first haemal arch and anterior curvature of the ventral tip of the premaxilla warrant further investigation. A similar modified haemal arch occurs in several species of Trimma, the expanded area being utilized by the swim bladder (Winterbottom 1984, 2007). The swim bladder of A. nikkiae also occupies this expanded area in the specimen examined. The anterior curvature of the ventral tip of the premaxilla occurs in Antilligobius and several species of Microgobius. Birdsong (1975) observed sexually dimorphic differences in the curvature of the ventral tip of the premaxilla, with the tip curving anteriorly in females of several species of Microgobius. Differences were also observed between small and large-mouthed species of Microgobius, regardless of sex (Birdsong 1975). The significance and distribution of these characters within the Gobiosomatini and Gobiiformes awaits further studies. Within the Microgobius group Antilligobius is most similar morphologically to Bollmannia and Parrella. Antilligobius shares with those genera I,11-15 elements in the second dorsal fin (I,14-20 in Microgobius, Akko, Palatogobius); I,10-15 elements in the anal fin (I,14-21 in Microgobius, Akko, Palatogobius); mostly ctenoid scales on the trunk (few ctenoid scales present in Microgobius, and cycloid scales present in Palatogobius and Akko); scales present on the breast, belly and pectoral fin base (no scales in Microgobius, Akko, Palatogobius); scales in the lateral series ( in Microgobius and Akko, in Palatogoaqua vol. 18 no April

33 James L. Van Tassell, Luke Tornabene, Patrick L. Colin bius); and 6-8 transverse scale rows (19-20 in Microgobius, Akko and none in Palatogobius). Antilligobius and Bollmannia both possess scales on the cheek and a spatulate basihyal, features not present in the other genera. In Parrella sensory papillae row n is not continuous across the nape, whereas it is continuous or almost so in the other genera. The foramen on each anterior vertebrae of Antilligobius (Fig. 8C) is much larger than that in B. litura, B. communis and B. boqueronensis, but is similar in size to that of B. eigenmanni. Parrella maxillaris vertebrae are unlike those of Antilligobius and Bollmannia in that there are numerous small foramina on the anterior vertebrae. Antilligobius also lacks the collagenous supports on the pelvic frenum that are present in Bollmannia (Fig. 12), which may be homologous with the extensive fimbriae found on the posterior margin of the pelvic frenum of Parrella. Antilligobius also differs from Parrella and Bollmannia ecologically, as it inhabits rock-reef talus slopes while both Bollmannia and Parrella are burrow dwellers in soft mud-sand substrate. Elongate first dorsal filaments are found in nearly all species of Bollmannia, variable within Microgobius and Parrella and absent in Palatogobius. In Antilligobius the first four dorsal spines are all very elongate, giving the fin a sabre-like appearance. Microgobius, Bollmannia and Antilligobius all possess 2-3 pores in the preopercular canal (pores absent in Parrella maxillaris, Akko, Palatogobius) and have a sagittal crest on the frontal bone (absent in Parrella, Akko, Palatogobius). The sagittal crest was considered a specialized character within the Microgobius group by Birdsong (1975). Antilligobius shares the presence of a ventral postcleithrum with Bollmannia and Palatogobius. The very narrow metapterygoid in Antilligobius, somewhat similar but wider in Palatogobius, is broad in the remaining genera. Molecular phylogenies by Rüber et al. (2003) and Rüber & Van Tassell (unpublished, presented in 2006) suggest a sister relationship between Antilligobius and Bollmannia. Rüber et al. (2003) recovered a strongly supported clade comprising Antilligobius + a monophyletic group of four Bollmannia species. That clade is sister to a clade containing eight species of Microgobius and Akko. Parrella and Palatogobius were not included in their study. With additional species of Microgobius and Parrella lucretia incorporated in the tree, the phylogeny reconstructed by Rüber & Van Tassell (unpublished, presented in 2006) showed an identical relationship between Bollmannia and Antilligobius (labeled as New Genus in the tree), but Parella lucretia and Akko now formed a distinct clade sister to Microgobius. While several key morphological characters support a close relationship between Antilligobius, Bollmannia and Parrella, the existence of other characters shared between Antilligobius and other Microgobius group genera cloud this relationship. The large number of autapomorphic specializiations and losses within the group coupled with the absence of DNA sequence data for many species in the Microgobius group adds to the ambiguity regarding the relationships among species. A robust phylogenetic analysis of all Microgobius group species using morphological and molecular data will help clarify the relationships between Antilligobius and allied species. Appendix I Gobiosomatini material examined: Akko dionaea USNM 32524, holotype, 1, male, 78.1 mm SL, east of Isla de Maraca, Mapa Province, Brazil, 25 May 1976, Oregon II. Akko brevis EL (JVT collection), 2 cleared and stained, 20.0 m, shrimp trawler, N, W to N, W, El Salvador, J.L. Van Tassell. Aruma histrio CAS 79602, 1, Guaymas, 2-3 miles north of Bahia San Carlos, Mexico, 25 May 1954, C. Limbaugh. Barbulifer pantherinus SU 49929, 52, Ensenada San Francisco, flat reef 0.3 miles north-west of - tip of Punta de las Cuevas, Puerto San Carlos, Mexico, 6 April 1955, Dickinson et al. Barbulifer ceuthoecus ANSP , 7, shore at International Airport, Key West, Florida, 21 March 1958, Cohen et al. Bollmannia chlamydes CAS 42777, 11, off Ballenas Bay, Gulf of Nicoya, N, W, Costa Rica, 25 February 1938, New York Zoological Society Eastern Pacific Expedition. Chriolepis minutillus LACM 20148, 4, Gulf of California, Angel de la Guarda Island, Puerto Refugio, Mexico, 27 January 1940, R/V Velero; UF 72110, 3 plus 1 cleared and stained, Gulf of Mexico, Florida, 3 October 1970, R/V Tursiops. Chriolepis vespa UF 72110, 1 cleared and stained, Gulf of Mexico, Florida, 3 October 1970, R/V Tursiops. Elacatinus horsti UF 26237, 4, Isla de Providencia, Colombia, 29 August 1971, Gilbert and Anderson. Elacatinus oceanops UF 16169, 23, 91 aqua vol. 18 no April 2012

34 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Hawk Channel, Florida Keys, Florida, 6 August 1967, Gilbert and Starck. Elacatinus xanthiprora UF 24797, 2, Isla de Providencia, Colombia, western Atlantic, 30 August 1971, P. Colin. Eleotrica cableae USNM , 47, Indefatigable Island, Galapagos Islands, S, W, 18 May 1966, H. A. Fehlmann. Enypnias seminudus USNM , 2 cleared and stained, Miraflores Locks, Panama Canal, Panama, 28 April 1937, S.F. Hildebrand; FMNH 8457, 1, tidepools, Panama, 24 March 1912, Meek and Hildebrand. Evermannichthys spongicola AMNH 82393, 3 cleared and stained, Gulf of Mexico, 25 July 1975, S. Bortone. Ginsburgellus novemlineatus UF , 8, Bethel Channel, Andros Islands, Bahama Islands, June 1951, E.C. Jones. Gobiosoma chiquita SU 66911, 1 cleared and stained, 1.5 m, Outer San Carlos Bay, shore of island, Gulf of California, Mexico, 2 June 1950, Wilimovsky et al. Gobiosoma homochroma USNM , 9, Rio Pirre above tidal zone, Darien Prov., Panama, 17 February 1985, B. Chernoff et al. Gobiosoma longipala GCRL 815, 26, station 896, Gulf of Mexico, Mississippi, 3 November Gobiosoma nudum GCRL V73:11523, 11, station 1607, Venado Beach, Pacific Panama, 4 April 1973, Dawson et al. Gobulus hancocki SIO61-265, 4, mm SL, Espiritu Santo Islands, El Cardonal Cove, N, W, Mexico, 23 June 1961, R/V Orca, B. Walker et al. Gymneleotris seminudus -MCZ 44759, 2, Rey Island, Perlas Islands, Panama, 27 September 1964, I. Rubinoff. Microgobius miraflorensis FMNH 8489, 2, Rio Culebra, Culebra, Panama, 19 March 1911, Meek and Hildebrand. Microgobius signatus USNM 35164, 1 of 12, Havana, Cuba; MCZ 30602, 3, Havana, Cuba, 1913, T. Barbour. Nes longus ANSP , 2 cleared and stained, Castle Harbor, Bermuda, 10 August 1975, W. F. Smith-Vaniz. Ophiogobius jenynsi USNM , 1, Ancud, Golo de Quatalmaue, Pullingua, Chile, 7 February 1945, M. J. Lobell. Palatogobius grandoculus USNM , holotype, 1, male, 33.0 mm SL, off Cozumel, N, W, Mexico, 11 April 1976, Marine Biomedical Institute. Palatogobius paradoxus ANSP , holotype, female, 25.5 mm SL, St. John, Virgin Islands, 30 June 1961, J.E. Randall; UF , 2, Gulf of Mexico, N, W, Florida, 20 July 1975, SA Bortone and Nester; UF , 7, Gulf of Mexico, N, 86.1 W, USA, 3 June 1974, R/V Mafla. Pariah scotius AMNH 26096, 3, Conception Island, Bahamas, 6 March 1966, C.L. Smith. Parrella macropteryx UF , cleared and stained, PIL751 Field number, Caribbean Sea, Venezuela, 26 July 1968; UF , 1, 36 m, Caribbean Sea, 9.52 N, W, Colombia, 13 July 1966, R/V Pilsbury. Parrella maxillaris USNM , Paratype, 1, La Paz Bay, Gulf of California, Mexico, 21 February 1936, Hancock Expedition; USNM , 82, Punta San Ignacio, Sonora, N, 110, 25 W, Mexico, 19 June 1990, G. Allen and D. Robertson; UF , 22.9mm SL, 1 female, cleared and stained, m, 7 miles north of Manta, bearing 270 deg, along 20 fm curve, S, W, Ecuador, 1 October 1961, R/V Argosy. Robinsichthys arrowsmithensis UF , paratype, 1 female, cleared and stained, 23.5 mm SL, m, Quintana Roo, station 893, N N, Mexico, 10 Sept 1967, R/V Gerda. Tigrigobius dilepis UF 14032, 2, Exuma Sound, Exuma Cays, Bahama Islands, 29 August 1966, C.R. Gilbert. Tigrigobius gemmatum AMNH 26076, 6, cleared and stained, Hog Cay, Ragged Islands, Bahamas, 9 July 1965, C. L. Smith & H. Tischler. Tigrigobius janssi LACM , 1, Bahia Herradura, at tip of outer reef on N side of bay, Costa Rica, 9 March 1972, Bussing et al. Tigrigobius macrodon ANSP , 36, mm, Baileys Bay, Bermuda, 23 July 1981, W. F. Smith-Vaniz et al. ACKNOWLEDGEMENTS Thanks to D. Ross Robertson, Ben Victor, and Ernesto A. Peña E. along with the crew of the RV Urraca (Smithsonian Tropical Research Institute) for their aid in obtaining the Bollmannia litura specimens. Frank Pezold provided valuable comments throughout this study. Funding for collections in Panama was provided by Smithsonian Tropical Research Institute to D. Ross Robertson. B. Brown and R. Arrindell at AMNH, M. S. Pérez aqua vol. 18 no April

35 James L. Van Tassell, Luke Tornabene, Patrick L. Colin at ANSP, S. LeCroy at GCRL, K. Conway and H. Prestridge at Texas A&M University College Station, R. Robins at UF, M. Craig at UPRM, J. Williams, D. Pitassy and S. Raredon at USNM, and D. Skelly at YPM provided assistance with museum specimens. G. D. Johnson provided assistance with digital photography of cleared and stained specimen and helpful discussions on osteology. D. Smith provided assistance with unpublished Ginsburg material at USNM. C. Baldwin provided photographs of Antilligobius habitat. F. Young provided habitat information for Antilligobius specimens in Curaçao. A. Schrier provided critical comments, photographs and specimens of Antilligobius. R. G. Gilmore provided the specimen from Cuba, illustrations and detailed records of Antilligobius from Cuba and other localities. REFERENCES AKIHITO, P., HAYASHI, M. & YOSHINO, T Suborder Gobioidei. In: The Fishes of the Japanese Archipelago. (Ed. K. Masuda, C. Amaoka, C. 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36 Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species MILLER, P. J. & WONGRAT, P A new goby (Teleostei: Gobiidae) from the South China Sea and its significance for gobioid classification. Zoological Journal of the Linnean Society 67: MORENO-MENDOZA, R., GONZALEZ-SALAS, C., AGUILAR- PERERA, A., GALLARDO-TORRES, A. & SIMOES, N First record of the white-eye goby, Bollmannia boqueronensis (Teleostei: Perciformes: Gobiidae) along the coast of the Yucatan Peninsula (Gulf of Mexico). Marine Biodiversity Records 4: 1-4. MURDY, E. & HOESE, D Gobiidae. In: The living marine resources of the Western Central Atlantic. Bony fishes part 2 (Opistognathida to Molidae). FAO species identification guide for fishery purposes and American Society of Ichthyologist and Herpetologists Special Publication No. 5. (Ed. K. Carpenter.): FAO, Rome. MYERS, G. S Isaac Ginsburg. Copeia 1796: NELSON, J., CROSSMAN, E., ESPINOSA-PÉREZ, H., FINDLEY, L., GILBERT, C., LEA, R. & WILLIAMS, J Common and Scientific Names of Fishes from the United States, Canada, and Mexico. American Fisheries Society, Special Publication 29, Bethesda, 386. PEZOLD, F Evidence for a monophyletic Gobiinae. Copeia 3: ROBINS, C. & RAY, G A Field Guide to Atlantic Coast Fishes of North America. Houghton Mifflin Company, Boston, 368 pp. RÜBER, L. & VAN TASSELL, J. L Small fish in a big clade: a molecular perspective on gobioid phylogeny. Presented at Joint Meeting of Ichthyologists and Herpetologists; July, 2006, New Orleans. RÜBER, L., VAN TASSELL, J. L. & ZARDOYA, R Rapid speciation and ecological divergence in the American seven spined gobies (Gobiidae: Gobiosomatini) inferred from a molecular phylogeny. Evolution 57: SABAJ PÉREZ, M Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 1.5 (4 October 2010). Electronically accessible at American Society of Ichthyologists and Herpetologists, Washington D.C. SANZO, L Distribuzione delle papille cutanee (organi ciatiformi) e suo valore sistematico nei Gobi. Mitteilungen aus der Zoologischen Station zu Neapel 20: SARUWATARI, T., LOPEZ, J. A. & PIETSCH, T. W Cyanine blue: a versatile and harmless stain for specimen observations. Copeia 1997: SMITH, C. L National Audubon Society Field Guide to Tropical Marine Fishes: Caribbean, Gulf of Mexico, Florida, Bahamas, Bermuda. Alfred A. Knopf, New York. 720 pp. SMITH, D. G. & BALDWIN, C. C Psilotris amblyrhynchus, a new seven-spined goby (Teleostei: Gobiidae) from Belize, with notes on settlement-stage larvae. Proceedings of the Biological Society of Washington 112: THACKER, C. E Molecular phylogeny of the gobioid fishes (Teleostei: Perciformes: Gobioidei). Molecular Phylogenetics and Evolution 26: THACKER, C. E. & HARDMAN, M Molecular phylogeny of basal gobioid fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei). Molecular Phylogenetics and Evolution 37: VAN TASSELL, J. L Phylogenetic relationships of species within the gobiid genus Gobiosoma sensu Böhlke and Robins (1968) with comments on their relationships to other genera in the tribe Gobiosomini. PhD Dissertation, City University of New York Graduate Center, 283 pp. VAN TASSELL, J. L. & BALDWIN, C. C A review of the gobiid genus Akko (Teleostei: Gobiidae) with description of a new species. Zootaxa 462: VAN TASSELL, J. L., MILLER, P. J. & BRITO, A A revision of Vanneaugobius (Teleostei: Gobiidae), with description of a new species. Journal of Natural History 22: WINTERBOTTOM, R A review of the gobiid fish genus Trimma from the Chagos Archipelago, central Indian Ocean, with description of seven new species. Canadian Journal of Zoology 62: WINTERBOTTOM, R Three new species of genus Trimma from Palau, western Pacific (Percomorpha: Gobiidae). International Journal of Ichthyology 13 (1): aqua vol. 18 no April

37 aqua, International Journal of Ichthyology Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae) Rüdiger Riesch 1*, Ryan A. Martin 1, David Bierbach 2, Martin Plath 2, R. Brian Langerhans 1 and Lenin Arias-Rodriguez 3 1) North Carolina State University, Department of Biology & W. M. Keck Center for Behavioral Biology, 127 David Clark Labs, Raleigh, NC , USA: ramarti2@ncsu.edu; langerhans@ncsu.edu 2) J. W. Goethe-University of Frankfurt, Department of Evolutionary Ecology, Max-von-Laue Strasse 13, D Frankfurt am Main, Germany. David.Bierbach@gmx.de; plath.ecology.evolution.behavior@googl .com 3) División Académica de Ciencias Biológicas, Universidad Juárez Autónoma de Tabasco (UJAT), C.P Villahermosa, Tabasco, México: leninariasrodriguez@hotmail.com *Corresponding author s address: North Carolina State University, Department of Biology, 127 David Clark Labs, Raleigh, NC , USA Tel: Fax: rwriesch.evolutionarybiology@gmail.com Received: 22 October 2011 Accepted: 05 January 2012 Abstract We report on basic natural history, life history, and diet of Priapella chamulae (Poeciliidae) from Arroyo Tres, a small creek in Tabasco, southern México. The tertiary (adult) sex ratio was heavily female-skewed, female P. chamulae produced medium-sized offspring (~2.3 mg), one clutch at a time (i.e., showed no superfetation), and relied predominantly on yolk for embryo provisioning (Matrotrophy Index: 0.71). Furthermore, P. chamulae at Arroyo Tres had relatively short guts, were highly carnivorous, and preyed mainly on terrestrial arthropods (especially ants). Zusammenfassung Wir berichten über grundlegende Naturgeschichte, Nahrung und Lebenszyklusdaten von Priapella chamulae (Poeciliidae), die im Arroyo Tres, einem kleinen Bach in Tabasco in Südmexiko gefangen wurden. Das tertiäre (adulte) Geschlechterverhältnis war weibchenlastig, weibliche P. chamulae produzierten nur eine Brut zur Zeit (d.h., sie zeigten keine Superfetation), die Embryonen wurden während Ihrer Entwicklung überwiegend durch Dotter ernährt (Matrotrophieindex: 0.73) und die Nachkommen waren von mittlerer Größe (ca. 2.3 mg). Außerdem hatten P. chamulae im Arroyo Tres relativ kurze Därme, waren karnivor und erbeuteten bevorzugt terrestrische Arthropoden (insbesondere Ameisen). Résumé Nous abordons l histoire naturelle de base, l histoire de la vie et le régime alimentaire de Priapella chamulae (Poeciliidae) d Arroyo Tres, un petit cours d eau de Tabasco, Mexi - que méridional. Le sex ratio tertiaire (adulte) était surtout constitué de femelles ; la femelle P. chamulae produisait une descendance de taille moyenne (~ 2.3 mg), une ponte à la fois (càd. ne montrant pas de superfétation) et consi - stant surtout en jaune d oeuf pour l alimentation des embryons (Matrotrophy index: 0,71). En outre, P. chamulae d Arroyo Tres avait des intestins relativement courts, était essentiellement carnivore et prélevait surtout des arthropodes terrestres (principalement des fourmis). Sommario Riportiamo la storia naturale, il ciclo vitale e la dieta di Priapella chamulae (Poeciliidae) da Tres Arroyo, un piccolo torrente in Tabasco, nel sud del Messico. Il rapporto tra i sessi nella terza fase del ciclo vitale (adulto) era fortemente spostato verso la femmina, la femmina produceva prole di taglia media (~ 2.3 mg), una nidiata alla volta (cioè senza alcuna produzione sovrapposta di embrioni), che si basava prevalentemente sul tuorlo per il nutrimento degli embrioni (Indice di matrotrofia: 0.71). Inoltre, P. chamulae a Tres Arroyo avevano un intestino relativamente breve, erano prettamente carnivori predando principalmente artropodi terrestri (soprattutto formiche). INTRODUCTION Life-history evolution has been an integral part of evolutionary research over the past few decades, and livebearing fishes (Poeciliidae) have proven to be excellent models for studying life history adaptations (e.g., Reznick & Miles 1989; Magurran 95 aqua vol. 18 no April 2012

38 Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae) 2005). Originally, the more than 200 species and genera of poeciliids were endemic to the Americas, but due to human introductions they are now found worldwide (Lucinda 2003). Within the Poeciliidae, at least three characters evolved to give rise to the present diversity within the family: (1) internal fertilization using a transformed anal fin in males, the gonopodium, for sperm transfer, (2) livebearing, and (3) different degrees of maternal provisioning for the developing young (Constantz 1989; Reznick & Miles 1989). With the sole exception of Tomeurus gracilis Eigenmann, which is characterized by oviparous egg retention, internal fertilization and viviparity are found in all species of poeciliids (Rosen 1964). Furthermore, some poeciliids are able to accommodate several clutches simultaneously, so-called superfetation (e.g., Turner 1937; Thibault & Schulz 1978; Reznick & Miles 1989). Priapella chamulae Schartl, Meyer & Wilde (2006), is a medium-sized livebearing fish inhabiting the waters of the Río Grijalva basin in Tabasco, México (Schartl et al. 2006). The genus Priapella comprises at least six species (e.g., Miller 2005; Schartl et al. 2006; Meyer et al. 2011) and is considered the least derived (i.e., most basal) group within the tribe Gambusiini (Hrbek et al. 2007). The other genera in this tribe are Scolichthys, Carlhubbsia, Xiphophorus, Heterandria, Belonesox, Heterophallus, and Gambusia (Miller 2005; Hrbek et al. 2007). The natural history of the members of the genus Priapella and several other genera in this tribe remains largely unknown (see Miller 2005; Schartl et al. 2006; Meyer et al. 2011; Riesch et al. 2011a). However, to fully understand the evolution of reproductive strategies in livebearing fishes (e.g., Reznick & Miles 1989; Pires et al. 2011), it is of utmost importance to also collect data on smaller and lesser-known clades. We report on basic natural history of P. chamulae as observed over the last five years while conducting fieldwork in Tabasco, southern México. We also report on life history and diet data derived from specimens collected in January Furthermore, we describe the fish species communities in the lotic environments they inhabit. Fig. 1. Overview of the general study area in México with reference cities in gray (modified after Riesch et al. 2011a). Numbers indicate rivers and creeks; all Priapella chamulae analyzed in the present study were collected at Arroyo Tres (#5). aqua vol. 18 no April

39 Rüdiger Riesch, Ryan A. Martin, David Bierbach, Martin Plath, R. Brian Langerhans and Lenin Arias-Rodriguez Table I. Raw values for male and mean±sd of female life history traits for reproductively active Priapella chamulae caught in Arroyo Tres (Tabasco, México) in January Values in parentheses provide minimum-maximum. GSI: gonadosomatic index; RA: reproductive allocation; MI: matrotrophy index. males females Sample size a 1/6 34/49 SL [mm] ±4.20 (24-40) Somatic dry mass [mg] ±76.76 (72-386) Fat content [%] ±3.75 (0-14) Fecundity ±3.75 (2-16) Estimated embryo dry mass at birth b [mg] Embryo fat content [%] ±3.91 (3-23) GSI [%] RA [%] ±3.42 (4-23) MI a the numerator corresponds to reproductively active males and females & the denominator equals the total number of collected and dissected males and females b estimated dry mass at birth is calculated using the slope and intercept from the regression between log-transformed embryonic dry mass and stage of development. MATERIAL AND METHODS Study population and sampling procedure: We conducted life-history dissections on a total of 55 P. chamulae (6 males and 49 females). All fish were collected in January 2010 in the Arroyo Tres ( N, W; Fig. 1), a small creek and tributary to the Río Oxolotán (part of the Río Grijalva drainage) near the village of Tapijulapa in Tabasco (Fig. 1). Collections were made using seine nets, and fish were field-preserved in 10% formaldehyde. Life-history analysis: Following the protocol of Reznick & Endler (1982), all preserved fish were weighed and measured for standard length. Males were classified as mature based on the morphology of their modified anal fin (i.e. gonopodium; Fig. 2C). The anal fin in poeciliids undergoes a complex metamorphosis as fish attain maturity, and the endpoint of this metamorphosis provides a reliable index of sexual maturity, allowing us to determine the state of complete maturation based on external cues alone (e.g. Constantz 1989; Greven 2011; Langerhans 2011). We therefore consulted the illustration of the fully developed gonopodium of P. chamulae presented by Schartl et al. (2006). Unfortunately, almost all males that we collected turned out to be sexually immature (5 out of 6), and so their data had to be discarded for most life history analyses. However, male poeciliids are characterized by determinate growth and therefore almost completely cease growth after reaching sexual maturity (Snelson 1989). Since all immature males were within days of reaching sexual maturity (i.e., only the terminal structures of the gonopodium were not yet fully developed; Turner 1941) we did use their standard length (SL) data for statistical comparisons with females, and included them in our gut length analysis (see below). The reproductive tissue (i.e., testes for males and ovaries for females) and, if present, all developing offspring were removed. Offspring were counted and their stage of development determined (Haynes 1995; Reznick 1981). Somatic tissues, reproductive tissues, and embryos were then dried for 10 days at 40 C and weighed again. To assess male, female, and embryo condition, somatic tissues and embryos were rinsed six times for at least six hours in petroleum ether to extract soluble nonstructural fats (Heulett et al. 1995; Riesch et al. 2010a, b, 2011b) and were then dried again and reweighed. We thus measured standard length [mm], dry mass [mg], lean mass [mg] (i.e., weight after fat extraction), and fat content [%] for males and females, gonadosomatic index [GSI, %] for males only, and fecundity [number of developing embryo], reproductive allocation [RA, %], embryo dry mass [mg], offspring lean weight [mg], and embryo fat content [%] for females only. GSI is calculated as testis dry mass [g] divided by the sum of 97 aqua vol. 18 no April 2012

40 Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae) testis dry mass [mg] and somatic dry mass [mg]. RA, on the other hand, was calculated as the total dry mass of all developing embryos [mg] divided by the sum of the total dry mass of all developing embryos [mg] and somatic dry mass [mg]. Finally, we calculated the matrotrophy index [MI] as a measure to indirectly evaluate post-fertilization maternal provisioning (Marsh-Matthews 2011). The MI equals the estimated dry mass of the embryo at birth divided by the estimated dry mass of the oocyte at fertilization and is derived from a linear regression of log-transformed embryo dry mass against stage of development (Marsh- Matthews 2011). Thus, if the eggs were fully provisioned by yolk prior to fertilization (lecithotrophy), we would expect the embryos to lose 30-40% of their dry mass during development (MI 0.75; Marsh-Matthews 2011; Pires et al. 2011). On the other hand, in the case of continuous maternal provisioning even after fertilization (matrotrophy), one Fig. 2. Priapella chamulae SCHARTL, MEYER & WILDE. (A) Male (top) and female (bottom); (B) ready-to-be-born embryos; (C) tip of the gonopodium of a male P. chamulae. B and C were stacked using Helicon Focus V (2010 Helicon Soft Ltd.). aqua vol. 18 no April

41 Rüdiger Riesch, Ryan A. Martin, David Bierbach, Martin Plath, R. Brian Langerhans and Lenin Arias-Rodriguez Table II. Diet of Priapella chamulae as inferred by gut-content analysis. Diet categories Number of diet items Individuals with diet item [%]* Total diet [%] Arachnida Blattaria 1 1 < 1 Chironomidae larvae Coleoptera adults larvae Diptera adults Gastropoda 1 1 < 1 Hemiptera 1 1 < 1 Hymenoptera adult ants adult wasps Lepidoptera larvae 2 3 < 1 Oligochaeta 1 1 < 1 Orthoptera Unidentified insects adults larvae Plant material seeds 2 3 < 1 rootlets 18 * percentages may not sum to 100 due to rounding would expect the embryos to lose less mass (MI between 0.75 and 1.00) or even to gain mass during development (MI 1.00; Marsh-Matthews 2011; Pires et al. 2011). Gut content analysis: We inferred the diet of P. chamulae by dissecting the guts of all preserved fish and identifying the contents. We identified gut contents to Order in most cases. If this was not possible, we identified items to the lowest possible taxonomic level. We counted the number of gut contents within each diet category except for woody plant material and calculated the percentage of guts containing each diet category, and the proportion of each diet category relative to the total diet, and in total for all individuals. Additionally, before dissecting out gut contents we measured gut length [mm] with digital calipers. We used these gut length measures to calculate relative gut length (gut length/standard length) and tested for sexual dimorphism in relative gut length with a Mann-Whitney U-test. RESULTS General natural history: Over the last five years and while sampling other poeciliid fishes, we found P. chamulae at a variety of habitats in Tabasco, ranging from small creeks [Arroyo Tres, Arroyo Bonita ( N, W), Arroyo Cristal ( N, W), and Arroyo Tacubaya ( N, W)] to large rivers [Río Amatán ( N, W)] all near the village of Tapijulapa (Fig. 1). Surprisingly, we have so far never caught P. chamulae in the Río Oxolotán, but this is probably an artifact of our sampling efforts, as all other positive locations suggest that P. chamulae also inhabits that river (Fig. 1). Teleost species communities at these habitats are quite complex (Tobler et al. 2006; Riesch et al. 2009) and at Arroyo Tres P. chamulae share the habitat with Poecilia mexicana Steindachner, Xiphophorus hellerii Heckel, Heterophallus milleri Radda, Heterandria bimaculata Heckel (all Poeciliidae), Astyanax aeneus Günther (Characidae), Thorichthys helleri Steindachner, and Vieja bifasciata Steindachner (both Cichlidae) (see also Tobler et al. 2006). In some of the other habitats, species communities are even more complex (e.g., Riesch et al. 2011a). More generally, P. chamulae appears to prefer the slightly faster flowing parts (e.g., riffles) of these habitats, as we rarely catch significant numbers in the stagnant pools that are usually dominated by P. mexicana and A. aeneus. Life-history analysis: General life history data are compiled in Table I. Priapella chamulae at Arroyo Tres was characterized by sexual size dimorphism, with males being smaller than females (female 99 aqua vol. 18 no April 2012

42 Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae) SL = 30.30±4.21 mm; male SL = 26.02±2.93 mm; t-test: t 38 = 2.379, P = 0.022; Fig. 2A). The tertiary (adult) sex ratio was female-biased and approximately 1:8.2 (males:females; Table I). The majority of dissected females (69.3%) were reproductively active, but only one out of the six collected males was sexually mature (16.7%). Embryo weight decreased with developmental stage (R 2 = 0.292, P < 0.001) in a fashion corresponding to a predominantly lecithotrophic provisioning strategy (MI = 0.71; Fig. 2B, Fig. 3). We did not find any evidence for superfetation in P. chamulae, as all developing embryos of the same clutch were always of approximately the same developmental stage. Gut content analysis: Only one individual had an empty gut (except for some incidental woody plant material). Priapella chamulae at Arroyo Tres are almost exclusively carnivorous (Table II); on average, individual P. chamulae had 4.6±2.5 (mean±sd, range: 0-10) diet items in their guts, and the vast majority of their diet was made up by terrestrial arthropods (~98 %), with ants being the single most common identifiable diet item (Table II). Males and females did not differ in their relative gut length (females: median = 21.08%, IQR = 9.22%; males: median = 23.75%, IQR = 11.55%; Mann-Whitney U-test: U = , P = 0.545). DISCUSSION Our observations confirm previous accounts that described P. chamulae to prefer the medium to fast flowing parts of small creeks (Miller 2005; Schartl et al. 2006). Like many other life history traits, offspring size and fecundity are usually related to female body size (Reznick & Miles 1989). It is therefore slightly surprising that this medium-sized poeciliid produces only relatively few, average-sized offspring, which translates into low values for RA. For example, similar sized Poecilia mexicana from comparable creeks around Tapijulapa produce two to three times more offspring, which are also larger (Riesch et al. 2010b). However, RA-values of P. chamulae are surprisingly similar to those of P. mexi - cana (given the pronounced differences in fecundity and offspring size), while more closely related poeciliids like H. milleri or Gambusia sexradiata Hubbs (that are also considerably smaller in body size) from nearby habitats invest almost twice as much per Fig. 3. Scatter plot of mean embryo dry mass vs. stage of development. Stages are determined on a progressive scale with the earliest (5) being the neurula stage and the oldest (50) being equivalent to embryos that are ready-to-be-born (Riesch et al. 2011c; after Reznick 1981; Haynes 1995). aqua vol. 18 no April

43 Rüdiger Riesch, Ryan A. Martin, David Bierbach, Martin Plath, R. Brian Langerhans and Lenin Arias-Rodriguez clutch (i.e., RA around 20 %; Riesch et al. 2010a, 2011a). Nonetheless, several life history traits of P. chamulae are similar to those of other members of the tribe Gambusiini (e.g., no superfetation and lecithotrophy; but see Pires et al for citations on rare occurrences of superfetation in Gambusiini). Likewise, offspring fat content matches that found in other Gambusiini from the same drainage (i.e., G. sexradiata and G. eurystoma Miller, Riesch et al. 2010a; H. milleri, Riesch et al. 2011a). Furthermore, the sexual size dimorphism we report on here is typical for most poeciliids (Pires et al. 2011). Most natural populations of poeciliids are characterized by a female-skewed sex ratio (Snelson 1989; Magurran 2005) and P. chamulae from Arroyo Tres were no exception. Since secondary sex ratios (i.e., sex ratio at birth) are typically 1:1 in poeciliids, higher male mortality is usually thought to be the reason for a female-biased sex ratio at maturity (Snelson 1989). Our results clearly demonstrate that P. chamulae at Arroyo Tres are carnivorous. While there is at least one other purely carnivorous species in the tribe Gambusiini (i.e., Belonesox belizanus Kner; Miller 2005), the diet of P. chamulae more closely resembles that described for Heterandria bimaculata, Gambusia affinis (Baird & Girard), G. holbrooki Girard, G. sexradiata, and other Gambusia spp. that are known to prey heavily on insects and other invertebrates (Miller 2005; Trujillo-Jiménez & Beto 2007). Other close relatives, swordtails and platyfish of the genus Xiphophorus, however, are known to be mostly omnivorous with an emphasis on herbivory (e.g., Maddern et al. 2011). In agreement with this apparent carnivorous dietary specialization, P. chamulae had relatively small gut-to-sl ratios. This clearly sets them apart from other more herbivorous/omnivorous species like Poecilia latipinna (Lesueur) and allfemale Poecilia formosa (Girard) that have considerably longer gut-to-sl ratios (Scharnweber et al. 2011a, b). However, based on the predominance of terrestrial arthropods in the diet it is likely that P. chamulae is an opportunistic feeder that, rather than actively hunting for its prey, mainly waits for arthropods to fall or land on the water surface. This study represents the first diet and life history characterization of a poeciliid fish from the little investigated genus Priapella. While the present study already provides us with relevant insights into their general biology, it is important to keep in mind that the diet data and life history data we report on here were derived from specimens collected from a single point in space and time. Previous studies have clearly shown that diet and life histories vary considerably between habitats and seasons in livebearing fishes (e.g., Reznick & Endler 1982; Meffe & Snelson 1989; Reznick & Miles 1989; Johnson & Bagley 2011; Riesch et al. 2010a-c, 2011; Scharnweber et al. 2011b). However, water chemistry and the teleost community at Arroyo Tres are representative for a range of similar environments in this part of southern México (Tobler et al. 2006, 2008; Riesch et al. 2009). We are therefore fairly confident that the life history data presented in the current study are largely representative of P. chamulae life-history strategies in the other creek environments inhabited by this species (i.e., Arroyos Bonita, Cristal, and Tacubaya). Nonetheless, P. chamulae from larger rivers, such as the Ríos Amatan and Oxolotán, may differ in life histories (and potentially also diets) from those reported here, and more extensive studies (i.e., comparisons of P. chamulae from different sample sites, or of laboratory-reared specimens with field-caught fish) are clearly warranted to help characterize the full dietary niche width and breadth of life-history strategies exhibited by this species. ACKNOWLEDGEMENTS We thank two anonymous reviewers for improving a previous version of this manuscript and the Deutsche Forschungsgemeinschaft (DFG) for financial support (PL 470/2-1). This research was conducted under the Mexican authorization from the Municipio de Tacotalpa in Tabasco and SEMARNAT. REFERENCES CONSTANTZ, G. D Reproductive biology of poeciliid fishes. In: Ecology and Evolution of Livebearing Fishes (Poeciliidae) (Eds. G. K. Meffe & F.F. Snelson Jr.): Prentice Hall, Englewood Cliffs. 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45 aqua, International Journal of Ichthyology Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea Shima Bakhshalizadeh 1*, Shahram Abdolmalaki 2, Ali Bani 1 1) Fisheries Department, Faculty of Natural Resources, University of Guilan, PO Box 1144, Sowmeh-Sara, Iran 2) Inland Water Aquaculture Research Institute, PO Box 66, Bandar Anzali, Iran *Corresponding author: Shima Bakhshalizadeh; Fisheries Department, Faculty of Natural Resources, University of Guilan, PO Box 1144, Sowmeh-Sara, Iran Phone: Fax: shima_ba83@yahoo.com Received: 27 August 2011 Accepted: 19 January 2012 Abstract The age and growth of the starry sturgeon, Acipenser stellatus, obtained from Iranian coastal waters of the Caspian Sea, were studied through analysis of the pectoral fin ray section from 69 specimens, ranging in fork length (FL) from 83 to 173 cm. The specimens were obtained from commercial fisheries between October 2008 and June The interpretation of growth bands in the pectoral fin ray sections was carried out objectively using the direct reading of thin sections and by image analysis. The maximum age recorded for the specimens of starry sturgeon was 29 years. The estimates of asymptotic length (L ) and the growth coefficient (K) of females were cm and 0.08 per year respectively. Males had an asymptotic length of cm and a growth coefficient of 0.15 per year. The total mortality coefficient (Z) for females and for males was estimated to be 0.79 and 1.08 per year respectively. Annual mortality rates were calculated at 55 percent for females and 66 percent for males. This study revealed differences in the life history parameters of the starry sturgeon compared with previous study results which may be associated with increased fishing pressure and the degradation of environmental conditions. Zusammenfassung Altersentwicklung und Wachstum des Sternstörs, Acipenser stellatus, wurden durch Analyse des Brustflossenstrahlen- Abschnitts anhand von 69 Exemplaren von den iranischen Küstengewässern des Kaspischen Meeres untersucht, deren Länge mit Schwanzgabel (fork length, FL) zwischen 83 und 173 cm betrug. Die untersuchten Exemplare wurden zwischen Oktober 2008 und Juni 2010 vom kommerziellen Fisch fang zur Verfügung gestellt. Die Beurteilung der Wachstumsbänder in den Abschnitten mit Brustflossenstrahlen erfolgte objektiv: durch direktes Ablesen an dünnen Abschnitten und durch Bildanalyse. Das ermittelte maximale Alter bei den Exemplaren des Sternstörs betrug 29 Jahre. Als asymptotische Länge (L ) und als Wachstumskoeffizient (K) wurden bei den Weibchen durchschnittlich 153,69 cm bzw. 0,08 pro Jahr berechnet. Die Männchen hatten durchschnittlich eine asymptotische Länge von 131,02 cm und einen Wachstumskoeffizienten von 0,15 pro Jahr. Als Sterblichkeits-Koeffizient (Z) konnten die Werte 0,79 bei Weibchen und 1,08 bei Männchen ermittelt werden. Als jährliche Sterblichkeitsrate wurden 55 Prozent bei den Weibchen und 66 Prozent bei den Männchen berechnet. Die deutlichen Unterschiede bei den Parametern der Lebensgeschichte im Vergleich zu früheren Untersuchungen dürften mit der Zunahme des Fischereidrucks und der Verschlechterung der Umweltbedingungen zu erklären sein. Résumé L âge et la croissance de l esturgeon, Acipenser stellatus, prélevé dans les eaux côtières de la Caspienne en Iran, ont été analysés par la coupe du rayon de la pectorale de 69 spécimens, de longueur variable (FL) de 83 à 173 cm. Les spécimens ont été fournis par des pêcheries commerciales d octobre 2008 à juin L interprétation des anneaux de croissance dans les coupes de la pectorale à été effectuée objectivement par la lecture directe de fines sections et par analyse de l image. L âge maximal observé sur lesdits spécimens était de 29 ans. Les estimations de la longueur asymptotique (L ) et le coefficient de croissance (K) des femelles étaient respectivement de 153,69 cm et de 0,08 par an. Les mâles avaient une longueur asymptotique de 131,02 cm et un coefficient de croissance de 0,15 par an. Le coefficient total de mortalité (Z) des femelles et des mâles était estimé respectivement à 0,79 et 1,08 par an. Le taux de mortalité annelle a été calculé à 55% pour les femelles et à 66% pour les mâles. Cette étude a révélé des différences dans les paramètres de vie de cet esturgeon, en comparaison avec des résultats d études antérieures, ce qui peut être associé avec l augmentation de la pression de pêche et la dégradation des conditions environnementales. 103 aqua vol. 18 no April 2012

46 Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea Sommario L'età e la crescita dello storione stellato, Acipenser stellatus, ottenuto da acque costiere iraniane del Mar Caspio, sono state studiate attraverso l'analisi della sezione del raggio della pinna pettorale da 69 esemplari, la cui lunghezza all inforcatura (FL) variava da 83 a 173 cm. I campioni sono stati ottenuti da attività di pesca commerciali tra ottobre 2008 e giugno L'interpretazione delle bande di crescita nelle sezioni del raggio della pettorale è stata effettuata utilizzando la lettura diretta di sezioni sottili e mediante analisi di immagine. L'età massima registrata per gli esemplari di storione stellato era di 29 anni. Le stime di lunghezza asintotica (L ) e il coefficiente di crescita (K) di femmine erano cm e 0.08 all'anno, rispettivamente. I maschi avevano una lunghezza asintotica di cm e un coefficiente di crescita di 0.15 all'anno. Il coefficiente di mortalità totale (Z) per le femmine e i maschi è stato stimato a 0.79 e 1.08 per anno, rispettivamente. Tassi annui di mortalità sono stati calcolati al 55 per cento per le femmine e 66 per cento per i maschi. Questo studio ha rivelato differenze nei parametri del ciclo vitale dello storione stellato rispetto ai risultati degli studi precedenti, che possono essere associate con l aumento della pressione di pesca e il degrado delle condizioni am - bientali. INTRODUCTION The starry sturgeon, Acipenser stellatus, (Pallas 1771), limited to the Black, Azov and Caspian Seas, is one of the most important sturgeon species. The largest populations are now concentrated in the Caspian Sea, from which they migrate into the Volga, Kura, Ural, Terek, Sulak and Samur rivers (Berg 1948). They also live in the Iranian rivers Sefid-Rud and Gorgan-chaii (Rostami 1961). This species is commercially one of the most common sturgeons in catches from Iranian coastal waters of the Caspian Sea, comprising about 45% of the total sturgeon catch (Moghim 2003). However, official reports of the sturgeon catch in the Caspian Sea show that landings have sharply decreased from 24.8 thousand tons in 1975 to 0.74 thousand tons in 2004, as was confirmed by the Caspian Research Institute of Fisheries (Casp- NIRKH) (Pourkazemi 2006). The starry sturgeon is a highly valuable species, a long-lived fish that grows and matures slowly with a low rate of natural mortality (Billard & Lecointre 2001). These characteristics, coupled with high and unregulated commercial fishing, habitat loss and environmental degradation (such as the accumulation of pollutants in sediments, the damming of rivers, and the restriction of water flows) have negatively influenced the migration and reproduction of these fish (Birstein et al. 1997; Billard & Lecointre 2001). Their stocks have been reduced, as have some other marine and freshwater fish species world wide (Myers & Worm 2003; Pauly et al. 2003; Safina et al. 2005). Quantifying the nature and magnitude of the differences in the life history parameters of the starry sturgeon, as an example of a fish species that is under fishing pressure, warrants further investigation into the age structure and mortality rates. The period between spawning is shorter for males than for females (Makarov 1970; reviewed by Holčík 1989) and males reach sexual maturity 2-3 years earlier than females (Holčík 1989). Hence, because males appear more in inshore and estuarine waters for the spawning activity than females, they are more susceptible to fishing. The late age at maturity and the 2-year spawning interval in the starry sturgeon probably inhibits population recovery (Pikitch et al 2005). Effective management of the starry sturgeon fishery would be sensitive to biases in the estimation of their life history traits (Brennan & Cailliet 1989; 1991). The estimation of sturgeon age is commonly made from cross sections of the pectoral fin rays (Koch et al. 2008). These bony structures provide the greatest precision for age estimation, and unlike other structures such as opercles, clavicles, cleithra, and medial nuchals, they can be obtained without killing the fish (Brennan & Cailliet 1989). Pectoral fin sections of the starry sturgeon are suitable for aging since they are easily collected, processed and have legible, precisely interpretable growth zone (Brennan & Cailliet 1989; Stevenson and Secor 2000). The objectives of this study were to estimate ages at size, somatic growth, population age structure and mortality of starry sturgeons collected in to determine whether changes in life history parameters have occurred during this recent period of increased fishing mortality. MATERIAL AND METHODS Study sites and sampling A total of 69 specimens of the starry sturgeon, Acipenser stellatus, were obtained from Iranian coastal waters of the Caspian Sea between October 2008 and June 2010 (Fig. 1). Sixteen samples were obtained from commercial catches of Iranian fisheries (for the restocking program) using anchored gill nets. Additionally, 53 specimens were obtained from beach seine fisheries to provide a broader range of fish. The gill nets used to collect the samples measured 18 m long by 2.1 m deep with a mesh size (stretched knot to knot) of 100 mm. aqua vol. 18 no April

47 Shima Bakhshalizadeh, Shahram Abdolmalaki, Ali Bani Gutted weight (W) to the nearest kilogram, and total length (TL) and fork length (FL) to the nearest centimeter were recorded after sacrificing all samples. Fish were sexed and staged by macroscopic examination of the gonads. The gender of three fish could not be determined and therefore they were classified as immature. The right pectoral fin rays were removed for the purposes of age analysis (Chugunova 1959; Rien & Beamesderfer 1994). Processing of the pectoral fin ray The fin rays were placed in water at C for 10 min (Jearld 1992) to separate soft tissues. They were then defleshed with a stiff brush and placed on filter papers to dry. Transverse sections were obtained using a fret saw and were polished with 250 and 400 grit sandpapers successively until a thickness of between mm was achieved. Samples were cemented to a glass slide with clear nail polish to keep them immobilized. Glycerol was used to enhance the differentiation between the rings and to aid in the examination of growth increment formation under transmitted light using a microscope system and a camera (Motic China Group Company) that displayed the image on a video monitor. The final magnification used varied from 10-40x. Bias and precision analyses All sections were read twice by the first reader (the first author) and 17% of the samples were randomly chosen for a second reading by an expert on sturgeon aging to determine the percentage agreement (PA). All sections were read blind without reference to fish size. Age estimates were accepted if counts were identical between readings. If counts differed by 1 year, fish were allocated the higher age on the basis of the conservative assumption that a reader was more likely to underestimate age by overlooking a partly obscured section than to overestimate age by counting an anomaly like false bands (Brennan & Cailliet 1989). If counts differed by 2 years or more, sections were read again until agreement was within one year. To assess the precision of ring counts of the fin ray sections provided using images, the ages of all 69 samples were estimated by the first reader from the one section provided, using both the direct reading and estimates from the images. The average percent error (APE) between the ring counts using the two techniques was calculated using the following formula (Beamish & Fournier 1981): where N is the number of fish aged in the subsample, R is the number of times the age of each fish was estimated, Xij is the ith determination for the jth fish, and Xj is the average estimated age of the jth fish. A paired sample T-test was used to determine if there was a difference between the ages estimated using the two techniques. In addition, the precision of age estimates was also assessed between the two readers by the coefficient of variation (CV) as indices of precision (Chang 1982). The CV is described as the ratio of the standard deviation of age estimates to the mean: where Xij is the ith determination for the jth fish, Xj is the mean age of the jth fish, and R is the number of times the age of the fish was estimated. The CV was averaged across the samples for each reader to derive a mean CV. The Chi-square test of independence was used to examine differences in age frequency distributions between the sexes. The data from four seasons were pooled for age frequency distributions between the sexes. Fig. 1. Location of the sampling areas (red closed circles) of sturgeon in Iranian coastal waters of the Caspian Sea. Growth estimates and curves Parameters of the length-weight relationship were 105 aqua vol. 18 no April 2012

48 Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea Table I. Descriptive length and weight composition of the male and female starry sturgeon from Iranian waters of the Caspian Sea in Size Sex N Mean S.E. of mean Minimum Maximum FL (cm) Female Male W(kg) Female Male achieved by fitting the power function to length and weight data (Ricker 1975): W = a FL b where W is the gutted weight, a and b are regression constants and FL is the fork length. The condition factor (CF) was calculated by CF=W/ FL b, and the comparison of CF between the genders was performed by applying the ANOVA test (Saborowski & Buchholz 1996). The length and weight of the starry sturgeon estimated in this study were compared with the mean length and weight obtained from a previous study (Taghavi Motlagh 1996) in the south Caspian Sea, using the Student s one sample t-test (Zar 1996). Linear, logarithmic, power, Von Bertalanffy, and exponential equations were tested to acquire the best fit of size-at-age data. Finally, growth was characterized using the von Bertalanffy growth function, fitting to size-at-age data using standard nonlinear optimization methods (least square method). The von Bertalanffy growth function is explained as: L t = L (1- e -K (t-t 0 ) ) where L t is length at age t, L is the asymptotic length, K is the growth coefficient and t 0 is the hypothetical age at which length is equal to 0. The comparison of von Bertalanffy growth curves of the two genders was done using an analysis of the residual sum of squares (ARSS) (Chen et al 1992). Shabany et al (2003) revealed that there is only one population of the starry sturgeon in Iranian coastal waters of the Caspian Sea and therefore data from different locations in the south Caspian Sea were pooled in this study. Mortality For an estimate of the annual instantaneous total mortality coefficient (Z), we used the formula (Gulland 1983): where Z is the instantaneous total mortality coefficient, K is the shape parameter from the von Bertalanffy growth equation, L is length at infinity from the von Bertalanffy growth equation, L is the mean fork length at capture and L is length for which all fish of that length and longer are under full exploitation. Following the calculation of Z, annual mortality can be calculated through the formula A = 1 e -z ; survival rate through the formula S=1-A (Ricker 1975). All statistical analyses were performed with the SPSS (Version 13) software package. RESULTS Starry sturgeon females had a higher fork length and body weight (Table I) compared to males. The relationship between length and weight was best expressed using a geometric mean functional regression (Fig. 2) as described by Ricker (1973). The condition factor (CF), expressing the feeding activity during the year, was not significantly different between the sexes (F = 0.49, df 1, P = 0.83). Bands were present in all the fin sections examined. Growth bands (Cailliet & Goldman 2004) were more widely spaced near the origin and in older fish were usually more tightly grouped toward the outer edge. Tight groupings of growth rings were a common problem in age determination, and they occurred even in the youngest samples. Moreover, incorporation of secondary fin rays into the posterior lobe of the first ray could potentially lead to errors in age determination. Two independent readers agreed on the same age 58% of the time, and differed within ± 1 band counts 17% of the time. Because of high agreement and precision between the two readers on the subsample (PA = 75%, CV= 4.55), only one reader continued with the age estimates for all the samples. There was no significant difference between the counts taken by the two techniques (the direct reading and readings from the images) (t=-1.33, df 68, P=0.188). The average peraqua vol. 18 no April

49 Shima Bakhshalizadeh, Shahram Abdolmalaki, Ali Bani cent error (APE) was very low (0.95%), suggesting no difference between the profiles of pectoral fin rays and their images. An APE of less than 5% is indicative of consistent interpretation of age (Morison et al 1998). The estimated ages for females caught in ranged from 9 to 29 years with a median age of 17 and for males the ages ranged from 9 to 17 years with a median age of 13 (Fig. 3). Males and females had dissimilar age- frequency distributions in Iranian coastal waters of the Caspian Sea (χ 2 =36.63, df 14, P<0.001) The Iranian coastal Fig. 2. Length-weight relationship for female (open circles and dotted line) and male (closed circles and solid line) starry sturgeon in Iranian coastal waters of the Caspian Sea, (W = gutted weight (kg), FL = fork length (cm)). Fig. 3. Age composition for female and male starry sturgeon between 2008 and 2010 in Iranian coastal waters of the Caspian Sea (n= 69). Arrows indicate the direction in which the observed frequencies differed from expected frequencies under the assumption that age class was independent of sex. 107 aqua vol. 18 no April 2012

50 Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea Fig. 4. The von Bertalanffy growth function fit of size-at-age relationships for female (open circles and dotted line) and male (closed circles and solid line) starry sturgeon in Iranian coastal waters of the Caspian Sea, (L t = fork length (cm)). Fig. 5. Confidence regions (95%) of mean for growth parameter estimates (K and L ) for males and females of the starry sturgeon in and in in Iranian coastal waters of the Caspian Sea. aqua vol. 18 no April

51 Shima Bakhshalizadeh, Shahram Abdolmalaki, Ali Bani waters of the Caspian Sea starry sturgeon had a higher proportion of females in the 18 and 19 year classes as compared to the males. Approximately 68% of females and 97% of males were less than 17 years old. The frequency of the starry sturgeon younger than 12 years is low (Fig. 3), suggesting that these age classes were under sampled, and the maximum age recorded for the starry sturgeon was 29 years. Growth trajectories were significantly different between the sexes for the starry sturgeon (F=3.79, df 3, 67 P < 0.05). The growth curves for both sexes therefore were plotted separately (Fig. 4). The predicted length at age for females was greater than for males (Fig. 4). For the starry sturgeon in Iranian coastal waters of the Caspian Sea, the mortality rates (Z) for females and for males were calculated as 0.79 and 1.08 per year and the annual mortality (A) as 55% and 66% respectively. Additionally, the survival rates of the starry sturgeon in Iranian coastal waters of the Caspian Sea were found to be 45% for females and 34% for males. DISCUSSION Life history characteristics can be used to classify the vulnerability of a species to fishing pressure and to judge the level of productivity within a population (Musick 1999; Roberts & Hawkins 1999). The slow growth and high longevity of the starry sturgeon in combination with their big size and the late achievement of sexual maturity cause much variation in their life-history characteristics. The age estimate precision from pectoral fin ray for the starry sturgeon is acceptable compared to the Atlantic sturgeon (Acipenser oxyrinchus) (Mitchill 1814) (Stevenson & Secor 2000) and the white sturgeon (Acipenser transmontanus) (Richardson 1836) (Rien Starry sturgeon, Acipenser stellatus, live, not preserved. Photo by S. Bakhshalizadeh 109 & Beamesderfer 1994) that reported higher CV for these long-lived species. Nevertheless, difficulties in reading the fin sections were caused by damaged sections, abnormal or compressed bands on the anterior fin ray margin of older fish. It is assumed that the size and associated age structure were representative of the study populations because samples were obtained throughout the year. Part of the existing starry sturgeon population in the south Caspian Sea originated from hatcheries, but due to their long life, these fish may be considered to represent a wild population. The maximum recorded age for the starry sturgeon in a study made in (Pirogovskii & Fadeeva 1982) was 35 years. Makarova and Alekperov (1988) found a similar age structure to that in the present study which was same as the record previously by Taghavi Motlagh (1996). The ages of the starry sturgeon captured in the Iranian coastal water of the Caspian Sea fishing sites were 4 to 27 years, in which the highest proportion of age classes for males and females were 11 to 14 (76%) and 9 to 12 (75%) years old respectively (Taghavi Motlagh 1996). The predominance of females in the older age classes accords with observations made by Pirogovskii and Fadeeva (1982) in the whole Caspian Sea, and appears to be a life history strategy (Roff 1984) because of the bigger maximum length and probably longer life span of females. The number of females aged 18 years and over exceeds that of males (Fig. 3) which can be a result of the more frequent spawning migrations of males to inshore waters (Scarnecchia et al 2007). This result seems to indicate indirectly that males, which migrate to spawn at a younger age and return to spawn at more frequent intervals, have a higher mortality coefficient. The combination of the higher mortality rate of males and the greater years of vulaqua vol. 18 no April 2012

52 Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea nerability to fishing is reflected in the much higher prevalence of females aged 18 years and more. The mean fork length of females was the same as that reported for females in (Taghavi Motlagh 1996). However, males showed a decreased fork length in and also the mean fork length of both females and males was lower than that reported by Veshchev and Novikova (1986) in the Volga. These differences might be related to the capture of predominantly the mature migrating fish in this area, while the catches in the southern part contained both mature and immature fish (Taghavi Motlagh 1996). Moreover, the average weight of starry sturgeon of both sexes showed a decrease compared to the fish studied in (Taghavi Motlagh 1996) and were lower in weight than the larger, mature fish which enter the Volga river (Veshchev 1979). The weight-length relationship of the starry sturgeon appears to be dependent on age, sex, and the feeding conditions in the habitat (Babushkin & Borzenko 1951; Putilina 1981). Although the type of sampling gear may introduce a bias in the size structure of the fish, in this current study the use of both gill nets and beach seine nets allows for representation of all exiting sizes of the starry sturgeon. The length and weight-at-age relationships were asymptotic in form, with the majority of growth being achieved early in life. There was a comparatively small increase in size after that, similar to the growth scenario for the white sturgeon, Acipenser transmontanus (Brennan & Cailliet 1989). The reduction in growth rate coincided with the age at sexual maturity, suggesting a physiological shift from somatic growth to reproductive development. The von Bertalanffy growth parameters estimated in this study ( ) were different from those in the study made in (Taghavi Motlagh 1996). Bias in individual L and K estimates can be overcome by considering the parameter estimates jointly (Sainsbury 1980). In both time periods, the predicted length at age in different size classes and asymptotic length (Fig. 5) for females was greater than for males. This difference has been associated with the faster growth of females of a brood year compared with males of that brood year which have undergone sexual maturation (Rosen et al 1982; Alexander et al 1985; reviewed by Stamps 1993) and the greater age at maturity of females (Russell 1986; Scarnecchia et al 1996). Estimates of the asymptotic length for females in was much lower than for females in , and there was an increase in the growth coefficient, indicating a decrease in size both overall and at similar ages. This may be a result of increased fishing pressure on the large (female) component of population, as males in the study approached a smaller asymptotic size with an increase growth rate compared to the males of the study. Harvesting pressure could generate an evolutionary response towards slower growth and smaller size; this has been studied mainly by quantitative genetic methods (Favro et al 1979; Law & Rowell 1993) in the effects of harvesting pressure reported in plaice (Pleuronectes platessa L.) (Linnaeus 1758) (Rijnsdorp 1989; Barot et al 2005), Atlantic cod (Gadus morhua) (Linnaeus 1758) (Barot et al 2004; Olsen et al 2005) and in small mouth bass (Micropterus dolomieu) (Lacépède 1802) (Dunlop et al. 2005). However the effect of fishing on the starry sturgeon population can be determined where estimates of the age at first sexual maturity exist, and there may be a minimum threshold age below which sexual maturation does not occur. Overfishing, which specifically increased after the breakdown of the ex-soviet Union (Pourkazemi 2006) could be one factor influencing the severe depletion of the starry sturgeon stock. In addition, several environmental conditions such as pollution (from factories and oil rigs) can be influential on life history variables (Ivanov 2000) since pollutants affect both the gills and liver of the starry sturgeon (Halajian et al. 2006). The mortality coefficient of starry sturgeon females (Z = 0.65 per year) and males (Z = 0.92 per year) in (Taghavi Motlagh 1996) was considerably lower than for the period , for both females and males, which indicates that intensive commercial fishing may be a causative factor in the sharp decline of the starry sturgeon stock. The increase in the annual mortality and decrease in the survival rate of the starry sturgeon force the conclusion that there is indeed fishing pressure in the Caspian Sea. The current fishing pressure on the starry sturgeon, as one of the most precious species in the Caspian Sea, is high and with its life history characteristics makes this species vulnerable. For a better understanding of the magnitude and mechanisms of the processes affecting demographic parameters, analysis of fish stocks by continual sampling is necessary to assess the status of the stock of starry sturgeon, with an intent to rationalize and to restructure state fishing regulations so that fish mortality may be reduced. aqua vol. 18 no April

53 Shima Bakhshalizadeh, Shahram Abdolmalaki, Ali Bani ACKNOWLEDGEMENTS We would like to thank R. Nahrevar for a second reading of fin ray sections. We also thank R. Rastin for his assistance in the preparation of fin ray sections. Sampling authorization was given by the Madar Khaviari sector of the Iranian fisheries organization. We would also thank H. Dadari, A. Alinejad, M. Ataee, A. Qavidel, F. Shakori, A. Kor and G. Salehi for their assistance in the collection of fish. We greatly appreciate the constructive comments and suggestions provided by G. Cailliet. We also wish to acknowledge Dr. Margaret Nelson for her assistance in editing the English language text of this manuscript. We also thank K. Bakhshalizadeh, S. Bakhshalizadeh and O. Arshad for their spiritual support. REFERENCES ALEXANDER, C. M., MYHR, A. I. & WILSON, J. L Harvest potential of paddlefish stocks in Watts Bar Reservoir, Tennessee. Proceed. 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HOLÍK, J., BANARESCUE, P. & EVANS, D Systematic part. In: The freshwater fishes of Europe. (Ed. J. Hol ík.): AULA-Verlag, Wiesbaden. IVANOV, V. P Biological Resources of the Caspian Sea. CaspNIRKH, Astrakhan, 96 pp. JEARLD, A Age determination. In: Fisheries Techniques. (Eds. L. A. Nielson., & D. L.Johnson): The American Fisheries Society, Bethesda, Maryland. KING, M Fisheries Biology, Assessment and Management. Fishing News Books, Oxford, 341 pp. KOCH, J. D., SCHRECK, W. J. & QUIST, M. C Standardised removal and sectioning locations for shovelnose sturgeon fin rays. Fisheries Management and Ecology 15: LAW, R., & ROWELL, C. A Cohort-structured populations, selection responses, & exploitation of the North Sea cod. In: The Exploitation of Evolving Resources. (Eds. T. K. Stokes., J. M. McGlade. & R. Law): Springer- Verlag, Berlin. MAKAROV, E. V Otsenka dinamiki I struktur stada azovskikh osetrov kh. Trud VNIROU 7 1: MAKAROVA, I. A. & ALEKPEROV, A. 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54 Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea An integrated system for production fish aging: image analysis and quality assurance. North American Journal of Fisheries Management 18: MUSICK, J. A Criteria to define extinction risk in marine fishes. Fisheries 24: OLSEN, E. M., LILLY, G. R., HEINO, M., MORGAN, M. J., BRATTEY, J. & DIECKMANN, U Assessing changes in age & size at maturation in collapsing populations of Atlantic cod (Gadus morhua). Canadian Journal of Fisheries and Aquatic Sciences 62: PAULY, D., ALDER, A., BENNETT, E., CHRISTENSEN, V., TYEDMERS, P. & WATSON, R The future of fisheries. Science 302: PIKITCH, E. K., DOUKAKIS, P., LAUCK, L., CHAKRABATY, P., & ERICKSON, D. L Status, trends and management of sturgeon and paddlefish fisheries. Fish and fisheries 6: PIROGOVSKII, M. I. & FADEEVA, T. A The size age composition of the Stellate Sturgeon, Acipenser stellatus, during the marine period of life. Vopr Ichthyology 5: POURKASEMI, M Caspian Sea sturgeon Conservation and Fisheries: Past present and Future. Journal of Applied Ichthyology 22: PUTILIN, L. A Qualitative structure of the spawning part of the Persian sturgeon population of the Volga. Rational Principles of Sturgeon Farming, Volgograd, 210pp (in Russian). RIEN, T. A. & BEAMESDERFER, R. C Accuracy and precision of white sturgeon age estimates from pectoral fin rays. Transactions of the American Fisheries Society 123: RICKER, W. E Linear regressions in fishery research. Journal of the Fisheries Research Board of Canada 30: RICKER, W. E Computation and Interpretation of Biological Statistics of Fish Population. Bulletin Fisheries Research Board of Canada 191, 382 pp. RIJNSDORP, A. D Maturation of male & female North Sea plaice (Pleuronectes platessa L). Journal du Conseil International pour l explor mer 46: ROBERTS, C. M. & HAWKINS, J. P Extinction risk in the sea. Trends in Ecology & Evolution 14: ROSEN, R. A., HALES, D. C. & UNKENHOLZ, D. G Biology & exploitation of paddlefish in the Missouri River below Gavins Point Dam. Transactions of the American Fisheries Society 111: ROFF, D. A The evolution of life history parameters in teleosts. Canadian Journal of Fisheries and Aquatic Scien - ces 41: ROSTAMI, I Biologie et exploitation des sturgeons (Acipenserides) Caspians. Narledus (meuse), France, 210 pp, (In French). RUSSELL, T. R Biology and life history of paddlefish a review. In: ThePaddlefish: Status,Management & Propagation. (Eds. J. G. Dillard., L. K. Graham., & T. R. Russell): American Fisheries Society, Bethesda, Maryland. SABOROWSKI, R. & BUCHHOLZ, F Annual changes in the nutritive state of North Sea. Journal of Fish Biology 49: SAFINA, C., ROSENBERG, A., MYERS, R., QUINN, T. & COL- LIE, J. U. S Ocean fish recovery: staying the course. Science 309: SAINSBURY, K. J Effect of individual variability on the von Bertalanffy growth equation. Canadian Journal of Fisheries and Aquatic Sciences 37: SCARNECCHIA, D. L., STEWART, P. A. & POWER, G. J Age structure of the Yellowstone-Sakakawea paddlefish stock, , in relation to reservoir history. Transactions of the American Fisheries Society 125: SCARNECCHIA, D. L., RYCKMAN, L. F., LIM, Y., SCHMITZ, B. J. & FIREHAMMER, J. A Life history and the costs of reproduction in northern Great Plains Paddlefish (Polyodon spathula) as a potential framework for other acipenseriform fishes. Journal of Fisheries Science 15: SHABANY, A., POURKAZEMI, M., REZVANI, GHILKOLAHI, S., KAMALI, A. & VITESKAIA, L Study of mtdna variation of stellate sturgeon population from Northern Caspian Sea (Volga River) and Southern Caspian Sea (Gorganroud river) using RFLP analysis of PCR amplified ND5/6 gene regions. Iranian Journal of Marin Science and Technology 2: STAMPS, J. A Sexual size dimorphism in species with asymptotic growth after maturity. Biological Journal of the Linnean Society 50: STEVENSON, J. T. & SECOR, D. H Age determination and growth of Hudson River Atlantic sturgeon (Acipenser oxyrinchus) Fisheries Bulletin 98: TAGHAVI MOTLAGH, A Population dynamics of Sturgeon in the southern part of the Caspian Sea. Dissertation, University of Wales, Swansea, 300pp. VESHCHEV, P. A Composition of the spawning population of the sturgeon, Acipenser guldesnstadti and Acipenser stellatus, in the spawning grounds of Akhtuba River. Vopr Ichthyology 19: VESHCHEV, P. V. & NOVIKOVA, A. S Biological characteristics of sevryuga, Acipenser stellatus, on the spawning grounds of the lower Volga. Vopr Ichthyology 5: ZAR, J. H Biostatistical analysis. Prentice Hall, New Jersey, 123 pp. aqua vol. 18 no April

55 aqua, International Journal of Ichthyology Scleropages inscriptus, a new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes) Tyson R. Roberts Smithsonian Tropical Research Institute, Balboa, Panama. tysonscleropages@gmail.com Received: 19 March 2012 Accepted: 09 April 2012 Abstract Scleropages inscriptus, a new species of bony-tongue fishes, is described from the Tenasserim or Tananthayi River basin on the Indian Ocean coast of peninsular Myanmar. It differs from the previously known Southeast Asian and Australian members of the genus in having the bones of the circumorbital and opercular series and all or most of the scales on the sides of the body densely covered with complex maze-like markings. In morphology and in meristic and morphometric characters it is closer to the other Asian species of Scleropages, S. formosus, than to S. leichhardti or S. jardinii, the two species currently recognized from the Australian Region; it is therefore referred to the subgenus Delsmania Fowler 1934 (type species S. formosus). Zusammenfassung Beschrieben wird Scleropages inscriptus, eine neue Art der Knochenzüngler, vom Einzugsgebiet des Flusses Tenasserim oder Tananthayi an der Küste des Indischen Ozeans auf der Halbinsel in Myanmar. Von den bisher bekannten südostasiatischen und australischen Angehörigen der Gattung unterscheidet sich die neue Art dadurch, dass die Knochen der zirkumorbitalen und opercularen Reihen und alle oder die meisten Schuppen auf beiden Rumpfseíten dicht mit komplexen, wirr verteilten Markierungen versehen sind. Nach Morphologie und meristischen wie morphometrischen Merkmalen steht die neue Art der anderen asiatischen Scleropages-Art: S. formosus, näher als den beiden gegenwärtig aus dem australischen Raum bekannten Arten, S. leichhardti oder S. jardinii; sie wird deshalb der Untergattung Delsmania Fowler 1934 zugeordnet (Typenart S. formosus). Résumé Scleropages inscriptus, une nouvelle espèce d Ostéoglossidé, est décrit en provenance du Tenasserim ou du bassin de la rivière Tananthayi, sur la côte de l Océan Indien de la péninsule de Myanmar. Il se distingue des membres du genre déjà connus du sud-est asiatique et d Australie par les os des séries circumorbitales et operculaires et par toutes ou la plupart des écailles des côté du corps densément couverte de marques complexes en labyrinthe. Au point de vue de la morphologie et des caractéristiques méristiques et morphométriques, l espèce est plus proche de l autre espèce asiatique de Scleropages, S. formosus, que de S. leichhardti ou de S. jardinii, les deux autres espèces asiatiques actuellement identifiées dans la région australienne ; pour cette raison, elle est référée au sous-genre Delsmania Fowler 1934 (espèce-type S. formosus). Sommario Scleropages inscriptus, una nuova specie di pesci lingua d osso, è descritta dal bacino del fiume Tenasserim o Tananthayi sulla costa dell'oceano Indiano della regione peninsulare del Myanmar. Si differenzia dai membri già noti del genere del sud-est asiatico e dell Australia per avere le ossa della serie circumorbitale e opercolare e tutti o la maggior parte delle scaglie ai lati del corpo densamente coperti di complessi labirinti. In morfologia e in caratteri meristici e morfometrici è più vicina alla specie asiatica di Scleropages, S. formosus, che a S. leichhardti o S. jardinii, le due specie attualmente riconosciute dalla regione australiana; la nuova specie è pertanto inclusa nel sottogenere Delsmania Fowler 1934 (specie tipo S. formosus). INTRODUCTION About ten years ago word reached Bangkok that Scleropages occurs in the Myanmar part of the Malay Peninsula. As Scleropages formosus was well known from the Malaysian part of the peninsula and neighboring areas to the east and southeast, it was assumed that the population in Myanmar was the same species. When Bangkok aquarists saw specimens in the Yangon aquarium trade, however, they recognized immediately that it was an undes - cribed species with a highly distinctive color pattern unlike anything previously known (Anon., 2011; 2012). 113 aqua vol. 18 no April 2012

56 Scleropages inscriptus, a new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes) There is a substantial gap in distribution between the known localities for S. formosus in the Malay Peninsula and the Tenasserim or Tananthayi River basin, presently considered the only river system inhabited by the new species. Geographical distribution of S. formosus lies within the area formerly occupied by the Greater Sunda River system and in lowland areas immediately adjacent to it, while the Tenasserim or Tananthayi River system of Myanmar lies outside it and separated from it by lowlying but biogeographically significant mountain ranges. The holotype and paratype of S. inscriptus, specimens obtained dead from vendors in Meik, are deposited in the fish collection of the Thailand Natural History Museum of the National Science Museum in Pathum Tani. Photographs are presented of the preserved holotype and of a larger live fish not designated as a type specimen. SYSTEMATICS Scleropages Günther 1864 Scleropages Günther, 1864: 196 (type species Scleropages leichardti Günther 1864, type locality Burdekin River, Queensland, Australia, by monotypy and original designation. Delsmania (subgenus of Scleropages) Fowler, 1934: 243 (type species Osteoglossum formosum Müller & Schlegel 1844, type locality Borneo, Barito River, by monotypy and original designation. The genus Scleropages was described by Albert Günther for Scleropages leichhardti Günther 1864, the first known osteoglossoid fish from the Australian Region. The second known species from the Australian Region is S. jardinii (Saville-Kent 1892). Henry Weed Fowler distinguished the only previously known Asian species of Scleropages as a subgenus of Scleropages, which he named Delsmania. This taxon has not been generally applied but is available at generic or subgeneric level. The new species S. inscriptus clearly belongs in Delsmania rather than Scleropages. Delsmania differs from Scleropages most obviously in having only instead of scales in the lateral line series; it also has a less elongate body, longer snout and longer pectoral and pelvic fins (Pouyaud, et al., 2003: 293). The osteoglossid genus most closely related to Scleropages, Osteoglossum, lives in South America. Müller and Henle (1844) originally placed S. formosus in Osteoglossum. Fig. 1. Scleropages inscriptus, holotype, THNHM-F-01521, 385 mm, presumed locality Tananthayi River system, Myanmar. Right side is shown because left side is missing many scales. Scleropages formosus (Müller & Schlegel 1844) Osteoglossum formosum Müller & Schlegel, 1844: 7, pl. 1 (type locality Barito River, Borneo). Scleropages aureus Pouyaud, Sudarto & Teugels, 2003: 298, figs. 10, 11A-B (type locality Pekanbaru market, wild specimen from Siak River, Sumatra, Indonesia). Scleropages legendrei Pouyaud, Sudarto & Teugels, 2003: 300, figs. 12, 13 A-C (type locality Semiaqua vol. 18 no April

57 Tyson R. Roberts tau, area of Sentarum Lake, West Kalimantan, Indonesia). Scleropages macrocephalus Pouyaud, Sudarto & Teugels, 2003: 296, Figs. 8, 9A-B (type locality Pinoh market, wild specimen from Melawi River, West Kalimantan, Indonesia). Scleropages inscriptus, n. sp. Figs 1-2 Holotype: THNHM-F-01521, 385 mm standard length, sex undetermined, supposedly from Myanmar, Tananthayi district, Tananthayi River basin, obtained dead from aquarium fish vendor at Meik, Paratype: THNHM-F-01522, 290 mm standard length, sex undetermined, same locality data as holotype. Diagnosis: Scleropages inscriptus differs from all other Scleropages in having highly distinctive angulated linear markings on dermal bones of circumorbital and opercular series on sides of head and on scales on sides of body. It differs from the two Australian species of the subgenus Scleropages in meristic and morphometric characters, while at the same time agreeing closely in these characters with its Southeast Asian congener S. formosus of the subgenus Delsmania. Description: Broad dorsal surface of head and body and scales of predorsal scale row dark, without maze-like markings; background color of sides of head and body slightly iridescent, varying from bronze, pearly or pinkish to white or silvery; in fully grown fish dark lines, straight or curved, often joining at right angles, form a maze-like pattern on dermal bones of circumorbital and opercular series and on all scales on sides of body; similar markings on small scales on base of median fins but not on fin rays extending beyond them; pectoral fins with dark rays and pale membranes; pelvic fins pale like background color of body; median fins uniformly dark (not rosy or pinkish); barbels dark dorsally and white ventrally, sometimes with red streak; opening of pupil with red border, eye otherwise dark. Meristic and morphometrics: The following counts and measurements are from the holotype (and paratype), the only preserved specimens presently available. Measurements are expressed as times in standard length: gill rakers on first gill arch 6+9=15 (6+10=16); pored scales in lateral line scale row 23 (23); median predorsal scales 17 (18); circumferential scales (in front of pelvic fins) 13 (13); circumpeduncular scales 13 (13); peduncular scales (from base of last anal fin ray to end of hypural plate) 9 (9); dorsal fin rays 20 (20); anal fin rays 27 (28); pectoral fin rays 7 (7); pelvic fin rays 5 (5); and caudal fin rays i7+7i (i8+8i); head 3.5 (4.0); snout 16.0 (20.7); horizontal diameter of eye 21.4 (17.1); interorbital width 11.3 (12.1); barbel length 64 (58); greatest depth of body 2.4 (4.0); predorsal length 1.2 (1.3) ; pre-pelvic length 2.0 (2.1); pre-anal length 1.35 (1.6); depth of cau- Fig. 2. Scleropages inscriptus, live non-type specimen, about 600 mm, Yangon aquarium trade, aqua vol. 18 no April 2012

58 Scleropages inscriptus, a new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes) Fig. 3. Dot map with distribution records of the two currently recognized species of Scleropages subgenus Delsmania (after Pouyaud, et al., 2003). Records of S. formosus for peninsular Thailand are from Vidthayanon et al., 1997, and Vidthayanon, 2002; 2004). The single blue dot for S. inscriptus is placed near middle of the Tananthayi River system. aqua vol. 18 no April

59 Tyson R. Roberts dal peduncle 12.8 (12.6); length of caudal peduncle 21.4 (17.1). Ontogenetic variation: Young fish up to 200 mm lack maze-like markings on the head and body and have overall silvery or whitish coloration and plain color pattern nearly identical to immature S. formosus which differ mainly in having pinkish or rosy median fins (Pouyaud et al., 2003: figs. 7, 8, 11A, 13C). The distinctive markings of S. inscriptus appear first on the bones of the gill covers at about 200 mm. At progressively larger sizes they extend to the circumorbital bones, the lower jaw, the gill cover membrane, and the scales on the sides of the body. At 300 to about 500 mm the markings cover nearly all of the scales on the sides of the body except the lowermost two rows on the abdomen. Only in specimens of mm does the mazelike pattern extend to all of the scales on the body except the darkened uppermost median pre-dorsal row of scales. Distribution: No precise locality data is available for any specimens of this new species. Thus far it is known only from the aquarium trade in Meik and Yangon. According to a 2006 website posting of the Myanmar fish exporting company Hein Aquarium, the fish comes from the Tananthayi or Tenasserim River basin in Tananthayi district of peninsular Myanmar (Fig. 3). No other localities have been reported. Etymology: The Latin trivial or species name inscriptus, an adjective meaning inscribed, refers to the distinctive markings on the scales and facial bones. Discussion: Sufficient numbers of live specimens of S. inscriptus have been photographed for it to be noted that the maze-like markings are never exactly the same on any two individuals of the species, on both sides of an individual fish, or on any two bones or scales. Nothing quite like this color pattern has been reported in any other osteoglossoid fish or for that matter in any other bony fish. Overall morphology and nearly identical meristic and morphometric characters indicate that the new species S. inscriptus is most closely related to S. formosus. Based on the information presently available, the three closely related new species or color varieties of Southeast Asian Scleropages proposed by Pouyaud et al. (2003), as recommended by Kottelat and Widjanarti (2005: 145), are not regarded as valid species. Data from partial sequences of DNA coding for cytochrome b and the nearest neighbor dendrogram constructed from it (Pouyaud, et al., 2003: , table 1, fig. 1) are consistent with the following taxonomic concepts or hypotheses: 1) Scleropages and Delsmania are distinct subgenera; 2) Scleropages leichardti and S. jardinii are distinct species; and 3) Scleropages macrocephalus, S. aureus, and S. legendrei are conspecific with S. formosus. Having lived in Bangkok for much of the past 40 years, mostly in Sapan Khwai only fifteen minutes walk from the most important Thai tropical fish emporium in Chatuchak Park, I have seen many color variations of S. formosus. All of the so-called arowanas or dragon fish are expensive, but unusually colored ones may fetch ten times or more the price of normally colored ones. Any aquarist would of course add brine shrimp to fish diets if the fish liked the food and it enhanced their coloration. Hobbyists have been doing this with S. formosus for decades. But commercial dealers reportedly have gone far beyond that, adding to the food artificial colors including, but not limited to, those used to produce the colors of popular soft drinks such as nam daeng, nam kheao, and nam see sohm (red drink, lime green drink, and orange-colored drink) to produce new and more valuable color varieties. Of course this is a trade secret, and my information on such practices is hearsay or inference. The problem is not mentioned by Pouyaud et al. (2003), so perhaps they did not know about it. This presumably has contributed to rejection of their paper by other ichthyologists. Nothing like this has happened yet with the new species of Scleropages, because too few of them have been marketed and the species probably has yet to be bred in captivity. Bangkok aquarists have succeeded in producing great differences in color pattern in the featherfin Chitala ornata (Osteoglossiformes, Notopteridae), but this has been achieved by selective breeding. The differences in coloration in S. formosus never involve color pattern, only coloration. Thus the striking differences in color pattern in S. inscriptus strongly indicates that it is not the same species as S. formosus. Scleropages formosus is distributed extensively in mainland and insular Southeast Asia almost entirely within the geographical area of the former Greater Sunda River basin (Fig. 3). During periods of lower sea level this area was continuous land drained mainly by a single great river system, now represented by several large rivers separated by shallow oceanic gaps including the Mekong, Chao Phraya, Meklong and Tapi on the mainland, Kapuas of western Borneo, and Djambi and 117 aqua vol. 18 no April 2012

60 Scleropages inscriptus, a new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes) Batang Hari of Sumatra. Numerous freshwater fish genera and species are common to this area. The Tenasserim or Tananthayi River basin of peninsular Myanmar lies outside the area formerly occupied by the Greater Sunda River. Most of its freshwater fish genera and species are shared with the Salween and Irrawaddy river basins of central Myanmar. Only Scleropages and relatively few other genera are shared with Sundaic rivers and not with the Salween or Irrawaddy. S. formosus occurs in the Thai and Malaysian parts of the Malay Peninsula, in which the main river systems are the Tapi and Perak. The Tapi and other river systems of peninsular Thailand are separated from the Tenasserim and other river systems of peninsular Myanmar by mountain ranges that apparently have been a barrier especially for large lowland freshwater fish genera such as Scleropages. Scleropages inscriptus is one of the most charismatic and attractive of the freshwater fishes endemic to Myanmar. Its continued availability to aquarium fish enthusiasts may depend on successful efforts to cultivate it in captivity. Scleropages formosus has been maintained and successfully bred in varying degrees of captivity and semi-captivity in Thailand, Malaysia and Indonesia for many years. It should be possible to duplicate this success with the new species. REFERENCES ANON Aktuelles kurz gemeldet: Ein neuer Arowana aus Myanmar. Amazonas (German edition) 7 (6): 3. ANON Aquatic Notebook: A new arawana from Myanmar. Amazonas (USA edition) 1 (2): 6. FOWLER, H. W Descriptions of new fishes obtained 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Proceedings of the Academy of Natural Sciences of Philadelphia 85: GÜNTHER, A On a new generic type of fishes discovered by the late Dr. Leichardt in Queensland. Annals and Magazine of Natural History series 3 14 (81): , pl. 7. KOTTELAT, M. & WIDJANARTI, E The fishes of Danau Sentarum National Park and the Kapuas Lake Area, Kalimantan Barat, Indonesia. Raffles Bulletin of Zoology Supplement No. 13: MÜLLER, S. & SCHLEGEL, H Beschrijving van een nieuwen Zoetwater-visch van Borneo, Osteoglossum formosum. Verhandelingen over de natuurlijke geschiednis der Nedertaldsche overzeesche bezittinen, door de Leden der Natuurkundige Commissie in Indies. Leiden, 1-7. POUYAUD, L., SUDARTO & TEUGELS, G. G The different colour varieties of the Asian arowana Scleropages formosus (Osteoglossidae) are distinct species: morphologic and genetic evidences. Cybium 27(4): SAVILLE-KENT, W Description of a new species of true Barrimundi, Osteoglossum jardinii, from northern Queensland. Proceedings of the Royal Society of Queensland 8 (3): VIDTHAYANON, C Peat Swamp Fishes of Thailand [in Thai and English]. Office of Environmental Policy and Planning, Bangkok, 136 pp. VIDTHAYANON, C Handbook of Freshwater Fishes [in Thai]. Sarakadee Press, Bangkok, 232 pp. VIDTHAYANON, C., KARNASUTA, J. & NABHITABHATA, J Diversity of Freshwater Fishes in Thailand [in Thai]. Office of Environmental Policy and Planning, Bangkok, 102 pp. aqua vol. 18 no April

61 Book review THE BANGGAI CARDINALFISH: Natural History, Conservation and Culture of Pterapogon kauderni Alejandro A. Vagelli Wiley-Blackwell, John Wiley and Sons, Ltd. 2011, 224 pages ISBN (Hardcover) $ ISBN (ebook) $ There are three online excerpts at: CDA/WileyTitle/productCd html, in addition to a brief description and table of contents of what is in the book. Dr. Vagelli divided his treatise on the single species in Pterapogon into four parts: 1) a historical review of previous research, overview of the geography, ecology, and human activities of the Banggai archipelago; 2) an in-depth description of distribution, morphology, reproduction, ecology, and genetics followed by a review and hypotheses for systematics and evolution of Pterapogon including possible fresh-water origin of the Apogonidae; 3) conservation and regulation of the wild harvest of Pterapogon for sale in the world-wide aquarium trade; and 4) captive breeding and rearing of Pterapogon. Scientific knowledge of Pterapogon is mostly after the 1992 rediscovery apart from the original description in 1933 and an osteological description in 1972 based on the type specimens. By 1998 scientists expressed concerns about threats of overfishing due to demands by the aquarium trade based on the known limited distribution of the species and its low spawning potential. The book contains significant information about other members of the Apogonidae in Part 2, but this is not indicated in the title or in the online descriptions. Chapter 3 contains a geologic history of the Banggai Archipelago with 5 maps summarizing existing knowledge. The island complex inhabited by Pterapogon originated near New Guinea. This fact coupled with the analysis of known distribution and direct development of young leads the author to interesting hypotheses about vicariance of the species and evolution of the Apogonidae. Chapter 5 describes the methods used to determine island distributions within the natural (native) geographic range (see map 3.2). Three island groups isolated by deep-water barriers were proposed. Known genetic information suggests that even small spatial distances of 2-5 km show genetic isolation (Chapter 9). Direct development, shallow water preferences and a body shape indicative of slower swimming ability all must play a role in reducing ability to disperse. The author concludes that the present fine scale native distribution was the result of localized colonization from sea level fluctuations and the species geographic distribution with respect to other apogonids was from geologic vicariance at least since the Oligocene. There has been a non-native range expansion of Pterapogon by human introductions, likely as part of the aquarium trade. In Chapter 6 the author presents new information about the morphology of Pterapogon with some comparisons to other apogonids. His main intent is to provide morphological aspects useful in taxonomy, ecology and phylogenetic studies. Table 6.1 displays meristic characters with variation among the islands. For example, contrast specimens from Banggai and Seku for gill raker counts: Upper Gill Rakers Lower Gill Rakers Banggai Seku If the spot polymorphism is stable and variable across geographic populations (Chapter 9.3), then it may be likely that specimens from Banggai and Seku could be completely separated as if they were different species by mid-body spot pattern (Fig. 9.1) and gill raker counts. Add the genetic evidence, populations of Pterapogon may be acting like incipient species (reviewer s observation). Aspects of the reproduction of apogonids are discussed in Chapter 7 with a comprehensive table (7.1) summarizing 45 species in at least 12 (22 by newer information). There are significant gaps in available facts, but this work is an excellent place to start. Dr. Vagelli rightly questions reported cases of female egg brooding, internal fertilization and multiple egg ball brooding, all needing better documentation. Male brooding of the egg ball has been 119 aqua vol. 18 no April 2012

62 documented many times. Sexual dimorphism in apogonids appears to be confined to small species of Siphamia and temporarily to buccal cavity volume of male apogonids just before and during brooding the egg ball. A number of apogonids display temporary dichromatism during courtship, others do not. I know of at least one species, Ostorhinchus lateralis, with permanent dichromatism involving the presence or absence of a small spot on the side of the head. Other suspected examples involve the presence or absence of a black spot surrounding the genital area of a species of Cheilodipterus and a species of Ostorhinchus. Pterapogon has large eggs brooded by the male, exhibits no known dimorphism or dichromatism and is characterized by courtship and spawning taking place during the day. There is a comprehensive section (7.2) on almost all aspects of the reproductive processes from courtship and spawning, the morphology and development of ova and sperm, early ontogeny of fertilized eggs and evidence for considering Pterapogon as having the lowest fecundity in the family. Morphological information did not extend to the micropyle, ridge pattern or filaments on the ova or eggs. The estimated age and growth was from captive fish. The author inferred that in nature the maximum age was ~2.5-3 years. Batch spawning may occur at intervals of ~25 days (captive specimens) in adult females greater than 40 mm standard length year round. Female fish appear to choose a mate. Figure 7.3a-f depicts female egg ball extruding, spawning and transfer action by the male. Habitat preferences and associations with living organisms begin chapter 8. The author missed a review article by J.E. Randall (2005) describing a number of cases of mimicry for apogonids. A complete description is given of three kinds of hosts, sea urchins, anemones and branching corals, that Pterapogon associates with during the day (color plates ). Diet consisted mostly of harpacticoid, calanoid and cycloid copepods and other small crustaceans. Feeding begins at first light and ceases at dusk. Ectoparasites are briefly touched upon. Systematics and evolution of Pterapogon and its nearest relative are in flux. The author discusses at length the evidence for direct development, bypassing larval stages, for Pterapogon, Glossamia, Quinca, and Vincentia. Species of the latter three genera are confined to the main Australian-New Guinea land mass. None has been found on intermediate islands between the Bird s Head of New Guinea and the Banggai Island complex. The author sets up two hypotheses for the nearest living relatives: 1) Pterapogon is sister to Sphaeramia and 2) Pterapogon is sister to the only other genera known to have direct development. The first hypothesis gives no clue to whether the ancestor of Pterapogon originated before or after complete separation from the Australian-New Guinea land mass and may imply multiple origins/lineages for large eggs and direct development to juveniles. (The reviewer favors the first alternative that they are sister genera and both are sister to Eosphaeramia, a well-preserved fossil about 50 million years old from Monte Bolca, Italy, based on shared morphologic characters.) The second hypothesis implies the ancestor existed on part of the Australia-New Guinea land mass and gave rise to three other genera with direct development the uniting factor. In the author s scenario of freshwater origin of the Apogonidae, direct development would be basal, not a synapomorphy. Holapogon, mentioned only briefly, would support a marine origin of the family, having fewer specializations than Glossamia or Vincentia. Published partial molecular hypothetical trees to date for cardinalfish do not support, in part, either hypothesis. The author clearly admits...the suggestion of an ancestral freshwater apogonid lineage is particularly speculative. Dr. Vagelli discusses some of the literature, pre-2010, about interrelationships among the Gobioidei, Kurtidae and Apogonidae. The search (section 10.5) for convincing morphological evidence to agree with partial molecular trees is not materially advanced with his review of literature. The idea that an ancestor to gobioids, kurtids and apogonids originated somewhere on the Australian-New Guinea landmass in freshwater is new. Chapters 11 to 13 examine various aspects of the aquarium trade fishery for Banggai cardinalfish. The species is on the International Union for the Conservation of Nature s Red List as endangered, but failed to obtain protection by the Convention on International Trade in Endangered Species. Chapter 14 on captive breeding probably is the best hope for blunting overexploitation of natural populations. Undoubtably, there will be much more research associated with this unique apogonid. There is a very complete reference list and a well-documented index. I had no problems navigating around the book. There are a few typing errors, but they are not enough to be distracting. The online companion website did not work for me. The author is willing to send electronic figures and tables if the site is unavailable. It is a beautiful, pleasing book for those who collect fish books. The Banggai Cardinalfish will be a primary source for students, ichthyologists (molecular and morphologic), hobbyists and professional aquarists. Literature Cited: RANDALL, J. E Review of mimicry in marine fishes. Zoological Studies, 44 (3): , 124 figs. Thomas H. Fraser Ichthyology Florida Museum of Natural History, University of Florida- Gainesville, Florida 32611, USA aqua vol. 18 no April

63 Guidelines for Authors 1. Manuscript preparation: manuscripts must be submitted in English. In exceptional cases aqua may provide translations of manuscripts written in French, German, Italian, or Spanish. Manuscripts must be word-processed in Microsoft WORD and submitted in an electronic form. Generic, specific, and sub-specific names must be italicised. All papers must conform to the International Code of Zoological Nomenclature. Authors are strongly advised to follow the format set out in previous publications of aqua. 2. Title: the title must be short and should precisely identify the main topic of the article. Names of genera or species are followed by the systematic group to which they belong. Author name(s) are given in full beneath the title, followed by the complete mailing and address(es). 3. Abstract: the abstract should not exceed 250 words and draw attention to the principal conclusions. It should not contain any uncommon abbreviations or literature citations. The inclusion of abstracts in other languages is optional. 4. Subject matter: the text of the manuscript is usually arranged in four main sections: Introduction, Materials and methods (including a key to abbreviations), Results, and Discussion. Other subdivisions may be chosen depending on the material presented. Acknowledgements should be placed between the text and references. Scientific names of genera, species, and subspecies should be followed by the name(s) of author(s) and the year of publication on first mention. A description of a new taxon must contain the following sections: Material, Diagnosis, Description, and Affinities. Synonyms must be arranged in chronological order. Identification keys must be dichotomous. Holotype and paratypes must be clearly identified, the institution in which they have been deposited named, and the catalogue numbers given. Private collections are not acceptable as repositories for holotypes. DNA sequences must be archived in GenBank, DNA Databank of Japan or European Molecular Biology Laboratory. Voucher specimens associated with DNA sequences must be deposited in a recognized research collection accessible to the professional community. Accession numbers for sequences and catalog number for vouchers must be included as a table in an appendix to the final manuscript. The metric system and SI units must be used. Temperatures are given in C. Fractions should not be used. 5. References to literature: the name-year system must be used. The list of references should be placed at the end of the paper, alphabetically arranged according to author name. Only those publications cited in the paper may be included. Titles of journals must be given in full. Examples of correct reference formats: BLABER, S. J. M Fish of the Trinity inlet system of North Queensland, with notes on the ecology of fish faunas of tropical Indo-Pacific estuaries. Australian Journal of Marine and Freshwater Research 31: DAY, J. H., BLABER, S. J. M., & WALLACE, J. H Estuarine fishes. In: Estuarine Ecology with Particular Reference to Southern Africa. (Ed. J.H. Day.): A. A. Balkema, Rotterdam. DIMMICH, W. W Ultrastructure of North American cyprinid maxillary barbels. Copeia 1988 (1): TREWAVAS, E Tilapiine Fishes of the Genera Sarotherodon, Oreochromis and Danakilia. British Museum (Natural History), London, 583 pp. 6. Submission of manuscript and illustrations: The manuscript and illustrations must be submitted digitally to the Scientific Editor: Dr Frank L. Pezold College of Science & Engineering Texas A&M University Corpus Christi 6300 Ocean Drive Corpus Christi, TX frank.pezold@tamucc.edu to whom all subsequent correspondence shall be addressed. Texts, tables, and graphs should be in Microsoft-compatible electronic form. After the paper has been accepted for publication, illustrations as high-resolution TIF files or original photographs (ideally transparencies; otherwise glossy prints, preferably in the size in which they will appear - the type area of aqua is 158 x 224 mm, one column is 76 mm wide) must be sent to: Aquapress, The Managing Editor Via G. Falcone 11, Miradolo Terme (Pavia), Italy aquapress@aquapress-bleher.it Authors should retain copies of all materials for reference. Proofs of accepted papers will be sent as PDF files by e- mail attachment to the corresponding author. 7. Evaluation of manuscripts: manuscripts will be evaluated by the editors and referees. Papers are accepted on the understanding that they have not and will not be published elsewhere. 8. Reprints: Authors will receive one free copy of the issue in which their paper appears and an e-print in PDF format. Additional copies may be ordered from Aquapress.

64 aqua International Journal of Ichthyology Vol. 18 (2), 15 April 2012 Contents: James L. Van Tassell, Luke Tornabene, Patrick L. Colin: Review of the western Atlantic species of Bollmannia (Teleostei: Gobiidae: Gobiosomatini) with the description of a new allied genus and species Rüdiger Riesch, Ryan A. Martin, David Bierbach, Martin Plath, R. Brian Langerhans and Lenin Arias-Rodriguez: Natural history, life history, and diet of Priapella chamulae Schartl, Meyer & Wilde 2006 (Teleostei: Poeciliidae) Shima Bakhshalizadeh, Shahram Abdolmalaki, Ali Bani: Aspects of the life history of Acipenser stellatus (Acipenseriformes, Acipenseridae), the starry sturgeon, in Iranian waters of the Caspian Sea Tyson R. Roberts: Scleropages inscriptus, a new fish species from the Tananthayi or Tenasserim River basin, Malay Peninsula of Myanmar (Osteoglossidae: Osteoglossiformes) Book review: The Banggai Cardinalfish: Natural History, Conservation and Culture of Pterapogon kauderni. Alejandro A. Vagelli Papers appearing in this journal are indexed in: Zoological Record; BioLIS Biologische Literatur Information Senckenberg; Cover photo: Scleropages inscriptus n. sp., semi-adult in aquaria. Photo courtesy by J. Tan. Pseudanthias bicolor, 83 mm SL, male, Mauritius. Photo by J. E. Randall from an upcoming issue of aqua.