aqua, International Journal of Ichthyology Vanderhorstia lepidobucca, a new species of shrimpgoby from Sulawesi, Indonesia Gerald R. Allen 1, Teguh Peristiwady 2 and Mark V. Erdmann 3,4 1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: tropical_reef@bigpond.com 2) Technical Implementation Unit for Marine Biota Conservation, Indonesian Institute of Sciences, Tandurusa, Bitung, North Sulawesi, 95227 Indonesia 3) Conservation International Indonesia Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235, Indonesia 4) California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, USA Received: 15 January 2014 - Accepted: 29 March 2014 Abstract A new species of gobiid fish, Vanderhorstia lepidobucca, is described from Lembeh Strait, North Sulawesi, Indonesia on the basis of eight specimens, 23.5-40.4 mm SL. The new taxon is the only member of the genus possessing scales on the preoperculum (cheek) and also three scales on the upper anterior corner of the operculum. Other diagnostic features include 13 dorsal soft rays, 14 anal soft rays, first dorsal fin relatively short with no elongate rays, 51-55 longitudinal scales, 17-18 predorsal scales, and a lanceolate caudal fin. The head and body are generally light brown with yellow vermiculated bands on the head, prominent yellow and blue-margined black spot on the middle of the operculum, and about 16-18 short brown bars along the middle of the body. The median fins are generally bluish with yellow spots, stripes, or streaks, except the anal fin is mainly yellow with blue on the distal third. Zusammenfassung Beschrieben wird eine neue Art der Gobiiden: Vanderhorstia lepidobucca, von der Lembeh Strait, Nordsulawesi, Indonesien, auf der Grundlage von acht Exemplaren mit 23,5 bis 40,4 mm Standardlänge. Die neue Art ist die einzige Vertreterin der Gattung mit Schuppen auf dem Prä-Operculum (Wangenbereich) und außerdem drei Schuppen auf der obersten, vorderen Stelle des Kiemendeckels. Weiterhin gibt es folgende Unterscheidungsmerkmale: 13 weiche Rückenflossenstrahlen, 14 weiche Afterflossenstrahlen, erste Rückenflosse verhältnismäßig kurz ohne verlängerte Flossenstrahlen, 51-55 Schuppen in der Längsrichtung, 17-18 prädorsale Schuppen sowie eine lanzettförmige Schwanzflosse. Kopf und Rumpf zeigen eine hellbraune Grundfarbe mit gelben geschlängelten Streifen am Kopf, einem auffälligen schwarzen Fleck mitten auf dem Kiemendeckel mit gelbem und blauem Rand sowie 16-18 kurzen braunen Streifen auf der Mittellinie des Rumpfes. Die mittleren Flossen sind grundsätzlich bläulich mit gelben Flecken, Streifen oder Strichen; die Afterflosse aber ist hauptsächlich gelb mit blauer Tönung auf dem hinteren Drittel. Résumé Une nouvelle espèce de gobie, Vanderhorstia lepidobucca, est décrite, originaire de Lembeth Strait, North Sulawesi, Indonésie, sur base de huit spécimens, de 23,5-40,4 mm de LS. Le nouveau taxon est le seul membre du genre à possé - der des écailles sur le préopercule (joue) ainsi que trois écailles sur l angle supérieur antérieur de l opercule. D autres don nées diagnostiques comprennent 13 rayons dorsaux mous, 14 rayons mous à l anale, une première dorsale relati - vement courte sans rayons allongés, 51-55 écailles longitudi - nales, 17-18 écailles prédorsales et une caudale lancéolée. La tête et le corps sont généralement brun clair avec des bandes vermiculées jaunes sur la tête, une tache noire évidente bordée de jaune et de bleu au centre de l opercule et de 16-18 petites barres brunes le long du centre du corps. Les nageoires médianes sont généralement bleuâtres avec des ta - ches jaunes, des lignes ou des raies, sauf l anale qui est surtout jaune avec du bleu sur le tiers distal. Sommario Una nuova specie di gobide, Vanderhorstia lepidobucca, è descritta da Lembeh Strait, Nord Sulawesi, Indonesia, sulla base di otto esemplari di 23,5-40,4 mm SL. Il nuovo taxon è l'unico membro del genere con scaglie sul preopercolo (guancia) e anche tre scaglie all'angolo antero-superiore dell'opercolo. Altre caratteristiche diagnostiche includono 13 raggi molli dorsali, 14 raggi molli anali, prima pinna dorsale relativamente breve, senza raggi allungati, 51-55 scaglie longitudinali, 17-18 scaglie predorsali e pinna caudale lanceolata. La testa e il corpo sono generalmente di colore marrone chiaro con bande gialle vermiculate sulla 81 aqua vol. 20 no. 2-15 April 2014
Vanderhorstia lepidobucca, a new species of shrimpgoby from Sulawesi, Indonesia testa, una prominente macchia nera con margine giallo e blu al centro del opercolo e circa 16-18 brevi barre marroni lungo la parte centrale del corpo. Le pinne mediane sono generalmente bluastre con macchie, strisce o striature gial - le, tranne la pinna anale che è prevalentemente di colore giallo con il terzo distale di colore blu. INTRODUCTION The Indo-Pacific gobiid fishes of the genus Vanderhorstia Smith 1949 live symbiotically with alpheid snapping shrimps. There are 25 currently recognized species (Senou et al. 2004; Shibukawa & Suzuki 2004; Greenfield & Longenecker 2005; Winterbottom et al. 2005; Iwata et al. 2007; Randall 2007a and b; Randall & Munday 2008; Allen & Erdmann 2012; Eschmeyer 2013). The present paper describes a new species that was collected by the authors at Lembeh Strait, North Sulawesi in 2013. This distinctive new taxon differs from all known members of the genus in having a fully scaled preopercle (cheek) and three scales on the upper anterior corner of the operculum (both areas scaleless in other Vanderhorstia). MATERIALS AND METHODS Type specimens are deposited at Museum Zoologicum Bogoriense, Cibinong, Java, Indonesia (MZB), the Indonesian Institute of Sciences Bitung Field Station collection (LBRC-F) and the Western Australian Museum, Perth (WAM). Lengths are given as standard length (SL), measured from the median anterior point of the upper lip to the base of the caudal fin (posterior end of the hypural plate); body depth is measured at both the origin of pelvic fins and the origin of the anal fin, and body width at the origin of the pectoral fins; head length is taken from the upper lip to the posterior end of the opercular membrane, and head width over the posterior margin of the preopercle; orbit diameter is the greatest fleshy diameter, and interorbital width the least fleshy width; snout length is measured from the median anterior point of the upper lip to the nearest fleshy edge of the orbit; upper-jaw length is from the same anterior point to the posterior end of the maxilla; cheek depth is the least depth perpendicular from the ventral edge of the suborbital to the fleshy edge of the orbit; caudal-peduncle depth is the least depth, and caudal-peduncle length the horizontal distance between verticals at the rear base of the anal fin and the caudal-fin base; lengths of spines and rays are measured to their extreme bases; caudal- and pectoral-fin lengths are the length of the longest ray; pelvic-fin length is measured from the base of the pelvic spine to the tip of the longest pelvic soft ray. Morphometric data presented as percentages of the standard length is indicated in parentheses in the following description. Proportional measurements in the text are rounded to the nearest 0.05. Terminology and abbreviations for cephalic pores and papilla rows follow those presented by Akihito (1984) and Shibukawa & Suzuki (2004). The count of scales in longitudinal series is made from above the dorsal end of the gill opening to the base of the caudal fin; scales in transverse series are counted from the origin of the anal fin anterodorsally to the base of the first dorsal fin; gill rakers are counted on the first gill arch, those on the upper limb listed first; rudiments are included in the counts. Vanderhorstia lepidobucca, n. sp. Scalycheek Shrimpgoby (Figs 1-3; Table I) Holotype: MZB 21691, female, 40.3 mm SL, Aw Shucks dive site (1 30.013 N, 125 14.573 E), Lembeh Strait, North Sulawesi, Indonesia, silty sand bottom, 40 m, clove oil and hand net, M. V. Erdmann, 24 July 2013. Paratypes: MZB 21692, female, 1 specimen, 34.0 mm SL, collected with holotype; LBRC-F 3515, female, 1 specimen, 23.5 mm SL, collected with holotype; WAM P33894-009, 2 specimens, 28.7-29.7 mm SL, collected with holotype. WAM P.33940-001, 3 specimens, 30.7-37.6 mm SL, same data as holotype except collected on 8 September 2013. Diagnosis: Dorsal rays VI-I,13; anal rays I,14; pectoral rays 18; scales in longitudinal series 51-55; predorsal scales 17-18; 3 scales on upper anterior corner of operculum; cheek fully scaled; pre-pectoral scales present; scales ctenoid posteriorly, cycloid on head and anterodorsal portion of body; gill rakers 3-5 + 12-15; gill opening extending forward nearly to a vertical at posterior edge of orbit; dorsal fin relatively low, without elongate spines, longest spine 1.9-2.8 in HL; caudal fin long and pointed, 2.2-2.8 in SL; no sexual dichromatism; mid-dorsal, neon blue stripe on head; irregular orange bands on side of snout and cheek; prominent yellow and blue-margined black spot on middle of operculum; about 16-18 short brown bars along middle of side; first dorsal fin blue except pale yelaqua vol. 20 no. 2-15 April 2014 82
Gerald R. Allen, Teguh Peristiwady and Mark V. Erdmann low basally, row of large, irregular yellow spots (one between each spine) just below middle of fin, and tip of each spine yellow; second dorsal fin blue with pair of yellow stripes; anal fin mainly yellow except blue on distal third; caudal fin blue with large yellow spot medially at base from which pale yellow rays radiate posteriorly; pectoral fins with pair of large yellow spots at base and neon blue marking on uppermost part of base. Largest specimen, female holotype, 40.3 mm SL. Description: Dorsal rays VI-I,13; anal rays I,14; all dorsal and anal soft rays branched, last to base; pectoral rays 18, upper two and lower two unbranched; pelvic rays I,5, all soft rays branched, fifth rays joined medially with membrane; segmented caudal rays 17; branched caudal rays 14 (except one paratype with 15); upper and lower unsegmented caudal rays 5 and 4 respectively (5-7 and 4-6 respectively); longitudinal scale series 53 (51-55); transverse scale rows 10 (10-12) ; median predorsal scales 18 (17-18), median prepelvic scales 11 (11-12); circumpeduncular scales 14 (14-15); gill rakers 5 + 13 (3-5 + 12-15), total rakers 18 (16-18); pseudobranch with 7 (7-8) short fleshy lobes. Body elongate, depth at pelvic-fin origin 6.6 (5.7-6.2) in SL; body compressed, width at pectoral-fin origin 1.4 (1.4-1.7) in body depth; head length 4.4 (3.8-4.4) in SL; head compressed, width 1.4 (1.3-1.5) in body depth at pelvic origin; snout short, length 5.7 (5.0-6.7) in head length; orbit diameter 2.6(2.6-3.2) in head length; interorbital space narrow, least width 11.3 (4.6-6.0) in head length; caudal-peduncle depth 2.7 (2.1-2.9) in head length; caudal-peduncle length 1.6 (1.4-1.7) in head length. Mouth oblique, forming an angle of about 38 to horizontal axis of body, lower jaw projecting; mouth large, maxilla reaching vertical at level of middle of orbit, upper-jaw length 2.3 (2.2-2.7) in head length; upper jaw with 14-15 sender, slightly incurved teeth on each side, progressively larger towards front of jaw; also inner row of about 10 slender, teeth and pair of large, retrorse canines at front of upper jaw; front of lower jaw with 2-3 rows of incurved teeth nearly as large as those of upper jaw, ending in pair of large recurved, well-spaced canines about one-third back in jaw; remainder of Fig. 1. Lateral view of head of Vanderhorstia lepidobucca, paratype (MZB 21692), 34.0 mm SL, with sensory pores and main rows of papillae indicated by solid dots and broken lines respectively. Photo by G. R. Allen. 83 aqua vol. 20 no. 2-15 April 2014
Vanderhorstia lepidobucca, a new species of shrimpgoby from Sulawesi, Indonesia jaw with single row of small incurved teeth; no teeth on vomer or palatines; roof of mouth with prominent papillae; edge of lips smooth; tongue tip truncate to broadly triangular; no distinct mental flap. Gill opening broad, extending forward to vertical at posterior edge of orbit; gill membranes attached only anteriorly to isthmus, with no free fold; gill rakers slender, longest about equal in length to longest gill filaments of first gill arch. Posterior nostril a large, nearly round aperture in front of center of eye at fleshy edge of orbit; anterior nostril a short membranous tube, anteroventral to posterior nostril just above edge of upper lip. Cephalic pores and papilla rows (Fig. 1) well developed, consisting of: large pore (B ) immediately adjacent to each posterior nostril, two unpaired pores (C and D) at anterior and posterior interobital, a pore (E), behind rear, upper corner of orbit, a postocular pore (F) followed by pair of pores (G and H ) above preoperculum, pair of pores above operculum (K and L ), and three pores (M, N, and O ) along posterior margin of preopeculum. Scales on body progressively larger posteriorly, scale rows irregular, especially anteriorly; cheek fully covered with small scales; opercle naked except three scales at upper anterior corner; scales cycloid on head and anterodorsally on body, ctenoid posteriorly; scales ventrally on abdomen and chest cycloid; cycloid scales on pectoral-fin base; no scales on fins except for about three rows at base of caudal fin, smaller than last row on caudal peduncle. Origin of first dorsal fin above rear base of pelvic fins, predorsal length 3.5 (3.2-3.4) in SL; dorsal spines slender and flexible, none filamentous; first dorsal spine 3.4 (2.5-3.5) in HL; second to fifth dorsal spines subequal, longest, 2.2 (1.9-2.8) in HL; last membrane of first dorsal fin ending about one-scale width from origin of second dorsal fin; spine of second dorsal fin 2.8 (2.7-3.3) in HL; penultimate dorsal soft ray longest, 1.8 (1.5-2.0) in HL; origin of anal fin below base of first or second dorsal soft ray, preanal length 1.8 (1.8-1.9) in SL; anal spine 4.0 (3.3-4.6) in HL; penultimate anal soft ray longest, 1.9 (1.6-1.9) in HL; caudal fin long and pointed (lanceolate), 2.8 (2.2-2.8) in SL; pectoral fins pointed, eleventh or twelfth ray longest, reaching to slightly beyond level of pelvic fin tips, 5.4 (4.5-5.2) in SL; prepelvic length 3.6 (3.2-3.7) in SL; pelvic fins tips falling well short of Fig. 2. Underwater photograph of live Vanderhorstia lepidobucca, male paratype (WAM P.33940-001), 37.6 mm SL, Lembeh Strait, North Sulawesi, Indonesia. Photo by M. V. Erdmann. Fig. 3. Preserved paratype of Vanderhorstia lepidobucca, MZB 21692, female, 34.0 mm SL, Lembeh Strait, North Sulawesi, Indonesia. Photo by G. R. Allen. aqua vol. 20 no. 2-15 April 2014 84
Gerald R. Allen, Teguh Peristiwady and Mark V. Erdmann Table I. Proportional measurements of type specimens of Vanderhorstia lepidobucca expressed as percentage of the standard length. Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype MZB WAM WAM MZB WAM WAM WAM LBRC 21691 P.33940 P.33940 21692 P.33940 P.33894 P.33894 F3515 Sex female male female female male male male female Standard length (mm) 40.3 37.6 35.7 34.0 30.7 29.7 28.7 23.5 Body depth (P2 origin) 15.1 16.8 17.6 16.2 16.9 17.5 17.4 16.2 Body depth (A origin) 14.6 14.6 16.2 14.7 16.0 16.8 17.4 14.5 Body width 10.7 10.9 12.0 11.5 11.4 11.1 11.1 9.8 Head length 22.6 22.6 23.5 25.0 23.5 25.9 26.5 25.5 Head width 11.2 12.0 11.8 12.6 12.1 12.8 12.5 11.5 Snout length 4.0 4.5 4.2 3.8 3.9 4.4 4.2 3.8 Orbit diameter 8.7 7.2 8.1 8.2 9.1 9.8 8.4 9.8 Interorbital width 1.0 2.1 2.2 2.6 2.6 2.4 2.1 2.1 Cheek depth 6.7 7.2 6.4 6.5 7.5 6.7 6.6 6.8 Upper-jaw length 9.7 10.1 10.1 9.1 9.8 10.4 10.5 9.8 Caudal-peduncle depth 9.4 10.6 10.1 9.4 10.1 10.4 9.1 9.4 Caudal-peduncle length 15.4 16.2 15.7 15.9 16.6 18.5 16.0 15.7 Predorsal length 28.3 29.0 30.5 29.4 29.6 31.3 29.3 31.1 Preanal length 55.8 54.8 56.3 53.8 55.4 55.9 55.4 54.5 Prepelvic length 27.8 27.4 28.3 29.7 28.3 31.3 30.3 29.4 Base of dorsal fins 56.3 57.2 52.4 53.8 55.7 55.9 54.7 56.2 First dorsal spine 6.7 8.5 8.1 8.2 9.4 9.4 8.4 7.2 Third dorsal spine 10.2 10.4 10.6 11.2 12.1 11.8 11.1 11.5 Fifth dorsal spine 8.2 9.3 10.1 11.8 12.4 12.8 12.5 12.8 Spine of second dorsal fin 7.9 6.9 7.6 8.8 8.8 9.4 8.7 8.1 Longest dorsal ray 12.4 14.1 13.4 13.2 15.6 15.2 13.9 12.8 Base of anal fin 31.0 34.6 31.7 32.4 34.2 35.0 34.8 39.1 Anal spine 5.7 6.1 5.6 7.1 7.2 7.7 7.0 5.5 Longest anal ray 12.2 13.8 14.3 14.7 15.0 16.2 13.9 13.2 Caudal-fin length 35.5 36.2 38.9 40.9 37.8 45.5 38.3 40.0 Pectoral-fin length 18.4 19.1 19.9 20.3 21.5 21.9 20.6 22.1 Pelvic-spine length 6.7 6.9 6.4 7.1 6.8 7.4 7.3 7.2 Pelvic-fin length 15.6 19.1 17.9 18.2 18.6 18.5 20.2 19.1 anal fin origin, length of pelvic fin 6.4 (4.9-5.6) in SL; pelvic spine about one-third length of longest pelvic ray; pelvic frenum thin, membrane nearly reaching tip of pelvic spines. Colour in life (from underwater photographs including Fig. 2): head and body generally pale grey to pale tan; mid-dorsal stripe of neon blue from snout tip to dorsal fin origin; head with relatively broad, vermiculated yellow bands, including one from side of snout to anterior edge of eye, several behind eye and on cheek, and several others on side of nape; prominent black spot on middle of operculum, greater than pupil size, with inner margin of yellow anteriorly and dorsally, and outer margin of neon blue; neon blue markings also evident on cheek, preorbital, and upper lip; iris bright yellow except upper and lowermost portions neon blue; mid-lateral scale row of body with about 16-18 faint brown bars; uppermost scale row of back with small yellow dots and usually with blue stripe along base of second dorsal fin; first dorsal fin blue except pale yellow basally, row of large, irregular yellow spots (one between each spine) just below middle of fin, and tip of each spine yellow; second dorsal fin blue with pair of yellow stripes; anal fin mainly yellow except blue on distal third; caudal fin blue with large yellow spot medially at base from which pale yellow rays radiate posteriorly; pectoral fins with pair of large yellow spots at base and neon blue marking on uppermost part of base. Color in alcohol (Fig. 3): overall brown, darker on back, grading to white on ventral surface of head and abdomen; prominent black spot on middle portion of operculum, also apparent in gill cavity on inner surface of operculum; midlateral 85 aqua vol. 20 no. 2-15 April 2014
Vanderhorstia lepidobucca, a new species of shrimpgoby from Sulawesi, Indonesia row of 16-18 faint brown spots/blotches on body; fins translucent without markings. Remarks: The species is currently known only from the type locality at Lembeh Strait of far northern Sulawesi. The habitat consists of siltysand bottoms in sheltered water exposed to periodic strong currents at depths ranging from about 30-40 m. The fish live in burrows and presumably, like other members of the genus, are associated with snapping shrimps of the genus Alpheus, although we did not observe this behaviour. The new species differs from all other members of the genus, including the 15 species reported by Allen & Erdmann (2012) from the East Indian region in having a full complement of cycloid scales on the preoperculum as well as a small patch of three scales on the upper operculum. Both areas are entirely scaleless in other Vanderhorstia. In addition, the new species has well-developed median predorsal scales, a feature found only in V. attenuata Randall 2007a, V. hiramatsui Iwata et al. 2007, V. kizakura Iwata et al. 2007, V. macropteryx (Franz 1910), V. opercularis Randall 2007b, and V. papilio Shibukawa & Suzuki 2004. It also differs from other congeners in having a combination of 13 dorsal and 14 anal soft rays. Although anal fin-ray counts for the genus range from 11-18, the count of 14 rays, typical for V. lepidobucca, is unique for this genus. Finally, the colour pattern is highly distinctive, particularly the large yellow and blue margined black spot on the operculum. Dark opercular spots are also present in V. auronotata (Molucca Islands and North Sulawesi), V. opercularis (Red Sea), and V. papilio (Indonesia to Japan), but they are not as intense and lack the bright yellow and blue margins. The operculum of V. hiramatsui from southern Japan is largely dark brown, but is not in the shape of a spot, but rather part of a prominent bar extending from the nape to the throat. Although the last-mentioned four species all possess a dark opercular marking, they differ noticeably from V. lepidobucca in lacking scales on the head, and counts of soft dorsal and anal rays (respective counts of 14 and 15, 12 and 12, 11 and 11, and 12 and 11 for V. auronotata, V. opercularis, V. papilio, and V. hiramatsui). Etymology: The species is named lepidobucca (Latin: scaly-cheek), with reference to the unique head scalation. ACKNOWLEDGEMENTS We thank Renny Hadiaty (MZB), Sue Morrison (WAM), and the staff of the LIPI Bitung field station for curatorial assistance and providing museum registration numbers. We also thank Bruce Moore and the attentive dive staff at Black Sand Dive Retreat for hosting us and providing outstanding dive support for the photographic survey that led to the discovery of this new species. Finally, we thank Conservation International s Indonesia marine program for the financial support for this work. REFERENCES AKIHITO. 1984. Suborder Gobiodei. In: Fishes of the Japanese Archipelago (English text). (Eds. Masuda H., Amaoka, K., Araga, C., Uyeno, T. & T. Yoshino). Tokai University Press, Tokyo, pp 236-238. ALLEN, G. R. & ERDMANN, M. V. 2012. Reef fishes of the East Indies. Volumes I-III. Tropical Reef Research, Perth Australia. 1260 pp. ESCHMEYER, W. N. (ed). Catalog of fishes: genera, species, references. (http://research.calacademy.org/research/icht hyology/catalog/fishcatmain.asp). Electronic version accessed 20 December 2013. GREENFIELD, D. W. & LONGENECKER, K. R. 2005. Vanderhorstia bella, a new goby from Fiji (Teleostei: Gobiidae). Proceedings of the California Academy of Sciences, 56 (2): 619-623. IWATA, A., SHIBUKAWA, K. & OHNISHI, N. 2007. Three new species of the shrimp-associated goby genus Vanderhorstia (Perciformes: Gobiidae: Gobiinae) from Japan, with re-descriptions of two related congeners. Bulletin of the National Museum of Nature and Science (Ser. A) Supplement 1: 185-205. RANDALL, J. E. 2007a. Descriptions of four new shrimpgobies of the genus Vanderhorstia from the western Pacific. aqua, International Journal of Ichthyology 12 (3): 89-100. RANDALL, J. E. 2007b. Vanderhorstia opercularis, a new shrimp goby from the northern Red Sea. Electronic Journal of Ichthyology, Bulletin of the European Ichthyology Society 3 (1): 18-25. RANDALL, J. E. & MUNDAY, P. L. 2008. Vanderhorstia steelei, a new shrimpgoby from the Society Islands. aqua, International Journal of Ichthyology 14 (1): 35-41. SENOU, H., SUZUKI, T., SHIBUKAWA, K. & YANO, K. 2004. A Photographic Guide to the Gobioid Fishes of Japan. Heibonsha, Ltd., Tokyo. 534 pp. (in Japanese). SHIBUKAWA, K. & T. SUZUKI. 2004. Vanderhorstia papilio, a new shrimp-associated goby from the Ryukyu Islands, Japan (Perciformes: Gobiidae: Gobiinae), with comments on the limits of the genus. Ichthyological Research, 51: 113-119. WINTERBOTTOM, R., IWATA, A. & KOZAWA, T. 2005. Vanderhorstia nannai, a new species of burrow-associated goby from Palau and the Philippines (Pisces: Gobiidae). aqua, Journal of Ichthyology and Aquatic Biology 9 (3): 109-114. aqua vol. 20 no. 2-15 April 2014 86