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1 Zootaxa 3947 (2): Copyright 2015 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Description of a new genus of ponyfishes (Teleostei: Leiognathidae), with a review of the current generic-level composition of the family JOHN S. SPARKS 1,3 & PROSANTA CHAKRABARTY 2 1 American Museum of Natural History, Department of Ichthyology, Division of Vertebrate Zoology, Central Park West at 79th Street, New York, NY 10024, USA. jsparks@amnh.org. 2 Museum of Natural Science (Fish Section), Department of Biological Sciences, Louisiana State University, 119 Foster Hall, Baton Rouge, LA 70803, USA. prosanta@lsu.edu. 3 Corresponding author Abstract In order to recognize a monophyletic taxonomy for Leiognathidae based on unique features of the bacterially mediated light-organ system, we describe a new sexually-dimorphic genus of elongate, shallow-bodied ponyfishes within the tribe Equulitini, which itself is recovered within the subfamily Gazzinae. Photolateralis, new genus, is unique among ponyfishes in possessing a translucent mid-lateral flank stripe, which depending on the species, may be either a composite stripe comprised of numerous independent translucent windows (P. stercorarius and P. moretoniensis), or a continuous translucent lateral stripe (P. antongil). This translucent lateral stripe is either lacking entirely in females, or is considerably less well developed. In contrast, males in its sister taxon, Equulites, are characterized by the presence of an expansive, triangular, translucent lateral flank patch that also exhibits species-specific morphology. Internally, in Photolateralis the light organ system is characterized by a moderately enlarged, donut-shaped, and conspicuously spotted light organ in males that extends only slightly posteriorly into the gas bladder. In contrast, in members of Equulites the dorsolateral lobes of the light organ in males are greatly enlarged, heavily pigmented, and extend posteriorly well into the gas bladder. In addition, internally male members of Photolateralis exhibit lateral clearing of the silvery gas bladder lining posteriorly, corresponding in extent to the external translucent lateral stripe or windows on the flank. Including the new genus described herein, we now recognize 10 monophyletic genera of extant ponyfishes. Key words: bioluminescence, Equulitini, Gazzinae, light organ, new genus, Photolateralis, ponyfish Introduction Several recent family-level phylogenetic hypotheses have been generated for Leiognathidae, each building on the previous hypothesis of relationships as taxonomic sampling and molecular and morphological data collection has progressed. These phylogenetic hypotheses and faunal surveys have greatly increased our knowledge of leiognathid diversity, and have resolved both intrafamilial and intrageneric relationships (Sparks and Dunlap, 2004; Sparks et al., 2005; Chakrabarty et al., 2011a, 2011b). As a result, based on whether internal and external features of the bacterially-mediated light-organ system (LOS) are sexually dimorphic or not, we now recognize two monophyletic genera that are not sexually dimorphic, Aurigequula and Leiognathus, and the subfamily Gazzinae, comprising eight monophyletic genera whose members all exhibit a sexually-dimorphic LOS (Fig. 1). To achieve a monophyletic taxonomy for Leiognathidae in light of these new phylogenetic hypotheses, and as a result of extensive Indo-Pacific faunal surveys (e.g., Chakrabarty et al., 2009, 2010a) conducted by our group that have resulted in the discovery and description of several new species (Sparks, 2006a, 2006b; Sparks and Chakrabarty, 2007; Chakrabarty and Sparks, 2007; Chakrabarty et al., 2010b; Baldwin and Sparks, 2011), a number of new ponyfish genera have recently been described or resurrected from synonymy with the historically poorly diagnosed, catch-all genus Leiognathus. Herein we discuss the current generic-level composition of Leiognathidae and describe a new genus on the basis of apomorphic features of the sexually-dimorphic light-organ Accepted by M.R. de Carvalho: 20 Mar. 2015; published: 15 Apr

2 system. Including the new genus described herein, we now recognize 10 monophyletic genera of extant ponyfishes on the basis of unique anatomical features of the light-organ system, internal and external morphology, and molecular synapomorphies (Fig. 1; Sparks et al., 2005; Chakrabarty et al., 2011b). FIGURE 1. Generic level phylogeny for Leiognathidae after Chakrabarty et al. (2011b) indicating placement of Photolateralis, new genus (in boldface type). Materials and methods Morphology of the light-organ system was examined in situ or in isolation via dissection to permit detailed anatomical comparisons. Osteological and soft tissue features were analyzed using standard film and digital radiographs, high-resolution digital images, examination of whole alcohol-preserved specimens, specimens cleared-and-stained (C&S) for bone and cartilage (following the protocol of Taylor & Van Dyke, 1985), and dry skeletal (S) preparations. Specimens examined are listed under Comparative material. Institutional abbreviations are as listed in Leviton et al. (1985) and Sabaj Pérez (2012). Photolateralis, new genus (Figures 2 4A,B) Photoplagios in part: Sparks et al., 2005; Sparks, 2006b; Sparks and Chakrabarty, 2007 Equulites in part: Kimura et al., 2008a; Chakrabarty et al., 2011a, 2011b Type species: Photolateralis stercorarius (Evermann & Seale, 1907) 182 Zootaxa 3947 (2) 2015 Magnolia Press SPARKS & CHAKRABARTY

3 FIGURE 2. Internal and external features of light-organ system (LOS) in members of Photolateralis. (A) Male P. stercorarius external LOS anatomy. (B) Male P. stercorarius internal LOS anatomy. (C) Female P. stercorarius internal LOS anatomy. Figure abbreviations: es = esophagus, g = gut, gb = gas bladder, lo = circumesophageal light organ, mls = translucent midlateral stripe. Stippled region on posterolateral region of gas bladder corresponds to clearing of silvery guanine lining in region internal to translucent mid-lateral stripe. Yellow region on light organ corresponds to window that is directed and opens into the gas bladder. Other included species: Photolateralis moretoniensis (Ogilby, 1912), P. antongil (Sparks, 2006) Diagnosis. Members of Photolateralis are distinguished from all other leiognathids by the presence of a translucent flank stripe, which depending on the species, may be comprised of either a continuous mid-lateral stripe (P. antongil), or a composite stripe comprised of numerous independent translucent windows (P. stercorarius and P. NEW GENUS OF PONYFISHES Zootaxa 3947 (2) 2015 Magnolia Press 183

4 moretoniensis). In Photolateralis, the translucent lateral stripe is either lacking entirely in females, or is considerably less developed. Internally and externally members of Photolateralis can be distinguished from their sister genus, Equulites, by morphology of the light-organ system. Externally, male members of Equulites are characterized by an expansive translucent triangular, cornucopia-shaped, or trapezoidal patch on the flank, in contrast to a continuous or composite mid-lateral stripe comprised of numerous independent translucent windows in Photolateralis. Although both genera share lateral clearing of the guanine-lined internal gas bladder surface corresponding to the location of the translucent external patch or stripe, clearing in Equulites is more extensive, extending the full length of the gas bladder, whereas clearing in Photolateralis is restricted to the posterior region of the gas bladder. The light organ in male members of Equulites is greatly enlarged, with paired dorsolateral lobes extending posteriorly well into the gas bladder (less E. leuciscus), in contrast to a moderately enlarged donutshaped light organ in male members of Photolateralis that extends at most only slightly posteriorly into the gas bladder. FIGURE 3. Species of Photolateralis illustrating phenotypic variability of translucent mid-lateral stripe characteristic of genus. (A) P. stercorarius, USNM 55906, holotype, 83.8mm SL, Philippines. (B) P. moretoniensis, AMS I , non-type, 76.9 mm SL, Australia. (C) P. antongil, AMNH , holotype, 75.0 mm SL, Madagascar. 184 Zootaxa 3947 (2) 2015 Magnolia Press SPARKS & CHAKRABARTY

5 FIGURE 4. Comparison of internal and external anatomy of LOS in sister genera Equulites and Photolateralis, new genus: (A) Translucent lateral flank stripe comprised of discrete windows in male Photolateralis stercorarius (SIO 96-96, 98.2mm SL, Philippines). (B) Internal anatomy of LOS in male P. stercorarius (AMNH uncat., [VN ], 73.1 mm SL, Vietnam). Note comparatively small donut shaped, spotted light organ and posterior lateral clearing of silvery gas bladder lining. (C) Expansive translucent, triangular lateral flank patch characteristic of male members of Equulites (E. elongatus, UMMZ uncat., [PVD 8206/19a LE], 76.4 mm SL, Japan). (D) Internal anatomy of LOS in male E. elongatus (UMMZ uncat., 68.0 mm SL, Japan). Note greatly enlarged dorsolateral light-organ lobes in Equulites that extend posteriorly well into gas bladder, and lateral clearing of silvery guanine layer running entire length of gas bladder. Light organs circled in yellow. Figure abbreviations: gb = gas bladder, mls = translucent mid-lateral stripe, tp = translucent, triangular lateral flank patch. NEW GENUS OF PONYFISHES Zootaxa 3947 (2) 2015 Magnolia Press 185

6 Discussion and comparisons. Aside from features of the light-organ system (LOS), leiognathids are, in general, morphologically conservative and non-descript silvery fishes with an external appearance that superficially resembles mojarras (Gerreidae), with which they are often confused. However, all leiognathids possess a circumesophageal light organ that contains symbiotic luminescent bacteria (Photobacterium), whose luminescence the fish co-opt for photic communication and predator avoidance (McFall-Ngai and Dunlap, 1983), whereas gerreids are not bioluminescent. In a majority of leiognathid species, comprising all members of the subfamily Gazzinae (Fig. 1), the light organ and associated internal and external anatomical features (e.g., translucent windows/stripes and gas bladder clearing) are sexually dimorphic and both intra- and intergenerically variable in morphology (Sparks et al., 2005: Figs. 4 8). This integrated LOS provides a rich set of phylogenetically informative characters for diagnosing both genera and species (Fig. 1). The unique anatomical features of the LOS facilitate the transmission of bacterially-generated luminescence (Sparks et al., 2005; Chakrabarty et al., 2011a) via a species-specific translucent lateral patch (or patches) in males of most species that is located in the head region or on the flank (Sparks et al., 2005). All members of Photolateralis were formerly placed in Equulites (Kimura et al., 2008; Chakrabarty et al., 2011a,b). Photolateralis shares the presence of translucent lateral flank regions with its sister genus Equulites (Figs. 2 4). In Photolateralis, however, these translucent areas form a mid-lateral stripe, either continuous or comprised of discrete windows (Figs 2 and 3), whereas in Equulites, comprising E. elongatus, E. rivulatus, E. leuciscus, E. klunzingeri, E. laterofenestra, and E. absconditus, males are characterized by the presence of a species-specific triangular, translucent lateral flank patch (Fig. 4; Sparks and Chakrabarty, 2007: Fig. 1). Although Equula lineolata Valenciennes, in Cuvier and Valenciennes, 1835 is placed within Equulites (Eschmeyer, 2015), with the new placement attributed to Kimura et al. (2008a), in fact Kimura et al. (2008a) never mention Equula lineolata. Sparks (2006b) discussed the taxonomic status of Equula lineolata and concluded the name was a nomen dubium of uncertain placement beyond the family level. In both Photolateralis and Equulites, the lateral gas bladder lining exhibits clearing of the silvery guanine layer that corresponds specifically to the external translucent region characteristic of that taxon (Figs. 2 and 4; Sparks et al., 2005). Males of both genera also possess enlarged light organs (moderately so in Photolateralis vs. greatly enlarged in Equulites) compared to conspecific females and in which the dorsal lobes extend posteriorly into the gas bladder (Figs. 2 and 4). However, both the degree of enlargement and posterior extension of the light organ are considerably muted in Photolateralis (Figs. 2B and 4B) as compared to Equulites (Fig. 4D). The light organ in male members of Photolateralis is more or less donut shaped (Figs. 2B and 4B), lacking the greatly enlarged dorsolateral extensions characteristic of male members of Equulites (Fig. 4D). In addition to Photolateralis, new genus, we currently recognize the following nine extant leiognathid genera, in chronological order of their description, whose phylogenetic relationships are shown in Figure 1 (after Chakrabarty et al., 2011b): 1) Leiognathus Lacepède, Type species: Leiognathus argenteus Lacepède, 1802, by monotypy. Equula Cuvier, 1815 was resurrected from synonymy with Leiognathus Lacepède, 1802 by Chakrabarty and Sparks (2008) to encompass E. fasciata and E. longispinis. Chakrabarty and Sparks (2008:5) treated Equula as valid, however, they incorrectly assigned the wrong type species to Leiognathus. The correct type species is Leiognathus argenteus Lacepède 1802 (see Sparks and Dunlap, 2004; Eschmeyer, 2015). As the same specimen was used to describe the type species of both Equula and Leiognathus, these genera are objective synonyms and identical. The genus Equula is therefore a synonym of Leiognathus (corrected in Chakrabarty et al., 2009). Leiognathus is recovered as the sister group to the remaining members of Leiognathidae less Aurigequula. Members of Leiognathus, comprising Leiognathus equula and L. robustus, plus a number of undescribed species (Chakrabarty and Sparks, 2008; Chakrabarty et al., 2011b), are not sexually dimorphic with regard to features of the light-organ system. 2) Gazza Rüppell Type species: Gazza equulaeformis Rüppell 1835, by monotypy. Gazza is recovered as the sister taxon to Deveximentum and placed within the tribe Gazzini, subfamily Gazzinae. 3) Deveximentum Fowler, Type species: Zeus insidiator Bloch 1787, by original designation. The species now placed in this genus were formerly placed in Secutor Gistel, Kottelat (2013) considered Secutor a replacement name for Equula Cuvier 1815, which is itself a synonym of Leiognathus Lacepède, 1802 (see above for Leiognathus). Deveximentum is recovered as the sister taxon to Gazza and placed within the tribe Gazzini, subfamily Gazzinae. 4) Equulites Fowler, Type species: Leiognathus vermiculatus Fowler 1904, by original designation. 186 Zootaxa 3947 (2) 2015 Magnolia Press SPARKS & CHAKRABARTY

7 Equulites was described as a subgenus of Leiognathus. Equulites was removed from synonymy with Leiognathus and elevated to generic rank by Chakrabarty and Sparks (2008). Equulites currently comprises E. leuciscus, E. klunzingeri, E. elongatus, E. rivulatus, E. absconditus, and E. laterofenestra. Photoplagios was described by Sparks et al. (2005), but is a synonym of Equulites Fowler 1904, given that Equulites, an older name, was available for one of the species included in Photoplagios (Kimura et al., 2008a; Chakrabarty and Sparks, 2008). Equulites is recovered as the sister taxon to Photolateralis, new genus, and placed within the tribe Equulitini, subfamily Gazzinae. 5) Eubleekeria Fowler, Type species: Equula splendens Cuvier 1829, by original designation. Eubleekeria was described as a subgenus of Leiognathus. Eubleekeria was removed from synonymy with Leiognathus and elevated to generic rank by Chakrabarty and Sparks (2008). Eubleekeria encompasses the Leiognathus splendens species complex (E. splendens, E. jonesi, and E. kupanensis; Kimura et al., 2005), and also includes multiple undescribed species. Eubleekeria is recovered as the sister group to Photopectoralis and placed within the tribe Eubleekerini, subfamily Gazzinae. 6) Aurigequula Fowler, Type species: Clupea fasciata Lacepède, 1803, by original designation. Aurigequula was described as a subgenus of Leiognathus. Aurigequula was mistakenly placed in Equula by Chakrabarty and Sparks, (2008). Aurigequula was resurrected from synonymy with Leiognathus Lacepède, 1802 by Chakrabarty et al. (2009) to replace a previously established invalid name, and comprises A. fasciata (type species), A. longispina, and A. striata, plus a number of undescribed species. For additional comments regarding generic status see heading for Leiognathus above and Chakrabarty et al. (2009). Aurigequula is recovered as the sister group to all other members of Leiognathidae. Like Leiognathus, members of Aurigequula do not appear to be sexually dimorphic with regard to features of the light-organ system (Sparks and Dunlap, 2004). 7) Nuchequula Whitley, Type species: Equula blochii Valenciennes, in Cuvier and Valenciennes, 1835, by original designation. Nuchequula was described as a subgenus of Eubleekeria. The subgenus Nuchequula was elevated to generic rank and revised by Chakrabarty and Sparks (2007). Nuchequula was also considered to warrant generic rank and was subsequently revised again the following year by Kimura et al. (2008b). Nuchequula is recovered as the sister taxon to Karalla and placed within the tribe Nuchequulini, subfamily Gazzinae. 8) Photopectoralis Sparks, Dunlap, and Smith, Type species: Leiognathus aureus Abe and Haneda, 1972, by original designation. Photopectoralis currently comprises P. aureus, P. bindus, P. hataii, and P. panayensis, plus at least two undescribed species. Photopectoralis is recovered as the sister group to Eubleekeria and placed within the tribe Eubleekerini, subfamily Gazzinae. 9) Karalla Chakrabarty and Sparks, Type species: Equula daura Cuvier 1829, by original designation. Karalla currently comprises K. daura and K. dussumieri. Karalla is recovered as the sister taxon to Nuchequula and placed within the tribe Nuchequulini, subfamily Gazzinae. Comparative material Aurigequula fasciata: AMNH 15520; AMNH ; AMNH ; CAS 1872; UMMZ ; AMNH ; AMNH uncat.; USNM ; USNM ; WHT PC-195; WHT PC-196; WHT PC-197. Aurigequula longispina: MNHN A-0579, holotype; AMNH ; AMNH ; AMNH uncat. (SI-87); AMS I ; AMS I ; AMS ; AMS ; USNM ; ZRC Aurigequula striata: SU 22909; WHT PC-183; WHT PC-184; WHT PC-192; WHT PC-193; WHT PC-194. Deveximentum hanedai: FUMT P 8701, paratype, 1 ex. Deveximentum indicium: AMNH ; AMNH ; AMNH ; AMNH ; ANSP 76147; ANSP 87160; UMMZ ; USNM ; USNM Deveximentum insidiator: ZMB 1676, syntypes, 2 ex.; AMNH ; ANSP 51324; ANSP 62128; ANSP 87838; CAS 29894; UMMZ uncat. Deveximentum mazavasaoka: AMNH , holotype; AMNH , paratype, 1 ex., AMNH , paratype, 1 ex., AMNH , paratypes, 43 ex., AMNH , paratypes, 12 ex., AMNH , paratype, 1 ex., AMNH , paratypes, 10 ex., AMNH , paratype, 1 ex., AMNH , paratypes, 5 ex., ANSP 53115, paratype, 1 ex., ANSP 54891, paratypes, 5 ex., ANSP 88786, paratype, 1 ex., USNM , paratypes, 7 ex., USNM , paratypes, 17 ex. Deveximentum megalolepis: UMMZ Deveximentum ruconium: AMNH ; AMNH ; AMNH ; ANSP 77437; ANSP 87141; CAS-SU 29895; UMMZ ; UMMZ uncat.; USNM NEW GENUS OF PONYFISHES Zootaxa 3947 (2) 2015 Magnolia Press 187

8 Equulites absconditus: AMNH , holotype; AMNH , paratype, 1 ex.; AMNH , paratypes, 97 ex.; LSUMZ 13236, paratype, 1ex.; LSUMZ 13339, paratypes, 21 ex.; NTUM 5698, paratypes, 2ex. Equulites elongatus: BMNH , holotype; CAS 52602; LACM ; LACM ; SIO 83-55; USNM 55613; UMMZ ; UMMZ ; UMMZ uncat. Equulites klunzingeri: NMW 68277, syntypes, 4 ex.; NMW 68280, syntypes, 2 ex.; NMW 76008, syntypes, 4 ex.; NMW 76009, syntypes, 4 ex.; AMNH 44488; AMNH 44491; AMNH Equulites laterofenestra: USNM , holotype; USNM , paratypes, 4 ex.; AMNH , paratypes, 2 ex.; AMNH ; AMNH ; AMNH ; AMNH ; WHT PC-132; WHT PC-133; WHT PC-188. Equulites leuciscus: BMNH , holotype; AMNH ; AMNH ; AMNH ; AMNH ; AMNH ; AMNH ; AMNH ; AMS I ; AMS I ; AMS I ; ANSP (holotype of Leiognathus vermiculatus); UMMZ ; UMMZ uncat.; USNM 76609; USNM ; USNM ; USNM ; USNM ; QM I.878 (holotype of Equula longispina). Equulites rivulatus: AMNH 34850; UMMZ ; UMMZ ; UMMZ uncat. Eubleekeria jonesi: ZSI (CMFRI.T 100/1), uncatalogued holotype; AMNH ; AMNH ; AMNH ; UMMZ ; UMMZ ; UMMZ ; UMMZ uncat. Eubleekeria kupanensis: UMMZ Eubleekeria splendens: AMNH ; ANSP 59963; ANSP 27530; ANSP (holotype of Leiognathus philippinus); ANSP 47487; CAS 1485; CAS 38789; CAS 56438; CAS 56441; MNHN A- 6724; UMMZ ; UMMZ , UMMZ uncat.; USNM ; USNM ; WHT PC-136; WHT PC-137; WHT PC-166; WHT PC-191. Gazza achlamys: CAS-SU 21652, paratype, 1 ex.; CAS-SU 22853, paratype, 1 ex.; UMMZ ; UMMZ ; UMMZ Gazza dentex: MNHN A-578, lectotype. Gazza minuta: AMNH ; AMNH ; UMMZ ; UMMZ ; UMMZ ; UMMZ ; UMMZ uncat. Gazza rhombea: USNM , paratype, 1 ex.; USNM , paratype, 1 ex. Gazza squamiventralis: USNM , holotype; USNM , paratype, 1 ex.; AMNH Karalla daura: AMNH ; ANSP 89582; USNM ; USNM ; WHT PC-131; WHT PC-170; WHT PC-187. Karalla dussumieri: MNHN A-6721, syntype, 1 ex.; AMNH ; WHT PC-130; WHT PC- 134; WHT PC-135; WHT PC-161; WHT PC-162; WHT PC-190. Leiognathus equulus: ZMUC P48219, lectotype (dry skin; photographs and radiographs examined); ZMUC P48220, paralectotype (dry skin, photograph and radiograph examined); ZMB 8756 (holotype of Leiognathus edentulus); AMNH 59535; AMNH 88039; AMNH ; AMNH ; CAS 57306; CAS-SU 35627; CAS-SU 38781; MNHN A-6723; UMMZ ; UMMZ ; UMMZ (in part); UMMZ ; UMMZ ; UMMZ ; UMMZ uncat. Leiognathus robustus: UMMZ , holotype; AMNH , 1 ex., paratype; UMMZ , 1 ex, paratype.; UMMZ Nuchequula blochii: MNHN A-6757, syntype, 1 ex.; MNHN A-6759, syntype, 1 ex. Nuchequula decora: AMNH , AMS I Nuchequula mannusella: AMNH , holotype; AMNH , paratypes, 15 ex.; AMNH , paratypes; 16 ex.; AMNH , paratypes, 15 ex.; AMNH , paratypes, 14 ex.; AMNH , paratypes; 20 ex.; AMNH , paratypes, 9 ex.; AMNH , paratypes, 13 ex.; AMNH , paratypes, 1 ex.; AMNH , paratype,1 ex.; AMNH , paratypes; 1 ex.; AMNH , paratype,1 ex.; AMNH , paratypes, 29 ex.; ASIZP , paratype, 1 ex.; ASIZP , paratype; 1 ex.; ASIZP , 2 ex.; ASIZP Nuchequula nuchalis: RMNH 1287, holotype; AMNH 26819; AMNH 34861, AMNH 26819; AMNH ; CAS-SU 4757; UMMZ Nuchequula pan: CUMZ , holotype; CUMZ , paratype, 1 ex.; CUMZ , paratype, 1 ex.; USNM , paratype, 1 ex. Photolateralis antongil: AMNH , holotype; AMNH , paratypes, 3 ex. Photolateralis moretoniensis: QM I.1583, syntype, 1 ex.; AMS I ; AMS I ; LSUMZ 13646, LSUMZ 13647, LSUMZ 13648, LSUMZ 13656, LSUMZ 13657, LSUMZ Photolateralis stercorarius: USNM 55906, holotype; USNM , cotype; ANSP 33289, paratype, 1 ex.; CAS 42171, paratype; CAS 17678; CAS-SU 20004, paratype; AMNH uncat.; SIO 00-13; SIO 96-96; UMMZ ; UMMZ uncat.; USNM ; ZRC Photopectoralis aureus: SIO ; UMMZ ; UMMZ ; UMMZ uncat.; USNM Photopectoralis bindus: AMS I , CAS 51097; UMMZ ; UMMZ ; UMMZ uncat.; USNM ; MNHN A-6763 (syntype of Equula brevirostris). Photopectoralis cf. bindus: AMNH 188 Zootaxa 3947 (2) 2015 Magnolia Press SPARKS & CHAKRABARTY

9 Photopectoralis hataii: UMMZ uncat. Photopectoralis cf. hataii: AMNH Photopectoralis panayensis: UMMZ , holotype; UMMZ , paratypes, 4 ex.; UMMZ , paratypes, 5 ex.; UMMZ , paratypes, 8 ex.; UMMZ , paratypes, 16 ex.; UMMZ ; UMMZ uncat. Acknowledgments For the loan of specimens in their care, we are grateful to L. Parenti, E. Wilbur, and J. Williams (USNM), J. Leis, M. McGrouther, and T. Trnski (AMS), J. Johnson (QM), E. Mikschi (NMW), W. Fink, G. Smith, and D. Nelson (UMMZ), P. Dunlap (UM), M. Sabaj and J. Lundberg (ANSP), H. Walker and P. Hastings (SIO), M. van Oijen (RMNH), and D. Catania (CAS). Thanks to B. Andriamihaja (MICET, Madagascar), D. Meyers (WCS, Madagascar), N. Raminosoa (University of Antananarivo, Madagascar), H.C. Ho (NMMBA, Taiwan) and H. Mok (NSYSU, Taiwan), N. Van Quan (IMER, Vietnam), H.H. Ng and P. Ng (RMBR, Singapore), K. Tang (UMF), W.L Smith (KU), and R. Schelly (AMNH) for their hospitality and considerable assistance in the field. Thanks to B. Brown, R. Schelly, and R. Arrindell (AMNH) for assistance with loans and radiographs. Fishes were collected in accordance with AMNH IACUC guidelines. This work was supported by grants from the National Science Foundation (DEB , IOS , DEB ) and the Niarchos Expedition Fund of the American Museum of Natural History to JSS. References Abe, T. & Haneda, Y. (1972) Descriptions of two new species of the ponyfish genus Leiognathus from Indonesia. Science Report of the Yokosuka City Museum, 19, 1 7. Baldwin, Z.H., & Sparks, J.S. (2011) A new species of Secutor (Teleostei: Leiognathidae) from the Western Indian Ocean. Zootaxa, 2998, Bloch, M.E. (1787) Naturgeschichte der ausländischen Fische. Berlin. v. 3, i xii [Plates ] Chakrabarty, P. & Sparks, J.S. (2007) Phylogeny and taxonomic revision of Nuchequula Whitley 1932 (Teleostei: Leiognathidae), with the description of a new species. American Museum Novitates, 3588, Chakrabarty, P. & Sparks, J.S. (2008) Diagnoses for Leiognathus Lacepède 1802, Equula Cuvier 1815, Equulites Fowler 1904, Eubleekeria Fowler 1904, and a new ponyfish genus (Teleostei: Leiognathidae). American Museum Novitates, 3623, Chakrabarty, P., Amarasinghe, T. & Sparks, J.S. (2009) Redescription of ponyfishes (Teleostei: Leiognathidae) of Sri Lanka and the status of Aurigequula Fowler, Ceylon Journal of Science (Biological Sciences), 37 (2), Chakrabarty, P., Sparks, J.S. & Ho, H.-C. (2010a) Taxonomic review of the ponyfishes (Perciformes: Leiognathidae) of Taiwan. Marine Biodiversity, 40, Chakrabarty, P., Chu, J., Nahar, L., Baldwin, Z.H. & Sparks, J.S. (2010b) Geometric morphometrics reveals a new species of ponyfish (Teleostei: Leiognathidae: Equulites), with comments on the taxonomic status of Equula berbis Valenciennes. Zootaxa, 2427, Chakrabarty, P., Davis, M.P., Smith, W.L., Berquist, R., Gledhill, K., Frank, L. & Sparks, J.S. (2011a) Evolution of the light organ system in ponyfishes (Teleostei: Leiognathidae). Journal of Morphology, 272, Chakrabarty, P., Davis, M.P., Smith, W.L., Baldwin, Z.H. & Sparks, J.S. (2011b) Is sexual selection driving diversification of the bioluminescent ponyfishes (Teleostei: Leiognathidae)? Molecular Ecology, 20, Cuvier, G. (1815) Observations et recherches critiques sur différents poissons de la Méditerrannée, et à leur occasion sur des poissons d'autres mers, plus ou moins liés avec eux. Memoirs du Museum National d'histoire Naturelle. v. 1, , , , [Plates. 16, 23] Cuvier, G. (1829) Le Règne Animal, distribué d'après son organisation, pour servir de base à l'histoire naturelle des animaux et d'introduction à l'anatomie comparée. Edition 2. Règne Animal 2, i xv Cuvier, G. & Valenciennes, A. (1835) Histoire naturelle des poissons. Tome dixième. Suite du livre neuvième. Scombéroïdes. Livre dixième. De la famille des Teuthyes. Livre onzième. De la famille des Taenioïdes. Livre douzième. Des Athérines. Histoire naturelle des poissons. v. 10, i xxiv pp. [Plates ] Eschmeyer, W. (2015) Catalogue of fishes. Available from: fishcatsearch.html (Accessed 24 February 2015) NEW GENUS OF PONYFISHES Zootaxa 3947 (2) 2015 Magnolia Press 189

10 Evermann, B.W., & Seale, A. (1907) Fishes of the Philippine Islands. Bulletin of the United States Bureau of Fisheries, 26 (1906), Fowler, H.W. (1904) A collection of fishes from Sumatra. Journal of the Academy of Natural Sciences, Philadelphia, 12 (pt 4), [Plates. 7 28] Fowler, H.W. (1918) New and little known fishes from the Philippine Islands. Proceedings of the Academy of Natural Sciences of Philadelphia, 70, Gistel, J. (1848) Naturgeschichte des Thierreichs, für höhere Schulen. Stuttgart, i xvi, [Plates. 1 32] Kimura, S., Ito, T., Peristiwady, T., Iwatsuki, Y., Yoshino, T. & Dunlap, P.V. (2005) The Leiognathus splendens complex (Perciformes: Leiognathidae) with the description of a new species, Leiognathus kupanensis Kimura and Peristiwady. Ichthyological Research, 52, Kimura, S., Motomura, H. & Iwatsuki, Y. (2008a) Equulites Fowler 1904, a senior synonym of Photoplagios Sparks, Dunlap, and Smith 2005 (Perciformes: Leiognathidae). Ichthyological Research, 55, Kimura, S., Kimura, R. & Ikejima, K. (2008b) Revision of the genus Nuchequula with descriptions of three new species (Perciformes: Leiognathidae). Ichthyological Research, 55, Kottelat, M. (2013) The fishes of the inland waters of Southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. The Raffles Bulletin of Zoology, Supplement 27, Lacepède, B.G.E. (1802) Histoire naturelle des poissons 4. Paris: Chez Plassan, Imprimeur-Libraire. Lacepède, B.G.E. (1803) Histoire naturelle des poissons. Histoire naturelle des poissons 5. Paris: Chez Plassan, Imprimeur- Libraire. Leviton, A.E., Gibbs, Jr., R.H., Heal, E. & Dawson, C.E. (1985) Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985, McFall-Ngai, M.J. & Dunlap, P.V. (1983) Three new modes of luminescence in the leiognathid fish Gazza minuta: discrete projected luminescence, ventral body flash, and buccal luminescence. Marine Biology, 73, Ogilby, J.D. (1912) On some Queensland fishes. Memoirs of the Queensland Museum, 1, Rüppell, W.P.E.S. (1835) Neue Wirbelthiere zu der Fauna von Abyssinien gehörig. Fische des Rothen Meeres. Siegmund Schmerber, Frankfurt am Main, i ii [Plates. 1 7] Sabaj Perez, M.H. (editor) (2014) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 5.0 (22 September 2014). Electronically. Available from: American Society of Ichthyologists and Herpetologists, Washington, DC. Sparks, J.S. (2006a) Leiognathus longispinis (Valenciennes, in Cuvier and Valenciennes, 1835), a senior synonym of Leiognathus smithursti (Ramsay and Ogilby, 1886) (Teleostei: Leiognathidae). Copeia, 2006, Sparks, J.S. (2006b) A new species of ponyfish (Teleostei: Leiognathidae: Photoplagios) from Madagascar, with a phylogeny for Photoplagios and comments on the status of Equula lineolata Valenciennes. American Museum Novitates, 3526, Sparks, J.S. & Dunlap, P.V. (2004) A clade of non-sexually dimorphic ponyfishes (Teleostei: Perciformes: Leiognathidae): phylogeny, taxonomy, and description of a new species. American Museum Novitates, 3459, Sparks, J.S. & Chakrabarty, P. (2007) A new species of ponyfish (Teleostei: Leiognathidae: Photoplagios) from the Philippines. Copeia, 2007, Sparks, J.S., Dunlap, P.V. & Smith, W.L. (2005) Evolution and diversification of a sexually dimorphic luminescent system in ponyfishes (Teleostei: Leiognathidae), including diagnoses for two new genera. Cladistics, 21, Taylor, W.R. & Van Dyke, G.C. (1985) Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium, 9, Valenciennes, A. (1835) Des Equula. In: Cuvier, G. & Valenciennes, A. (Eds.), Histoire naturelle des poissons. Tome dixième, Paris: Chez F.G. Levrault, pp [plates ] Whitley, G.P. (1932) Some fishes of the family Leiognathidae. Memoirs of the Queensland Museum, 10, Zootaxa 3947 (2) 2015 Magnolia Press SPARKS & CHAKRABARTY