A New Species of Ponyfish (Teleostei: Leiognathidae: Photoplagios)
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PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3526, 20 pp., 7 figures, 2 tables September 8, 2006 A New Species of Ponyfish (Teleostei: Leiognathidae: Photoplagios) from Madagascar, with a Phylogeny for Photoplagios and Comments on the Status of Equula lineolata Valenciennes JOHN S. SPARKS ABSTRACT A new species of ponyfish in the genus Photoplagios is described from material collected in coastal waters of northeastern Madagascar. Photoplagios antongil, new species, is distinguished from congeners by the presence of a broad midlateral stripe and two darkly pigmented flank patches located ventral to the lateral midline, which are presumably translucent in life but darkly pigmented in preservative due to a concentration of melanophores. The new species is further distinguished from P. leuciscus, the only externally similar species occurring in the region, by the absence of a large translucent triangular patch on the flanks, a much shorter second dorsal-fin spine, a straight predorsal profile, pigmentation pattern on the upper flanks, absence of black pigment in the pectoral-fin axil, and exposed conical oral dentition in two distinct rows. A phylogeny for Photoplagios is provided based on the simultaneous analysis of anatomical features of the light-organ system and nucleotide characters. The taxonomic statusofEquula lineolata Valenciennes, in Cuvier and Valenciennes, 1835 is discussed, and the species is herein concluded to be a nomen dubium of uncertain placement beyond the family level. INTRODUCTION olatus (Valenciennes, in Cuvier and Valen- ciennes, 1835), P. moretoniensis (Ogilby, Photoplagios Sparks, Dunlap, and Smith, 1912), P. rivulatus (Temminck and Schlegel, 2005 comprises eight species: P. elongatus 1845), P. stercorarius (Evermann and Seale, (Gu¨nther, 1874), P. klunzingeri (Steindachner, 1907), and a new species from the coastal 1898), P. leuciscus (Gu¨nther, 1860), P. line- waters of Madagascar described herein. 1 Division of Vertebrate Zoology (Ichthyology), American Museum of Natural History ([email protected]). Copyright E American Museum of Natural History 2006 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3526 Although P. klunzingeri and P. moretoniensis Discussion); each of their putative ranges is were not included in the family-level phyloge- nearly as great as that of the genus in its netic analysis of Sparks et al. (2005), they entirety. As Gill and Kemp (2002) stressed, possess the following apomorphic features of however, putatively wide distributional ranges the light-organ system (LOS) that diagnose for many Indo-Pacific shore-fishes may simply the genus. Members of Photoplagios are reflect poor taxonomic understanding. This strongly sexually dimorphic, and males are claim is corroborated for ponyfishes in general distinguished from all other members of by a recent study using nucleotide characters Leiognathidae by the presence of an expansive to resolve phylogenetic intrarelationships translucent lateral flank patch or stripe, dorso- (Sparks et al., 2005); the study indicated that lateral lobes of the light organ that are many putatively widespread ponyfish species hypertrophied and extend posteriorly into the (e.g., Leiognathus equulus, L. fasciatus, gas bladder (extensively in P. elongatus and P. Photoplagios leuciscus, Photopectoralis bindus, rivulatus, less so in the remaining congeners), and Gazza minuta) may actually comprise and lateral clearing of the silvery lining of species assemblages. the gas bladder, which is directly correlated Luminescence in leiognathids is produced with species-specific transparent lateral flank by large numbers of the symbiotic luminous patches or stripes (Sparks et al., 2005: figs. bacterium Photobacterium leiognathi, which 3–5). are harbored in an internal circumesophageal A comprehensive phylogenetic and taxo- light organ (Boisvert et al., 1967; Hastings and nomic study of Leiognathidae is needed, Mitchell, 1971; Bassot, 1975; Reichelt et al., including detailed comparisons of the LOS, 1977; Dunlap, 1984). Collectively with the to establish the generic limits of Photoplagios light organ, the leiognathid LOS comprises and the other genera. For example, based on reflectors and chromatophore-embedded remarks in the original description, which light-organ shutters, reflective and translucent refer to an elongate body and possession of a tissues of the gas bladder, and translucent midlateral stripe, Equula berbis Valenciennes, bone, musculature, and skin (Sparks et al., in Cuvier and Valenciennes, 1835 also is 2005). These highly modified accessory LOS potentially a member of Photoplagios, al- tissues function to regulate, direct, and diffuse though no types are known and the original the intense blue-green bacterial light over the description (and earlier comments relating to ventrum (Harms, 1928; Ahrens, 1965; Bassot, this taxon by Forsska˚l, 1775: 58) does not 1975; McFall-Ngai, 1983; Dunlap and serve to diagnose this species on the basis of McFall-Ngai, 1987). Additionally, males of unique anatomical features or to distinguish it most species possess the ability to emit light in from congeners on the basis of a combination rapid flashes from translucent patches located of unique attributes. Likewise, based on either on the flanks, along the opercular similar remarks in the original description margin, in the buccal cavity, or some combi- and an external examination of the holotype, nation of these regions (Haneda, 1940; Equula oblonga Valenciennes, in Cuvier and Hastings, 1971; Herring and Morin, 1978; Valenciennes, 1835 is likely also to be a mem- McFall-Ngai and Dunlap, 1983; McFall-Ngai, ber of Photoplagios; however, the holotype is 1991; Woodland et al., 2002; Sasaki et al., in poor condition with regard to pigmentation 2003; Sparks et al., 2005). Ventral lumines- pattern, and the LOS cannot be studied. cence is hypothesized to provide camouflage Members of Photoplagios are biolumines- illumination against bottom-dwelling piscivo- cent schooling fishes and are distributed in rous fishes, whereas lateral, buccal, and shallow coastal habitats from the east coast of opercular flashing may function in prey Africa and the Red Sea (including a single attraction, other forms of predator avoidance, species, P. klunzingeri, which has dispersed schooling behavior, and sex-specific signaling into the Mediterranean Sea via the Suez (Hastings, 1971; Herring and Morin, 1978; Canal) to Japan and Australia. Within McFall-Ngai, 1983; McFall-Ngai and Dunlap, Photoplagios, P. leuciscus and P. lineolatus 1983, 1984; Dunlap and McFall-Ngai, 1987; are reported to have extensive ranges (but see McFall-Ngai and Morin, 1991; Woodland et 2006 SPARKS: NEW MALAGASY PONYFISH 3 al., 2002; Sasaki et al., 2003; Sparks and defined as the first vertebra bearing a definite Dunlap, 2004; Sparks et al., 2005). The degree hemal spine. Vertebral and fin spine/ray of species-specific anatomical specialization counts were obtained from radiographs. The and strong sexual dimorphism of the light terminal dorsal-fin and anal-fin rays, which organ and associated structures of the LOS are branched to the base of the fin, are observed throughout the family suggest a sys- counted as a single element. Pored scales in tem of mate recognition based on male the lateral line are counted in series from the species-specific luminescent signaling (Sparks dorsal margin of the gill opening to the caudal et al., 2005). flexure. Scale counts should be interpreted as In the present study, a new species belong- approximations, due to high intraspecific ing to the recently described genus variability, irregular arrangement, the decidu- Photoplagios Sparks, Dunlap, and Smith, ous nature of ponyfish scales in preservation, 2005 is described from material collected in and because small scale size and the degree to northeastern Madagascar. A phylogeny for which scales are embedded make accurate Photoplagios is presented based on a reanalysis counts problematic. of the morphological transformations and nucleotide characters (comprising both mito- INSTITUTIONAL ABBREVIATIONS chondrial and nuclear genes) analyzed by Sparks et al. (2005), with the addition of two Institutional abbreviations follow Leviton new taxa, P. klunzingeri and P. moretoniensis, et al. (1985): and additional anatomical features of the AMNH American Museum of Natural His- LOS. In the context of this phylogeny, tory, New York placement of the new species within AMS Australian Museum, Sydney Photoplagios is discussed with reference to ANSP Academy of Natural Sciences, homologous features of the LOS. In addition, Philadelphia the taxonomic status of Equula lineolata BMNH Natural History Museum, London Valenciennes, in Cuvier and Valenciennes, CAS California Academy of Sciences, San 1835, an allegedly widespread taxon to which Francisco several distinct species seem to have been LACM Los Angeles County Museum of Na- historically attributed, and presumably in- tural History cluding the new species, is discussed in detail. MNHN Museum National d’Histoire Na- turelle, Paris NMW Naturhistorisches Museum, Wien MATERIALS AND METHODS QM Queensland Museum, South Bank, Australia MORPHOLOGY SIO Scripps Institution of Oceanography, Osteological features of the new species and Marine Vertebrates Collection, La related taxa were examined using radiographs, Jolla specimens cleared and stained for bone and UMMZ University of Michigan, Museum of Zoology, Ann Arbor cartilage (C&S) using a protocol based