Zootaxa, 3642 (1): 1-105

Zootaxa 3642 (1): 001–105 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Monograph ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3642.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:CAFF80B0-1737-4DAE-BA43-C11D07EE1357

ZOOTAXA

3642

Phylogenetic systematics of the family Sillaginidae (Percomorpha: order Perciformes)

TATSUYA KAGA Laboratory of Marine Biology and Biodiversity (Systematic Ichthyology), Graduate School of Fisheries Sciences, Hokkaido University, 3-1-1 Minato-cho, Hakodate, Hokkaido 041-8611, Japan. Present address: Osaka Animal Plants Ocean College, 1-7-3 Sangenyahigashi, Taisho, Osaka 551-0002, Japan. E-mail: [email protected]

Magnolia Press Auckland, New Zealand

Accepted by M. R. de Carvalho: 4 Jan. 2013; published: 30 Apr. 2013 TATSUYA KAGA Phylogenetic systematics of the family Sillaginidae (Percomorpha: order Perciformes) (Zootaxa 3642) 105 pp.; 30 cm. 30 Apr. 2013 ISBN 978-1-77557-144-5 (paperback) ISBN 978-1-77557-145-2 (Online edition)

FIRST PUBLISHED IN 2013 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

Abstract ...... 4 Introduction ...... 4 Material and methods ...... 5 Sillaginidae ...... 5 Comparative material (dissected) ...... 6 Systematic procedures ...... 7 Comparative anatomy of Sillaginidae ...... 7 1.Osteology ...... 7 Orbital bones ...... 7 Neurocranium ...... 10 Jaws ...... 18 Suspensorium and opercular bones ...... 21 Hyoid arch ...... 25 Branchial arch ...... 30 Pectoral girdle ...... 33 Pelvic girdle ...... 37 Axial skeleton ...... 40 Pterygiophores and fin rays of dorsal and anal fins ...... 44 Caudal skeleton ...... 50 2. Myology...... 54 Eye muscles ...... 54 Cheek muscles...... 56 Cephalic muscles between neurocranium and suspensorium-operculum ...... 57 Ventral muscles of head...... 61 Branchial muscles ...... 64 Pectoral fin muscles ...... 73 Pelvic fin muscles ...... 75 Caudal fin muscles ...... 78 Body muscles ...... 80 3. Swimbladder ...... 82 4. External morphology ...... 87 Monophyly of the family Sillaginidae...... 87 Interrelationships of the family Sillaginidae ...... 88 Classification ...... 93 Family Sillaginidae Richardson, 1846...... 94 Key to the genera of Sillaginidae ...... 95 Genus Sillago Cuvier, 1817...... 95 Genus Sillaginopsis Gill, 1861 ...... 96 Genus Sillaginodes Gill, 1862...... 97 Genus Sillaginopodys Fowler, 1933 ...... 97 Genus Sillaginops gen. nov...... 98 Questionable genus: Neosillago Castelnau, 1875 ...... 98 A species discussion ...... 99 Discussion of evolutionary trends in Sillaginidae ...... 100 Acknowledgements ...... 102 References ...... 102

The phylogenetic relationships of the family Sillaginidae are inferred based on morphological characters. The Sillaginidae is revealed as being a monophyletic group supported by 16 synapomorphies such as shortened segments on the first soft ray of pelvic fin and the adductor mandibulae section A2 covering section A1 laterally. The characters recognized in 24 transformation series were used for the phylogenetic analysis to reconstruct the interrelationships of the family. Twelve equally most parsimonious trees were obtained in the analysis. A strict consensus tree from the 12 trees was adopted as representing the phylogenetic relationships of the family Sillaginidae. Although reversals and character changes to other derived conditions are recognized, it is inferred that five characters additionally support the monophyly of the Sillaginidae. The family Sillaginidae is redefined based on the synapomorphies supporting its monophyly for the first time. In the new classification proposed, the Sillaginidae comprises the following five genera: Sillago, Sillaginopsis, Sillaginodes, Sillaginopodys, and Sillaginops gen. nov. A key to identification of the genera of Sillaginidae is provided. The family Sillaginidae contains two groups having different evolutionary trends in the reconstructed phylogeny of the family. The first group, including Sillaginopsis, Sillaginodes, Sillaginopodys, and Sillaginops gen. nov., has a trend towards reducing the swimbladder; and the second group, including only Sillago, has a trend towards further refining the swimbladder.

Key words: Phylogeny, Sillaginidae, Monophyly, Evolutionary trend, Sillaginopsis, Sillaginodes, Sillaginopodys, Sillaginops, new genus

Introduction

The family Sillaginidae Richardson, 1846, commonly known as sand whitings or sand borers (McKay, 1992), belongs to the perciform suborder Percoidei (e.g., Greenwood et al., 1966; Johnson, 1993; Nelson, 2006). According to McKay (1992), Nelson (2006), Kaga et al. (2010), and Gao et al. (2011), the family consists of three genera and about 33 species. Sillaginids inhabit shallow marine waters, estuaries, and rarely into freshwaters of the Indo-West Pacific (McKay, 1992; Nelson, 2006). Several members are important inshore fishery and recreational sport fishing species, including Sillaginodes and Sillago (Parasillago) japonica (McKay, 1992; Konishi & Nakabo, 2007). Golani et al. (2011) rehabilitated Sillago erythraea Cuvier in Cuvier & Valenciennes, 1829, from the synonym of S. sihama (Forsskål, 1775), and the species S. sihama was restricted to the southern Red Sea. However, the status of S. erythraea and S. sihama is still uncertain. The family Sillaginidae is characterized by the following characters: body elongated; mouth small; two dorsal fins with little or no interspace, the first dorsal fin bearing 10–13 spines, the second dorsal with one slender spine and 16–27 soft rays; the anal fin with two small spines and 14–26 soft rays; three supraneural bones; swimbladder absent or vestigial to highly complex with various extensions; and 32–44 vertebrae (Nelson, 2006). The systematic position of the family Sillaginidae is still uncertain, although Springer and Orrell (2004) showed the close relationships with Rhyacichthyidae and Percidae by the phylogenetic analysis based on dorsal gill-arch muscles and skeleton. The Sillaginidae has been variously classified by previous ichthyologists. Gill (1861) recognized two genera, including Sillago Cuvier, 1817 and Sillaginopsis Gill, 1861, but subsequently, Gill (1862) added a new genus Sillaginodes. Bleeker (1874) followed Gill (1862) in recognizing the three genera. In contrast, Boulenger (1901) recognized only Sillago. Fowler (1933) recognized two subgenera (Sillago and Sillaginopodys) in the genus Sillago. McKay (1985) divided the genus Sillago into three subgenera based on morphology of the swimbladder: the subgenus Sillaginopodys Fowler, 1933 with a reduced swimbladder, the subgenus Sillago Cuvier, 1817 with two posterior extensions of the swimbladder, and the subgenus Parasillago McKay, 1985 with one simple posterior extension and duct-like process. Recently Sillago (Parasillago) indica McKay, Dutt & Sujatha, 1985 was reassigned to the subgenus Sillago from the observation of the swimbladder by Kaga & Ho (2012). However, some researchers have not supported McKay’s subgenera (e.g., Nelson, 2006). Fowler (1933) separated the Sillaginidae into two subfamilies, Sillaginae (including Sillago and Sillaginodes) and Sillaginopsinae (including Sillaginopsis). However, recent authors have not recognized Fowler’s subfamilies (e.g., McKay, 1985, 1992; Nelson, 2006). Such taxonomic disagreements of the composition of the Sillaginidae have arisen because of the lack of understanding of its phylogenetic relationships, which can provide the basis for classification.