AMERICAN MUSEUM Novltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2979, 33 pp., 27 figs., 4 tables August 7, 1990 Notes on the Anatomy and Relationships of the Bedotiid Fishes of Madagascar, with a Taxonomic Revision of the Genus Rheocles (Atherinomorpha: Bedotiidae) MELANIE L. J. STIASSNY1 ABSTRACT In a wide-ranging comparative study the anat- an alternative scheme for a monophyletic Perco- omy and relationships of the bedotiid fishes of morpha (including the mugiloids) is also pre- Madagascar are investigated. Evidence supporting sented. the following hypotheses is presented and dis- There follows a taxonomic revision ofthe genus cussed: (1) the mugiloids (sensu Nelson, 1984) are Rheocles Jordan and Hubbs, 1919. Four species the sister group ofthe Atherinomorpha (sensu Ro- of Rheocles are recognized in the present study, sen, 1964), (2) the Atherinomorpha are mono- one of which, Rheocles wrightae, is newly de- phyletic, (3) the Bedotiidae are monophyletic, (4) scribed. The intrarelationships ofRheocles are re- Bedotia is monophyletic, (5) Rheocles is mono- solved and a species-level cladogram is presented. phyletic. Counterevidence to claim (1), supporting INTRODUCTION In their analysis of atherinomorph mono- stethidae, and Telmatherinidae) ofunresolved phyly and relationships, Rosen and Parenti relationships. Their scheme has been amend- (1981) challenged earlier notions of a mono- ed recently by Parenti (1984b) with the phyletic Atherinoidei, preferring instead to formal recognition ofa monophyletic Phallo- recognize a division I atherinoid group con- stethoidea (Phallostethidae + Dentatherini- sisting of six families (Atherinidae, Bedoti- dae), but as yet there exists no consensus re- idae, Isonidae, Melanotaeniidae, Phallo- garding the interrelationships of this diverse I Assistant Curator, Department of Herpetology and Ichthyology, American Museum of Natural History. Copyright © American Museum of Natural History 1990 ISSN 0003-0082 / Price $5.25 2 AMERICAN MUSEUM NOVITATES NO. 2979 atherinomorph division (see Patten, 1978; study. Historically one acanthomorph group, Allen, 1980; Parenti, 1984b; White et al., the Mugiliformes, has been closely allied with 1984; Ivantsoff et al., 1987). various of the atherinomorph lineages (Jor- Members of the Madagascan bedotiid ra- dan and Hubbs, 1919b; Myers, 1928; Berg, diation are generally considered to represent 1940; Gosline, 1968, 1971). Following Rosen the most generalized or "primitive" ofliving (1964) and Nelson (1984) I adopt the ver- atherinoids (e.g., Jordan and Hubbs, 1919b; nacular names "mugiloid," "sphyraenoid," Rosen, 1964). Three bedotiid genera have and "polynemoid" for the mugiliform com- been described to date; Bedotia Regan, 1903, ponents of interest here, and have empha- Rheocles Jordan and Hubbs, 1919, and Rhe- sized the inclusion ofrepresentatives ofeach ocloides Nichols and LaMonte, 1931, but of these lineages in comparative study. Se- despite their suspected key phylogenetic po- lection from among the numerous other per- sition in relation to the remaining Atherino- comorph lineages has necessarily been some- morpha (Rosen and Parenti, 1981; Parenti, what arbitrary although I tended to 1984b), very little is known of the structure concentrate on the more morphologically or interrelationships of this assemblage. generalized taxa (Rosen, 1973; Johnson, 1980, Much of the reason for the dearth of in- 1984; Stiassny, 1981). Among other more de- formation on bedotiid anatomy results from rived atherinomorphs a range of atherinoids their poor representation in museum collec- have been incorporated for study, as have tions. Fortunately, a recent ichthyological representative cyprinodontiforms and belo- survey ofthe fresh waters ofthe eastern high- niforms. lands of Madagascar (Reinthal and Stiassny, A list ofthe acanthomorph species includ- in rev.) has made available a good size col- ed in this study follows: lection enabling a detailed anatomical and taxonomic study to be undertaken. The re- Polymixia lowei (AMNH 49674), Percopsis sults of that study form the nucleus of the omiscomaycus (AMNH 41145, 42032), Aphredo- present paper, and the impetus for the higher derus sayanus (AMNH 33540, 55089), Pollachius level systematic investigations reported virens (AMNH 40584), Gadus morhua (AMNH herein. 2972), Molva molva (AMNH uncat.), Merluccius bilineuris (AMNH 55086), Opsanus tau (AMNH OUTGROUP SELECTION AND 73813), Hoplostethus mediterranus (AMNH ASSOCIATED PROBLEMS 49718), Anomalops katopteron (AMNH 37949), Gibberichthys pumilus (AMNH 49679), Scopelo- In seeking to resolve the status and rela- beryx sp. (AMNH 49710), Zeus faber (AMNH tionships of the basal atherinomorph clades 29458), Polynemus approximans (AMNH 73388), ofMadagascan bedotiids, a number ofprob- Polynemus opercularis (AMNH 16003), Polydac- lems have been encountered. The first con- tylus virginicus (AMNH 74250), Sphyraena bo- cerns the selection of appropriate outgroups realis (AMNH 4339), Sphyraena barracuda (AMNH 20609), Agonostomus monticola (AMNH for analysis. 31538), Mugil curema (AMNH 39162), Mugil In broad outline, figure 1 represents the cephalus (AMNH 15481), Liza macrolepidotis current state of our knowledge of acantho- (AMNH 88044), Centropomus ensiferus (AMNH morph interrelationships. In view of the du- 35244), Morone americana (AMNH 44691), Am- bious status of the Paracanthopterygii (Ro- bassis urotaenia (AMNH 88082), Kurtus gulliveri sen, 1973, 1985; Stiassny, 1986), selection of (AMNH 43396), Micropterus dolomieui (AMNH "appropriate" outgroups from this hetero- 68385), Lutjanus fulviflamma (AMNH 88161), geneous assemblage is problematical. Where Serranus tigrinus (AMNH 43172), Gerres rappi possible I have examined a range of para- (AMNH 88058), Ptychochromis oligacanthus canthopterygian taxa with an emphasis on (AMNH 88117, 88102), Monodactylus argenteus (AMNH 58794), Hypseleotris tohizonae (AMNH members of the percopsiform "clade" (Ro- 85099, 88048), Deltentosteus sp. (AMNH uncat.), sen, 1962; Patterson and Rosen, 1989). The Melanotaenia maccullochi(AMNH 44401), Mela- position ofPolymixia seems more firmly es- notaenia nigrans (AMNH 55067, 20574), Mela- tablished as the sister group to the remaining notaenia goldiei (AMNH 13900), Chilatherina acanthomorpha (Rosen, 1985; Stiassny, lorentzi (AMNH 15028), Telmatherina ladigesi 1986), and this taxon has been included for (AMNH 35378), Atherinapinguis (AMNH 9434), 1 990 STIASSNY: BEDOTIID FISHES OF MADAGASCAR 3 Acanthomorpha I r Paracanthopterygi - I Acanthopterygi I Atherinomorpha F- Percomorpha I Fig. 1. Cladogram depicting the interrelationships ofacanthomorph fishes. After Rosen, 1973, 1985; Rosen and Parenti, 1981; Stiassny, 1986. Teramulus keineri(AMNH 88141), Craterocepha- Bedotiid taxa: Bedotia madagascariensis lus cuneiceps (AMNH 43184), Craterocephalus (MNHN 1942-81, 1963-169), B. longianalis stercusmuscarum (AMNH 43186), Atherinomorus (MNHN 1914-6,1936-147,148), B. geayi (AMNH stipes (AMNH 2357, 53025), Quirichthys stra- 28132, 57453, MNHN 1968-132), B. tricolor mineus (AMNH 20571), Pseudomugil tenellus (MNHN 1932-1, 2, 1934-190) B. spp. (AMNH (AMNH 36598), Menidia menidia (AMNH 40696, 88074, 88011), Rheocles sikorae (MNHN 1891- 40592), Menidia beryllina (AMNH 43043), Me- 727, MNHN 1962-188,1966-914, AMNH 28127), lanorhinus microps (AMNH 25878), Atherinops R. alaotrensis (MNHN 1913-328, 329, 330, 331, affinis (AMNH 51144), Hemiramphus brasiliensis 332,333, MNHN 1966-1074, MNHN 1966-1075, (AMNH 30478), Hemiramphus balao (AMNH MNHN 1962-187, MNHN 1934-275, 276, 55601), Arrhamphus sclerolepis (AMNH 40002), MNHN 1932, 28, 29, MNHN 1919-10, MNHN Stronglura marina (AMNH 36060), Scomberesox 1913-334, BMNH 1920.3.2:33-36, AMNH 28135, saurus (AMNH 76818), Exocoetus volitans AMNH 88171, AMNH 88001), R. wrightae n. sp. (AMNH 73177), Pachypanchaxplayfairi (AMNH (MNHN 1942-77,1989-1614, AMNH 58908), R. 20637), Cyprinodon variegatus (AMNH 196494), pellegrini (AMNH 9696, AMNH 11699). Xenotoca variata (AMNH 178760), Fundulus het- eroclitus (AMNH uncat.), Orestias ispi (AMNH A second, and perhaps more vexing, prob- 48643). lem involves the noncomparability of much .4 AMERICAN MUSEUM NOVITATES NO. 2979 FPERCOMORPHA rFMUGLOMORPHAA I ATHERNOMORPHA I Fig. 2. Cladogram of acanthomorph relationships, summarizing the results of the present study. Characters are: 1. Subdivision ofthe pharyngocleithralis muscle. 2. Pharyngohyoideus reduced to a small fan-shaped muscle with an elongate tendon. 3. Levator externus 1 separated from levatores externi 2-4 by the levatores interni 1-2. 4. Anterior neural arches expanded. 5. The A, section of the adductor mandibulae with a tendon to the medial face ofthe lachrymal bone (tA,iac). 6. The fifth ceratobranchials with well-developed ventral process. 7. The lateral horns ofthe fifth ceratobranchials with well-developed muscular processes. 8. Supraneural bones absent. 9. Both PU3ds and PU3vt radial cartliages in the caudal fin skeleton absent. 10. Posterior six or seven caudal vertebrae are markedly thickened. 11. A single elongate infraorbital element behind the lachrymal. 12. Medial processes of the pelvic girdle asymmetrical. 13. Absence ofanteromedial spines, and reduction ofposteromedial spines on the medial processes of the pelvic girdle. 14. Pelvic girdle united medially, anteromedial processes ventrally dis- placed. 15. Third basibranchial toothplates paired. 16. Premaxilla with a deep notch on dentigerous face. 17. Parhypural fused