Review of the Asiatic Freshwater Puffers of the Genus Tetraodon Linnaeus, 1758 (Pisces, Tetraodontiformes, Tetraodontidae)
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Review of the Asiatic freshwater puffers of the genus Tetraodon Linnaeus, 1758 (Pisces, Tetraodontiformes, Tetraodontidae) by W.J. Dekkers Institute of Taxonomie Zoology, University of Amsterdam, The Netherlands Contents Abstract 88 Introduction 88 Methods 89 Abbreviations 91 Acknowledgements 91 Taxonomie part 92 Genus Tetraodon 92 Key to the species-groups, species and subspecies 92 cutcutia-group 93 Tetraodon cutcutia Hamilton, 1822 95 Tetraodon lorteti Tirant, 1885 97 Tetraodon travancorius Hora & Nair, 1941 101 leiurus-group 102 Tetraodon palembangensis Bleeker, 1852 104 Tetraodon leiurus sensu lato 107 Tetraodon leiurus Bleeker, 1851 108 Tetraodon fangi Pellegrin & Chevey, 1940 112 erythrotaenia-group 114 Tetraodon erythrotaenia Bleeker, 1853 115 fluviatilis-group 117 Tetraodon fluviatilis sensu lato 120 Tetraodon kretamensis Inger, 1953 121 Tetraodon nigroviridis Marion de Procé, 1822 123 Tetraodon fluviatilis fluviatilis Hamilton, 1822 127 Tetraodon fluviatilis sabahensis nov. subsp 130 Tetraodon steindachneri nom. nov 132 waandersii-group 134 Tetraodon waandersii Bleeker, 1853 134 References 136 Index to the taxonomie part 141 Downloaded from Brill.com10/11/2021 05:35:07AM via free access 88 W. J. DEKKERS - REVIEW OF TETRAODON Abstract terial is of not readily recognizable because scanty labelling and of not being kept apart. The prob- The present review of the Asiatic freshwater puffers of lems evolving from this are treated De the Tetraodon because the by genus was thought necessary Beaufort (1964), Boeseman (1972), Hubrecht only preceding review (Le Danois, 1959) disregards some major taxonomic rules and methods. The material studied (1879), Rofen (MS.), and Whitehead et al. (1966). originates from existing museum collections and comprises Amongst the 19th century French ichthyolo- material of of the concerned. The type most species gists, most work on Tetraodon was carried out concept of the genus Tetraodon is adapted after Fraser- exclusion of by Bibron. This, however, was never Brunner (1943) with T. (Chelonodon) patoca, published still in the which is not considered. as such; it rests as a manuscript One new subspecies ( T. fluviatilis sabahensis) is described, Muséum National d'Histoire Naturelle, Paris. one new name (T. steindachneri) is introduced, two neo- Much of the information in it, however, was sub- types (T. fluviatilis fluviatilis Hamilton, 1822 and T. sequently published by Duméril (1855), including nigroviridis Marion de Procé, 1822) are designated, and numerous new generic names which have not one lectotype (T. lorteti Tirant, 1885) is selected. For the sake of convenience in discussing the taxonomic value of stood the test of time well at all. the characters used the species are grouped in five Other 19th workers the like century on group, species-groups: Cantor, De Castelnau, Marion de Procé and cutcutia-group (T. cutcutia Hamilton, 1822, T. lorteti Tirant, present comparable complications. De Tirant, 1885, T. travancorius Hora & Nair, 1941) Castelnau leiurus-group (T. palembangensis Bleeker, 1852, T. only left a sketchbook concerning leiurus T. & Bleeker, 1851, fangi Pellegrin Chevey, Tetraodon, the material of Marion de Procé was 1940) for the destroyed greater part during a revolt, erythrotaenia-group (T. erythrotaenia Bleeker, 1853) and neither Cantor nor Tirant left illustrations of fluviatilis-group (T. kretamensis Inger, 1953, their new Tetraodon Like most 19th cen- T. nigroviridis Marion de Procé, 1822, species. scientists had to much the T. fluviatilis fluviatilis Hamilton, 1822, tury they rely very on T. sabahensis fluviatilis nov. subsp., descriptions of their colleagues from abroad, as T. steindachneri nom. nov.) they had little access to material from colonies waandersii-group (T. waandersii Bleeker, 1853). other than their homeland's. In useful our century a compilation of knowl- edge on Indonesian species was given by De INTRODUCTION Beaufort (1962). Some years before, in 1959, Le Danois had published the only attempt at a real Asiatic freshwater Although the taxonomy of the review on the specific level. Her extensive mono- puffers has been studied quite intensively from the graph on what she calls the "Orbiculates" turned beginning of the 19th century, much systematic out, however, to make matters even more com- confusion is left to be cleared. Partly this is due plicated. She did a bad service to science by ne- to the peculiarities of the group, partly to the glecting other than material in Paris, by neglect- peculiarities of the scientists studying it. ing the internationally approved type concept, by neglecting illustrations accompanying original de- Hamilton, the first major contributor to our scriptions, by exhuming forgotten names of un- of the Asiatic Tetraodon based knowledge species, recognizable nominal species, and by raising his fishes from the river study (1822) on Ganges myological characters to the highest ranks of He made descriptions and did not system. poor taxonomically important characteristics. Her views leave collections. As he points out himself, the on osteology were conclusively refuted by Tyler illustrations therefore are of uttermost importance. (1963). One quotation of him (: 205) probably Unfortunately, the illustrations of three out of will do: "the line drawings of the skulls are in- his six new Tetraodon species were lost at the accurate, for they show many more bones than time of printing, of which one was refound. only actually exist." The problems connected with this are discussed by Gudger (1924), Hora (1929), and others. The taxonomie confusion due to the peculiari- The second major contributor, Bleeker, in his ties of the fishes themselves originates in their the publications on genus (1849-1868) is mainly polymorphism. Besides the known variability con- concerned with Indonesian fishes. He had no nected with geographical distribution, aquarium material means to compare his with the species studies by Benl (1957 a & b), Benl & Chlupaty described He and Klausewitz & and field by Hamilton from Bengal. did leave (1957), (1957 a b), collection, but a huge fish most of his type ma- studies by Sontirat et al. (1971) have shown con- Downloaded from Brill.com10/11/2021 05:35:07AM via free access BUDRAGEN TOT DE DIERKUNDE, 45 (1) - 1975 89 fluviatilis- siderable changes in colour and shape depending group would have to be regarded as a single counts: D 10-14; on age, sex, mood, habitat type, and lighting. species with fin ray 10-16; A shown in P 18-24. variations in fin Striking sexual dimorphism was T. lor- Although ray counts teti and perhaps more sexual dimorphism will be like this might occur in a single archipelagic found in the group as the phenomenon is not rare species — certainly when one regards it as a super- in the Tetraodontiformes. It was shown in — I consider it more to dis- e.g. species opportune the another family of the group, the Ostraciontidae, tinguish species and subspecies as long as in the species Ostracion meleagris Shaw, 1796, available material prohibits a more advanced and O. solorensis (Bleeker, 1853) by Fraser- taxonomie approach. Brunner (1940). Variations with age could be in in It be said that the of the even stronger nature than aquarium life must scope present because most species are recorded from fresh and study does not go beyond classical taxonomy: brackish waters and some even from marine the classical taxonomie problems inherited from This that reckon the 19th be before waters. implies one must with century must cleared starting variation due to ecological and migratorial cir- ecological, population dynamical, or experimental cumstances. The latter proposition could not be research. verificated in the this would The is based of present paper; need present study on comparison far more material from exact localities. Varia- existing museum series and data recorded in tions with mood include the inflating and deflating publications. Some authors intended such com- of the belly region. Body depth and body width parisons, but local or temporary circumstances actual action. for therefore taxonomically are almost useless. The prevented Bleeker (1865a: 68) distribution after 16 of de- apparent size and of the spines also instance, years contemplating, vary with mood, that is to say, the spines can be clared his T. potamophilus a junior synonym of be- retracted to a great extent under the skin, often Crayracion fluviatilis (Hamilton, 1822), not in dermal he of it but be- specialized pits or under dermal papillae. cause was convinced being so, Records of early authors, considering size and cause he did not have material for comparisons. distribution of the In the both nominal spines a major character, are present paper species are reliable in this T. and T. not respect. separated again (as fluviatilis nigro- On the other hand, some characters proved to viridis). Many type specimens are compared for the first time with other and be so constant throughout the genus that they type specimens neo- could not be used either. This in the first place types are designated when necessary to prevent concerns form and position of the fins, mouth future confusion. width, diameter. Besides the mentioned and eye approach above, zoo- After variable and data eliminating the characters over- geographical arguments aquarium were lapping too much between species and the con- used — although with caution. While Tetraodon characters of the the freshwater stant genus, only following species are by no means primary characters proved to