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

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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-

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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 be useful: fishes, they follow to some extent the zoogeo-

— structure of the nasal organ graphical distribution patterns of the latter as dis-

— relative snout length cussed by De Beaufort (1926, 1951) and reviewed

— relative head length by Inger & Chin (1962). The Wallace Line, for

— form does divide of the interorbital region instance, Tetraodon species: T. ery-

— position of the mouth throtaenia is the only one living east of it. Another

— colour pattern remarkable zoogeographical hypothesis can be

— fin counts variable demonstrated in Tetraodon viz. ray (although within one species, the strong species). variability of the six species living in Borneo.

Colour pattern is incorporated in the diagnostic This variability supports the thesis of De Beaufort characters when it showed and that least only remarkably con- others, assuming at two separate stant differences, and when no individual colour immigration flows occurred in the island and that

pattern contradicted the proposed discrimination. both flows were mixed only partially, due to the

What happens when colour pattern is not in- barriers formed by the prominent mountain ridges. corporated is shown by Le Danois, who in doing so reduced the number of Orbiculate speoies from METHODS

320 to 50. Without splitting according to colour

the named in this For the various of the pattern, group paper aspects species treated,

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the methods and of transcribed in following ways representation were mostly a more recent geo- were used: graphical equivalent when appropriate. If a re-

corded locality is based on material reported on

and references. — References scien- another it is Synonyms to earlier by the same or author, not tific publications aim at completeness; aquarium listed. If all localities and material were listed publications are referred to only when important earlier, this is indicated by "no additional ma- for taxonomie if there is indication that material purposes. "Figure" always repre- terial"; no sents lateral depictions or photographs of living other than living specimens were recorded this is specimens, unless stated otherwise between brack- indicated by "no material". If it is known or sus-

The "listed" with in which ets. word indicates that no descrip- pected great probability museum tion was given. Localities of recorded material collection the material recorded is or was present,

Figs. 1—2. Measurements as used in the present

publication:

1 standard length

2 total length

3 fin, longest ray

shortest the 4 fin, ray (always 1st)

5 fin, base

6 depth, maximum 7 depth, at pectoral fin base

8 depth, at end of dorsal fin base

9 width, maximum

10 head length

11 snout length

12 interocular distance

13 horizontal diameter eye,

14 distance from dorsal eye, profile

15 nasal organ, length

16 nasal distance from of snout organ, tip

17 nasal organ, distance from eye

18 mouthwidth

19 upper lip depth

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this is all indications checked: indicated. Not were IMC Indian Museum (Calcutta) (= ZSI) IOI Institut Océanographique de l'Indochine they merely are given for the sake of convenience (Cauda) the to reader. If the material recorded represents ITA Instituut voor de Tropen (Amsterdam) contains this is such. or type material indicated as (ex KMA)

KMA Koloniaal Museum (Amsterdam) (= ITA)

LOZB Laboratorium voor het Onderzoek der Zee Descriptions. — Measurements were taken with

(Batavia = Djakarta) dial callipers, in accordance with figs. 1-2. With MHNL Muséum d'Histoire Naturelle de la ville de the exclusion of total length measurements, the Lyon anteriormost of the teeth was considered the point MNH Hungarian Natural History Museum (Budapest) anteriormost point of the body. Width and depth (= TMKB)

MNHN Muséum National d'Histoire Naturelle (Paris) were taken at three differentpoints to compensate MPD Muséum de la Porte Dorée for the inexactitudes due to the various degrees MSNG Museo Civico di Storia Naturale "Giacomo of inflation of the belly Morphometries of region. Doria" (Genova) individual when characters are listed only they were NAM Natura Artis Magistra (Amsterdam) taken in NMB Naturhistorisches at least 50 percent of the specimens Museum (Basel) NMS National Museum examined. If (Singapore) a separate description of type ma- NMW Naturhistorisches Museum (Wien) terial is given, the data on this nevertheless are PBSF Philippine Bureau of Science, Section of in the of the incorporated general description Fisheries (Manila) species, unless stated otherwise. Fin ray counts PFCM Philippine Fisheries Commission (Manila)

RMNH Rijksmuseum van Natuurlijke Historie (Leiden) were made with the aid of a binocular dissecting RSM Royal Siamese Museum (Bangkok) microscope. In case of doubt whether or not the SM Selangor Museum foremost structure in a fin should be regarded SMF Natur-Museum und Forschungs-Institut a true fin or whether or not the last ray ray(s) Senckenberg (Frankfurt am Main) is (were) divided to the base and should be SMM Zoologische Staatsammlung des Bayerischen Staates (München) counted separately, the maximum possible count TF Tropicarium Frankfurt is given in the description. Only the first one or TMKB Természettudomânyi Müzeum Könyvtara unbranched indicated in more rays are as such, (Budapest) (= MNH) words or in roman numerals, while the last one ZMA Zoologisch Museum Amsterdam — Instituut

voor Taxonomische Zoölogie (Amsterdam) or more unbranched rays are not indicated as such. ZMH Zoologisches Staatsinstitut und Zoologisches

Museum (Hamburg) Figures. — If possible and opportune, the nasal ZMUG Zoology Museum of the University of and both the lateral and dorsal view organ are Gorakhpur lack of of these in earlier de- ZSI given, since one Zoological Survey of India (Calcutta) scriptions represents a major source of confusion.

ACKNOWLEDGEMENTS

ABBREVIATIONS

I would like to thank the and following persons hi head length institutions who made the present study possible:

to m mean value Dr. H. Nijssen for giving access all facilities of ZMA

and in other C. si standard length helping many ways; Dr. J. Tyler of the tl total length Lerner Marine Laboratory of the AMNH for carefully

reading the manuscript and giving valuable suggestions

A anal fin for alterations; Dr. M. Boeseman for giving access to the

D dorsal fin collections of RMNH, loaning specimens and giving

N number of specimens examined valuable information on Bleeker Collection specimens;

P pectoral fin Dr. P. H. Greenwood, Mr. J. Chambers, and Mr. G.

Howes for giving access to the collections of BMNH,

AMNH American Museum of Natural History and loaning specimens; Dr. M. L. Bauchot (MNHN) for

(New York) loaning specimens and giving valuable collecting data;

AMS Australian Museum (Sydney) Miss C. Smulders and colleagues (ITA) for giving access

ANSP Academy of Natural Sciences (Philadelphia) to the files and cartographic collections of KMA; Dr.

BFB Bureau of Fisheries (Bangkok) E. R. Alfred (NMS), Dr. J. E. Böhlke (ANSP), Dr. L.

David BMNH British Museum (Natural History) (London) (MHNL), Dr. A. Kapur (ZSI), Dr. P. Kähsbauer

DNHM Darjeeling Natural History Museum (NMW), Dr. W. Klausewitz (SMF), Dr. W. Ladiges

FFKU Faculty of Fisheries, Kasetsart University (ZMH), Dr. A. G. Menon (ZSI), Dr. I. A. Ronquillo (Bangkok) (PFCM), Dr. E. Sutter (NMB), Dr. F. Terofal (SMM),

FMNH Field Museum of Natural History (Chicago) and Dr. L. P. Woods (FMNH) for loaning specimens and

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information. for nominal inten- providing A grant the publication of so high that this species needs an this review from the Netherlands' Organization for the sive study of its own. Advancement of Pure Research (ZWO) is gratefully The definition of the genus, modified from acknowledged. Fraser-Brunner (1943), thus becomes:

of the Tetraodontidae with the A genus family sphenotic expanded laterally beyond frontal to

form flattened orbital roof scarce- TAXONOMIC PART a broad, lobe;

ly arched, the lateral ethmoid not bent down be-

fore the an forameh; eye, not forming olfactory reach- Genus Tetraodon mesethmoid broad; upper lateral line not

ing end of tail, curved down above anal fin to

The genus was established by Linnaeus in 1758 meet lower line, which extends forward beyond

(later frequently misspelled Tetrodon). According the point of union; 19 vertebrae; dermal ossifica-

to Tyler (1964: 121) the type species is generally tions of back bearing simple prickles, evenly

taken to be T. lineatus, an African representative. scattered; spines on side small, closely set; dorsal

have made to Several authors attempts split fin maximally counting 16 rays; nasal organ an

The of their nominal two the genus. validity genera elevated tube, terminally often divided into

and is a affair. The con- lobes which reach the basis. subgenera complicated lips or may nearly is cept presently adopted by most ichthyologists This definition includes all Asiatic Tetraodon-

Herein the the one by Fraser-Brunner (1943). tidae occurring (if not exclusively) in fresh wa-

is subdivided in three Mono- genus subgenera: ters, with the exception of Chonerhinus naritus

tretus Bibron, 1855, Chelonodon Müller, 1839, (Richardson, 1848), C. modestus (Bleeker, 1850), and the last Chelonodon Monotreta Tetraodon, subgenus only comprising or patoca (Hamilton,

African species. This subdivision, other than the 1822), and Sphoeroides ocellatus (Linnaeus, 1766). distinction between based The first be their genera, was mainly on two species can recognized by

distribution the form of the nasal organ and the high (25-38) number of dorsal fin rays, the third of does the prickles. Fraser-Brunner (1943: 14) by its nasal organ consisting of a depression, sur-

consider the form of the nasal im- into not organ an rounded by a low rim, produced a posterior portant enough character to establish genera and an anterior flap, the fourth by its nasal

series "forming as they do a developmental among organ being a low papillum, pierced by two open-

of which are otherwise of a group species closely ings. similar structure". Unfortunately, the Fraser- For the sake of convenience the species in this

Asiatic Brunner concept for the two subgenera paper are grouped into the following species-

125- seems to be wrong, according to Tyler (1964: groups:

126). He considers Monotreta a synonym of - cutcutia-group: T. cutcutia, T. lorteti, T. tra-

Chelonodon and concludes by stating that "the vancorius present concept of Chelonodon will undoubtedly - leiurus-group: T. palembangensis, T. leiurus, have to be changed". T. fangi

As the present study does not consider osteo- - erythrotaenia-group: T. erythrotaenia logical characters, it is not proper to interfere in - fluviatilis-group: T. kretamensis, T. nigrovi- the discussion of and sub- on the validity generic ridis, T. fluviatilis fluviatilis, T. f. sabahensis, generic names. In this paper the Fraser-Brunner T. steindachneri

definition of Tetraodon is followed with one ex- — waandersii T. waandersii. group: is ception: T. patoca Hamilton, 1822, not con- sidered, both for taxonomie and practical reasons.

T. patoca is quite different from the other KEY TO THE SPECIES-GROUPS, SPECIES AND

SUBSPECIES Tetraodon species listed by Fraser-Brunner, in its general body form (which is much more

— Nasal rounded tube with terminal 1 organ a short, a elongate), the form of the nasal (which organ opening, not or only slightly two-lipped (cutcutia- does not fit in the "developmental series"), and group) 2

— Nasal short oblong, or the distribution pattern (ranging from Africa to organ a or slightly strongly compressed tube, the distal 1/3—5/6 of which is East Asia). The practical reason is that the dis- divided into two lips or lobes 4 tribution area is of such a great dimension and 2 — D 7—8; A 8; P 16—17 T. travancorius (p. .. 101) the number of closely allied nominal species is D 10—13; A 10—12; P 14—21 3

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— 3 Height of nasal tube 2 or more in its diameter; sides 11 — Back with dark transverse bands, often including

of with dark colour behind surfaces body mostly a ocellus; pattern a V-shaped marking eyes; apposed

not strikingly sexually dimorphic; D 10—11; A 10; of nasal lobes always without spongy tissue in all

P 18—21 T. cutcutia (p. 95) specimens > 60 mm si; often a broad dark

— Height of nasal tube nearly 1 in its diameter; sides longitudinal band separating dark colour of upper

of body without a dark ocellus; colour pattern parts from light colour of belly; D 11—13; A 10;

strikingly sexually dimorphic; D 11—13; A 10—12; P 18—19 T. kretamensis (p. 121)

P 14—16 T. lorteti (p. 97) — Back with 3—4 dark blotches in longitudinal row;

4 — Nasal tube divided surfaces of nasal lobes with tissue over 1/3—1/2 of its length; apposed spongy

in 85% of apposed surfaces of nasal lobes never with spongy specimens > 60 mm si; never a

sides with dark D 13 A 11 P tissue; many polygonal dark spots longitudinal band; —16; —14;

19—24 leaving only lighter network or with wide-meshed 12

12 — Head 2.4—3.1 in D 13—16; A 11—14; P 19— dark network enclosing some dark rounded spots; si;

head deflated {leiurus-group) 5 24; India, Burma .. T. fluviatilis fluviatilis (p. 127)

— — Head 2.1—2.3 in si; 13—15; A 11—12; P 21— Nasal tube divided over (1/3)—1/2—4/5 of its D

length; apposed surfaces of nasal lobes sometimes 22; East Malaysia.. T. fluviatilis sabahensis (p. 130)

with sides coloured head spongy tissue; otherwise;

mostly rounded 7

5 — Sides with wide-meshed dark network, 3—18 of the cutcutia-group meshes enclosing one, sometimes two, rounded,

dark spots or ocelli; body spines often in deep Diagnosis. — Tetraodon species with the nasal dermal pits, with small dermal papillae; D 12—14; formed with terminal A 10—12; P 21—23 .. T. palembangensis (p. 104) organ by a cylinder one

— Sides with the border of which is many polygonal spots leaving lighter opening, not or scarcely

network; back often with one or two dark patches, two-lipped. D 7-13; A 8-12; P 14-21. Three species: sometimes with many oval, whitish spots; body T. cutcutia, T. lorteti, T. travancorius. spines mostly in only superficial dermal pits;

D 12—15; A 10—12; P 21—24 6 Discussion. The is 6 — One of the lateral several spots always formed into a group recognized by

darker and bigger ocellus; authors distinct snout 2.7—2.9 in hi; as a genus or subgenus, by the maximum known si 61 mm; D 12—14; A 10—11; name of Leiodon Swainson, 1839, Leiosomus P 21—23 T. fangi (p. 112) Swainson, 1839, or Monotreta Bibron, 1855. The

— Sides with or without lateral ocellus; snout 2.0 — difference with other Tetraodon species, notably 2.4 in hi; maximum known si 153 mm; D 13—15; those of the is A 10—12; P 21—24 T. leiurus (p. 108) leiurus-group, however, gradual,

7 — Sides with a single or double longitudinal band as some of the latter species have their nasal

separating an uniformly dark dorsal region from a organs two-lipped for only one-third of their length, differently coloured ventral region; D 9 —11; A Benl (1957b) proposed the new genus Carinote- 8—9; P 18—19 (erythrotaenia-group)- traodon for his species chlupatyi (a junior T. erythrotaenia (p. 115) syn- l:: .

— of T. Sides sometimes with a lorteti) based on the of the simple longitudinal band, onym ability spe- but otherwise cies differently coloured; D 11—16; A to erect a low mid-dorsal and mid-ventral

P 16—24 10—14; 8 In ridge. my opinion one should await the results

8 — Sides with about 35 dark transverse bandelets on of osteological studies before adopting this view. upper parts; back dark; D 12; A 11; P 16 (waan- dersii-group -T. waandersii (p. 134)

— and back with dark Within the at least 13 Sides spots and/or blotches group names were pro- (fluviatilis-group); D 11—16; A 10—14; P 18—24.. 9 of these posed. Three were published as early 9 — Sides with 2—4 dark ocellated spots and with as 1822 by Hamilton, from the Ganges River whitish irregular lines, some of them broken into system: T. cutcutia, T. caria, and T. The spots; back with similar lines and with three dark gularis. first of these first patches; D 13—15; A 11—13; P 18—21 also printed as of the three

T. steindachneri 132) is easily recognizable from the (p. figures accompa-

— Sides without whitish back with markings; or the and is nying description generally accepted as without whitish markings and dark patches; D valid a species. The drawings of the latter two 11—16; A 10—14; P 18—24 10 were retained by the — and 10 Back and sides with more or less regular, rounded Bengal government

could not be in Hamilton's spots, which are more or less evenly scattered, and incorporated publi-

only occasionally confluent, never broad cation. Of these forming two only one later was refound. surfaces of nasal lobes blotches; apposed spongy It then was published by Hora (1929: pl. 16, in 50% of specimens > 60 mm si; D 12—14; A figs. 2-3). According to Hora it represents 10—12; P 18—23 T. nigroviridis (p. 123) T. caria because the carried the — name Back with dark transverse markings or with large drawing dark "Tetradon the words blotches; nasal lobes with or without spongy Kariya Phoksa", last being,

tissue; D 11—16; A P 18—24 recorded 10—14; 11 as by Hamilton in his original notes,

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the local equivalent of T. caria. This view, how- my concept is T. lorteti, Tirant, 1885. ever, is contradicted by the description by Hamil-

valid after ton, in which T. caria is characterized by the T. lorteti proved a species exami- lack the of a lateral black spot; figures repro- nation of the syntypic material. Important charac- the duced by Hora show a distinct spot. In Day's ters discriminating it from T. cutcutia are the fin 18-21 in opinion, as cited by Hora, figures represent pectoral ray counts (15-16 against

T. cutcutia. This would imply that Hamilton had T. cutcutia). the length of the nasal organ (1

in diameter in and the lack commissioned four drawings for a single species, against 2 diameter)

of In scientific the a less probable proposition. To me it seems most a lateral ocellus. circles species likely that the figures represent T. gularis as this is better known as T. borneensis, while in aquari- would match the description without the improb- um circles it is popular under the names of Tetrao- abilities emanating from Hora's and Day's views. don somphongsi Klausewitz, 1957, or Carinote-

Anyway, differences between the four figures and traodon chlupatyi Benl, 1957. Both the first and between the three descriptions are too slight to the last names proved to be junior synonyms of T.

distinguish separate species. Hamilton himself re- lorteti after examination of the type material. T. caria "I Carinotetraodon declared marks on (1822: 9): cannot take upon chlupatyi was a syn-

myself to say, whether it is a distinct species or a onym of T. somphongsi described only some

in 1959. mere variety", and on T. gularis he admits (1822: months earlier by its own author, Benl,

"Its a mere is still more In observations both nominal 10) being ... variety prob- aquarium species able". Given these statements, the absence of and a third form, provisionally named T. werneri,

type material and the loss of exactly definable proved to be physiological forms of a single illustrations, it seems most reasonable to synony- species, the differences being due to age, sex, mize the three nominal species under the heading food conditions, type of vegetation, and compo- of T. cutcutia, as has been done ever since 1822 sition of the aquarium community. T. werneri

— until In that latter Danois the and females, 1959. year Le resur- proved to represent juveniles rected the T. the males and name T. caria (as Monotreta caria) somphongsi representing younger

for T. lorteti Tirant, 1885, without giving any T. chlupatyi the mature males. This was confirm- arguments for doing so. According to Hamilton ed by sex determination on 200 specimens by

T. caria found in River in Differences is the Ganges system, has Somphongs (cited Benl, 1959). about 18 pectoral fin rays, and has jaws of near- in colour pattern although not related to sex ly equal length. T. lorteti, in contrast, is not known differences were earlier recorded by Hora from India, fin identifiedhis has (14)-15-(16) pectoral rays (1923b, 1924), who, however, wrongly

(N=42), and has its mouth directed upwards. specimens as T. palembangensis. Probably not ac-

The name gularis (also in the genus Mono- quainted to the work of B®nl and later authors,

also resurrected Le Danois the de- treta) was by (1959) female-juvenile form of T. lorteti was and used for T. the of the latter Monotreta cutcutia, name scribed as a separate new species species being reserved by her for T. leiurus and tiranti by d'Aubenton & Blanc (1966), who

T. palembangensis. These two Bleeker species, named the male form Monotreta caria, following dissimilar however, prove to be from T. cutcutia Le Danois (1959). The incorrectness of this view by taking a second's look at the figures in the was recently emphasized by Sontirat et al. (1971)

"Gangetic Fishes" (Hamilton, 1822) and those in who found the colour pattern during field obser- the "Atlas Ichthyologique" (Bleeker, 1865a). vations to be strongly dependent on surroundings,

The name Leiodon marmoratus or Leiosomus behaviour and light intensity. marmoratus Swainson, 1839, is a nomen nudum;

The third and last valid it was synonymized by Bleeker (1865a) with T. species recognized as de- cutcutia. Some time later (1865c) Bleeker in the cutcutia-group, T. travancorius, was

— & Nair from India. Al- himself attributed a nomen nudum Chelono- scribed by Hora (1941)

— and all don dumerili when he promised to describe a though the holotype paratypes are less

than in doubt new species from Thailand, a promise which an inch length, there can be little

be- never was redeemed. According to the publica- about them representing a separate species low fin D 7-8; tion the material was deposited in MNHN. Le cause of the strikingly ray counts:

Danois (1959, 1961) recognized it in the collec- A 8 (T. cutcutia: D 10-11; A10; T.lorteti: D 11-13; tions of MNHN as no. 2007. Her redetermination A 10-12). Although in the original description reads Monotreta caria Hamilton, 1822, which in the collector of the specimens, K. B. Nair, is

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quoted stating (1941: 393): "I am sure bigger Calcutta, Hooghly River; material partly in RMNH

in reg. no. 7345, partly ZMA, reg. no. 110.225). specimens, if there were any, would not have been Monotrète cutcut ia; overlooked", I am quite sure the species can grow BIBRON (in: DUMÉRIL), 1855: 280 (listed as type species bigger as the relative is eye-diameter quite high, for Monotrète; India or Bangla Desh: Ganges River

which in Tetraodon indicates that the material in B generally system; MNHN, reg. no. 1470).

Arothron ? specimens are juvenile. A juvenile state of the cutcutia;

BLEEKER, 1859a: 267 (listed; no additional material). type series cannot, however, be a strong argument Leiosomus cutcutia; against the separate specific status as the number BLYTH, 1860: 173 (listed; Burma: Moulmein). of fin in is rays my experience not or scarcely Leiodon cutcutia;

1892: 710 of related with age in Tetraodon. On the other hand, GILL, —711 (listed as representative genus;

no material). the supposed youthfulness most probably indicates Tetraodon cutcutia; that the colour description as given by Hora & PRASHAD & MUKERJI, 1929: 162, 163, 166, 169, 223 Nair is not useful as a specific character. This last (description; Burma: Indawgyi Lake and associated impression is strengthened by the fact that the rivers; material in ZSI).

show in colour SHAW & SHEBBEARE, 1938: 8, 124—125, pi. 4 figs. 12—13, paratypes a considerable variability text-fig. 126 (description; North Bengal: Terai & pattern. North Bengal: Duars; material in ZSI and DNHM). Finally it must be remarked that the species HORA & NAIR, 1941: 388—391 (listed; India: Travancore; recorded Axelrod al. by et (1967) to be T. cut- material in ZSI) (in subgenus Monotretus).

cutia is not that, but T. fangi. FRASER-BRUNNER, 1943: 14 (listed as representative of

and additional sub- genus subgenus; no material) (in

genus Monotretus).

& 1947: 308 BREDER CLARKE, (note on reproduction; no Tetraodon cutcutia Hamilton, 1822. additional material). 3-5. Figs. BENL, 1956: 144—146, 175, 202 (description, aquarium

data; no material).

Tetrodon cutcutia BENL, 1957a: 65, Abb. 4 (in discussion on differences

1822: 8 18 3 HAMILTON, —9, 362, pi. fig. (original descrip- with T. leiurus brevirostris Benl, 1957; no material).

lateral and dorsal view tion, figured; India, West MERCKENS, 1958: 52, 53, 54 (description, aquarium data;

Desh: River Bengal or Bangla Ganges system; no no material).

material known). STERBA, 1959: 634, 637, Taf. 256, 273 (description, 1870: 290 India: Calcutta; material GÜNTHER, (description; aquarium data; no material). in BMNH) (in subgenus Monotretus). HERVEY & HEMS, 1963: 385 (description, aquarium data; DAY, 1878: 703, 182 5 India: Orissa pi. fig. (description; no material).

& India: Assam & Bangla Desh) (in subgenus Mono- Srivastava, 1968: 146—147, fig. 86 (description; India:

tretus). Gorakhpur, Ramgarh Tal; material in ZMUG).

V INC1GUERRA, 1883: 660 (description; Burma: Irrawaddy Monotretus cutcutia;

River material in near Minhla; MSNG). MUNRO, 1955: 282, pi. 55 fig. 820 (description; Ceylon;

1889: 493 and material after DAY, (description Day, 1878). no material).

V 1890: 231 Burma: & INCIGUERRA, (description; Rangoon Monotreta gularis;

Burma: & Burma: Bhamo & Burma: Mandalay LE DANOIS, 1959: 21, 152—153, 246—247, 250—251, 253, material in Meetan; MSNG). 84E nasal discussion fig. (description, organ figured, D'ABREU, 1925: 711 India: River (listed; Ganges system, on osteology, evolution; Ganges River system; River, Ghogra Newajitola). material in MNHN, including MNHN B 1470). RANDOW, 1934: 561—563, Abb. 1 (field observations, LE DANOIS, 1961: 465 (in type list of MNHN, listed as aquarium data; Ceylon; no material). type species for Monotreta). BECK, 1950: 240, fig. 101 (description, aquarium data;

no material). General description. — LADIGES, 1954: 67, fig. 64 (aquarium data; no material).

Tetrodon caria

General morphology. — Body compressed laterally; HAMILTON, 1822: 9—10, 362 (original description, discus- dorsal rectilinear to midst of sion on differences with T. cutcutia; India: Bihar, profile rising nearly

Ganges River system, Kosi River; no material known). back, from there sloping gradually to caudal fin;

Tetrodon gularis interorbital flat; body spines apparently absent; HAMILTON, 1822: 10, 362 (original description, discussion mouth terminal, directing forwards or downwards; on differences with T. cutcutia and T. caria; India, lower border of above River no prominent chin; eye West Bengal or Bangla Desh: Ganges system,

level of mouth border not inter- including Kosi River; no material known). corner, upper

Leiosomus marmoratus fering with dorsal profile; nasal organ a very short 1839: 328 of SWAINSON, (listed as representative genus; no with tube, height 2 or more in diameter, one material) (nomen nudum). terminal opening, at the most only slightly lobed; Tetraödon cutcutia;

skin folds on back or BLEEKER, 1853a: 14, 29, 78—79, 160 (description; India: no belly.

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Figs. 3—5. Tetraodon cutcutia Hamilton, 1822.

Fig. 3. Lateral view of “iconotype” after Hamilton (1822:pl. 18, fig. 3). Dorsal view of Fig. 4. “iconotype” after Hamilton (1822: pl. 18, fig. 3). 5. Nasal of ZMA Fig. organ 110.225, sl 45 mm.

Colour in alcohol. — Ground colourof Hooghly upper parts River; coll. T. Cantor, ca. 1849; Bleeker

lower back with Collection. grey or greenish, parts whitish; SMM 1 si 60 India: Calcutta 23917; specimen, mm; dark cloudy or reticulated markings, sometimes live imported. descending to sides; a light interocular band; sides mostly with a dark meshwork and with a dark Note. — Descriptions were chiefly made from ocellus just in front of dorsal and anal fins; literature. sometimes a spot (red in life) on chin; caudal fin bordered with a darker band (red in life) or all

of the — dusky; other fins plain. Description holotype.

A holotype in the sense of the International Code Morphometries. — Maximum si ca. 95 mm; dorsal of Zoological Nomenclature does not exist, as fin rays 10—11; anal fin rays 10; pectoral fin rays of Hamilton did not leave any known specimen 18—21; head length ca. 2.2 in si; snout length Ham- the species. Generally, however, one accepts ca. 2.2 in hi; mouth width ca. 3.6 in hi. ilton's figures as the "holotype" ("iconotype").

As T. cutcutia is the first species of the cutcutia- Material examined. — and Ham- described in Hamilton, 1822, as ZMA 110.225; 1 specimen, si 45 mm; India: Calcutta, group

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97

ilton is the first author after 1758 describing Klausewitz, 1957b; no additional material).

Indian BENL, 1957b: 1—4 (in discussion on differences with Tetraodon species there can be only slight Carinotetraodon chlupatyi Benl, 1957; no additional doubt about the identity of the species illustrated material).

as T. cutcutia. of a is not Designation neotype BENL & CHLUPATY, 1957: 227—229 (ethological and

necessary at the moment and thus prohibited by morphological comparisons with Carinotetraodon

chlupatyi 1957 and T. werneri the Code. Hamilton's figures (pi. 18 fig. 3) — Benl, provisionary

name; no additional material). reproduced hereby in adaptated form — and his OSTERMÖLLER, 1957: 288 —290, 1 fig. (aquarium data; no description remain the basis for the species. material).

MERCKENS, 1959: 171, 1 fig. (description, aquarium data; Distribution. — T. cutcutia is known with cer-

no additional material). tainty from East India, Ceylon, Bangla Desh, and SONTIRAT et al., 1971: 5—6 (description of living and Burma. preserved specimens, discussion on variability of

colour pattern; Thailand: 50 km S. of Krung Thep;

— Like most of the of Etymology. names species material in FFKU). described by Hamilton, the name cutcutia is Carinotetraodon chlupatyi

derived from local BENL, 1957b: 1—4, 1 fig. (original description of a name, being, according to genus

and species, discussion on colour changes and Hora (1929), "kotkotiya". morphology; Thailand: near Krung Thep; material

living at time of description, at present preserved in Tetraodon lorteti Tirant, 1885. SMM, reg. no. 15419, holotype).

Figs. 6-9. BENL & CHLUPATY, 1957: 227—229, Abb. 1, 2 (aquarium

discussion data, on differences with T. somphongsi

Klausewitz, 1957 and T. werneri Chelonodon Dumerili provisionary name;

emendated type locality: Thailand: 400 km N. of BLEEKER, 1865C: 33 (no description; Thailand; material, Krung Thep; material including described to Le Danois, 1959 & 1961 in holotype according MNHN, reg. alive). no. 2007, from Ayuthia) (doubtful synonym). Tetraodon werneri BLEEKER, 1865d: 172 (listed; no additional material). BENL & 1957: 227—229 Tetrodon Lorteti CHLUPATY, (description, aquarium

not data; explicitly used as a provisionary as TIRANT, 1885: 175 (in ed. 1929: 96) (original description; name,

a new species material). Vietnam: surroundings of Thu dau mot; material in name; no Carinotetraodon MHNL, reg. no. 3907, lectotype and paralectotypes, somphongsi; BENL, 1959: 42—44 (lumping of T. somphongsi Klause- and in MNHN, reg. no. 37—60, paralectotype) (in T. werneri subgenus Crayracion). witz, 1957, provisionary name and Carino- tetraodon Tetrodon borneensis chlupatyi Benl, 1957 based on aquarium

observations concerning individual REGAN, 1902: 303, pi. 24 fig. 3 (original description; variability; emen- dated Thailand: Chao River East Sarawak; material in type locality: Phaya Malaysia: BMNH, reg. no.

Tachin = Nakon Chaisi 38 km N. of 1894.1.20: 16—17, syntypes). system, River, Tetraodon palembangensis; Krung Thep; no additional material). STERBA, 1959: Taf. 276 HORA, 1923b: 183—184 (description of two colour phases; 635—636, (description, aquarium

Thailand: data; no material). Nontaburi; material at least in part in ZSI,

F VAN DER VLUOT, 1968: 218—224, 219, 221 reg. no. 10612/1). figs. p. (de- discussion scription, aquarium data, on colour HORA, 1924: 499, fig. 9a, b (emendated description of changes; Thailand & East two colour phases, both phases figured; Thailand: Malaysia, Sarawak; material im-

Nakon Lampang; material at least in part in ZSI, portedalive). Monotreta caria; reg. no. F 10471/1).

Tetrodon lorteti; Le Danois, 1959: 132, 152—154, 246, 247, 251, 254, fig. 84F, 107, 108 (description; skull, nasal and BONNET, 1927: 367 (publication not seen by me). organ, lateral view discussion CHEVEY, 1932: 31 (listed; no additional material). figured; on osteology, zoogeo- Tetraodon borneensis; graphy, and evolution; no localities mentioned; mate-

rial should include material of Chelonodon FRASER-BRUNNER, 1943: 14 (listed as representative of type

dumerili nudum and genus and subgenus; no additional material) (in sub- Bleeker, nomen T. lorteti).

LE DANOIS, 1961: 466 (in list of MNHN, listed genus Monotretus). type as senior DE BEAUFORT, 1962: 398 (description after syntypes, synonym of Chelonodon dumerili Bleeker,

discussion nomen nudum, on differences with original description; no and T. lorteti; no additional material).

additional material) (in subgenus Monotretus). D'AUBENTON & BLANC, 1966: 555 —556, fig. 1 (description;

Tetraodon somphongsi Cambodia: Prek Tamen, Snoc Trou; material in

KLAUSEWITZ, 1957b: 205—208,Abb. 1, 2 (original descrip- MNHN, reg. no. 66—52).

tion, discussion on ethology, physiological colour Monotreta tiranti and differences changes with T. cutcutia and T. leiurus; D'AUBENTON & BLANC, 1966: 556—558, fig. 2 (original Thailand, live material imported; in SMF, reg. no. description, discussion on differences with T. palem-

4083, holotype, and 4084—4086, paratypes; 2 topo- bangensis; Cambodia: Tonle Sap km 9; material in

still at time of types living description). MNHN, reg. no. 66—48, holotype, and 66—49/51, KLAUSEWITZ, 1957C: 115—116, 2 figs, (compilation of paratypes).

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General description. — of anal fin base; mouth terminal, directed up-

wards; a heavy chin; lower border of eye slightly

General — above level of mouth morphology. Body oblong, compressed corner, upper border not

most at caudal with dorsal nasal short laterally, strongly compressed pe- interfering profile; organ a

duncle; dorsal profile arched, highest at origin cylinder, nearly as high as broad, with one ter-

in minal at the of dorsal fin; interorbital convex, laterally opening, most only scarcely lobed, shrunken specimens with a medial groove; lateral in laterally shrunken specimens situated in a lon- line system indistinct; body spines small, covering gitudinal depression; males often with a ventral back, sides and from nasal end and dorsal median skinfold 1—4 belly organs to (depth mm)

Figs. 6—9. Tetraodon lorteti Tirant, 1885.

6. Lateral view of ZMA Fig. 110.223, sl 42 mm.

Fig. 7. Dorsal view of ZMA 110.223, sl 42 mm.

Fig. 8. Nasal organ of MHNL 3907, lectotype, sl 31 mm.

Fig. 9. Lateral view of lectotype, sl 31 mm.

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between tip of snout and dorsal fin. Anal fin (m 2.7) in si, at end of dorsal fin base 3.2—5.2

sometimes accompanied ventrally by shorter, par- (m 4.3) in si; width maximum 3.0—4.2 (m 3.6)

allel skinfolds on throat. in si and 1.1—1.8 (m 1.4) in maximum depth, at

pectoral fin base 3.2—4.5 (m 3.7) in si and 1.0—

Colour in alcohol. — Dependent upon age, sex and 2.0 (m 1.4) in corresponding depth, at end of

physiological conditions at time of killing. In dark dorsal fin base 5.6—10.6 (m 7.5) in si and 1.3—3.0

colour phase both sexes irregularly dark. Light (m 1.8) in corresponding depth; head length 2.3—

colour phase sexual dimorphic: 2.8 (m 2.5) in si; snout length 2.0—2.9 (m 2.4) in males: colour of Mature Ground upper parts hi; eye horizontal diameter 2.4—4.2 (m 3.3) in hi,

tan, of lower parts lighter, border between both distance from dorsal profile 5.0—17.6 (m 12.1) in

colours running from chin to end of anal fin base; hi; interocular distance 1.5—2.4 (m 1.9) in hi;

band between bordered nasal in a light eyes, posteriorly organ length 10.9—19.6 (m 15.2) hi,

by a trapezoid or triangular dark patch reaching distance from tip of snout 3.3—5.9 (m 4.7) in hi,

to midst of back or snout dark; distance —- beyond; dorsally from eye 4.4—6.7 (m 5.8) in hi and 0.9 dark throat and chin, connected with a patch on 2.0 (m 1.3) in distance from tip of snout; mouth

a dark band; a dark band from posterior width 3.3—5.7 in 6.8— eyes by (m 4.0) hi; upper lip depth

border of eye to origin of dorsal fin, broadening 24 (m 16.0) in hi; lower lip depth 11.3—24 (m

and sometimes branching posteriorly; caudal fin 17.9) in hi.

with narrow white terminal border, preceded by

Material examined. —• a dark transversal band of varying width some- BMNH 1894.1.20: 16—17; 2 specimens, si 43, 44 mm; times the of caudal occupying larger part fin; East Malaysia: Sarawak; coll. Bartlett (syntypes of T. other fins plain. borneensis Regan, 1902). Females and juveniles: Patches and bands like BMNH 1904.7.25: 49; 1 specimen, si 53 mm; West

in males; back and sides posterior of connecting Malaysia: Kuala Lumpur, Selangor; ex SM.

BMNH 1906.10.29: 41—42; 2 specimens, si 38, 43 mm; band between eye and dorsal fin reticulated; caudal East Malaysia: Sarawak, Sibu; coll. C. Hope. fin reticulated or densely spotted with dark, with- MHNL 3907; 13 specimens, si 20—31 mm; see lecto- out dark band; dorsal and anal fins with a a type description and paralectotypes description. dark rounded their of spot at bases; light parts MNHN 37—60; 1 specimen, si 25 mm; see paralecto-

ventral surface often with few small dark types a spots description. MNHN si 56—41; 1 specimen, 31 mm; Indochina; coll or longitudinal stripes. Arnoult.

MNHN 66 si —52; 1 specimen, 30 mm; Cambodia: Prek

Note. — Colour in life is strikingly different Tamen, Snoc-Trou; coll. F. d'Aubenton, 12-VII-1962.

(Klausewitz, 1957b; Benl, 1957b), the most strik- MNHN A 5072; 6 specimens, si 39 —50 mm; Thailand:

Chao Phaya River system; coll. Harmand. ing differences being red (males) or partly reddish MNHN A 5099; 1 specimen, si 40 mm; Thailand; coll. (females) dorsal and anal fins and a red ventral Harmand.

median stripe in many specimens. MNHN A 5100; 1 specimen, si 40 mm; Thailand;

coll. Harmand.

SMM 1 si 52 — si — 15419; Thailand; Morphometries (based on 42 specimens). 20 specimen, mm; imported alive, 29-1-1957; died in aquarium of P. Chlupaty, Mün- 54 (m 36) mm; tl 26—68 (m 45) mm; dorsal fin chen, V-1958 (holotype of Carinotetraodon chlupatyi rays 11—13 (m 12.0) of which 1—4 (m 2.7) un- Benl, 1957). branched, longest (the 6 —10th) 6.4—9.1 ray ZMA 110.223; 112 specimens, si 28 —54 mm; Thailand;

(m 7.6) in si, shortest ray (the 1st) 10.8—21 (m imported alive; died 1965. ZMA 110.224; 1 si 15.0) in si, base 5.8—9.8 (m 7.1) in si and 0.7—1.2 specimen, 43 mm; imported alive; died in NAM, 1963. in anal fin 10—12 (m 0.9) longest ray; rays (m Thailand: ZSI F 10471/1; 1 specimen, si 28 mm; 11.1) of which 1—3 (m 2.1) unbranched, longest Nakon Lampang; coll. N. Annandale. (the 6—9th) 6.8—9.1 in si, shortest ray (m 7.7) ray ZSI F 10612/1; 1 specimen, si 39 mm; Thailand: (the 1st) 10.3—17.6 (m 14.4) in si, base 7.6—12.9 Nontaburi; coll. M. Smith, 10-XII-1923.

(m 9.2) in si and 0.9—1.6 (m 1.2) in longest ray;

pectoral fin rays 14—16 (m 15.0) of which 1—2 Description of the lectotype. —

(m 1.3) unbranched, longest ray (the 5—7th) 7.4

in si, shortest — —11.1 (m 9.2) ray (the 1st) 8.3—16.7 Designation. There can be no doubt about the

in base —11.6 si series used for (m 12.7) si, 7.7 (m 8.8) in and selecting a lectotype being the

0.8—1.3 in maximum series of Tirant, as it is (m 1.0) longest ray; depth original syntypical pre-

2.0—3.0 (m 2.7) in si, at pectoral fin base 2.0—3.2 served in Lyon (MHNL), the place where he

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worked. all be males. Paris (MNHN) only possesses a single specimens to specimen, donated by MHNL. The lectotype was

chosen out of the series on base of having the Morphometries. — si 20—29 (m 25) mm; tl 26—36

of greatest si. (m 31) mm; dorsal fin rays 12—13 (m 12.3)

which 3—4 (m 3.3) unbranched, longest ray (the

General morphology. — In spite of the small size 8 —9th) 6.0—9.0 (m 7.6) in si, shortest ray (the and the bad state of preservation, all characters 1st) 12.5—19.1 (m 15.6) in si, base 5.8—7.7 (m

mentioned in the general description — including 6.7) in si and 0.7—1.1 (m 0.9) in longest ray; anal

skinfolds — middorsal and midventral are present. fin rays 11—12 (m 11.4) of which 1—3 (m 2.5)

—9.1 unbranched, longest ray (the 6—9th) 7.0

Colour in alcohol. — Due to bad preservation, (m 7.9) in si, shortest ray (the 1st) 13.8—17.3 (m colour pattern only on lateral parts of head and 15.8) in si, base 7.7—10.0 (m 9.0) in si and 0.9— dorsal fin; remains of black, white 1.4 in fin 15—16 from there to (m 1.2) longest ray; pectoral rays bordered band caudal fin the 7.4—10.7 in on determine lecto- (m 15.1), longest ray (m 9.0) si, type to be a male. shortest ray (the 1st) 8.3—12.4 (m 10.7) in si, base

7.7—11.1 (m 9.0) in si and 0.8—1.2 (m 1.0) in

— tl dorsal fin 2.5 in si, Morphometries. si 31 mm; 38 mm; longest ray; depth maximum —2.9 (m 2.7)

iii.10, 4.5 mm in at fin base 2.6—2.9 in si, at end rays longest ray (the 9th) (6.9 si), pectoral (m 2.8) base dorsal in width shortest ray (the 1st) 2.2 mm (14.1 in si), of fin base 3.9—5.2 (m 4.5) si;

in si and in fin in si and —1.5 4.4 mm (7.0 1.0 longest ray); anal maximum 3.5—4.2 (m 3.9) 1.2 (m

3.7 in in maximum at fin base 3.5— rays ii.9, longest ray (the 8th) mm (8.4 si), 1.4) depth, pectoral

shortest ray (the 1st) 2.8 mm (11.1 in si), base 4.4 (m 3.9) in si and 1.2—1.5 (m 1.4) in corre-

3.7 mm (8.4 in si and 1.0 in longest ray); pectoral sponding depth, at end of dorsal fin base 7.0—9.6 fin 3.0 in in si and 1.6—2.1 in rays 15, longest ray (the 5th) mm (10.3 (m 8.3) (m 1.8) corresponding

si), shortest ray (the 1st) 2.5 mm (12.4 in si), base depth; head length 2.3—2.6 (m 2.5) in si; snout

2.8 in and 1.1 in 2.4—2.9 in horizontal dia- mm (11.1 si longest ray); depth length (m 2.6) hi; eye

fin base 2.4—3.3 in from maximum 11.1 mm (2.8 in si), at pectoral meter (m 2.9) hi, distance dorsal

in end of dorsal fin base 8.5—21 in interocular distance 10.6 mm (2.9 si), at profile (m 12.6) hi;

7.0 mm (4.4 in si); width maximum 7.4 mm (4.2 1.9—2.4 (m 2.2) in hi; nasal organ length 10.9— in si and 1.5 in maximum depth), at pectoral fin 15.0 (m 13.0) in hi, distance from tip of snout 4.0

base (4.4 in si and 1.5 in —5.9 in distance from 5.2—7.0 7.0 mm corresponding (m 4.7) hi, eye

depth), at end of dorsal fin base 3.5 mm (8.8 in (m 5.9) in hi and 1.1—1.5 (m 1.3) in distance

si and 2.0 in corresponding depth); head length from tip of snout; mouth width 5.2—5.7 (m 5.5)

in 5.1 in hi. 12.6 mm (2.5 si); snout length mm (2.5 in

hi); interocular distance 5.6 mm (2.3 in hi); eye

in distance — horizontal diameter 3.9 mm (3.2 hi), Material examined. MHNL 3907 12 si 20—29 dorsal in nasal A-L; specimens, mm, para- from profile 0.6 mm (21 hi); organ lectotypes; South Vietnam: surroundings of Thu dau mot length 0.7 mm (18 in hi), distance from tip of River; coll. G. Tirant, 1897. snout 2.4 in distance from 2.1 mm (5.3 hi), eye 37 1 si 25 MNHN —60; specimen, mm; paralectotype; distance from of mm (6.0 in hi and 1.1 in tip South Vietnam: surroundings of Thu dau mot River; coll.

to be the Le snout); mouth width 2.2 mm (5.7 in hi). Gaillard, MHNL (considered holotype by Danois, 1961).

Material examined. —

MHNL 3907 1 si 31 T. lorteti is known from ex parte; specimen, mm, lectotype; Distribution. Thailand,

mot River; South Vietnam: surroundings of Thu dau Cambodia, South Vietnam and East and West

coll. G. Tirant, 1897. Malaysia. Thus, it is geographically isolated from

the other members of the cutcutia-group, T. Description of the paralectotypes. — travancorius living in India, Travancore, T. cut-

cutia living in East-India, Burma, Ceylon and — inter- General morphology. Faint groove on Bangla Desh. orbital always present; body spines always dis-

tinct; otherwise like general description.

— — writ- Etymology. The name lorteti originally

— Lorteti — Tirant in honour of Colour in alcohol. Like lectotype or even worse ten was given by the time de la Faculté preserved; caudal fin colour pattern determines Dr. L. Lortet, at Doyen

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de Médicine de Lyon and Directeur du Muséum only distalmost part of which is formed into two

de Lyon. very small lips which are bent inwards, giving

a key-hole appearance to opening of nasal organ.

Colour in alcohol. — Ground colour of upper parts Tetraodon travancorius Hora & Nair, 1941. grayish, of lower parts much lighter; usually two Figs. 10-12. of black, oval patches on upper lateral surface

body in front of dorsal fin, the situated Tetraodon travancorius patches in much of HORA & NAIR, 1941: 391—393, figs. 3—4 (original a lighter area; posterior patches a

discussion description, and figures on variation of dark, broad band running to caudal fin, partly colour discussion differences with T. pattern, on central dark in continuing on rays; usually a spot cutcutia; India: Travancore, Pamba River; material in the middle of course of band; other dark spots ZSI, reg. no. F 13601/1, holotype and paratypes) (in at base of caudal fin and at base of subgenus Monotretus ). posterior-

MENON & 1963: 162 list of dorsal YAZDANI, (in type ZSI; no most two fin rays; a dark patch above additional material) (in subgenus Monotretus). pectoral fin and a spot behind it; back with a

narrow, light interocular band, two irregular dark Description of holotype and paratypes (after Hora patches posterior to V-shaped marking behind

& Nair, 1941; no material examined). — patches, an irregular band in front of dorsal fin

and triangular patches in front of or behind dorsal

General morphology. — Body oblong, compressed fin; fins plain. laterally; dorsal profile arched, highest at midst of back; interorbital flat; body spines inconspicu- Morphometries (based on 5 specimens). — si 15.7 ous; skin reticulated finely; origin of anal fin — 17.0 (m 16.5) mm; tl 20.8—22.0 (m 21.4) mm; situated beneath anterior third of dorsal fin base; dorsal fin rays 7—8 (m 7.8), longest ray 6.5—8.0 mouth terminal, directed forwards; lower border in anal fin fin (m 7.3) si; rays 8; pectoral rays of below level of mouth 16—17 (m 2.2—2.6 (m 2.4) in si; eye slightly corner, upper 16.8); depth border not interfering with dorsal profile; nasal head length 2.3—2.5 (m 2.4) in si; eye diameter

hollow hi. organ a cylinder, nearly as high as wide, 2.4—2.7 in

Figs. 10—12. Tetraodon travancorius Hora & Nair, 1941.

Fig. 10. Lateral view of ZSI F 13601/1, holotype, after Hora & Nair (1941: fig. 3b).

Fig. 11. Dorsal view of ZSI F 13601/1, holotype, after Hora & Nair (1941: fig. 3a).

12. Nasal of ZSI F after Hora & Nair Fig. organ 13601/1, holotype, (1941: fig. 3c).

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Material examined by Hora & Nair. — deeper dermal pits in T. palembangensis. ZSI F 13601/1; 1 specimen, holotype; India: Travan- Le Danois (1959) considers T. leiurus merely core, Pamba River; coll. K. B. Nair. the male of a sexual dimorphic Monotreta cut- ZSI F 13601/1; 9 specimens, paratypes; India: Travan- cutia palembangensis. She does not give core, Pamba River; coll. K. B. Nair. any evidence for this opinion other than the exami-

— This has hitherto been Distribution. species only nation of "les types des Bleeker: Arothron pa-

recorded from the India: in type locality: Travancore, lembangensis et Tetraodon leiurus”, present accord- Pamba River, where it occurs abundantly MNHN. According to her type list (1961) these

of the K. B, ing to the collector type material, "types" should be MNHN 2310: paratype of A.

Nair. palembangensis, 160 mm; and MNHN 03-184:

As the first paratype of T. leiurus, 130 mm. species

— The named after the Etymology. species was was described originally by Bleeker from a single

India: Travancore. type locality, specimen, tl 210 mm, and the second species from

indication Le 5 specimens, tl 60-98 mm, the by

Danois is incorrect and her opinion on sexual

leiurus-group dimorphism thus has no ground.

nasal Diagnosis. Tetraodon species with the or- In 1902 Vaillant described a new species, close

hollow which is gan formed by a cylinder distally to T. palembangensis: T. pinguis. After compar-

one-third with divided into two lips over to one-half ing the single central Borneo specimen a of its length; head depressed; body spines often single T. palembangensis specimen he found the

embedded in a complicated papillated dermal following discriminating characters: a wider and

Three skin structure; D 12-15; A 10-12; P 21-24. shorter snout, bigger papillae, fins more fatty,

T. T. leiurus and fin D 9; A 9. In the holo- species: palembangensis, T. ray counts examining

fangi. type of T. pinguis I could not find any difference

with 23 T. palembangensis specimens, apart from

The leiurus- is fins indeed The Discussion. group intermediate the being more fatty. snout length

between the which has the nasal the value for cutcutia-group, even proved to equal exactly mean

cylinder not or scarcely two-lipped, and the fluvia- T. palembangensis, viz. 2.8 in hi.

tilis-group with its cylinders often deeply divided De Beaufort (1962) rejects the opinion of T.

distinct into flaps. Differences with the erythrotaenia- pinguis being a species. He, however, con- group include the fin ray counts, the form of firms the original description in stating that there

and that there the dorsal profile, and the colour pattern. The is only one lateral spot on each side, dorsal fin difficult group was considered a single species by Le are 9 rays, although they are to

indicates that the lateral Danois (1959). Her misuse of the name cutcutia count. My own experience

for this species is discussed with the cutcutia- spots are 4 (left side) plus 3 (right side) and X-ray

group. photographs showed 13 dorsal fin rays, hence normal for T. palembangensis.

The oldest of the Le Danois be mistaken when names proposed for members (1961) must

group are T. leiurus Bleeker, 1851, and T. palem- claiming two (sic) holotypes of T. pinguis for

bangensis, Bleeker, 1852. Both species were fig- MNHN (reg. no. 91-213, 214) as only one specimen

ured in Bleeker, 1865a. It should be noted that was originally described. Moreover, the type lo-

T. drawn from is Borneo: Mahakam the MNHN the figure of leiurus is a non- cality River,

River. typical specimen, as it shows many round, ocel- specimens originating from Borneo: Kapuas

lated spots, while all Bleeker material and other

The six in the be material examined show the dark lateral spots remaining names group can

more or less polygonal. Nevertheless, the difference ascribed to the closely allied species T. leiurus

in colour pattern between the two Bleeker-species Bleeker, 1851 and T. fangi Pellegrin & Chevey, 1940. The number of is is striking, T. leiurus always possessing many synonyms not surprising,

lateral dark the of the colour spots, leaving only a whitish network, considering great variability

T. this consists of palembangensis always showing a wide-meshed pattern. Basically an irregularly back of rounded lateral network, some of the meshes enclosing dark coloured and a great number or

the spots or ocelli. Other differences are a slightly polygonal dark spots on sides. Variations on 1-3 lower dorsal this include a interocular band, fin ray count, a shorter snout and pattern light

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dark dorsal and the of ocellus be due, De Beaufort, to the patches presence an as pointed out by ventral to the origin of the dorsal fin. Benl (1957a) same cause which made the skin form pustules

and claimed several other authors in aquarium journals on head and body. T. hilgendorfii was found the dorsal colour to with mood, to be different in different head pattern vary having a length, and the lateral ocellus to vary with age, at least a lower and more anteriorly placed eye, and the

in Klausewitz the bel- T. fangi. (1957a) found geo- lateral spots more elongate, descending to graphical variation of the dorsal pattern in the ly. Head length in 5 out of the 6 syntypes proved same species. A very striking colour variation is to be 2.3 in si, hence normal again. The slight found in the Cambodian differences in colour be of type specimens of T. pattern cannot system- cambodgiensis Chabanaud, 1923. Instead of a atic value, as pointed out before.

also greyish-tan dorsal ground colour with or without T. cambodgiensis Chabanaud, 1923, can dark retained author patches they show many oval or polygonal not be as a separate species. The whitish spots, leaving a dark network only. This gives as discriminating characters that the body dorsal pattern descends for some length along the spines are retractile and provided with a fleshy

that the lateral line the sides, which otherwise, at first glance, only are lobe, system encircles eye, marked by a big ocellus. The dark network, how- and that the snout length exceeds the distance ever, could also be found in some specimens from eye to gill opening. In my experience all

otherwise coloured well, al- among normally series from these characters apply to T. leiurus as

Singkarak (Sumatra) and Bogor (Java). One of though not in all specimens. Other characteristics the Bogor specimens (si 74 mm) gives extreme counted and measured by me did not show im-

of the of colour differences. values in cambod- proof unemployability pattern in portant Mean 4 T.

T. leiurus the black- D A P by possessing characteristic giensis syntypes were 14.0; 11.0; 23.8;

leiurus- spot pattern on the left side and parts of snout length 2.2 in hi; head length 2.6 in si. For the and leiurus P belly the characteristic white-spot cam- T. they are D 13.8; A 11.0; 23.0; snout bodgiensis pattern on the right side and the back. length 2.3 in hi; head length 2.5 in si. value The of the lateral ocellus as a systematic character was discussed e.g. by De Beaufort (1962).

In this discussion it is stated that the specimens The third species recognized in the leiurus- recorded Hora Deli by (1923b, 1924), and a (Su- group, T. fangi, was described by Pellegrin & matra) specimen in ZMA laterally only are marked Chevey (1940) from Vietnam. Benl (1957a) de- by an ocellus. This is not true: the specimens scribed the same species from aquarium imported show a faint indicationof the polygonal dark spots specimens as T. leiurus brevirostris, and in the

the The be for described it near ocellus. same proves to true same year Klausewitz (1957a) as two of the cambodgiensis syntype specimens. When T. ocellaris. Probably both Benl and Klausewitz examining 71 specimens from 19 series it became overlooked the publication of Pellegrin & Chevey clear that all intermediates between absence because in view there not be doubt occur my can any and presence of the ocellus, the frequency of pres- of the conspeoifity of the three nominal species.

to the south-west of & did not differences ence decreasing apparently Pellegrin Chevey give any

Borneo. with T. leiurus. They only compared T. fangi with

Having rejected colour pattern as a splitting T. patoca Hamilton, 1822, a species which can character in Bleeker's concept of T. leiurus, the easily be recognized by the different form of the problem has become clearer. In 1905 Popta con- nasal organ. tributed nominal from Klausewi'tz two species Borneo: T. only gives the presence of the lateral

and T. The taxonomie value ocellus character. bergii hilgendorfii. as a discriminating As pointed of both was discussed by De Beaufort (1962). out above, this can not be an important specific

character. The According to Popta T. bergii differs from typical characteristic T. fangi colour pat-

in head inter- also in T. leiurus. leiurus a different length, a wider tern occurs some typical Benl orbital, a lower position of the eyes and the colour gives three differences for T. fangi: less spinöse,

below. Ex- shorter and interorbital being plain tan above, plain cream a snout, a greater width amination of the unique specimen shows the head expressed in the distance between the tip of the

and the length to be 2.3 in si (exactly the mean value snout interocular band. Spinosity, however, for the interorbital in hi is doubtful T. leiurus), being 1.8 a character, as it usually is in the

(T. leiurus 1.7-2.2), and the position of the eyes Tetraodontidae in general and in small specimens being normal. The different colour pattern could in particular (as most T. fangi specimens are).

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Fig. 13. Relation between relative snout length and standard length in Tetraodon leiurus, showing discontinuity

from standard between T. fangi and the three other forms and showing independence of relative snout length

length (age).

Thailand: Small specimens always are less spiny than bigger near Patalung; Vietnam) they suggest

specimens (as most T. leiurus specimens are). that the species could be sympatric with T. leiurus

There difference in the Thailand: mouth of River and is however, a papillose (e.g. Patalung

structure hiding the spines, as this is more Patalung Lake; Cambodia).

elaborate in T. fangi (figured in Klausewitz The pioture given above of differences between

The T. leiurus and T. is obscured 1957a), but this is only a relative difference. fangi slightly by their quotient used by Benl was found by him to be the syntypic material of T. hilgendorfii as

1.30 and 1.31 in two T. fangi specimens compared relative snout length is slightly intermediate (see

to 0.90-1.17 (m 1.03) in 7 leiurus specimens fig. 13). I therefore hesitated whether to consider

T. reunite examined. My own measurements yielded ratios of hilgendorfii a separate subspecies or to

1.0-1.4 (m 1.15) in 15 T. fangi specimens and T. leiurus and T. fangi. The final argument in

T. hil- 0.8-1.2 (m 1.01) in 36 T. leiurus specimens, so my considerations was a Zoogeographie one: rich the difference is not striking. gendorfii was collected in East Borneo, a

of This leaves only one discriminating character: source biological variability, as was also the

length and form of the snout. Length of the snout case in species of the fluviatilis-group. So, I con-

sider T. a of T. leiurus. proves to be very constant as can be seen from hilgendorfii junior synonym for the sake of the material fig. 13. Snout length ratios given in the literature, However, certainty T. which are probably taken with methods different was not incorporated in the descriptions of

leiurus. from those employed here, even indicate values up to 3.3 in hi (Fowler, 1937). Fig. 13 also shows that the likely assumption of T. fangi being the

of T. leiurus is improbable: the relative young Tetraodon palembangensis Bleeker, 1852. snout is snout length independent of age. The short Figs. 14-15. and the of the presence ocellus proved constant not only in the specimens examined, but also in Tetraodon palembangensis the specimens referred to in the aquarium jour- BLEEKER, 1852a: 21, 25, 26 (original description, dis- nals. cussion on differences with T. reticulatus Bleeker, 1849

and T. lineatus Indonesia: Sumatra, An argument for giving T. fangi a different Bloch, 1785; holotype not recognizable within Bleeker rank instead of Palembang; specific a subspecific rank may Collection). be found in its geographical distribution. Al- BLEEKER, 1852C: 37, 38 (description and discussion after a few exact localities are known though only (e.g. Bleeker, 1852a; no additional material).

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BLEEKER, 1853e: 429, 435, 438 (listed; Indonesia: Borneo, Tetraodon palembagensis;

Pontianak; material not recognizable within Bleeker VAILLANT, 1893a: 36, 61 (listed; Indonesia: Borneo,

Collection). Kapuas River; material in MNHN, reg. no. 91—213/

Arothron palembangensis; 214).

BLEEKER, 1854b: 22, 28 (listed; Indonesia: Sumatra, VAILLANT, 1893b: 2 (listed; no additional material).

Muarakumpe; material not recognizable within Blee- Tetraodon pinguis

ker Collection). VAILLANT, 1902: 7, 24, 28, 38, 40 (original description,

additional discussion differences with BLEEKER, 1854d: 260 (listed; no material). on T. palembangensis;

at BLEEKER, 1855a: 9 (listed; no additional material). Indonesia: Borneo, banks of Mahakam River Tepu;

Indonesia: BLEEKER, 1859g: 340 (listed; Sumatra, Palem- material in RMNH, reg. no. 7928, holotype).

bang; material not recognizable within Bleeker Col- Tetraodon palembangensis;

lection). VAILLANT, 1902: 12, 14, 16, 28, 40 (listed, discussion on Indonesia: BLEEKER, 1859h: 4, 10 (listed; Indonesia: Sumatra, Musi differences with T. pinguis Vaillant, 1902;

material not at at Sintang and at River system near Palembang; recogniz- Borneo, Kapuas River Pontianak,

able within Bleeker Collection). Semitau; material in RMNH, reg. no. 7926-27).

? Arothron palembangensis; FOWLER, 1905: 507 (listed; Indonesia: Borneo, Kapuas

BLEEKER, 1857C: 12 (listed; no additional material). River) (doubtful reference). material). BLEEKER, 1858a: 7, 30, 35 (listed; Indonesia: Sumatra, SMITH, 1945: 576 (description; no additional

Palembang; material not recognizable within Bleeker TWEEDIE, 1952: 89 (description; West Malaysia: Kuala

Collection). Tahan & West Malaysia: Tasek Bëra; material in

1859a: 201 no additional material). BLEEKER, (listed; NMS, reg. no. 2316, 2317).

BLEEKER, 1859f: 266 (listed; Indonesia: Sumatra, Djambi DE BEAUFORT, 1962: 392—394 (description, discussion on

River; material not recognizable within Bleeker differences with T. pinguis Vaillant, 1902 and T .

additional Collection). palembangensis cf. Hora, 1923 & 1924; no

BLEEKER, 1860a: 9, 66 (listed; no additional material). localities) (in subgenus Monotretus).

BLEEKER, 1860c: 5, 7 (listed; Indonesia: Borneo, Sintang; Tetraodon pelambangensis;

material not recognizable within Bleeker Collection). FRASER-BRUNNER, 1943: 14 (listed as representative of

200 no additional material). and subgenus; no additional material) (in BLEEKER, 1860g: (listed; genus

Crayracion palembangensis; suhgenus Monotretus).

BLEEKER, 1865a: 66, 67, tab. 208 fig. 3 (emendated Monotreta cutcutia palembangensis;

differences with description, figured, discussion on LE DANOIS, 1959: 20, 152, 154, 156, 157, 246, 247, 251,

T. leiurus Bleeker, 1850; no additional localities). 254, figs. 84d, 112, 113, 114 (description, lateral view,

additional skull and nasal discussion BLEEKER, 1866: 36 (listed; no material) frontal view, organ figured, osteology and evolution; Tetrodon palembangensis; on sexual dimorphism, T. GÜNTHER, 1870: 288 (description; Indo-Malayan Archi- material in MNHN, should comprise types of

in pelago & Thailand & unde; material BMNH, reg. palembangensis, T. leiurus and T. pinguis Vaillant,

latter series no. of first series 1867.11.28.115, un- 1902, see holotype description). material registered) (in subgenus Crayracion). LE DANOIS, 1961: 466 (in type list of MNHN,

KAROLI, 1882: 187 (listed; Thailand; material in MNH, supposed to be types of T. leiurus, T. palembangensis

material not see holotype reg. no. 1606) (doubtful reference, and T. pinguis Vaillant, 1902, descrip-

available for study). tion).

WEBER, 1894: 458 (listed; no material). D'AUBENTON & BLANC, 1966: 557, 558, fig. 3 (description d'Aubenton & Blanc, 1966, VOLZ, 1903: 413 (listed; Indonesia: Sumatra, Banjuasin in discussion on M. tiranti

River; material probably in NMB). figured after Bleeker, 1865a; Laos: Luang Prabang; VOLZ, 1904: 483 (listed; Indonesia: Sumatra, Inderagiri material in MNHN).

River system & Inderagiri River at Prawap &

Sumatra, Wampu-Selapian River; material probably in General description. — NMB).

VOLZ, 1906: 234 (listed; no additional material).

General — Body oblong, anteriorly WEBER & DE BEAUFORT, 1912: 541 (listed; Indonesia: morphology. Sumatra, Kampar Kiri River at Gunung Sahilan & compressed ventrodorsally; dorsal profile rooflike

Sumatra, Inderagiri River at Taluk; material in ZMA, between often or arched, highest point pectorals 108.957 109.000 reg. no. and respectively). with bumplike; interorbital concave or flat, often CHEVEY, 1932: 31 (listed; unde; no material) (doubtful median lateral line system very con- reference). a groove; small, hidden under HARDENBERG, 1936: 254 (listed; Indonesia: Borneo, middle spicuous; body spines papillae

material probably lost). course of Kapuas River; in rounded dermal pits; dermal pits very con- Indonesia: Java DE BEAUFORT, 1939: 194 (listed; Sea, and spicuous and densely covering back, sides, Island; material in ZMA, reg. no. 108.968) Belitung fin belly between nasal organs and anal base; (in subgenus Crayracion). themselves only conspicuous if body inflated; Tetrodon palembangensisVariété spines and often TIRANT, 1885 (in ed. 1929): 95 —96 (description, discus- skin with dermal muscles thick, on belly

with T. sion on differences typical palembangensis on other parts of body loosely attached; origin S. Vietnam: and on lack of material for comparisons; of anal fin situated beneath posterior half of dorsal Thu dau mot; material not in MHNL) (in subgenus fin base; mouth terminal directed forward; lower Crayracion).

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Figs. 14—15. Tetraodon palembangensis Bleeker, 1852. Fig. 14. Lateral view after Bleeker (1865a: tab. 208, fig. 3).

15. Nasal of ZMA sl 117 Fig. organ 101.852, mm.

border of eye at or below level of mouth corner, base 5.8-8.5 (m 6.8) in si; width maximum 2.1-

dorsal in si in maximum upper border sometimes interfering with 3.2 (m 2.5) and 0.7-1.1 (m 0.9)

tentacle, less than distal fin base 2.1-2.9 in si profile; nasal organ a depth, at pectoral (m 2.6)

third of which is divided into two narrow lobes; and 0.7-1.0 (m 0.9) in corresponding depth, at

apposed surfaces of lobes smooth. end of dorsal fin base 5.8-9.1 (m 7.6) in si and

0.8-1.4 (m 1.1) in corresponding depth; head

length 1.9-2.2 (m 2.1) in si; snout length 2.7-3.1 Colour in alcohol. — Ground colour of upperparts in hi; interocular distance 1.7-2.5 tan, of belly yellowish or greyish white; back with (m 2.8) (m 2.1) in hi; horizontal diameter 4.5-9.3 in obscure mixture of lighter and darker in which eye (m 6.3) hi; nasal length 12.4-31 (m 18.3) in hi, sometimes a light interocular band or a dark organ

of head be distance from tip of snout 3.3-5.7 (m 3.9) in hi, triangular patch on posterior part can distance from 8.0-11.8 9.8) in hi and 1.8- recognized; sides and belly with a dark meshwork, eye (m

meshwork 4.5 (m 2.6) in distance from tip of snout; mouth most distinct and regular on belly; on width 2.9-4.6 in 9.8- sides has been described as light vermiculating (m 3.6) hi; upper lip depth 18.0 13.8) in hi; lower 12.2-40 (m lines and streaks; sides with 3-18 rounded spots, (m lip depth 21) in hi. darker than meshwork, in meshes if present giving

impression of ocelli, diameter 14-106 (m 35) in si;

Material examined. — diameter of meshes on 15 in si; average belly BMNH 1867.11.28: 115; 1 specimen, si 167 mm; Indo- caudal fin or distally sometimes darker; proximally Malayan Archipelago; Bleeker Collection. other fins plain. BMNH 1887.2.28: 3—4; 2 specimens, si 177—182 mm;

Malaysia: Malaya, Perak; coll. J. Anderson.

BMNH 1922.5.19: 114; 1 specimen, si 194 mm; Morphometries (based on 23 specimens). — si Malaysia: Malaya, Jahan River; coll. Kloss. 36-194 mm; tl 45-232 mm; (m 119) (m 141) BMNH 1 si 79 coll. no reg. no.; specimen, mm; unde; dorsal fin 12-14 of which 1-3 rays (m 12.7) (m College of Surgeons.

si Thailand: Thale 1.8) unbranched, longest ray (the 7-10th) 8.3-11.0 NMB 5176; 1 specimen, 147 mm;

coll. 1937. in shortest 18-25 Luang; H. Bernatzik, (m 9.5) si, ray (the 1st) (m 21) RMNH 7341; 5 specimens, tl 150—220 mm; Indo- in si, base 8.7-12.6 (m 10.7) in si and 0.9-1.3 Malayan Archipelago; Bleeker Collection. (m 1.1) in longest ray; anal fin rays 10-12 (m 10.9) RMNH 7926—27 in part; 1 specimen; Indonesia: of which 1-2 (m 1.3) unbranched, longest ray (the Borneo, Pontianak; coll. Moret, 1895.

RMNH 7926 —27 in 2 specimens; Indonesia: 7-8th) 8.7-10.5 (m 9.7) in si, shortest ray (the 1st) part; Borneo, Sintang; coll. J. Büttikofer, 1894. 15-49 (m 23) in si, base 12.5-17.7 (m 14.1) in si and RMNH 7926—27 in part; 4 specimens; Indonesia: 1.2-1.9 (m 1.5) in longest ray; pectoral fin rays Borneo, Kapuas River at Semitau; coll. J. Büttikofer, 21-23 (m of which 1-2 22.0) (m 1.7) unbranched, 14-XII-1893.

8.3-11.6 in short- si Indonesia: longest ray (the 6-9th) (m 10.0) si, RMNH 7928; 1 specimen, 182 mm;

banks of Mahakam River at Tepu; 4th coll. cut ray (ths 1st) 20-33 (m 24) in si, base 7.1-10.3 Borneo, A. W. Nieuwenhuis, 1896—1897 (holotype of T. pinguis in si and 0.7-0.9 (m 8.2) (m 0.8) in longest ray; Vaillant, 1902). maximum 2.1-3.5 in si, at depth (m 2.7) pectoral Indonesia: ZMA 101.852; 2 specimens, si 117—122 mm; fin base 2.3-3.5 at end of dorsal fin (m 2.9) in si, Sumatra, Palembang; coll. Salm, 16-XI-1908.

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the fact that the 5 ZMA 108.956; 1 specimen, si 150 mm; Indo-Malayan This view is supported by

Bleeker Collection. Archipelago; specimens must represent the most numerous lot of ZMA 108.957; 1 specimen, si 36 mm; Indonesia: the auction, being the A-lot, generally believed to Sumatra, Gunung Sahilan; coll. J. P. Kleiweg de Zwaan, contain the There does not, however, exist a 1907. types.

of which of the two is ZMA 108.968; 1 specimen, si 124 mm; Indonesia: Java way determining specimens

1937. Sea, Belitung Island; coll. Kuiper, the holotype. Moreover, two specimens auctioned

ZMA 108.999; 5 specimens, si 85—137 mm; Indonesia: in 1879 could not be traced and theoretically one Sumatra, Djambi River; coll. Moolenburgh, 1909. of them might be the holotype. ZMA 2 si 96—102 Indonesia: 109.000; specimens, mm;

Sumatra, Inderagiri River at Taluk; coll. J. P. Kleiweg de Zwaan, 1907. Given the above facts, the holotype can not be

Indonesia: ZMA 110.195; 1 specimen, si 133 mm; indicated. The International Code of Zoological Java, Bay of Djakarta; coll. C. P. Sluiters. for this Nomenclature does not carry a solution ZMA 110.196; 1 specimen, si 69 mm; Indonesia: situation. A lectotype cannot be chosen because Borneo, Putussibau; coll. H. A. Lorentz, 29-VI-1909.

ZMA 1 si 79 Indonesia: there is no series, while a selec- 110.197; specimen, mm; syntypical neotype

26-VI-1909. Borneo, Bunut; coll. H. A. Lorentz, tion is prohibited by art. 75. Whitehead etal. (1966)

nevertheless selected neotypes in comparable ca-

of the — Description holotype. ses. In this case I choose not to do so because the

examined homo- 7 Bleeker specimens I are very

Recognition. — Bleeker based his original descrip- geneous, in both morphology and colour pattern, tion 1 tl 210 from Indonesia: on specimen, mm, and perfectly match the figure given by Bleeker

Sumatra, Palembang, fresh water. Apparently of the Bleeker in his Atlas Ichthyologique, so any Bleeker mixed the specimen at a certain time with specimens can be used as a safe base for defining of which he, according to his conspecific material, the species. Atlas Ichthyologique, in 1865 in total possessed 9

tl 101-203 mm. After his death the same specimens, Distribution. — T. palembangensis is known from number of was auctioned in 1879. They specimens Malaya, Thailand, Laos, and Borneo. The record were divided in four lots of 6, 1, 1, and 1 spec- from Indonesia: Java, Bay of Djakarta seems

the first of which most was imens, probably doubtful, as all other records are from fresh

RMNH, the second by the precursor of bought by water and the specimen was sent from the Bogor ZMA. Museum which at the time was not always accu-

rate when labelling.

In search of the holotype I could trace 8 spec- imens of the Bleeker Collection, viz.: Etymology. — The name palembangensis refers to

the type locality, Sumatra: Palembang, from where

the also often recorded after the — BMNH 1867.11.28: 115; 1 specimen, tl 197 mm; species was origi-

indicated by Günther as "One of the typical nal description. specimens".

— MNHN 2310; 1 specimen, si 160 mm; indicated by

Le Danois, 1961 as "Paratype. Sumatra (Palembang)".

Tetraodon leiurus sensu lato — RMNH 7341; 5 specimens, tl 150, 190, 200, 220,

220 mm.

— ZMA 1 specimen, tl 183 mm. 108.956; The synonyms and references listed below could

apply both to T. leiurus and T. fangi.

The BMNH specimen most likely is not the holo- Arothron leiurus the type as it was bought, according to no., reg. BLEEKER, 1861C: 101, 102 (listed; Thailand: Chao Phaya

was alive in or before 1867, when Bleeker still River; no material, just seen in sketchbook of De

he of his Castelnau). and, as generally is believed, kept most 1865C: material The BLEEKER, 33 (listed; Thailand; probably type material in his personal collection. in MNHN). because MNHN specimen can not be the holotype Tetraodon leiurus; its si is too short; besides it can not be a paratype BLEEKER, 1864: 353 (listed after Bleeker, 1861c; no exist in described from material). as no paratypes a species VAILLANT, 1902: Indonesia: Borneo, 1 specimen. This leaves the RMNH and ZMA 20, 21, 28 (listed; Upper Kapuas River system, mouth of Raun River & with decrease of 10% series. Counting a possible Upper Kapuas River system, Mandai River & Borneo, of tl the most candidates to be the holo- probable Middle Mahakam River, Tepu; material in RMNH, the 190 and 200 RMNH type are mm specimens. reg. no. 7929-31).

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SuvATTi, 1936: 165 (listed; Thailand: Klong River at Indonesia: Java, Djakarta & Indonesia: Sumatra,

Ban & Thailand: & material in Pong Phayau Lake, Chieng-rai Solok; probably at least part in RMNH,

Thailand: Tale Sap, Songkhla & Thailand, Sikuk reg. no. 26477, see syntypes description).

River & Thailand: Chao Phaya River system, Hang BLEEKER, 1853e: 435, 438 (listed; no additional material). Kraben; material probably in BFB). Arothron leiurus;

TWEEDIE, 1940: 81 (listed; West Malaysia: Jelai River; BLEEKER, 1854b: 22, 28 (listed; Indonesia: Sumatra,

material in NMS, reg. no. 2318). Padang; material not recognizable within Bleeker (listed of FRASER-BRUNNER, 1943: 14 as representative Collection).

and no additional material) 1854d: 260 additional genus subgenus; (in BLEEKER, (listed; no material).

subgenus Monotretus). BLEEKER, 1855a: 9 (listed; no additional material).

discussion differences SMITH, 1945: 577 (description, on BLEEKER, 1855C: 260 (listed; Indonesia: Sumatra, Lahat;

with T. leiurus cf. Hora, 1923b and 1924; no material not recognizable within Bleeker Collection).

additional material). BLEEKER, 1857C: 12 (listed; no additional material).

1952: —89 West Jelai River TWEEDIE, 88 (listed; Malaysia: BLEEKER, 1858a: 30, 35 (listed; no additional material). in & West Malaysia: Kuala Tahan; material NMS, BLEEKER, 1858C: 86, 87 (listed; Indonesia: Java, Bogor &

reg. no. 2318, 2319, 2320). Java, Preanger; material not recognizable within

Crayracion leiurus; Bleeker Collection).

discussion distribu- BLEEKER, 1865a: 68 (ex parte) (in on BLEEKER, 1859a: 201 (listed; no additional material).

tion; after Bleeker, 1861c). BLEEKER, 1859e: 48 (listed; Indonesia: Java, Bogor;

Tetrodon liurus; material not recognizable within Bleeker Collection).

KÀROLI, 1882: 187 (listed; West Malaysia: Singapore & BLEEKER, 1859g: 340 (listed; Indonesia: Sumatra, Palem-

Indonesia: material in material within Bleeker Col- Borneo, Sanatbung; MNH, reg. bang; not recognizable

no. 1608, 1609, not available for study). lection).

VOLZ, 1904: 483 (listed; Indonesia: Sumatra, Upper BLEEKER, 1860a: 9, 66 (listed; no additional material).

Langkat, Wampu River). BLEEKER, 1860c: 7 (listed; no additional material).

VOLZ, 1906: 233 (listed; no additional material). BLEEKER, 1860e: 1 (listed; Indonesia: Sumatra, Lematang of skin River POPTA, 1906: 215, 218, 221, 232, 263 (description near Lahat; material not recognizable within

disease; Indonesia: Upper Kapuas River system, Bleeker Collection).

material in 1865d: additional Bongan River; RMNH, reg. no. 7656). BLEEKER, 172 (listed; no material).

Tetrodon leiurus; BLEEKER, 1866: 36 (listed; no additional material).

FOWLER, 1934a: 101 (listed; Thailand: Chiengmai; mate- Crayracion leiurus;

rial probably in ANSP). BLEEKER, 1865a: 66, 67, tab. 213 fig. 1 (description,

FOWLER, 1938: 237 (listed after Kâroli, 1882; no addi- discussion on differences with T. palembangensis and

tional material). T. fluviatilis; no additional localities).

Monotreta cutcutia cutcutia; Tetrodon liurus;

LE DANOIS, 1959: 154—156, 246, 247, fig. 109—111 GÜNTHER, 1870: 288 (description; Indonesia; material in

lateral frontal view and skull 1867.11.28; indicated "One of (description; view, BMNH, reg. no. 110, as

figured; discussion on osteology, sexual dimorphism the typical specimens") (in subgenus Crayracion).

and evolution; material in MNHN, including type WEBER, 1894: 429, 458 (listed; Indonesia: Sumatra,

material of T. cambodgiensis Chabanaud, 1923 and Singkarak Lake & Java, Bogor; material in ZMA,

T. 108.951 and fangi). reg. no. 108.967, respectively).

of LE DANOIS, 1961: 466, 467 (listed in type catalogue Tetrodon Hilgendorfii

valid for Indonesia: MNHN as name T. leiurus, T. cambodgiensis, POPTA, 1905: 185, 186 (original description;

and T. material in Mahakam River Boh fangi; MNHN, reg. no. 03-184, Borneo, Upper system, River;

indicated material in 1922-80/83, and 40-42, the first abusively RMNH, reg. no. 7658, holotype and

as paratype, see syntype description of T. leiurus). paratypes).

D'AUBENTON & BLANC, 1966: 555, 558, 560, fig. 4 POPTA, 1906: 211, 215, 221, 232, 263, 265, 296, pi. 10

(description, figured, discussion on differences with fig. 43 (emendated description, discussion on differ-

T. cambodgiensis Chabanaud, 1923; Cambodia: Tonle ences with T. leiurus; no additional material).

material in Tetrodon Sap; MNHN, reg. no. 66-53/54). Bergii

POPTA, 1905: 186 (original description; Indonesia: Borneo,

Upper Kapuas River system, Bongan River, Bulit;

material in RMNH, reg. no. 7657, holotype). Tetraodoii leiurus 1851. Bleeker, POPTA, 1906: 215—218, 221, 232, 263, 296, pl. 10 fig. 44

Figs. 16-20. (emendated description, discussion on differences with

T. leiurus; no additional material).

Tetraödon leiurus Tetrodon cambodgiensis

BLEEKER, 185 le: 97 (original description; Indonesia: Java, CHABANAUD, 1923: 137—140 (original description, dis-

with T. Djakarta, sea and river mouths; material in part in cussion on differences T. palembangensis,

no. 7342 ex descrip- Vaillant, 1902 and leiurus; Cambodia: RMNH, reg. parte, see syntypes pinguis T. 1922-80/83, tion). Phnom Penh; material in MNHN, reg. no. (in BLEEKER, 1852a: 18 (listed, discussion on differences with syntypes) subgenus Crayracion).

T. kappa Russell, 1803 and T. sans tache Lacépède, CHABANAUD, 1924: 581 (publication not -seen).

1798; no additional material). CHEVEY, 1936: 45 (listed; Cambodia: Tonle Sap; no

BLEEKER, 1852d: 410, 415, 440, 441 (emendated descrip- material).

tion; Indonesia: Borneo, Kusan River at Prabukarta & CHEVEY & LE POULAIN, 1940: 22, 78 (listed; Cambodia;

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no material) (in subgenus Crayracion). pattern of many oval or whitish spots (average Tetraodon liurus; diameter 20 in si), this pattern sometimes descend-

HORA, 1923b: 184 (description, discussion on differences ing on sides; upper part of snout irregularly dark; with T. leiurus cf. Günther, 1870; Thailand: Nonta- chin and white; fins material in F belly uniformly yellowish buri; ZSI, reg. no. 10613/1). defined similar caudal often darkest. HORA, 1924: 500, text-fig. 10 (specimen as plain, to Hora 1923b; Thailand: mouth of Patalung River

near Lampam; material in ZSI, ex parte reg. no. — si Morphometries (based on 57 specimens). F 10472/1). 30—153 tl —175 (m 80) mm; 38 (m 101) mm; Tetrodon leiurus; dorsal fin 13—15 of which 1—3 (m HARDENBERG, 1935: 239 (listed; Indonesia: Borneo, mouth rays (m 14.1) unbranched, 7 6.4—9.3 of Mahakam River near Samarinda; material probably 2.0) longest ray (the —10th)

lost). (m 7.9) in si, shortest ray (the 1st) 13.1—26 (m Tetraodon palembangensis; 18.4) in si, base 7.3—10.7 (m 9.2) in si and 1.0—1.5 SuvATTi, 1936: 166 (listed; Thailand: Upper Bangpakong (m 1.1) in longest anal fin rays 10—12 (m 11.0) River & Thailand: Thale Noi; material at least in part ray;

1897.10.8: of which 1—2 unbranched, in BMNH, reg. no. 148—149?). (m 1.1) longest ray (the

Tetraodon shortest leiurus; 6—9th) 6.8—10.0 (m 8.2) in si, ray (the data; BENL, 1956: 4, 6, 10, Abb. 5 (description, aquarium 1st) 12.9—26 (m 18.5) in si, base 10.6—15.8 (m

no additional material). 12.6) in si and 1.3—2.1 (m 1.6) in longest ray; DE BEAUFORT, 1962: 392, 394, 395 (description, discussion fin of which 1—2 and pectoral 21—24 (m 22.9) on differences with T. leiurus cf. Hora, 1923b rays

1924, T. hilgendorfii Popta, 1905, T. bergii Popta, (m 1.9) unbranched, longest ray (the 5 —8th) 7.7—

1905 and T. leiurus brevirostris Benl, 1957; Indonesia: 11.3 (m 8.9) in si, shortest ray (the 1st) 21—76 Sumatra, Deli & Indonesia: Borneo, Serawai River; (m 33) in si, base 7.2—10.1 (m 8.6) in si and 0.8 material in 101.891 and 108.959 ZMA, reg. no. —1.2 (m 0.9) in longest ray; depth maximum 2.2— respectively) (in subgenus Monotretus). 3.8 (m 3.1) in si, at pectoral fin base 2.5—4.1 (m

3.4) in si, at end of dorsal fin base 5.0—7.6 (m 6.4) General description. — in si, width maximum 2.4—3.9 (m 3.0) in si and

in maximum fin General morphology. — Body oblong, head region 0.8—1.1 (m 1.0) depth, at pectoral

base 2.4—3.8 in si and 0.8—1.1 depressed; dorsal profile arched or flattened, high- (m 3.0) (m 1.0)

in end of dorsal fin base est at midst of back; interorbital flat or convex, corresponding depth, at often with median lateral line 5.2—9.5 6.8) in si and 0.9—1.1 in a groove; system (m (m 1.1) often distinct; spines small, densely set, some- corresponding depth; head length 2.1 —2.5 (m

in 2.0—2.4 in times in part hidden under papillae, covering back, 2.3) si; snout length (m 2.2) hi; interocular distance 1.7—2.2 in sides and belly between nostrils and anus; origin (m 2.0) hi; eye of anal fin beneath posterior half of dorsal fin horizontal diameter 4.7—7.9 (m 5.8) in hi, distance from dorsal nasal base; mouth terminal, directed forwards or up- profile 13.5—61 (m 27) in hi;

11.7—30 (m in hi, distance from wards; lower border of eye above level of mouth organ length 20)

with dorsal of snout 2.6—3.7 (m in hi, distance from corner, upper border not interfering tip 3.1) in hi profile; nasal organ a tentacle, distal 1/3-1/2 of eye 6.3—13.9 (m 8.7) and 1.9—5.0 (m 2.6) which is divided into two flattened lobes; apposed in distance from tip of snout; mouth width 3.0— surfaces of lobes smooth. 5.3 (m 4.3) in hi; upper lip depth 7.3—21 (m

11.8) in hi; lower lip depth 12.9—29 (m 18.1) in hi.

— colour of Colour in alcohol. Ground upper parts

Material examined. — sides with tan, of lower parts yellowish white; BMNH 1867.11.28: 110; 1 specimen, si 91 mm; Indone- many polygonal or rounded dark spots (diameter sia; Bleeker Collection; indicated by Günther, 1870 as network 14-co in si) leaving mostly only a lighter "One of the typical specimens", see description of the of ground colour; sometimes one of these spots, syntypes.

BMNH 1897.10.8: situated beneath origin of dorsal fin at middle of 148—149; 2 specimens, si 92—124

mm; Thailand: Upper Bangpakong River; coll. S. S. side, markedly bigger and darker than adjacent Flower, III-1897; ex RSM. which around spots, are arranged circularly it, BMNH 1922.5.19: 110—112; 3 specimens, si 41—67 diame- as a distinct ocellus (horizontal West Jahan coll. eventually mm; Malaysia: River; Kloss.

least in of back BMNH 1922.5.19: 1 si 63 ter at 11.8 si); colour variable, 113; specimen, mm; West

Malaysia: Sembeling River; coll. Kloss. often with a light interorbital band, bordered pos- MNHN 1922-80/83; 4 specimens, si 122—153 midst mm; teriorly by a dark trapezoid patch reaching to Cambodia: Phnom Penh; coll. La Résidence Française, of back sometimes a second dorsal or beyond; 1915 coll. A. and/or Krempf, 1921 (syntypes of T. cam- before dorsal sometimes back dark patch just fin; bodgiensis Chabanaud, 1923).

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Figs. 16—20. Tetraodon leiurus Bleeker, 1851.

Fig. 16. Lateral view of ZMA 108.951, sl 96 mm.

of Fig. 17. Dorsal view ZMA 108.951, sl 96 mm.

Nasal of Fig. 18. organ MNHN 22—80, syntype of T. cambodgiensis Chabanaud, 1923, sl 153 mm,

Fig. 19. Lateral view of MNHN 22—80, syntype of T. cambodgiensis Chabanaud, 1923, sl 153 mm.

of Fig. 20. Dorsal view MNHN 22—80, syntype of T. cambodgiensis Chabanaud, 1923, sl 153 mm.

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NMB 2 si 93—106 Thai- 5174—5175; specimens, mm; or without one or more of the 1, 1, 1, 1 specimen land: Thale coll. H. Bernatzik, 1937. Luang; lots. Out of these 14 specimens 4 could without RMNH 14 si 30—105 7342; specimens, mm; Indonesia; much doubt be recognized as syntypes on the basis Bleeker Collection; probably containing 4 out of the 5 of the characteristics given by Bleeker, 185 le and syntypes, see syntypes description. (10 non-typical speci- 1852d: indistinctness of mens now catalogued RMNH 26477). flabbyness, colourlessness,

RMNH 7657; 1 specimen, si 65 mm; Indonesia: Borneo, the lateral line and a matching actual tl: 59-97 mm. Upper Kapuas River system, Bongan River, Bulit; coll. The remaining 10 specimens most probably do not A. W. Nieuwenhuis, VII-1898 (holotype of T. bergii contain the shows missing syntype as none of them Popta, 1905).

si Indonesia: the same characteristic colour as the other 4 RMNH 7658; 6 specimens, 48—80 mm; spec-

Mahakam River Boh coll. imens. Borneo, upper system, River; Probably the 10 specimens form, in part,

A. W. Nieuwenhuis, V/VIII-1900; (holotype and para- the material on which Bleeker based his emendated types of T. hilgendorfii Popta, 1905) (morphometries not description, their actual maximum tl (129 mm) used in general description). being too high and their aotual minimum tl (38 ZMA 101.886; 1 specimen, si 69 mm; Indonesia: low for Sumatra, Pangian River; coll. E. Jacobson, III-1914. mm) being too considering all of them to

ZMA 1 si 111 Indonesia: 101.891; specimen, mm; belong to the series used for the emendated diag- coll. de de Sumatra, Deli, coast; L. P. Cosquino Bussy, nosis. 1908/09. It must be noted that both the indications "para- ZMA 102.308; 1 specimen, si 67 mm; Indonesia; Bleeker " type" and Central Borneo" as given to the Collection; possibly completing the syntypes series in MNHN RMNH 7342 (see description of the syntypes). specimens must be incorrect for a speci-

ZMA 108.951; 10 specimens, si 37—114 mm; Indonesia: of which described men a species was from Sumatra, Sinkarak Lake; coll. M. Weber, 1888. syntypic material collected in Java. These indica- ZMA 108.959; 4 specimens, si 64—83 mm; Indonesia: tions were the major ground on which Le Danois Borneo, Serawai River, Lebang Hara; coll. Witkamp. based her that T. leiurus is the male ZMA 108.967; 11 specimens, si 71—93 mm; Indonesia: (1959) opinion

Java, Bogor; coll. M. Weber, 1888. of T. palembangensis.

ZSI si 115 Thailand: F 10472/1; 1 specimen, mm; I did consider it not opportune to select a lecto- Annan- mouth of Patalung River near Lampam; coll. N. since the type, syntypic specimens were already dale. colourless and flabby in Bleeker's time and the ZSI F 10613/1; 1 specimen, si 51 mm; Thailand: in fact is Nontaburi; coll. M. Smith. species based on the emendated diagnosis

which incorporates material possessing the discrim-

— colour be served Description of the syntypes. inating pattern. Science can not

by selecting a lectotype out of 4 badly preserved

and selection of Recognition. — T. leiurus was originally described syntypic specimens specimens from

the emendated is the Inter- by Bleeker from 5 specimens, tl 60-98 mm. The diagnosis prohibited by

national Code of Zoological Nomenclature. emendated description a year later was based on

11 specimens, tl 60-115 mm. In 1865, according

General — to the Atlas Ichthyologique, Bleeker possessed 19 morphology. Specimens in poor state of specimens, tl 46-130 mm. The catalogue of the preservation, flabby; spines small, not papillated,

in retracted, area between nasal and auction 1879 indicates the same total number, covering organs

end of dorsal fin divided into lots of 15, 1, 1, 1, 1 specimens. The base; eyes high on head, not

with dorsal lateral line syntypes, as usual, are not labelled as such and interfering profile; system otherwise like had to be refound within the Bleeker Collection. inconspicuous; general description.

In search of Bleeker Collection specimens the

Colour in alcohol. — following series were traced: Completely disappeared, as it was already at time of original description.

Material probably used for emendated — non- diagnosis RMNH 7342; 14 specimens, tl 38—129 mm (10 material typical specimens now catalogued RMNH 26477). (see examined) shows colouration in 6

— BMNH 1867.11.28: 110; 1 tl 111 indi- of 10 rounded dark specimen, mm; out specimens, viz. lateral spots cated Günther "One of the by as typical specimens". all 6 (in specimens), a lateral ocellus (in 3 speci- — MNHN 03—184; 1 specimen, si 130 mm; Indonesia: and two dorsal blotches, and before mens) on nape Central Borneo; ex RMNH; indicated by Le Danois dorsal fin (in 1 (1961) as "paratype". specimen).

— ZMA 102.308; 1 specimen, tl 82 mm.

Morphometries (based on 4 specimens). — si 49-

The RMNH series undoubtedly represents the 79 mm; tl 59-97 mm; dorsal fin rays 13-14 (m greater part of the 15 specimens auction lot, with 13.8) of which 2-3 (m 2.3) unbranched, longest

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6.4-7.5 in base 8.8-9.8 in Tetraodon ocellaris ray (m 9.4) si, (m 9.4) KLAUSEWITZ, 1957a: 201, 202, Abb. 4, 5 (original de- si and 1.2-1.5 (m 1.4) in longest ray; anal fin rays scription, discussion on differences with T. leiurus and in i.10, longest ray 7.2-8.3 (m 7.6) si, base 11.3- T. cutcutia; Thailand: near Patalung & Thailand: near in si and 1.6-1.8 in 14.5 (m 1.7) longest material in (m 13.0) Krung Thep; SMF, reg. no. 3975, holotype,

ray; pectoral fin rays ii.21, longest ray 7.9-8.8 (m and 3976 & 3827, paratypes).

Tetraodon 8.3) in si, base 7.9-8.8 (m 8.4) in si and 1.0 in leiurus; JOHNSEN, 1964: 262 (description in accordance with T. longest ray; depth maximum 3.0-3.4 (m 3.2) in si; leiurus cf. Hora, 1923b and 1924, and in disaccordance width maximum 3.2-3.9 (m 3.6) in si and 0.8- with T. leiurus cf. Fowler, 1938 and cf. Inger & Chin, 1.1 in maximum head 2.4 (m 1.0) depth; length 1962; Thailand: Kwhae Noi at Ban Kao & Kwhae

-2.5 (m 2.5) in si; snout length 2.2-2.5 (m 2.3) Noi at Sai Yok). Tetraodon cutcutia; in hi; interocular distance 1.8-2.2 (m 2.0) in hi; AXELROD 1967: F-579.00 et al., F-579.00, fig. p. (de- distance from snout 2.6-3.4 nasal organ tip of (m scription, aquarium data; no material). 3.1) in hi, distance from eye 6.3-9.1 (m 7.7) in hi

and 2.6 in distance from tip of snout. General description. —

Material examined. — General morphology. — Body oblong, greatest RMNH 7342 ex parte; 4 specimens, si 59—79 mm; part compressed dorsoventrally; dorsal profile syntypes; Indonesia: Java, Djakarta; Bleeker Collection. arched, highest at midst of back; interorbital con-

without lateral line dis- vex, a groove; system Distribution. — T. leiurus is known from Thai- tinct; body spines small, soft, covering back, sides land, East and West Malaysia, Sumatra, Java,

and belly between nostrils and anus, hidden un- Borneo and Cambodia. der white papillae; papillae giving smooth impres-

sion, especially on belly arranged in netlikepattern;

— The leiurus is derived from Etymology. name origin of anal fin situated beneath anterior half of the greek "leios", meaning "smooth". The name dorsal fin mouth directed forwards; is base; terminal, incorrect in this regard, for some T. leiurus lower border of above level of mouth eye slightly specimens show distinct spines; in general these corner, upper border higher than lowest point of spines are smaller than those of fluviatilis- interorbital, sometimes even interfering with dor- group specimens.

sal nasal distal - profile; organ a tentacle, 1/3 1/2

of which is divided into two flattened lobes; ap- posed surfaces of nasal lobes smooth. Tetraodonfangi Pellegrin & Chevey, 1940.

Figs. 21-23. Colour in alcohol. — Ground colour of upper

parts tan, of belly yellowish white; whole of sides Tetrodon fangi and lateral of back covered with PELLEGRIN & CHEVEY, 1940: 157, 158, fig. 2 (original parts many polyg-

discussion differences with T. onal dark description, on patoca spots (diameter 20-co in si), leaving only

Hamilton, 1822; South Vietnam: Xano Canal, Vi- a lighter network of ground colour; back in part material in holo- thanh; MNHN, reg. no. 40—42, with same spots, in part uniformly tan; a light type) (in subgenus Leiodon). band between posteriorly often bordered by Tetrodon leiurus; eyes,

dark to level of FOWLER, 1937: 264, figs. 299, 300 (description, lateral a trapezoid patch reaching pecto-

view of body and dorsal view of head figured; Thai- ral fins or beyond; sometimes a second, smaller land: Pitsanulok; material in ANSP). dark patch just anterior of dorsal fin; snout dor- Tetraodon leiurus brevirostris with dark sally spots or uniformly dark; chin dark; BENL, 1957 a: 63—65, Abb. 1, 2, 3 (original description,

a ocellus diameter 6.4-11.1 in aquarium data, discussion on differences with T. leiurus big (horizontal si)

cf. Bleeker, 1851, T. cutcutia and T. palembangensis on sides, horizontally on level with pectoral fin, steindachneri live = T. nom. probably Thailand, with of dorsal Nov.; vertically on level origin fin, nucleus

imported; material in BMNH, no. 1956.1.7.1, reg. than lateral often darker spots; a light small central holotype, and 1956.1.7.2, paratype). in ocellus with surrounded 1958: spot nucleus; light ring, GEISER, 100—101 (aquarium data; no additional

material). by circularly arranged, and partly fused polygonal

MERCKENS, 1959: 170, 170 set contribute 171, fig. p. (description, spots; densely spinal papillae to

aquarium data, discussion on differences with general colour pattern; belly laterally with dark = T. steindachneri palembangensis” nom. Nov.; no spots, more oblong than lateral spots, centrally additional material). uniformly yellowish white; fins caudal dark- STERBA, 1959: 638, Taf. 273 (description, aquarium data; plain,

no additional material). est.

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Figs. 21—23. Tetraodon fangi Pellegrin & Chevey, 1940.

Fig. 21. Lateral view of MNHN 40—42, holotype, sl 55 mm.

Dorsal view of SMF of T. ocellaris Fig. 22. 3975, holotype Klausewitz, 1957, sl 48 mm.

23. Nasal of MNHN 40—42, mm. Fig. organ holotype, sl 55

— si in Morphometries (based on 15 specimens). (m 3.9) hl, distance from eye 8.2-12.5 (m 10.2)

in hl and 2.1-3.8 40-61 (m 50) mm; tl 49-74 (m 61) mm; dorsal (m 2.6) in distance from tip of

of which 1-2 un- mouth width 3.2-4.0 in fin rays 12-14 (m 13.2) (m 1.8) snout; (m 3.4) hi; upper 7.7-10.5 9.6-23 branched, longest ray (the 8th) (m 8.5) lip depth (m 13.9) in hi; lower lip depth

in si, 8.8-23 in hi. in si, shortest ray (the 1st) 20-22 (m 21) (m 16.0) base 7.9-10.2 (m 9.1) in si and 0.8-1.1 (m 1.0) in

of which Material examined. — longest ray; anal fin rays 10-11 (m 10.8) BMNH 1957.1.7: 2 si 1—2; specimens, 54—55 mm; 1-2 (m 1.3) unbranched, longest ray (the 7-8th) probably Thailand; imported alive through dealer in 6.5-10.7 (m 8.0) in si, shortest ray (the 1st) 13.5- Zürich; coll. G. Benl (holotype and paratype of T. leiurus 11.0-15.3 in si 17.0 (m 15.3) in si, base (m 12.4) brevirostris Benl, 1956).

in fin MNHN 40—42; 1 specimen, si and 1.1-2.2 (m 1.8) longest ray; pectoral 55 mm; see holotype description. rays 21-23 (m 22.0) of which 2 unbranched, lon- SMF 1 si 3975; specimen, 48 mm; Thailand: near in shortest gest ray (the 5-6th) 7.7-10.4 (m 8.4) si, Patalung; coll. E. Roloff, 21-IX-1956 (holotype of T. 14.0-18.5 in base 7.8- ray (the 1st) (m 16.6) si, ocellaris Klausewitz, 1957).

10.4 in si and 0.9-1.1 in (m 8.3) (m 1.0) longest SMF 3976; 1 specimen, si 58 mm; Thailand: near

coll. in at Patalung; E. Roloff, 21-IX-1956 (paratype of T. ray; depth maximum 2.5-4.0 (m 3.3) si, ocellaris Klausewitz, 1957). pectoral fin base 2.7-4.0 (m 3.3), at end of dorsal SMF 10 si 3827; specimens, 40—61 mm; Thailand: near fin base 4.5-8.1 (m 6.4) in si; width maximum Krung Thep; coll. E. Schmidt, 24-ÏV-1956 (paratypes of in si in maxi- 2.6-3.4 (m 2.9) and 0.8-1.1 (m 0.9) T. ocellaris Klausewitz, 1957). mum depth, at pectoral fin base 2.5-3.4 (m 3.0)

of the — in si and 0.9-1.0 (m 0.9) in corresponding depth, Description holotype. at end of dorsal fin base 5.4-8.4 (m 7.0) in si

General morphology. — Like general description. and 0.9-1.4 (m 1.1) in corresponding depth; head Specimen in poor state of preservation, deteriorat- length 2.1-2.6 (m 2.2) in si; snout length 2.7- ed, skinny, flabby; lateral line system indistinct for 2.9 (m 2.8) in hi; interocular distance 1.6-2.0 the larger part; whole of body densely papillose. (m 1.8) in hi; eye horizontal diameter 3.7-5.6

Colour in alcohol. — (m 4.4) in hi; nasal organ length 11.0-16.7 (m Like general description.

13.5) in hi, distance from tip of snout 3.1-4.6 Horizontal diameter of ocellular nucleus 8.5 and

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diameter of in the leiurus- are 8.5 mm (6.4 and 6.4 in si); horizontal pattern. Fin ray counts group

and 4.5 D A P in T. kretamensis central white spot in nucleus 3.5 mm 12—15; 10—12; 21—24; 11 P 18—19 and in T. stein- (15.7 and 12.2 in si); two dorsal patches present, D —13; A 10—10; median dachneri D A P 18—21. The but too faint for taking measurements; 13—15; 11—13; is within belly uniformly yellowish, laterally with some dark colour pattern of T. erythrotaenia unique

the in a dark back without spots. genus having plain further markings. Zoogeographically the species is

of the Wallace Line. — dorsal in living east Morphometries. si 55 mm; tl 67 mm; unique Weber fin ii. 10; fin It must be noted, as was done earlier by fin rays ii.10; anal rays pectoral rays Bleeker's in base 6.4 mm (1911) and De Beaufort (1962), that i-ii.20, longest ray 7.1 mm (7.7 si), maximum in the Atlas Ichthyologique (1865a) is (8.6 in si and 1.1 in longest ray); depth diagnosis

fin base 20 incorrect: the nasali" are not com- 22 mm (2.5 in si), at pectoral mm "papilla "apice

but bifida" as was correctly stated (2.8 in si); width maximum 20 mm (2.8 in si and pressa" "apice in the 1.1 in maximum depth), at pectoral fin base 20 original description (1853d).

mm (2.8 in si and 1.0 in corresponding depth);

in 8.8 The T. head length 25 mm (2.2 si); snout length only species resembling erythrotaenia

in the low fin counts are T. travancorius and mm (2.8 in hi); interocular distance 13.3 mm (1.9 ray

in T. kretamensis. T. travancorius has even lower in hi); eye horizontal diameter 4.5 mm (5.6 hi), D A P 16—17 but differs unmis- distance from dorsal profile 0.6 mm (42 in hi); counts: 7—8; 8; in the nasal characteristic nasal distance from of 6.3 mm takably possessing organ tip snout organ in hi of the cutcutia- being only slightly lipped (4.0 in hi), distance from eye 2.0 mm (12.5 group,

and 3.2 in distance from tip of snout); mouth at most. The latter species resembles erythrotae-

2.0 nia also in the other main characteristics: colour width 7.5 mm (3.3 in hi); upper lip depth mm

and distributional at (12.5 in hi); lower lip depth 2.6 mm (9.6 in hi). pattern pattern, living just

the east side of the Wallace Line and being the

lateral Material examined. — only other Tetraodon possessing a longitu-

South MNHN 40—42; 1 specimen, holotype, si 55 mm; dinal dark bar. Moreover, T. kretamensis like

Vietnam: Xano Canal, Vi-thanh; coll. IOI. T. erythrotaenia is small, not known to exceed

mention- si 64 mm. Differences in the characters

Distribution. — T. fangi is only known with than ed above however are greater they seem to in this certainty from the localities cited paper, be. The difference in mean value for dorsal fin viz. Thailand (Krung Thep, Ban Kao, Sai Yok, rays is 1.7, for anal fin rays 1.0. Zoogeographi- Patalung) and Vietnam (Vi-thanh). the cally the narrow Makassar Strait separating

is main barrier between two species a two regions.

Etymology. — Pellegrin & Chevey named the The apparant resemblance in colour pattern neither after M. P. W. Chinese species Fang, specialist on all is basic since the lateral bar does not occur in fishes. specimens of T. kretamensis, and Inger, although

having seen living specimens of his species, does

erythrotaenia-group not mention the second red band which is present

in most T. erythrotaenia specimens. Moreover, the

Diagnosis. Tetraodon species with the nasal absence of any special marking on the dorsal organ formed by a hollowcylinder which is distally surface in T. erythrotaenia seems to be a more

divided, over one-third to one-half of its length, fundamental difference, as it is a unique character,

two rounded head in kretamensis other into or triangular lips; slightly never occurring T. or any

D Tetraodon or not depressed; body spines not papillose; species.

9 —11; A B—9;8—9; P 18—19. One species: T. erythro- McCulloch (1922) and De Beaufort (1962) point-

taenia. ed out that the species described by Kner (1867)

is as Crayracion erythrotaenia not the present spe-

Discussion. Although T. erythrotaenia resem- cies, probably being referable to Sphoeroides

members to bles the of the leiurus-group, T. kreta- pleurogramma (Regan, 1902). Kner appears mensis and T. steindachneri considering the nasal have been followed by Ogilby (1886), who in turn

it is held to constitute on its followed Waite organ, a group was by (1904). basis of in own on the its extremely low fin ray Bleeker (1865a) identified specimens in a jar

Gernaerti T. counts, its colour pattern, and its distributional MNHN labelled Epipedorhynchus as

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erythrotaenia. This identification most probably is in part in ZMA, reg. no. 108.960). WEBER, 1913: 603—604, 606, 610 (description, discussion wrong: the jar was said to be from China and no with colour of cf. on differences pattern specimens reliable record of true Tetraodon from China is Weber, 1911; Indonesia: New Guinea, Lorentz River

known (neglecting a doubtful "China" locality at lower course and at Sabang; material in ZMA, reg.

for T. NMW Moreover Le nigroviridis, 64.297). no. 108.962 and 101.917, respectively).

Danois lists FOWLER, 1928: 471 (description after Günther, 1870; (1959) E. Gernaerti as a junior syn-

no material). of Geneion honckenii the onym (Bloch, 1787), so WHITLEY, 1957: 38 (in type list of AMS; a cotype claimed jar should have contained specimens with double- for AMS, reg. no. B 7629). nasal different from those in Te- pierced organs Tetraodon erythrotaenia;

traodon. DE BEAUFORT, 1913: 152, 159 (listed; Indonesia: Ambon

Island, Mirdika River; material in ZMA, reg. no.

108.964).

FRASER-BRUNNER, 1943: 14 (listed as representative of Tetraodon 1853. erythrotaenia Bleeker, and additional sub- genus subgenus; no material) (in

Figs. 24-25. genus Monotretus).

BENL, 1956: 144, 172, 202, Abb. 7 (description, aquarium

data; no material). Tetraodon erijthrotaenia MERCKENS, 1958: 54 (description, aquarium data; no BLEEKER, 1853C: 318, 319 (listed; Indonesia: Ambon material). Island; for material see syntypes description). data: STERBA, 1959: 637, Abb. 604 (description, aquarium BLEEKER, 1853d: 155 (listed; Indonesia: Celebes, Maros; no material). for material see syntypes description). DE BEAUFORT, 1962: 392, 397 (description, discussion on Tetraodon erythrotaenia colour variation in living specimens; no additional BLEEKER, 1853c: 328 (listed after 1853c: 319). localities) (in subgenus Monotretus). BLEEKER, 1853d: 154, 162, 174 (original description; Monotretus erythrotaenia; Indonesia: Celebes, Maros & Indonesia: Ambon MUNRO, 1958: 294 (listed; no material). Island; for material see syntypes description) (in MUNRO, 1967: 552 (in key; no material). subgenus Arothron).

Arothron erythrotaenia;

General description. — BLEEKER, 1854C: 477 (listed; no additional material).

BLEEKER, 1854d: 237, 260 (listed; no additional material).

BLEEKER, 1855b: 300 (listed; no additional material). General morphology. — Body oblong, anterior part BLEEKER, 1855e: 402 (listed; Indonesia: Ambon Island; slightly compressed dorsoventrally, posterior part material not recognizable within Bleeker Collection). dorsal 1856a: additional slightly compressed laterally; profile arched, BLEEKER, 22 (listed; no material).

BLEEKER, 1856b: 28 (listed; no additional material). highest at midst of back or behind; interorbital

BLEEKER, 1857a: 28 (listed; no additional material). lateral line in- convex, without a groove; system

BLEEKER, 1857b: 20 (listed; no additional material). distinct; spines short, slender, in skinny specimens BLEEKER, 1859a: 200 (listed; no additional material). distinctly two-rooted, scantly distributed on back BLEEKER, 1860d: 12 (listed; no additional material). sides and between midst of and Crayracion erythrotaenia; belly eyes origin

BLEEKER, 1865a: 67—69, pi. 214, fig. 4 (description, in of dorsal fin; origin of anal fin situated anterior

discussion on differences with key, Epipedorhynchus of dorsal fin or beneath anterior third of dorsal fin

gernaerti Bibron; no additional material). base; mouth terminal, directed forwards; lower BLEEKER, 1865b: 271 (listed; no additional material). border of eye at level of mouth corner or below, BLEEKER, 1866: 36 (listed; no additional material). with dorsal BLEEKER, 1868a: 19 (listed; Indonesia: Moluccas, Gebe upper border not interfering profile;

Island; material not recognizable within Bleeker tentacle, distal half less of nasal organ a or which Collection). is divided into two flattened, triangular or rounded Tetrodon erythrotaenia; lobes; apposed surfaces of lobes smooth. GÜNTHER, 1870: 298 (description; Indonesia: Ambon

Island; material in BMNH) (in subgenus Arothron).

WEBER, 1894: 458 Indonesia: Colour in alcohol. — Ground colour 432, 434, 443, (listed; of upper Ambon Island). tan or dark of whitish, parts grey, belly grey or WEBER, 1895: 261 (listed; no material). almost black; both colours usually separated by a WEBER, 1911: 9, 11, 12, 14, 15, 48—49, Taf. 2 fig. 1 whitish band in from snout head to I-VII discussion (red life) or (description, and figures on colour this band head often broken pattern variation, discussion on differences with caudal peduncle, on

descriptions by Bleeker and Günther; Indonesia: Kai in dots up or vertical lines; white band often Besar Warka & Indonesia: Wokam Island, near Island, bordered dark inferiorly by a second, band; upper Waskai River & Indonesia: Kobroör Island, Kololobo surface plain dark, sometimes with a light interocu- River near Seltutti & Kobroör Island, source of lar band; ventral surface almost Rubanratu River & Indonesia: Aru Islands, Tenuaborri whitish, grey or in the often River, mouth & Aru Islands, Bendjia River; material black, latter cases marked with whitish

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Figs. 24—25. Tetraodon erythrotaenia Bleeker, 1853.

Lateral view Fig. 24. of ZMA 108.962, sl 51 mm.

25. Nasal of ZMA Fig. organ 108.962, sl 51 mm.

lines and whitish fins caudal broken dots; plain, Bleeker Collection (containing type material, see lectotype in fin often darkest. description) (not incorporated morphometries due to bad state of preservation).

ZMA 101.917; 1 specimen, si 31 mm; Indonesia: New

— si 28— Morphometries (based on 27 specimens). Guinea, Lorentz River at Sabang; coll. H. A. Lorentz,

13/24-VI-1907. 55 (m 41) mm; tl 36—69 (m 52) mm; dorsal fin ZMA 108.960; 9 specimens, si 28—52 mm; Indonesia: rays 9 —11 (m 10.1) of which 2—3 (m 2.2) un- Aru Islands, Bendjia River; coll. H. A. Lorentz, 4-XII- branched, 6 5.6—7.2 longest ray (the —8th) (m 6.6) 1907. in 15.7—23 in si, shortest ray (the 1st) (m 19.4) si, ZMA 108.962; 8 specimens, si 30—51 mm; Indonesia: base 8.8—12.0 (m 10.7) in si and 1.3—1.9 (m 1.6) New Guinea, lower course of Lorentz River at ebbing; coll. A. Lorentz, anal fin of which H. 4-V-1907. in longest ray; rays 8 —9 (m 9.0) ZMA 108.964; 9 specimens, si 41 —55 mm; Indonesia: 1—2 5 (m 1.6) unbranched, longest ray (the —7th) Moluccas, Ambon Island, Mirdika River; coll. L. F. de

5.9—8.0 in si, shortest 12.5— (m 6.8) ray (the 1st) Beaufort, 11-1910. 25 (m 18.1) in si, base 10.2—12.9 (m 11.5) in si and 1.4—1.9 (m 1.7) in longest ray; pectoral fin Description of the lectotype. — rays 18—19 (m 18.3) of which 1—3 (m 2.0) un-

Designation. — Bleeker based his descrip- branched, longest ray (the 5 —6th) 5.9—8.3 (m 7.2) original tion tl 54-58 from In- shortest si, on 2 specimens, mm, one in si, ray (the 1st) 14.1—29 (m 19.5) in Maros and base 7.6—9.7 (m 8.6) in si and 1.1 —1.3 (m 1.2) in donesia: Celebes, one from Indonesia: Moluccas, Ambon. In the publication containing longest ray; depth maximum 2.7—3.8 (m 3.2) in si, the third Indone- at pectoral fin base 2.7—3.8 (m 3.3) in si, at end of original description a locality, dorsal fin base 5.1—7.0 (m 6.1) in si; width maxi- sia: Ceram, is referred to (Bleeker, 1853d: 155)

but the is mentioned mum 2.4—3.2 (m 2.9) in si and 0.8—1.0 (m 0.9) in material of this locality not is it mentioned in other maximum depth, at pectoral fin base 2.4—3.2 (m in the description, nor any On 2.9) in si and 0.8—1.0 (m 0.9) in corresponding article by Bleeker. the contrary, Bleeker states

depth, at end of dorsal fin base 6.1—8.7 (m 7.3) in in 1865a explicitly that T. erythrotaenia is only Maros and Ambon from si and 1.1— 1.3 (m 1.2) in corresponding depth; known from (aside a Chinese in head length 2.3—2.5 (m 2.4) in si; snout length supposed specimen MNHN), so most

the Ceram was due to a mistake 2.3—2.8 (m 2.6) in hi; interocular distance 1.5— probably locality

Bleeker or his and there are only two I.8 (m 1.7) in hi; eye horizontal diameter 2.7—4.2 by printer The Atlas in 1865 (m 3.4) in hi, distance from dorsal profile 11.7—27 syntypes. Ichthyologique men- tions 3 tl 41-58 third (m 19.5) in hi; nasal organ length 8.3—14.0 (m specimens, mm (a specimen

II.0) in hi, distance from tip of snout 3.1—4.2 (m was recorded from Ambon in Bleeker, 1855e). The

distance from small number of auctioned 3.6) in hi, eye 7.7—13.8 (m 10.1) same specimens was in in These auctioned do in hi and 2.1—4.1 (m 2.8) in distance from tip of 1879, one lot. specimens

not the three Atlas snout; mouth width 3.2—4.2 (m 3.7) in hi; upper necessarily represent specimens

additional material was recorded Bleeker lip depth 12.5—25.0 (m 16.4) in hi; lower lip as by from Gebe Island. depth 12.2—26 (m 17.6) in hi. (1868a) Indonesia:

In search for the I could trace 5 spec- Material examined. — syntypes from Bleeker Collection: RMNH 7356; 3 specimens, si 41—48 mm; Indonesia; imens

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— AMS B 7629; 1 specimen; recorded by Whitley (1957) Material examined. —

as "cotype". RMNH 7356 ex parte; 1 specimen, si 43 mm; Indonesia:

2 — 1 inch 64 recorded Indonesia: BMNH; specimen, tl 2 A = mm; Celebes, Maros or Moluccas, Ambon Island;

by Günther (1870) as "One of the typical examples". Bleeker Collection.

— RMNH 7356; 3 specimens, actual tl 41, 53, 58 mm.

Distribution. — As indicated by the synonyms and is not The AMS specimen most probably a very references, and the list of material, T. erythrotae- important one as Whitehead et al. (1966: 14) said nia is known from Celebes, Ambon, the Kai of the AMS Bleeker material in general: "These Islands, the Aru Islands, Gebe and Western New are almost certainly specimens from Bleeker's du- Guinea. plicate series". The BMNH specimen surely is too In the containing the original diminished paper description long, as tl only can have during the

Bleeker refers to a third locality, Ceram, but this The RMNH series almost is the years. certainly locality is not repeated in the original description, series auctioned in 1879 and as the series auctioned in On nor later publications. the contrary, Bleeker are generally bdieved to contain at least most of records in 1865a the species as only occurring on the type material, the RMNH specimens almost Celebes and Ambon, apart from a Paris series certainly incorporate both syntypes. said to be from China. The latter locality is doubt- It consider is very tempting to the RMNH se- ful as a true Tetraodon never again was recorded ries on base of the length of the specimens identical from China. The locality of Sydney, Australia, with the specimens mentioned in the Atlas Ich- given by Kner (1867), Ogilby (1886), and Waite thyologique. One has however to reckon with a (1904) probably is based on an erroneous deter- possible decrease in length of 10%, so the syntypes mination is out by Kner, as pointed in my discus- at present can be 49-54 and 52-58 mm. The only sion on the about from group. way to be sure selecting the lectotype

the syntype series is to designate the specimen with Etymology: The name erythrotaenia is derived actual tl 53 mm, as is done hereby. The other from the greek "erythros" meaning "red" and specimen with actual tl 58 mm most probably is "tainia" (latin "taenia") meaning "ribbon", refer- the paralectotype. ring to the red lateral head-tail band.

General in of morphology.— Specimen poor state

preservation, flabby; lateral line system inconspicu- fluviatilis-group ous; spines distinctly two-based (distance between

tips of roots ca. 1.1 mm), relatively densely covering Diagnosis. Tetraodon species with the nasal or- back, sides and belly from nasal organs to origin gan formed by a hollow cylinder, which is distally of dorsal fin; otherwise like general description. divided in two lobes over one-third to five-sixths

of its length; interorbital and dorsal profile of Colour in alcohol.— Completely disappeared; ac-

head not sides with cording to original description by Bleeker (1853d: convex; body spines papillose; dark rounded spots, variable in number and size, 174): "colore corpore superne olivaceo-fusco infer- but always one spot at base of dorsal fin and one ne flavo, fascia intermedia duplice maxillo-cauda- at midst of caudal fin base; back with dark li nigra et rubra, pinnis pectoralibus, dorsali cau- spot form three dark D dalique fusco-violaceis, anali flava." spots tending to patches; 11-16;

A 10-14; P 18-24. Four species, one subspecies:

Morphometries. — Actual si 43 mm; actual tl 53 T. kretamensis, T. nigroviridis, T. fluviatilis flu-

dorsal fin 6.0 T. T. steindachneri mm; rays ii.7, longest ray mm (7.2 viatilis, f. sabahensis, nom.

in si), base 3.4 mm (12.6 in si and 1.8 in longest Nov. ray); anal fin rays i.8, longest ray 4.3 mm (10.0 in

base si), 2.9 mm (14.8 in si and 1.5 in longest Discussion. Unlike the other groups, the fluvia- ray), pectoral fin rays ii. 16, longest ray 4.1 mm tilis-group is not homogeneous in the form of its

(10.5 in si), base 3.9 mm (11.0 in si and 1.1 in nasal organ. In T. kretamensis and T. steindachneri

head 18 in the nasal lobes longest ray); length mm (2.4 si); snout occupy only 1/3-2/3 of the organ's length 7 mm (2.6 in hi); interocular distance 9.5 length and are smooth; in T. fluviatilis and T. ni-

in nasal mm (1.9 hi); organ distance from tip of groviridis they occupy 1/2-5/6 of the length and snout 5.2 mm in distance from tissue the (3.5 hi), eye 1.6 possess spongy on apposed surfaces in in hi mm (11.3 and 3.3 in distance from tip of 50-90% of specimens > 60 mm si. Nevertheless I snout). prefer to take the four species together on the basis

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of similarities in colour pattern, all four species in statistically (D13 and All occurring in some in all specimens showing the spots at the base of the specimens of all species except T. kretamen- and both sis D A and the second dorsal and caudal fins T. steindachneri : 11-13; 10), proved

and T. fluviatilis possessing the dark dorsal to be valuable only in distinguishing the of The said patches, which may occur in T. kretamensis, and two subspecies T. fluviatilis. mor-

characters how- are absent in T. nigroviridis. phological discriminating proved,

ever, to be correlated with the colour pattern. Pec-

Most authors after Bleeker, 1865a consider the could be toral fin ray variation (18-24) not corre-

then known members of the a single species group lated, neither with fish characters nor with geo- and consider the in fin counts great variability ray graphic distribution, apart from T. kretamensis, and colour due to variation. pattern geographical showing a variation of 18-19 only.

I certainly do not exclude the possibility that they

but while and The in are right, examining comparing a first descriptions the fluviatilis-group

number of series I found discrimi- T. and T. relatively great are fluviatilis Hamilton, 1822, nigro- correlated with nating characters not directly geo- viridis Marion de Procé, 1822. Both species were graphical distribution. Very remarkable is the pres- described poorly and of neither of the two type

of all four here in Bor- known. The of ence species recognized material is description Hamilton, neo, mostly in variable series. This can be explain- however, was accompanied by excellent illustra- ed by the complicated Zoogeographie history of the tions, showing the form with three dark patches on

it is remarkable that T. steindach- the back and dark the sides. island. Anyway rounded, spots on neri is known from Thailand, Malaya and East The type locality is Bengal: Ganges River system. Java. Borneo, not from Sumatra or Likewise T. Marion de Procé did not even leave a drawing,

known Latin which fluviatilis is from India, Bangla Desh, only a short French and diagnosis,

Burma and Borneo, and not from the intermediate are so hard to obtain that I will give the Latin

Malaya, Sumatra, Java, Thailand, Cambodia or one in length (1822: 130): "T. corpore loevi; dorso

Vietnam, the intermediate countries being inhabit- viride splendente, maculis rotundis sicut ad latera.

ed by T. nigroviridis and T. steindachneri. P.18. D.12. A.10. C.8."; type locality: N.E. Coast

Of freshwater This is in course, one can explain the great variability of Sumatra, pool. description

in the with and the overlapping characters group by accordance the description (1849a) and figure assuming that speciation within the group is of (1865a) of T. potamophilus Bleeker and the

relatively recent origin. Perhaps a superspecies description of T. simulans Cantor, 1849, except for

but lack of material differences in fin P 22; D 14; concept would be very useful, ray counts (Bleeker: and field observations make this impossible at the A 12; Cantor: P 19-21; D 12-13; A 10-11). In

moment. Within the scope of the present study only his Atlas Ichthyologique (1865a) Bleeker, however, his three possibilities are open: either consider the gives P 18-22; D 12-14; A 11-12; in description

or subdivide which is based on group a polymorphic single species, of T. fluviatilis T. potamophilus

in the main material. Examination of two of T. simu- it many subspecies, or recognize syntypes

lans and Bleeker Collection of T. forms as species. specimens pota-

incorrect and The first possibility seems incon- mophilus convinced me of their conspecifity with venient as some constant discriminating charac- T. nigroviridis. ters can be given. The second possibility is unat- In 1852a — although noting the differences

— Blee- tractive as it would yield more than a dozen sub- in dorsal colourpattern and fin ray counts

So I choose third the that T. species on Borneo alone. the ker regards possibility potamophilus

in be of T. possibility, erecting a subspecies only the case merely might a climatological variety

of T. fluviatilis, as in this case two closely related fluviatilis. In 1865a he lumps all nominal species

— simu- populations with some constant discriminating T. nigroviridis, T. potamophilus and T.

characters are separated by some 3500 km, in lans — under the name of T. fluviatilis. In this which the Indonesian he followed since authors even Bleeker, combing out was uncritically ever by the and references listed Isles, could not find comparable specimens. copying synonymy by

earlier of mind in In an attempt to separate the species of the authors. Bleeker's change 1865a,

basis of characters however, was not based on additionalmaterial: he group on the morphological material of T. only the dorsal and anal fin ray counts and the still had no access to Indian

head length proved to be of taxonomie value, fluviatilis cf. Hamilton.

although the first proved to be valuable only In the present study comparisons were made

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with the following results: line system, but I found the given number of fin

rays (D 17-18; A 14) to be inaccurate. Examination T. fluviatilis fluviatilis (India, Ceylon, Bangla Desh, of the material fin syntypic yielded ray counts Burma): D 13-16 (m 14.9); A 11-14 (ml3.1); hi D 15-16 (m 15.4); A 12-14 (m 13.2), hence normal 2.4-3.1 (m 2.7) in si; nasal organ with spongy tissue for T. fluviatilis. The subspecies described in 1962 in 90% of specimens > 60 mm si; dark dorsal (Dichotomycterus rangoonensis dorsovittatus) nei- patches always present. ther showed discriminating characters in the T. nigroviridis: (Ceylon, Burma, Thailand, Malaysia, specimen indicated as "type" by Le Danois. Great Sunda Islands, S. Vietnam): D 12-14 (m In the introduction to the description of T. 13.1); A 10-12 (m 11.4); hl 2.1-2.6 (m 2.4) in si; rangoonensis, Le Danois states that T. viridipunc- nasal organ with spongy tissue in 50% of spec- tatus Day, 1865, in fact is T. fluviatilis cf. Hamil- imens > 60 mm si; dark dorsal patches always ton, without giving evidence supporting her absent. statement. Day's description (1865) and figure I do consider these differences of sufficient value (1878), however, show a strikingly differently

to re-establish T. with T. coloured and bands instead nigroviridis along fluvia- fish, bearing light spots tilis. of dark spots and patches.

In 1870 Steindachner described In 1865a Bleeker, in his synonymy of T. Crayracion fluviatilis, also listed Arothron semimaculatus fluviatilis var. ocellata. The type material can not

Riippell, 1852, a name for which a description be located at the moment, but the description and

allow identification the form well known never was given. It was only listed in a catalogue figures as of SMF with the annotations "Mare javanicum" by aquarists under the name of T. palembangensis:

small Tetraodon with dark dorsal and "Unter der Bezeichnung von T. fluviatilis a patches much

like in T. cf. but ornated H. Buchanan von Leyden erhalten" (Riippell, fluviatilis Hamilton, on

1852: 35). Given the locality it most probably back and sides with many light (yellow in life) broken concerns T. nigroviridis. lines and spots. The aquarists name is

Blyth (1860: 173), in searching for T. fluviatilis based on a wrong determination by Meinken in the who led Calcutta fish bazaars, discovered a nominal (1956), most likely was astray by a species "approximating it in appearance": Arothron doubtful collecting station (Sumatra). A similar

determination in scientific literature dorsovittatus, differing mainly in the distribution wrong was of the body spines, the absence of dark caudal bars made by Fowler (1934b, 1935) and probably Smith and a less typical dorsal colour pattern. Probably followed in part by (1945) and others.

after 1956 im- Blyth did not leave type material of his species, as Anyway, many specimens were

should be — Alfred traders and references Blyth's types according to Dr. ported by aquarium many

— made in literature cited in of NMS (in lit.) in BMNH, and no specimen were aquarium (not of the species could be located there. Judging from length), both sources indicating a maximum si (ca. the description the nominal species must be the 60 mm) far below the maximum si of T. fluviatilis,

and the lines same form as figured by Day, 1878, as T. fluviatilis, a very constant presence of light and

Annandale & Jenkins Examination of material shows these charac- which was distinguished by spots.

in (1910) as T. fluviatilis var. A and by Le Danois ters even to be constant the remote populations

of Thailand and Borneo. A (1959, 1962) as T. rangoonensis. T. fluviatilis cf. third discriminating

id. cf. Hamilton in character is the form of nasal Day and are distinguished the organ: flaps

broad literature mainly on base of the colour pattern of nearly as as long, not thickened, not provided

with tissue, of the the back, bearing more vague patches in the form spongy occupying 1/3-2/3 nasal cf. Day. Examination of both colour forms learnt organ length; this in contrast with the oblong, the colour differences not to be related with mostly thickened, often spongeous nasal flaps of morphological characters. Moreover, both forms T. fluviatilis and T. nigroviridis. were recorded from the same station (Ganges It is possible that the species is the same as

T. River Delta: Sattermukhi River) by Annandale & biocellatus Tirant, 1885. One can, however, not

be about that the Jenkins (1910). sure as description is very poor,

For T. rangoonensis (originally described in the no figures are known, and the type material could

not be located genus Dichotomycterus) Le Danois gives several (not present in MHNL; not indicat-

ed in list of discriminating characters. I can not judge the type MNHN). According to Le Danois differences with respect to osteology and lateral (1959) it concerns Tetraodon ocellatus Linnaeus,

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1758. This, however, is excluded the indication With the colour by 21.8). regard to pattern one might in the description of T. biocellatus: "deux tentacu- consider the Bornean T. fluviatilis the adult form les solides de chaque côté d'une fosse nasale of T. kretamensis. This, however, is contradicted

Linnaeus at is the imperforée"; T. ocellatus present by difference in fin ray counts. To consider the

forms placed in the genus Sphoeroides (sensu lato) two as two subspecies of a single species because of the double perforated nasal organs. is unprobable because Inger collected both forms

the ocellata the It proved necessary to replace name at same stations in the Pinang River. Dif- given by Steindachner by a new name to prevent ferences with the other intermediate form, T. confusion with the Linnaean as out include the fin the dorsal species, pointed nigroviridis, ray counts, colour in the etymology of T. steindachneri nom. nov. pattern and the nasal flap inner surface,

being spongy in 100% of the Bornean fluviatilis.

The last species recognized in the present group Affinities are closest with the Indian T. fluviatilis, North the was described by Inger (1953) from Borneo. colour pattern in some specimens being nearly

— kretamensis — is identical, This species T. mainly the spongy tissue occurring in nearly the discriminated from T. fin fluviatilis by lower ray same frequency and the number of caudal cross- counts (D 11-13, m 11.8; A 10; P 18-19, m 18.4), bars being the same. Main differences are the fin

surface of nasal and by the smooth inner the flaps. ray counts and head length, the latterin the Bornean

The colour pattern in some specimens resembles form being comprised 2.1-2.3 (m 2.2) in si, in the that of T. nigroviridis (mostly regularly rounded Indian form 2.4-3.1 (m 2.7). Because of the

of dark spots), in other specimens that T. fluviatilis arguments given above, I prefer to give the Bornean

dark chevron (a occipital followed posteriorly by form a subspecific rank rather than a specific one two great mid-dorsal spots), in still other specimens and name it T. fluviatilis sabahensis.

T. erythrotaenia (a ventrolateral dark band). Dif- ferences with T. erythrotaenia are discussed within Tetraodon fluviatilis sensu lato. the differences with T. erythrotaenia- group, fluvia- tilis are indicated above. Relationships with T. Below synonyms and references are listed which nigroviridis are less clear, the latter species oc- could apply to two or more species of the curring also in Borneo and showing strong fluviatilis-group. variation there, this variation including circular

head arrangement of the spots on the upper region Arothron semimaculatus and of ventrolateral longitudinal arrangement spots RÜPPELL, 1852: 35 (listed in catalogue of SMF) (nomen nudum). (tending to formation of a ventrolateral band). Crayracion fluviatilis; Moreover, in Borneo the lowest fin ray counts BLEEKER, 1866: 36 (listed; no additional material since in T. are found. Although does nigroviridis Inger Bleeker, 1865a). not consider the absence of dark caudal fin cross- BEAN & WEED, 1912: 611 (listed; Indonesia: Java, Bantam,

Welcome Bay). bars in his species an important character, it must Tetrodon fluviatilis; be noted that the original description is not correct GÜNTHER, 1870: 299—300 (description; India & Ceylon & in this after examination respect as two paratypes Singapore & Borneo; material in BMNH) (in sub- proved to these bars, though only faintly. possess genus Arothron).

On behalf of the facts described above it could be KÄROLI, 1882: 187 (listed; Singapore: Selita & Indonesia:

is isolated Borneo, Simunju; material in MNH, no. 1614— argued that T. kretamensis merely an reg. 1616, not available for study). variety or subspecies of T. nigroviridis. Since this VINCIGUERRA, 1890: 232 (listed; Burma: Rangoon; material form was already described as a species and no in MSNG). evidence is alter this it positive present to rank, WEBER, 1894: 458 (listed; no material).

1904: 191 West Kuala seems wise to keep things as they are. DUNCKER, (description; Malaysia: Selangor & West Malaysia: Kuala Langat, Muar River

Bandar material in at Maharani; SM, reg. no. 1097, The North Bornean situation is even more Selangor & in ZMH, reg. no. 8595 —6, Langat). the of complicated by presence a form considered VOLZ, 1904: 483 (listed; Indonesia: Sumatra, Lower to be T. Both in colour by Inger (1955) fluviatilis. Langkat, BatangSerangan). pattern and in fin ray counts this form is interme- DUNCKER, 1912: 270 (listed; Ceylon; no material men- diate between the latter and T. kretamensis. Dorsal tioned). HORA, 1923a: 766 (listed; India: Chilka Lake; material patches vary from the pattern typical of the first in ZSI). of the to the pattern latter, fin ray counts being VINCIGUERRA, 1926: 536 (description; East Malaysia: A 11-12 D 13-15 (m 13.8); (m 11.9); P 21-22 (m Sarawak; material in MSNG).

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HARDENBERG, 1931: 146 (listed; Indonesia: Sumatra, Philippines without exact record of such).

Rökan River mouth, Si Tsji Senebui). MERCKENS, 1958: 53—54 (description, aquarium data; no

DELSMAN & HARDENBERG, 1934: 374, fig. 264 (descrip- material).

tion, figure after Day, 1878; Indonesia: no material). STERBA, 1959: 634, 637, Taf. 274, 275 (in part) (descrip-

HARDENBERG, 1934: 5 (listed after Hardenberg, 1931). tion, aquarium data; no material).

HARDENBERG, 1935: 239 (listed; Indonesia: Borneo, DE BEAUFORT, 1962: 395 (in part) (description; ZMA

fish material in material of additional localities identified in present Samarinda, market; RMNH, reg. no. Monotretus). 15306). paper as T. nigroviridis) (in subgenus

HARDENBERG, 1936: 254 (listed; Indonesia: Borneo, Padangtikar Bay & Borneo, Terentang River & Dichotomycter fluviatilis; (listed; Sabah, Borneo, Pekadai Bay & Borneo, Ambawang River ABE, 1949: 130—131 East Malaysia: Sandakan & Sabah, Tawau River at Tawau & Sabah, mouth & Borneo, Peniti River; material not in RMNH Tawau or ZMA). Coast).

HARDENBERG, 1937: 13 (listed; Indonesia: Borneo, Kumai

River).

FOWLER, 1938: 236 (listed; no material). Tetraodon kretamensis Inger, 1953. PELLEGRIN & FANG, 1940: 123 (description; Laos: Ban Figs. 26-28. Nam Khueng, 30 km N.W. of Ban Houei Sai; material

in MNHN) (in subgenus Crayracion).

Tetraodon kretamensis WILLEM, 1947: 1—5, figs. 1—2 (description and figures INGER, 1953: 149 —152, fig. 27 (original description, of internal anatomy; no material).

nasal organ figured, discussion on differences with BECK, 1950: 240 (description, aquarium data; no material). T. leiurus and T. fluviatilis; East Malaysia: Sabah, LADIGES, 1954: 67, fig. 64 (aquarium data; no material). Kinabatangan District, Kretam Kechil River system, Tetraodon fluviatilis; Pinang River (type locality) and other stations of the SAUVAGE, 1883: 155 (listed; Thailand: Chao Phaya River). system (paratypes); material in FMNH, reg. no. 51558 VAILLANT, 1893a: 36 (listed; no material). (holotype) and 51562, 51559, 51560, 51561, 51563 VAILLANT, 1902: 28 (listed; Indonesia: Borneo, Pontianak, ). River material in (paratypes) Kapuas mouth; RMNH, reg. no. INGER, 1955: 82, 85 (description, notes on 7932). 52, 55, 81, additional ecology and food habits; no localities). JORDAN & RICHARDSON, 1909: 45 (listed; no material). INGER & CHIN, 1962: 192, 193, 209, 225, fig. 100B SEALE, 1910: 285 (description; East Malaysia, Sabah, dorsal colour pattern of head figured, material in (description, Sandakan; PBSF, reg. no. 2527, 2600, discussion on ecology; no additional localities). 2601, 2628, 2681, 2771). DE BEAUFORT, 1962: 398, 399 (description after Inger, HERRE, 1924: 497—498 (redescription of Seale, 1910 1953; no material seen by author) (in subgenus material).

discussion differences Monotretus). HORA, 1924: 499 (description, on

with T. fluviatilis cf. Day, 1878; Thailand: Thale Sap,

Songkhla; material in ZSI). Description of the paratypes. — HERRE, 1933: 5 (listed after Seale, 1910 and Herre, 1924).

INNES, 1935 (in ed. 1956): 496—497, fig. p. 496 (descrip-

General morphology. — Body oblong, cylindrical, tion, aquarium data; no material). head SUVATTI, 1936: 165 (listed; Thailand: Chanthaburi River; posteriorly compressed laterally, region some-

material in BFB). times slightly depressed; dorsal profile arched, HERRE & MYERS, 1937: 50 (listed; West Malaysia: Ubin highest at midst of back; interorbital convex, with- Island & Singapore & Indonesia: Sumatra Coast 100 out a groove; lateral line system mostly indistinct; miles W. of Singapore; material in NMS). small hidden ROXAS in 1 specimen, mostly & MARTIN, 1937: 257 (listed; no material). spines except

HERRE, 1940: 8 (listed; India: Andaman Islands, Middle under the skin, covering area between eyes and

Andaman, Long Island & Andaman Islands, South anus; origin of anal fin situated beneath anterior Andaman, Port Blair). half of dorsal fin base; mouth terminal, directed HERRE, 1941: 401 (listed after Herre, 1940). forwards; lower border of eye at level of mouth FRASER-BRUNNER, 1943: 14 (listed as representative of

and corner or lower, border not with genus subgenus; no material) (in subgenus upper interfering Chelonodon). dorsal nasal profile; organ a tentacle, more than

1947: 311 (listed; West Malaysia: Port Dickson; KOUMANS, distal half of which is divided in two flattened material in RMNH, reg. no. 17915). lobes; apposed surfaces of lobes always without BREDER & CLARK, 1947: 308 (reference to literature on tissue. spongy reproduction; no material).

SMITH, 1945: 577 (description; no additional material).

841 no additional Colour in alcohol. — colour HERRE, 1953: (listed; material). Ground of upper parts

1956: 10—12 BENL, 4, 6, 7, (description, aquarium data; tan, of belly yellowish white; both colours often no material). separated by an irregular dark band running from BOESEMAN, 1957: 77 (description; Ceylon, Colombo; mouth below and fin to meet band material in eye pectoral RMNH, reg. no. 20907).

of other side at anus or behind; back and sides HERRE, 1958: 80 (listed as known to occur on the

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Figs. 26—28. Tetraodon kretamensis Inger, 1953.

Fig. 26. Lateral view of FMNH 5161, paratype, sl 36 mm.

sl Fig. 27. Dorsal view of FMNH 5161, paratype, 36 mm.

Fig. 28. Nasal organ of FMNH 5161, paratype, sl 36 mm.

fin in end of with round and oblong dark spots of varying sizes; pectoral base 2.7-3.2 (m 2.9) si, at

back sometimes internasal dorsal fin base 5.3-6.7 in width spots on forming an (m 6.0) si;

band, an interocular band and/or — most often maximum 2.4-2.9 (m 2.6) in si and 0.8-1.1 (m 0.9)

— curved band in fin base 2.7-3.0 and most prominent a or V-shaped maximum depth, at pectoral

most of back between and in si and 0.9-1.1 in crossing eyes pectoral (m 2.8) (m 1.0) corresponding fins; lateral spots varying in size and number but depth, at end of dorsal fin base 6.6-8.1 (m 7.3) in always a spot at base of dorsal fin and at base of si and 1.1-1.4 (m 1.2) in corresponding depth; head

fin and chin in 2.6-2.9 caudal present; belly uniformly length 2.1-2.4 (m 2.2) si; snout length yellowish white; caudal fin plain or dusky, in 1 (m 2.7) in hi; interocular distance 1.6-2.0 (m 1.8) specimen with small dark spots suggesting cross in hi; eye horizontal diameter 2.8-3.6 (m 3.2) in bands; other fins plain. hi, distance from dorsal profile 8.6-21 (m 12.8) in

hi; nasal organ length 9.4-14.6 (m 12.1) in hi,

Morphometries (based on 11 specimens). — si 27- distance from tip of snout 3.4-4.2 (m 3.8) in hi,

45 (m 37) mm; tl 33-57 (m 45) mm; dorsal fin rays distance from eye 9.4-15.0 (m 12.1) in hi and 2.7-

11-13 (m 11.8) of which 2-3 (m 2.4) unbranched, 4.0 (m 3.3) in distance from tip of snout; mouth

6.5-7.7 in shortest width in longest ray (the 7-8th) (m 7.0) si, 3.0-3.6 (m 3.3) hi; upper lip depth 15.0- ray (the 1st) 18.3-29 (m 23) in si, base 8.2-10.7 21 (m 17.4) in hi; lower lip depth 12.9-21 (m 18.7) in si (m 9.3) and 1.2-1.5 (m 1.3) in longest ray; in hi. anal fin rays 10 of which 1-2 (m 1.4) unbranched,

Material examined. — longest ray (the 6-7th) 6.8-7.7 (m 7.2) in si, shortest

FMNH 51559; 5 specimens, paratypes, si 27—40 mm; ray (the 1st) 17.3-24 (m 20) in si; base 10.0-12.9 East Malaysia: Sabah, Kinabatangan District, Kretam (m 11.1) in si and 1.4-1.9 (m 1.6) in longest ray; Kechil River system, Pinang River; coll. R. F. Inger, pectoral fin rays 18-19 (m 18.4) of which 2 1950. FMNH 51560; 2 specimens, si 8.7 —9.6 unbranched; longest ray (the 5th) 7.1-8.7 in si; paratypes, mm; East Malaysia: Sabah, Kinabatangan District, Kretam shortest ray (the 1st) 15.7-25 (m 19.0) in si, base Kechil River; coll. R. F. Inger, 1950. 7.2-9.2 (m 8.6) in si and 1.0-1.3 (m 1.1) in longest FMNH 51561; 2 specimens, paratypes, si 11.7—36 mm; maximum 2.5-3.2 in at ray; depth (m 2.8) si, East Malaysia: Sabah, Kinabatangan District, Kretam

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Kechil River; coll. R. F. Inger, 1950. Tetrodon simulans

West FMNH 51562; 5 specimens, paratypes, si 33—45 mm; CANTOR, 1849: 1356—1357 (original description;

East Malaysia: Sabah, Kinabatangan District, Kretam Malaysia: Malayan Peninsula & West Malaysia: differences with Kechil River system, Gaja River; coll. R. F. Inger, 1950. Pinang & Singapore; discussion on

8.2—17.5 material least in FMNH 51563; 2 specimens, paratypes, si mm; T. fluviatilis Hamilton, 1822; at part

East Kretam in 60.3.19: 588 Malaysia: Sabah, Kinabatangan District, BMNH, reg. no. —589, syntypes) (in

Kechil River system, Gaja River; coll. R. F. Inger, 1950. subgenus Arothron).

BLEEKER, 1851b: 476, 478 (listed after Cantor, 1849; no

material).

Distribution. — T. kretamensis is known from only BLEEKER, 1852a: 26 (listed after Cantor, 1849; no Kechil in N. the Kretam River system Sabah, material).

Borneo, about 35 miles S.E. of Sandakan. It in- BLEEKER, 1852b: 60 (listed after Cantor, 1849; no material). habits the lower reaches of rivers, in brackish Arothron potamophilus; waters. The zoogeographical implications of the BLEEKER, 1854a: 314 (listed; Indonesia: Java, Perdana; discussed & who region are by Inger Chin, 1962, in 7357 material probably RMNH, reg. no. ex parte). point to the fact that the Labuk-Segama region, as BLEEKER, 1854b: 70, 76 (listed; Indonesia: Sumatra,

& Indonesia: Java & Indonesia: Borneo; they call it, contains most of the endemic species Sibogha material probably in RMNH, reg. no. 7357 ex parte). of primary freshwater fishes. Although kretamensis BLEEKER, 1854d: 260 (listed; Indonesia: Java & Indonesia: is not a freshwater fish it can thus be primary material in 7357 Borneo; probably RMNH, reg. no.

it is an endemic species — or perhaps argued that ex parte). a subspecies — of Tetraodon. BLEEKER, 1855a: 152 (listed; Indonesia: Borneo, Bandjer-

masin River; material probably in RMNH, reg. no.

7357 ex parte).

Etymology. — T. kretamensis was named after the BLEEKER, 1855d: 417 (listed; Indonesia: Borneo, Pontia-

nak & Borneo, material probably in type locality, Kretam Kechil River system. Bandjermasin; RMNH, reg. no. 7357 ex parte).

BLEEKER, 1856C: 419 (listed; Indonesia: Bangka, Muntok;

7357 material probably in RMNH, reg. no. ex parte).

BLEEKER, 1857C: 12 (listed; no additional material). Tetraodon nigro viridis Marion de Procé, 1822. BLEEKER, 1858a: 30, 35 (listed; no additional material). Figs. 29-31. BLEEKER, 1858b: 2 (listed; Indonesia: Borneo, Sinkawang;

material in 7357 probably RMNH, reg. no. ex parte).

1858C: 444, 445 (listed; Indonesia: Java, Dja- Tetrodon nigroviridis BLEEKER, karta & Java, Perdana & Java, Krawang & Java, MARION DE PROCÉ, 1822: 130 (original description; Tjikao & Java, Samarang & Java, Surabaya; material Indonesia: N.E. coast of Sumatra, freshwater pool; probably in RMNH, reg. no. 7357 ex parte). for material see neotype description). 1859a: 201 (listed; no additional material). Tetraödon potamophilus BLEEKER, BLEEKER, 1859b: 361, 374 (listed; Indonesia: Bangka, BLEEKER, 1849a: 16 (original description, discussion on Blinju, Klabat Bay & Bangka, Baturussak; material differences with T. fluviatilis Hamilton, 1822; Indone-

in no. 7357 ex parte). sia: Java Sea & Java, Strait Madura at Kamal and at probably RMNH, reg. BLEEKER, 1859c: 2 (listed; Indonesia: Borneo, Sinkawang; Surabaya & Java, river mouths near Djakarta and

material in no. 7357 ex material at least in in probably RMNH, reg. parte). Semarang; part RMNH, reg. BLEEKER, 1859d: 435 (listed; Indonesia: Borneo, Duri no. 7357 ex parte, syntypes). River; material probably in RMNH, reg. no. 7357 BLEEKER, 1849b: 12 (listed; no additional localities). BLEEKER, 1851a: 3, 5 (listed; Indonesia: Borneo, Banjer ex parte).

1860a: 66 no = BLEEKER, (listed; additional material). River Duson River, near Bandjermasin; material BLEEKER, 1860b: 447 (listed; Singapore; no material, just probably in RMNH, reg. no. 7357 ex parte) (in sub-

in sketchbook of De Castelnau). genus Chelonodon). seen

1860c: 7 no additional BLEEKER, 1851c: 60 (listed; no additional material). BLEEKER, (listed; material).

BLEEKER, 1860f: 238 (listed; Singapore; no material, just BLEEKER, 1851d: 262 (listed; no additional material). in sketchbook of De Castelnau). BLEEKER, 185If: 197 (listed; no additional material). seen material). BLEEKER, 1851g: 421 (listed; Indonesia: Borneo & Indone- BLEEKER, 1861b: 12 (listed; no additional BLEEKER, 1863: 73 Indonesia: Bangka; material sia: Sumatra & Indonesia: Java; probably no (listed;

in 7357 ex additional material). probably RMNH, reg. no. parte).

BLEEKER, 1852a: 4, 5, 7, 10, 17, 18 (description, discus- Dichotomyctère fluviatilis;

BIBRON 1855: 278 (listed as of sion on differences with T. fluviatilis Hamilton, 1822; (in DUMÉRIL), type genus;

Indonesia: material in B Java, Panimbang; material probably in MNHN, reg. no. 1500).

RMNH, reg. no. 7357 ex parte). Arothron simulans;

BLEEKER, 1852d: 408, 409, 415 (listed; Indonesia: Borneo, BLEEKER, 1858d: 251 (listed after Cantor, 1849; no

Pontianak & Borneo, Pamangkat). material).

BLEEKER, 1853e: 428, 429, 434, 438 (listed; Indonesia: BLYTH, 1860: 173 (listed; Burma: Moulmein).

Borneo, Sambas & Borneo, Pontianak; material prob- BLEEKER, 1861a: 69 (listed after Cantor, 1849; no

in 7357 material). ably RMNH, reg. no. ex parte).

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after material in BLEEKER, 1861b: 40 (listed Cantor, 1849; no phism; MNHN, not specified).

LE list material). DANOIS, 1961 (listed in type of MNHN as senior

Crayracion fluviatilis; synonym of material used by Bibron in establishing

1865a: tab. 210 4 the BLEEKER, 68, fig. (description, lumping genus Dichotomyctère; material in MNHN, reg.

of T. potamophilus Bleeker, 1849 with T. fluviatilis no. B1500).

Hamilton, 1822; no additional localities; material LE DANOIS, 1962: 694—700 (discussion on supposed

in 7357 and transitions between probably mostly RMNH, reg. no. ex parte). morphological osteological

KNER, 1867: 408 (description; Indonesia: Java; material D. rangoonensis Le Danois, 1959 and D. fluviatilis cf.

in Le 1959 T. probably NMW). Danois, = nigroviridis).

BLEEKER, 1868b: 290, 385 (listed; Indonesia: Bintang, Tetraodon fluviatilis;

in Rio; material probably RMNH, reg. no. 7357 ex MERCKENS, 1958: 54, 55 (in part), fig. p. 53 (description,

parte). aquarium data; no material).

Tetrodon fluviatilis; STERBA, 1959: 827 (in part), Taf. 274 (description,

GÜNTHER, 1870: 299—300 (in part) (description; material aquarium data; no material).

includes syntypes of T. simulans Cantor, 1849 and SCHNEIDER, 1964: 414—435, figs. 1, 3—9 (anatomical

Bleeker material of T. potamophilus Bleeker, 1849) studies of fin musculature).

(in subgenus Arothron). DE BEAUFORT, 1962: 392, 395—397 (in part) (description;

TIRANT, 1885 (in ed. 1929): 94—95 (description; South Indonesia: Java, Djakarta & Java, Tjilatjap &

Vietnam: surroundings of Saigon and Cho Lon) (in Indonesia: Sumatra, Deli; material in ZMA, reg. no.

subgenus Arothron). 108.952, 108.953, 108.955, respectively) (in subgenus

RANDOW, 1934: 561—563 (field observations and Monotretus).

aquarium data; Ceylon; no preserved material). Tetraodon leiurus;

Dichotomycter fluviatilis; INGER & CHIN, 1962: 191—192, fig. 102 (description,

FOWLER, 1937: 264, fig. 297 (listed; Thailand: Tachin; ecology; East Malaysia: Borneo, Sabah, Tawau

Kalabakan material in material probably in ANSP). District, River; FMNH, reg.

Dichotomycterus fluviatilis; no. 68480).

LE DANOIS, 1959: 132, 135—137, 246, 247, 250, 253—

255, figs. 84B, 89, 90, 91 (description; lateral view, General description. — dorsal skull and nasal discussion view, organ figured;

on differences with T. nigroviridis Marion de Procé,

General — Body oblong, posterior 1822 and Dichotomyctère javanicus Bibron, MS., morphology. dorsal evolution, zoogeography, osteology, sexual dimor- part compressed laterally; profile arched,

Figs. 29—31. Tetraodon nigroviridis Marion de Procé, 1822.

Fig. 29. Lateral view of ZMA 113.020, neotype, sl 64 mm.

Fig. 30. Dorsal view of ZMA 113.020, neotype, sl 64 mm.

31. Nasal of ZMA sl 64 Fig. organ 113.020, neotype, mm.

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highest at midst of back; interorbital convex, with- fin base 5.7-10.7 (m 7.6) in si and 1.0-2.2 (m 1.4) out a groove; lateral line system mostly indistinct; in corresponding depth; head length 2.1-2.6 (m 2.4)

small, often hidden under skin for in si; 2.3-2.9 hi; body spines snout length (m 2.7) in eye most part, not papillose, covering back, sides and horizontal diameter 2.8-5.4 (m 3.9) in hi, distance

from from belly eyes to anus; origin of anal fin situated dorsal profile 5.8-16.9 (m 8.5) in hi; inter-

anterior of dorsal fin beneath anterior ocular distance 1.5-2.0 in nasal slightly or (m 1.8) hi; organ half of dorsal fin base; mouth terminal, directed length 7.5-20 (m 10.2) in hi, distance from tip of forwards; lower border of eye slightly above level snout 2.9-4.2 (m 3.4) in hi, distance from eye 7.4- of mouth corner, upper border not interfering with 17.0 (m 10.7) in hi and 2.0-4.6 (m 3.1) in distance

from dorsal profile; nasal organ a tentacle, at least distal tip of snout; mouth width 2.8-4.4 (m 3.5) in half of which is divided into flattened hi; two and upper lip depth 7.5-22 (m 13.7) in hi; lower lip broadened lobes; apposed surfaces of lobes often depth 12.9-21 (m 15.5) in hi. with tissue 50% spongy (in of specimens > 60 mm

= in — si, N 19; 15% of specimens < 60 mm si, Material examined.

BMNH 1867.11.28: 108; 1 si 126 Indo- N = 29). specimen, mm; nesia; Bleeker Collection.

BMNH 60.3.19: 588 2 si 125 —589; specimens, 52, mm,

Colourin — Ground of alcohol. colour parts upper preserved dry; West Malaysia; coll. E. J. Comp, (syntypes tan, of lower parts yellowish white; back and sides of T. simulans Cantor, 1849).

BMNH, no reg. no.; 1 specimen, si 65 Philip- covered with many dark spots and rounded mm; pines; coll. H. 1836—1840. blotches diameter in Cuming, (maximal 10-100 si); spots FMNH 1 si 68480; specimen, 58 mm; East Malaysia: and blotches very variable in size and form, Borneo, Sabah, Tawau District, Kalabakan River; coll. sometimes sometimes encircled coalescent, by R. F. Inger, 2-VI-1956. thus sometimes stretch- lighter rings forming ocelli, MNHN 95—207, 208; 2 specimens, si 60, 60 mm; ed Thailand; coll. Bellanger. into thick longitudinal rows on caudal peduncle; NMW 64.297; 1 specimen, si 69 China: and blotches mm; Hongkong; spots of one specimen sometimes 1866. in size sometimes in all present one only, present RMNH 1 si Indonesia: 4580; specimen, 70 mm; possible sizes; ground colour often broken up by Borneo, Bandjermasin; coll. J. Semmelinck.

RMNH 7357; 61 tl 40—160 Indonesia; spots and blotches into spots and streaks; caudal specimens, mm; Bleeker Collection. fin mostly with 1-8 (m 3.8) dark transverse bands SMM 24403; 10 specimens, si 30—35 mm; live imported (in 65% of 60 specimens); dorsal fin sometimes from India: Calcutta; coll. A. Werner, 1956. with some small dark near base; other fins spots ZMA 102.307; 3 specimens, si 53—62 mm; Indonesia; plain; belly uniformly whitish. Bleeker Collection.

ZMA 108.950; 18 specimens, si 45 —57 mm; Indonesia

Borneo; coll. Teysman, 1877; ex KMA. Morphometries (based on 60 specimens). — si 26- ZMA 3 si 45—68 Indonesia: 108.952; specimens, mm; 143 (m 53) mm; tl 34-185 (m 70) mm; dorsal fin Java, Bay of Djakarta; coll. J. Buitendijk, 1-1910. 12-14 of which (m 13.1) 2-4 2.8) un- rays (m ZMA 108.953; 2 specimens, si 77 —79 mm; Indonesia: branched, longest ray (the 7-8th) 5.8-9.8 (m 7.3) in Java, Tjilatjap; coll. E. Jacobson, III-1911. si Indonesia: ZMA 108.955; 4 specimens, 56—143 mm; si, shortest ray (the 1st) 16.4-26 (m 20) in si, base Sumatra, Deli; coll. L. P. Cosquino de Bussy, 1905/ 6.7-11.0 (m 8.1) in si and 0.9-1.4 (m 1.1) in longest 1907. ray; anal fin rays 10-12 (m 11.4) of which 1-3 ZMA 110.211; 1 specimen, si 72 mm; Indonesia: South unbranched, 6.4-8.6 (m 1.9) longest ray (the 5-7th) Natuna Island; coll. Van Roosendaal; ex LOZB.

si 117 (m 7.5) in si, shortest ray (the 1st) 13.8-22 (m 18.1) ZMA 110.214; 1 specimen, mm; Indonesia:

Balikpapan; coll. Tissot van Patot, 1911. in si, base 7.5-12.5 (m 9.5) in si and 1.0-1.6 (m 1.4) Borneo, ZMA 110.217; 2 specimens, si 35, 36 mm; Indonesia: in longest ray; pectoral fin rays 18-23 (m 20.4) of Borneo, Upper Riko River; coll. Tissot van Patot, 1911. which 1-3 (m 2.1) unbranched, longest (the ray ZMA 110.375; 17 specimens, si 26—75 mm; Indonesia

5-7th) 6.0-10.7 (m 7.8) in si, shortest ray (the 1st) Borneo, Kota Baru; coll. H. A. Lorentz, 28-V-1909.

Indonesia: 15.2-23 (m 18.8) in si, base 6.7-11.0 (m 8.4) in si ZMA 112.470; 1 specimen, si 74 mm; Java, Djakarta; coll. J. Verwey; ex LOZB. and 0.9-1.3 (m 1.1) in longest ray; depth maximum ZMA 113.020; 1 specimen, si 64 mm; Indonesia: 1.9-3.2 (m 2.7) in si, at pectoral fin base 2.1-3.4 Sumatra, Deli; coll. L. P. Cosquino de Bussy, 1905/1907 in at end of dorsal fin base 4.2-7.1 (m 2.8) si, (neotype).

(m 5.5) in si; width maximum 2.1-3.4 (m 2.9) in

si and 0.9-1.5 (m 1.1) in maximum depth, at Description of the neotype. —

pectoral fin base 2.7-3.6 (m 3.1) in si and 0.9-1.4

(m 1.1) in corresponding depth, at end of dorsal Designation. — Marion de Procé based his original

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description on a single specimen collected during rays ii.9, longest ray (the 5th) 9.2 mm (7.0 in si),

a trip from France to Manila, Philippines. As shortest ray (the 1st) 4.6 mm (13.9 in si), base

in of the col- 6.3 in si stated an introductory note most mm (10.1 and 1.4 in longest ray); pectoral

lections made the lost in revolt. fin ii. 19, in during trip got a rays longest ray (the 6th) 9.0 mm (7.1

The descriptions were partly made on the basis of si), shortest ray (the 1st) 3.2 mm (20 in si), base

of drawings only. I assume the holotype T. 8.0 mm (8.0 in si and 1.2 in longest ray); depth

have been the of maximum 27 in fin base nigroviridis to among lost part mm (2.4 si), at pectoral

the collection, both because I could not trace it 26 mm (2.5 in si), at end of dorsal fin base 14.1

and "avait" in in because of the past tence of the word mm (4.5 si); width maximum 23 mm (2.8 si

in the next sentence accompanying the original and 1.2 in maximum depth), at pectoral fin base

description: "Un seul individu, observé dans une 23 mm (2.8 in si and 1.1 in corresponding depth),

mare d'eau douce sur la côte N.E. de Sumatra, at end of dorsal fin base 11.3 mm (5.7 in si and

avait environ de 1.3 in head 29 pouces de long" (Marion corresponding depth); length mm

Procé, 1822: 130). (2.2 in si); snout length 11.8 mm (2.4 in hi); eye

horizontal diameter The holotype being last and the species being 5.4 mm (5.4 in hi), distance

of from dorsal 2.8 in inter- part a very complicated species-group or super- profile mm (10.4 hi);

it ocular distance 16.8 in nasal species, I thought in accordance with the mm (1.7 hi); organ

International Code of Zoological Nomenclature to length 3.5 mm (8.3 in hi), distance from tip of

10.0 distance from designate a neotype. I choose it from a series snout mm (2.9 in hi), eye 3.1

collected in Sumatra, Deli, as Deli is an old mm (9.4 in hi and 3.2 in distance from tip of

"residentie" (province) of N.E. Sumatra (situated snout); mouth width 10.3 mm (2.8 in hi); upper

around Medan). From the series I choose the lip depth 1.7 mm (17.1 in hi); lower lip depth

1.9 specimen most closely matching the fin formula as mm (15.3 in hi). given by Marion de Procé (D 12; A 10; P 18).

Material examined. — The remaining difference compared to the formula ZMA 113.020; 1 specimen, si 64 mm; Indonesia: of the neotype (D 13; A 11; P21) can be explained Sumatra, Deli; coll. L. P. Cosquino de Bussy, 1905/1907 inaccurate Marion de Procé by counting by (most (neotype).

ichthyologists did not count the small foremost

Distribution. — As from the dorsal and anal fin rays) and, of course, by the seen synonyms,

in fin of references, and material examined the variability ray counts the species, being present is known from Burma, Thailand, from my experience D 12-14; A 10-12; P 18-23. species Ceylon, West and East Malaysia, Bintan, Sumatra, Bangka,

General — of lava, Borneo, Natuna and South Vietnam. morphology. Body posterior eyes Islands, compressed laterally; lateral line system distinct; The Chinese (Hongkong) specimen examined is body spines from nasal organs to anus, most no proof for the species occurring there, since the

densely distributed anteriorly and ventrally, most collector is unknown, the collecting date is long

two-rooted and and other from the distinctly anteriorly dorsally; origin ago (1866), no specimens region of anal fin slightly anterior of dorsal fin; nasal were subsequently recorded. The material imported organ flaps nearly reaching to base of nasal organ, from Calcutta is no proof either, as this town is a

surfaces with distinct well-known transit station in the apposed (left organ) or international indistinct (right organ) spongy tissue; otherwise like aquarium trade. general description. Herre, 1924, discusses the presence or absence of

the species on the Philippines and gives two

Colour in alcohol. — total with for its the Back and sides in arguments presence: presumed type

50 dark and recorded ca. spots (diameter 2-5 mm), mostly locality, Manila, a specimen by rounded, in 4 cases 2 spots coalescent; caudal fin Günther, 1870, which was collected by Hugh

first with 3 distinct and 1 indistinct transverse dark Cuming. The argument is based on persistent

other fins like authors of the lists of bars; plain; otherwise general copying by many synonymy description. earlier authors, as the type locality in fact is N.E.

The second is Sumatra. argument quite poor as

— is known careless Morphometries. si 64 mm; tl 84 mm; dorsal fin Hugh Cuming to have been quite

in in his Because other rays ii.ll, longest ray (the 7th) 10.1 mm (6.3 si), labelling collections. no since shortest ray (the 1st) 3.9 mm (16.4 in si), base material was recorded from Manila Cuming,

8.5 mm (7.5 in si and 1.2 in longest ray); anal fin it seems wise to await new material before

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of the distribu- rangoonensis; indicating the Philippines as part Dichotomycterus LE DANOIS, 1959: 21, 133—135, 246—248, 250, 251, tional area. 253, 254, figs. 84A, 85—88 (original description,

lateral view, dorsal view, frontal view, skull and nasal

— The name is derived discussion Etymology. nigroviridis organ figured; key; on osteology, evolution, from the Latin "niger", meaning "black" (refer- zoogeography, development, differences with T. fluviatilis cf. Hamilton, id. cf. 1878, and ring to the dark spots), and the Latin "viridis", 1822, Day, T. viridipunctatus Day, 1865; Burma: surroundings of meaning "green" (referring to the colour of the Rangoon & Bengal; material in MNHN, including parts in living specimens). upper B 1564).

LE DANOIS, 1961: 462 (in type list of MNHN; no ad- ditional material).

Tetraodon fluviatilis fluviatilis Hamilton, 1822. LE DANOIS, 1962: 694—700, figs. 1—11 (discussion with

figures on and transitions Figs. 32-34. morphological osteological between D. rangoonensis Le Danois, 1959 and D.

cf. T. fluviatilis Le Danois, 1959 = nigroviridis; Tetrodon fluviatilis material in MNHN, including reg. no. 61—1117). HAMILTON, 1822: 6—7, 362, pi. 30 fig. 1 (original de- Chelanodon fluviatilis; scription, lateral and dorsal view figured; India, West- MENON, 1961a: 50, 56, 61 (listed, description in key; Bengal or Bangla Desh: Ganges River system; no India: Chilka Lake off Bhasra Id; material probably material known, see neotype description). in ZSI). GÜNTHER, 1870: 299—300 (in part) (description; India: Dichotomycterus rangoonensis dorsovittatus; Malabar; material in BMNH) (in subgenus Arothron). LE DANOIS, 1962: 694—700, figs. 3, 5, 9 (description; DAY, 1878: 707, pi. 183 fig. 1 (description; discussion on Indo-Malayan Archipelago; material in MNHN, reg. Bleeker material from Indo-Malayan Archipelago; no. 61—1117 no. 3, indicated as "type"). India; material in part in BMNH) (in subgenus

Arothron).

1889: 496 after additional DAY, (description Day, 1878; General description. — locality from mutual communication: Burma: Amherst

District; no additional material).

General morphology. — Body oblong, compressed ANNANDALE & JENKINS, 1910: 15—16 (description of two in dorsal varieties: var. A after Hamilton, 1822 and var. B after laterally except fleshy specimens; profile

India: Day, 1878; A: Orissa Coast & India: Ganges arched, highest at midst of back; interorbital

River at Sara Ghat; India: Sattermukhi River; B: without lateral line convex, a groove; system material probably in ZSI). distinct; body spines small, sometimes very RANDOW, 1934: 561—563 (field observations, aquarium distinctly two-based, sometimes partly or wholly data; Ceylon; no material).

Tetraodon fluviatilis; indistinct, covering back, sides, and belly between

SWAINSON, 1839: 328 (listed as representative of genus; eyes and origin of dorsal fin; origin of anal fin

no material). situated beneath anterior half of dorsal fin base; INGER, 1953: 150—152 (in part) (in discussion on differ- mouth terminal, directed forwards; lower border ences with T. kretamensis and North Bornean speci-

of above or below level of mouth mens of T. fluviatilis; India: Ganges River at Allaha- eye slightly

material 139— bad; in BMNH, reg. no. 1934.10.17: corner, upper border not interfering with dorsal 149). profile; nasal organ a tentacle, more than distal BENL, 1956: Abb. 4 (figure illustrating description of T. half of which is divided in two flattened and fluviatilis sensu lato; no material). broadened lobes; apposed surfaces of lobes STERBA, 1959: Taf. 274 (figure illustrating description of with tissue 90% of T. fluviatilis sensu lato; no material). generally spongy (in specimens

1958: 54, 55 part), 51, 54 (descrip- = MERCKENS, (in fig. p. > 60 mm si, N 20; in 80% of specimens < 60

tion, aquarium data; no material). mm si, N=5). Tetraödon fluviatilis;

BLEEKER, 1853a: 78 (listed after Hamilton, 1822; no

Colour in alcohol. — colour material). Ground of upper parts

Arothron dorsovittatus tan, of belly yellowish white; three large yellowish-

BLYTH, 1860: 173—174 (original description; India: encircled dark patches on back between eyes and

Calcutta, fish bazaars; material not in ZSI or BMNH). dorsal fin: half anterior patch on posterior of head Crayracion fluviatilis; (length along midline of back 6.2-14.0 in si, and DAY, 1865: 314 (description of fin formulae only; no in material). 1.3-2.9 transverse length), middle patch between

Chelonodon fluviatilis; pectoral fins (length along midline of back 5.5-13.8

MUNRO, 1955: 282, pi. 55 fig. 822 (description; Ceylon; in si, and 0.8-1.8 in transverse length), posterior no material). patch in front of dorsal fin (length along midline of MENON, 1961b: 401 (listed; India: Sunnamber River at back 8.3-14.4 in si, and 0.6-1.3 in transverse Nonankuppam & India: Pondicherry & India: Tran- length);

material in middle dorsal broken into quebar; probably ZSI). patch occasionally up

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Figs. 32—34. Tetraodon fluviatilis fluviatilis Hamilton, 1822.

Fig. 32. Lateral view of BMNH 1934.10.17: 139, neotype, sl 93 mm

of Fig. 33. Dorsal view BMNH 1934.10.17: 139, neotype, sl 93 mm.

Fig. 34. Nasal organ of ZMA 108.954, sl 58 mm.

smaller anterior half of head anal fin of which two patches; variably longest ray; rays 11-14 (m 13.1)

coloured, often dark; sides with 5-28 2-4 unbranched, irregularly (m 2.1) longest ray (the 5-9th)

rounded, ocellated, dark (horizontal 6.1-8.1 in si, shortest 16-29 in mostly spots (m 7.1) ray (the 1st) base diameter 8-80 in si) of which always one at si, base 6.6-9.7 (m 8.3) in si and 0.9-1.4 (m 1.2) in

of dorsal fin the base of fin of (usually largest), one at longest ray; pectoral rays 19-24 (m 20.7)

caudal fin (usually the smallest), and one below or which 1-3 (m 2.0) unbranched, longest ray (the

against posterior dorsal patch; belly uniform 4-6th) 6.9-8.8 (m 7.9) in si, shortest ray (the 1st)

with 18.3-27 yellowish white or same but confluent spots (m 23) in si, base 7.1-9.1 (m 8.4) in si and

on sides in MNHN 0-9 0.9-1.3 in maximum as (especially 61-1117); (m 1.1) longest ray; depth

(m 3.8) crossbars on caudal fin; sometimes a small 2.4-3.2 (m 2.8) in si, at pectoral fin base 2.4-3.4

in dark spot on interior side of pectoral fin base; (m 2.9) si, at end of dorsal fin base 4.9-6.5

other fins plain. (m 5.7) in si; width maximum 3.1-4.0 (m 3.5) in si

and 1.1-1.5 (m 1.3) in maximum depth, at pectoral

— si 31- fin Morphometries (based on 25 specimens). base 3.1-4.6 (m 3.6) in si and 1.0-1.5 (m 1.3) in

108 80 tl end of fin base mm (m mm); 39-127 mm (m 96 mm); corresponding depth, at dorsal

dorsal fin rays 13-16 (m 14.9) of which 1-4 (m 3.1) 4.9-8.3 (m 7.2) in si and 1.1-1.5 (m 1.3) in unbranched, longest ray (the 6-10th) 6.0-7.5 (m 6.7) corresponding depth; head length 2.4-3.1 (m 2.7) in si, shortest ray (the 1st) 19.2-37 (m 23) in si, in si; snout length 2.3-2.9 (m 2.6) in hi; interocular base 5.9-8.0 (m 6.8) in si and 0.9-1.2 (m 1.0) in distance 1.5-2.0 (m 1.8) in hi; eye horizontal

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and the diameter 3.0-4.8 (m 4.3) in hl, distance from dorsal the colour pattern is well preserved, profile 4.8-9.7 (m 7.2) in hi; nasal organ length morphological characters fit very well with the

7.1-11.9 (m 8.9) in hi, distance from tip of snout general description.

2.8-3.6 (m 3.2) in hi, distance from eye 6.5-13.8

(m 9.8) in hi and 2.1-4.4 (m 2.8) in distance from General morphology. — Body spines covering tip of snout; mouth width 3.0-3.9 (m 3.4) in hi; back, sides and belly between eyes and anus,

10.4-14.7 in hi; lower of upper lip depth (m 12.3) lip distinctly two-rooted with exception spines on

14.4-26 in hi. anterior of nasal with depth (m 22) part back; organ spongy

tissue; otherwise like general description.

Material examined. —

BMNH 1911.12.6: 48; 1 specimen, si 78 mm; Kurrachu Colour in alcohol. — Anterior dorsal patch length (= India: Karachi ?); coll. J. W. Townsend. midline of back 24 in 2.1 in si along mm (3.9 si, BMNH 1934.10.17: 139; 1 specimen, 93 mm; see middle dorsal neotype description. transverse length); patch length

BMNH 1934.10.17: 140—149 + BMNH, no reg. no.; along midline of back 10.7 mm (8.7 in si, 1.8 in 10 9 si 75—108 India: River + specimens, mm; Ganges transverse length); posterior dorsal patch length at Allahabad; coll. Das (series to which neotype belongs). midline of back in in along 9.6 mm (9.7 si, 1.1 BMNH, no reg. no.; 1 specimen, si 78 mm; India: transverse 8 each side, horizontal Malabar; coll. F. Day. length); spots on

MNHN 61 —1117; 4 specimens, si 73—86 mm; Indo- diameter 4.4-7.5 mm (12-21 in si); caudal fin with

coll. MPD Malayan Archipelago; Arnoult; ex (described 4 dark transverse bars; other fins plain; otherwise by Le Danois (1962: 695) as: "types de transition entre like general description. Dichotomycterus rangoonensis et D. fluviatilis”).

MNHN B 1 si 48 India 1468; specimen, mm; or Bangla

— tl dorsal Desh: Ganges River; coll. Dussumier, ca. 1820. Morphometries. si 93 mm; 112 mm;

MNHN B 1564; 5 specimens, si 68—106 Burma: mm; fin rays iii. 12, longest ray (the 7th) 12.9 mm (7.2 mouth of coll. Expédition Rangoon; Reynaud, Chevrette, in si), shortest ray (the 1st) 4.4 mm (21 in si), base 1829 (syntypes of Dichotomycterus rangoonensis Le 14.2 mm (6.5 in si and 0.9 in longest ray); anal fin Danois, 1959).

ii.ll, 12.0 mm in NMW 64.048; 1 specimen, si 82 mm; Burma: Akyab; rays longest ray (the 5th) (7.8 si), coll. 1870. shortest Godeffroy, ray (the 1st) 5.0 mm (18.6 in si), base

NMW 64.292 2 specimens, si 31—42 mm; Burma: ; 11.3 mm (8.2 in si and 1.1 in longest ray); pectoral Rangoon; coll. Fea, 1893. fin ii. 11.5 rays 18, longest ray (the 5th) mm (8.1 ZMA 108.954; 2 specimens, si 58—59 mm; Bangla in si), shortest 4.0 mm in Desh: Ganges Delta at Khulna; coll. S. S. "Golden ray (the 1st) (23 si), base 11.3 in si in Crown"; coll. N. Annandale; ex IMC. mm (8.2 and 1.0 longest ray);

depth maximum 35 mm (3.0 in si), at pectoral fin

Description of the neotype. — base 35 mm (3.0 in si), at end of dorsal fin base

17.3 mm (5.4 in si); width maximum 29 mm (3.2 in

Designation. — As stated earlier, Hamilton did not si and 1.2 in maximum depth), at pectoral fin base leave materialof the species he described type many 28 mm (3.3 in si and 1.3 in corresponding depth),

(Gudger, 1924; Hora, 1929). According to general end of dorsal fin base in at 15.7 mm (6.0 si and custom the excellent drawings accompanying the 1.1 in corresponding depth); head length 31 mm

descriptions are taken as the basis for the species. in (3.0 si); snout length 10.7 mm (2.9 in hi); In the be insufficient. present case this seems to interocular distance 20 in mm (1.6 hi); eye Bleeker (1865a), Cantor (1849), Le Danois (1959) diameter horizontal 7.8 mm (4.0 in hi), distance and other authors in their efforts to discriminate from dorsal 6.3 in nasal profile mm (4.9 hi); organ related forms, suffered from lack of type material. length 4.0 mm (7.8 in hi), distance from tip of Because of and in connection with the this, snout 10.0 mm (3.1 in hi), distance from eye 3.8 (re)description of closely related species and mm (8.2 in hi and 2.6 in distance from tip of in consider the subspecies the present paper, I mouth width 10.3 in snout); mm (3.0 hi); upper selection of a a prerequisite for reducing neotype lip depth 2.4 mm (12.9 in hi); lower lip depth 2.1 taxonomical confusion within the There group. mm (14.8 in hi). were several reasons to choose the specimen here indicated as the neotype: it is collected from the Material examined. — river it is in locality a quite good state BMNH 1 si type system, 1934.10.17: 139; specimen, 93 mm; India: of preservation compared with the examined other Ganges River at Allahabad; coll. Das (neotype). series in American and European museums, it

Distribution. — belongs to a relatively large, homogeneous series, Considering the synonyms, refer-

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ences and material examined, T. fluviatilis fluviatilis Kretam Kechil River system, Pinang River; material

in is found in India, Ceylon, Bangla Desh and Burma. FMNH).

General description. — Etymology. — The name fluviatilis is Latin,

meaning of a river. It is biologically incorrect in General morphology. — Body oblong, posteriorly so far that the species is also recorded from compressed laterally, head region depressed in 2 brackish water with certainty and from salt water

larger specimens (90,108 mm si); dorsal profile with probability. arched, highest at midst of back; interorbital

convex, without a groove; lateral line system

distinct or indistinct; spines small (comparatively Tetraodon fluviatilis sabahensis nov. subsp. in 1 90 sometimes Figs. 35-37. large specimen, mm si),

indistinct, covering area between eyes and anus;

Tetraodon fluviatilis origin of anal fin situated beneath anterior half of

INGER, 1953: 150 —152 (in part) (in discussion on differ- dorsal fin base; mouth terminal, directed forwards;

ences with kretamensis T. and Indian specimens of lower border of eye at level of mouth corner, T. fluviatilis; East Malaysia: Sabah; material in upper border not interfering with dorsal profile; FMNH).

nasal a tentacle, more than distal half of INGER, 1955: 80, 81 (description; East Malaysia: Sabah, organ

Lower Gaja River & Sabah, Kretam Besar River which is divided in two flattened and broadened

mouth & Lower Pinang River; material in FMNH, lobes; apposed surfaces of lobes generally with

no. 51564, 51660—51662). reg. tissue all spongy (in specimens > 33 mm si). INGER & CHIN, 1962: 192, fig. 100A, C (description,

nasal organ and colour pattern of head figured,

Colour in alcohol. — Ground colour of ecology; East Malaysia: Sabah, Kinabatangan District, upper parts

Figs. 35—37. Tetraodon fluviatilis sabahensis nov. subsp.

Fig. 35. Lateral view of FMNH 51564 ex parte, holotype, sl 90 mm.

Fig. 36. view mm. Dorsal of FMNH 51564 ex parte, holotype, sl 90

Fig. 37. Nasal organ of FMNH 51564 ex parte, holotype, sl 90 mm.

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of lower st 36—90 East tan, parts yellowish white; four dark mm; Malaysia: Sabah, Kinabatangan marks District, Pinang River (05°30' N. 118°33' E.); coll. R. F. on back between eyes and dorsal fin: Inger, 10-V-1950. anteriormost mark a cross band between eyes, in FMNH 51660; 2 specimens, paratypes, si 103—108 mm; small specimens reduced to a second mark spot, East Malaysia. Mixed series: 1 specimen East Coast

most prominent, V-shaped, pointing backwards Residence, coll. R. F. Inger, 29-V-1950; 1 specimen

(length across back 2.8-5.7 in si and 2.4-4.8 in Kinabatangan District, Gaja River, Dewhurst Bay (05°30' N. 118°33' E.), coll. R. F. Inger, 14-V-1950. length along midline of back), third mark at midst FMNH 3 51661; specimens, paratypes, si 14—33 mm; of back, oblong (length across back 6.2-7.6 in si East Malaysia, East Coast Residence; coll. R. F. Inger, and 1.4-2.1 in length along midline of back), 25-V-1950.

mark in of dorsal posteriormost just front fin, FMNH 51662; 3 specimens, paratypes, si 36—90 mm;

East Malaysia, East Coast Residence; coll. R. F. Inger, rounded (length across back 9.0-15.7 in si and 10-VI-1950. 0.9-1.3 in length along midline of back); often a

dark band or spot connecting or between nostrils;

sides Description of the holotype. — with 7-16 rounded dark spots (horizontal in diameter 11-36 si), of which always one at base

General — of dorsal fin, one at base of caudal fin; belly morphology. Like general description. Head lateral line uniformly yellowish white or dirty; caudal fin with region depressed; system distinct;

1-7 dark bands between cross (m 3.9); other fins plain. spines comparatively big, covering area

eyes and anus. Morphometries (based on 10 specimens, holotype

and paratypes; 1 paratype, si 13.5 mm, excluded). Colour in alcohol. — All four dorsal marks

— si 30-108 40-135 (m 60) mm; tl (m 77) mm; present. Anteriormost mark length across back dorsal fin 13-15 (m of which 3-4 rays 13.8) (m 3.0) 16.7 in width mm (5.4 si), 2.7 mm (3.3 in si and unbranched, longest ray (the 6-9th) 6.5-8.3 (m 7.0) 6.2 in length); second mark length across back in shortest si, ray (the 1st) 18-31 (m 24) in si, base 22 mm (4.1 in si), width 6.6 mm (13.6 in si and 7.2-8.8 (m 8.1) in si and 1.0-1.3 (m in 1.2) longest 3.3 third mark in length); length across back 14.5 anal ray; fin rays 11-12 (m 11.9) of which 2-3 (m mm (6.2 in si), width 7.0 mm (12.9 in si and 2.1 2.2) unbranched, 6.4-7.7 longest ray (the 5-8th) in length); posteriormost mark length across back (m 7.0) in si, shortest (the 1st) 13-18 ray (m 15.8) 8.0 in width 7.5 mm (11.3 si), mm (12.0 in si and in si, base 9.4-10.9 (m 9.9) in si and 1.3-1.7 (m 1.4) 1.1 in between nostrils length); only a spot; 15 in pectoral fin 21-22 of longest ray; rays (m 21.8) lateral each spots on side; caudal fin with 5 cross which 2 unbranched, longest ray (the 4-6th) 6.2-7.0 bands made up of points. (m si, 6.9) in shortest ray (the 1st) 13-19 (m 16.5)

in si, base 7.6-8.4 (m 8.0) in si and 1.0-1.3 (m 1.2) Morphometries. — si 90 tl 118 dorsal in mm; mm; longest ray; depth maximum 2.0-3.0 (m 2.6) in fin 13.8 rays iii.ll, longest ray (the 6th) mm in at fin (6.5 si, pectoral base 2.4-3.1 (m 2.7) in si, at end shortest si), ray (the 1st) 5.0 mm (18.0 in si), base of dorsal fin base 4.6-6.0 (m 5.0) in si; width 11.8 mm (7.6 in si and 1.2 in longest ray); anal fin maximum 2.3-2.8 (m 2.6) in si and 0.9-1.1 (m 1.0) rays iii.9, longest ray (the 6th) 13.9 mm (6.5 in si), in maximum depth, at pectoral fin base 2.5-3.0 shortest ray (the 1st) 5.9 mm (15.2 in si), base (m 2.9) in si and 0.9-1.3 (m 1.1) in corresponding 9.5 mm (9.5 in si and 1.5 in longest ray); pectoral depth, at end of dorsal fin base 5.3-8.3 (m 6.8) in fin rays ii.20, longest ray (the 4th) 14.5 mm (6.2 si and 1.2-1.5 (m 1.3) in corresponding depth; head in shortest 6.1 si), ray (the 1st) mm (14.8 in si), length 2.1-2.3 (m 2.2) in si; snout length 2.2-2.9 11.9 in base mm (7.6 si and 1.2 in longest ray); (m 2.6) in hi; interocular distance 1.4-1.8 (m 1.6) maximum 44 depth mm (2.0 in si), at pectoral in hi; eye horizontal diameter 2.8-5.2 (m 3.7) in hi, fin base 38 mm (2.4 in si), at end of dorsal fin from distance dorsal profile 7.8-11.7 (m 9.6) in hi; base 19.8 mm (4.6 in si); width maximum 39 mm nasal organ length 7.3-12.4 (m 9.4) in hi, distance (2.3 in si and 1.1 in maximum at from depth), pectoral tip of snout 2.8-3.8 (m 3.2) in hi, distance fin base 36 mm (2.5 in si and 1.1 in from corresponding eye 6.6-20 (m 12.2) in hi and 2.3-5.3 (m 3.8) end of depth), at dorsal fin base 16.9 mm (5.3 in in distance from tip of snout; mouth width 2.6-3.5 si and 1.2 in head in corresponding depth); length (m 3.1) hi; upper lip depth 13.8-18.8 (m 16.4) 40 mm (2.3 in 17.2 si); snout length mm (2.3 in hi); in hi; lower lip depth 15.4-26 (m 21) in hi. interocular distance 28 in mm (1.4 hi); eye horizontal diameter 8.8 mm in Material examined. — (4.5 hi), distance

FMNH 3 from dorsal 51564; specimens, holotype and paratypes, profile 4.0 mm (10.0 in hi); nasal

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in material be located organ length 5.4 mm (7.4 hi), distance from tip pore: Pengulon Patie; can not at of snout in distance from present). 14.5 mm (2.8 hi), eye Tetrodon palembangensis; 4.7 mm (8.5 in hi and 3.1 in distance from tip of FOWLER, 1934b: 351, fig. 13 (listed; Thailand: Kratt; mouth width 14.5 in snout); mm (2.8 hi); upper material in ANSP, reg. no. 60239). in lower lip depth 2.9 mm (13.8 hi); lip depth FOWLER, 1935: 163 (listed; Thailand: Srisawat; material in

ANSP, can not be located at 2.6 mm (15.4 in hi). present). Tetraodon palembangensis;

MEINKEN, 1956: 61—63, fig. p. 61 (description, aquarium

Material examined. — live in data; unde, imported; material ZMH, reg. no. FMNH 51564 ex parte; 1 specimen, holotype, 90 mm 1252). si; East Malaysia: Sabah, Kinabatangan District, Pinang GROBE, 1956: 111 (aquarium data; no material). River (05°30' N. 118°33' E.); coll. R. F. Inger, 10-V-1950. BENL, 1956: 4, 6, 7, 10, Abb. 6 (description, aquarium

data;no material).

and distribution.— The name sabahen- Etymology MERCKENS, 1959: 170 (description, aquarium data; no sis refers to the impression that the subspecies is material). STERBA, 1959: 640, Taf. 256 (description, aquarium data; geographically confined to Sabah and perhaps to no material). other parts of Borneo.

General description. —

Tetraodon steindachneri nom. nov. General morphology. — Body oblong, slightly Figs. 38-40. compressed laterally; dorsal profile arched, highest

at midst of interorbital without back; convex, a Crayracion fluviatilis var. ocellata groove; lateral line system distinct; body spines STEINDACHNER, 1870: 640, Taf. 5 figs. 2, 2a (original sometimes description; lateral and dorsal views figured; Singa- small, partly or wholly indistinct,

Figs. 38—40. Tetraodon steindachneri nom. nov.

Fig. 38. Lateral view of syntype after Steindachner (1870: tab. 5, fig. 2).

after Steindachner tab. Fig. 39. Dorsal view of syntype (1870: 5, fig. 2a).

Fig. 40. Nasal organ of ZMA 112.580, sl 47 mm.

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covering back, sides, and belly between eyes and 0.8-1.2 (m 1.0) in longest ray; depth maximum 2.3-

origin of dorsal fin; origin of anal fin situated 3.3 (m 2.8) in si, at pectoral fin base 2.4-3.3 (m 2.8)

beneath anterior half of dorsal fin base; mouth in si, at end of dorsal fin base 5.5-7.1 (m 6.3) in

terminal, directed forwards; lower border of eye si; width maximum 2.6-3.3 (m 2.9) in si and 1.0-

border slightly above level of mouth corner, upper 1.3 in maximum depth, at pectoral fin base 2.6-

not interfering with dorsal profile; nasal organ a 3.5 (m 3.1) in si and 1.0-1.2 (m 1.1) in correspond-

tentacle, more than distal half of which is divided ing depth, at end of dorsal fin base 6.4-9.0 (m 7.3)

into two flattened and broadened lobes; apposed in si and 1.1-1.3 (m 1.2) in corresponding depth;

surfaces of lobes smooth. head length 2.3-2.5 (m 2.4) in si; snout length

2.4-2.9 (m 2.8) in hi; interocular distance 1.7-2.0

— colour horizontal Colour in alcohol. Ground of upper (m 1.8) in hi; eye diameter 2.9-3.9 (m

parts tan, of belly yellowish white; three large 3.3) in hi, distance from dorsal profile 7.4-25 (m

back yellowish-encircled dark patches on between 13.8) in hi; nasal organ length 7.6-13.3 (m 9.8) in

and dorsal fin: anterior distance in eyes patch on posterior hi, from tip of snout 3.1-3.9 (m 3.5) hi,

half of head midline of back 4.4-9.2 distance from 7.5-14.3 in hi and 2.4- (length along eye (m 11.3)

in si and 1.3-2.4 in transversal length), middle 4.3 (m 3.2) in distance from tip of snout; mouth

between fins midline width in hi; patch pectoral (length along 2.9-3.9 (m 3.4) upper lip depth 9.1- of back 4.3-5.7 in si and 0.7-1.1 in transversal 15.5 (m 12.2) in hi; lower lip depth 8.7-24 (m 16.7)

length), posterior patch in frontof dorsal fin (length in hi.

along midline of back 5.7-14.8 in si and 0.6-2.0 in

transversal length); sometimes (in 2 out of 7 Material examined. —•

in anterior specimens) a light centre patch (dia- ANSP 60239; 1 specimen, si 32 mm; Thailand: Kratt;

coll. R. M. de Schauensee of the Third De Schauensee meter 12-30 in si); sometimes (in 1 out of 7 Siamese Expedition, XII-1933. specimens) a light centre in middle patch (diameter RMNH 15306; 3 specimens, si 35—46 mm; Indonesia: 30 in si); a yellowish band like the ones encircling Borneo, Samarinda; coll. Witkamp, 29-111-1931; ex LOZB. dorsal between nasal patches organs giving impres- ZMA 112.580; 1 specimen, si 47 mm; unde, imported

sion alive; died at NAM, 1965. of a fourth dark patch between nasal organs

ZMH 1252; 2 specimens, si 34—59 mm; unde, imported and hind border of eyes; sometimes (in 1, the alive; donation Meinken, 1960. largest out of 7 specimens) 2 more yellowish bands,

the anterior of nasal sides crossing snout organs;

Description of the — with similar bands and lines, often bended or syntypes.

broken up in oblong or rounded spots; sides with

2-4 dark base of dorsal fin The first description of T. steindachneri was made ocellated spots: one at

by Steindachner (1870). The description was based (always present, the largest), one at base of caudal

fin base of fin on 4 specimens. These syntypes are most probably (always present), one at pectoral

in NMW, but can not be located there (present in 2 out of 7 specimens), one anterior of present they

at the moment (Kähsbauer, in lit., 1971). It is eyes (present in 5 out of 7 specimens); fins plain

for ocellus interior side of possible that Steindachner returned the specimens except an on pectoral

fin base. to the collector, Ransonnet, and that from there

elsewhere. they went Anyway, the syntypes can not

be located, but there is no proof that they are lost. Morphometries (based on 7 specimens). — si 32-59

So, a neotype can not be selected from other (m 42) mm; tl 41-70 (m 51) mm; dorsal fin rays material. Confusion, will arise be- 13-15 (m 14.0) of which 2-5 (m 2.9) unbranched, however, not of the 6.3-10.2 in si, cause excellent illustrations by Steindachner, longest ray (the 7- 10th) (m 7.6) which are reproduced hereby in adapted form shortest ray (the 1st) 12.1-24 (m 20) in si, base (figs. 38-39). 6.5-7.8 (m 7.2) in si and 0.7-1.1 (m 1.0) in longest ray; anal fin rays 11-13 (m 12.0) of which 1-2

(m 1.6) unbranched, longest ray (the 7-8th) 6.7-9.8 Distribution. T. steindachneri is known only

in shortest in from (m 8.0) si, ray (the 1st) 14-23 (m 18) West Malaysia (including Singapore), Thai-

si, base 8.0-9.8 (m 8.9) in si and 0.8-1.2 (m 1.1) in land and Borneo. The specimens described by fin longest ray; pectoral rays 18-21 (m 19.3) of Meinken (1956) were said to have been collected

which 2-3 unbranched, in Sumatra. This (m 2.1) longest ray (the locality was considered in the

5-8th) 7.4-9.8 (m 8.7) in si, shortest ray (the 1st) publication as doubtful, because the specimens

in base 16-27 (m 22) si, 7.7-8.9 (m 8.2) in si and were imported by aquarium trade from Penang.

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Both under the former is the Etymology. names which the species phrase more applicable to

been known until invalid. The name smaller the latter to the In has now are specimens, larger ...

T. palembangensis belongs to a distinctly different this statement Regan misses the point: in the first is nasal species. The name Crayracion fluviatilis var. ocel- place he obviously describing the organs lata is subject to the rules of the International of T. fluviatilis (he uses the plural form: specimens),

Code of Zoological Nomenclature as Steindachner in the second place there was no problem in the is considered to have described a subspecies (art. nasal organs being splitted only terminally or to

elevated and the but in 45d-i). After being to species rank base, being patoca-like or not. being placed in the genus Tetraodon by me, the A last clue to the form of the nasal organ could name becomes a junior secondary homonym of be the figure in the Atlas Ichthyologique. This

five but Tetraodon ocellatus Linnaeus, 1758, (art. 57) and shows the nasal organ possessing flaps,

and obvious whether these must be rejected replaced (art. 53) even though it is not represent only one the Linnaean species is at the moment generally or both of the nasal organs, although in the other different nasal believed to belong to a genus, Sphoeroides figures of Tetraodon species only one organ

additional is shown. in or Torquigener. An reason to rename T. patoca general only possesses two the species, not prescribed by the Code, is to avoid flaps. Most probably the figure is not correct in confusion with the name T. ocellaris Klausewitz, this aspect. the in honour of the first that the form of the 1957. I renamed species Concluding we can say author to it distinct from nasal is not known. Bleeker, in his recognize as typical organ Probably fluviatilis. emended description, was right and the species is

close to T. patoca. The latter possesses a totally

different colour pattern, consisting mainly of oval waandersii -group whitish spots on the upper half of the body, never

possessing the dark transverse bandelets like in

— with about Diagnosis. Tetraodon species 35 not in Sphoeroides oblongus, even young specimens. dark transverse bandelets on half of body; upper It is, however, known that T. patoca is very D12;A11;P16. One species: T. waandersii. variable in some characteristics, hence the great number of allied species described. This is

Discussion. — T. waandersii was de- originally understandable of a species having the widest scribed Bleeker in 1853 from by a single specimen, distribution area of all freshwater tetraodonts.

Island, fresh type locality Bangka Marawang, I consider it the best to await new material from water. No was recorded ever since, so specimen the Bangka Island before deciding on the affinities is check the of there reason to validity the of it in of its every T. waandersii, listing a group own.

species. The only other true Tetraodon known from the

Bangka Island is T. nigroviridis, a completely A in the great problem studying unique specimen differently coloured species.

is the state of the nasal as no poor organs, generic A strong argument for T. waandersii being a allocation be based this character. can anymore on of is the number of 16 young T. patoca P rays:

In literature there is confusion about the organ. 11 fin (not as stated by Bleeker) as the pectoral The original description (Bleeker, 1853b: 194) ray counts of T. patoca vary according to De

states: nasali latere "papilla utroque simplice Beaufort (1962) from 15 to 16, a number which is indivisa but his emended oblonga" in description unknown in Indonesian Tetraodon species.

of 1865a (: 77) Bleeker says: "Ces organes, en

la forme entonnoir à effet, ont en non perforé et

bord libre fimbrié comme le Leiodon patoca". The Tetraodon waandersii Bleeker, 1853. latter statement would imply that the species Figs. 41-42. to belongs the patoca-group, often considered a

separate subgenus or named Chelonodon. genus Tetraödon Waandersii however, Regan (1902: 302-303), synonymized BLEEKER, 1853b: 176, 182, 194 (original description; T. waandersii with T. fluviatilis Hamilton, 1822, Indonesia: Bangka Island, Marawang; material in

BMNH, no. 1867.11.28: 112, holotype). stating that "The supposed difference in the nasal reg. Rohita Waandersii; organs is non-existent, they might impartially be BLEEKER, 1853b: 185 (name misprinted). described either as a short tube with two terminal Arothron Waandersii; tentacles united lips, or as two basally, although BLEEKER, 1854d: 260 (listed; no additional material).

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1859a: 202 additional — BLEEKER, (listed; no material). Colour in alcohol. Ground colour of upper parts additional BLEEKER, 1859b: 374 (listed; no material). tan, of belly yellowish; about 35 transverse dark Leiodon Waandersii; bandelets half of on upper body from snout to BLEEKER, 1865a: 76—77, tab. 214 fig. 3 (emendated

caudal peduncle (still visible at a few of description; no additional material). present),

BLEEKER, 1866a: 38 (listed; no additional material). which have their ends branchedor mutually united;

Tetrodon waandersii; fins plain, caudal fin darkest.

GÜNTHER, 1870: 289 (description; no additional material)

(in subgenus Chelonodon).

additional — WEBER, 1894: 458 (listed; no material). Morphometries. tl 45 mm (41 mm at present); Tetrodon fluviatilis; caudal fin length 5 in tl, so si was 36 mm (35 mm REGAN, 1902: 302—303 (in part) (comparison of holo- fin at present); dorsal rays ii.10, longest ray longer of T. waandersii with of T. type young specimens than base; anal fin ii.9, longest longer fluviatilis sensu lato; no additional material). rays ray than fin ii.14 de- Tetraodonfluviatilis; base; pectoral rays (original DE BEAUFORT, 1962: 395—397 (in part) (listed in syno- scription; i.10); depth maximum 4 in tl, so 11 mm after additional nymy, probably Regan, 1902; no (3.3 in si); front part of body as deep as wide, so material). width maximum about 11 mm (3.3 in si); head

dia- length 3.3 in tl, so 13.6 mm (2.6 in si); eye

Description of the holotype. — meter 3.3 in hi, so 4.1 mm.

Recognition. — In all publications by Bleeker only

Material examined. — one specimen is recorded; no further specimens BMNH 1867.11.28: 112; 1 specimen, si 36 mm, holo- were recorded ever since. So, the specimen Indonesia: Bangka type (si 35 mm at present); Sumatra, recorded by Günther (1870: 289) as "Type of the Island, Marawang, fresh water; coll. H. L. van Bloemen species" must indeed be the holotype. The correct- Waanders, about 1853; presented to Bleeker about 1853).

underlined the fact ness of this view may be by that the catalogue of the auction of the Bleeker Note. — Due to the bad state of preservation of

Collection does not mention the made species, apparently the unique specimen the description was because the specimen was already incorporated in also using the descriptions and the figure provided the collections of BMNH. by Bleeker (1853b, 1865a).

— dorsal General morphology. Body oblong; from Distribution. — The species is known only profile of head rounded; interorbital convex, Indonesia: Sumatra, Bangka Island, from fresh without a groove; lateral line system indistinct; water. sides and from spines on back, belly, eyes to near origin of dorsal fin; origin of anal fin beneath

— The named in honour anterior half of dorsal fin base; for form of nasal Etymology. species is

waandersii- of the collector, H. L. van Bloemen Waanders. organ see group.

Figs. 41—42. Tetraodon waandersii Bleeker, 1853.

1867.11.28: sl 36 after Bleeker tab. Fig. 41. Lateral view of BMNH 112, holotype, mm; (1865a: 214, fig. 3).

Fig. 42. Lateral view of nasal organ(s) of BMNH 1867.11.28: 112, holotype, sl 36 mm; after Bleeker (1865a: tab. 214, fig. 3).

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REFERENCES

ABE, T., 1949. Taxonomie studies on the puffers (Tetrao- 1849b. tot de kennis —, Bijdrage der ichthyologische

dontidae, from Japan and regions. fauna van Teleostei) adjacent Midden- en Oost-Java, met beschrijving van

V. Synopsis of the puffers from Japan and adjacent eenige nieuwe species. Verh. batav. Genoot. Kunst.

regions. Bull, biogeogr. Soc. Japan, 14 (13): 89—140, Wet., 23 (12): 1—23.

1—2. 1851a. Bijdrage tot de kennis der pis. —, ichthyologische A list of fishes obtained in the ABREU, E. A. D', 1925. fauna van Borneo met beschrijving van 16 nieuwe

Saran District, Behar, chiefly from the Ghogra River soorten van zoetwatervisschen. Natuurk. Tijdschr. and in its near the town of Chapra vicinity. J. Bom- Ned.-Indië, 1: 1—16.

bay nat. Hist. Soc., 30 (3—4): 707—711. 1851b. de kennis der —, Bijdrage tot ichthyologische

ANNANDALE, N. & J. T. JENKINS, 1910. Report on the fauna van Riouw. Natuurk. Tijdschr. Ned.-Indië, 2:

fishes taken by the Bengal Fisheries Steamer "Golden 469—497.

Crown". Part III. Plectognathi and Pediculati. Mem. 1851c. Derde bijdrage tot de kennis der —, ichthyologi-

(1): — Indian Mus., 3 7—21, pis. 1 2. sche fauna van Borneo met beschrijving van eenige

AUBENTON, F. D' & M. BLANC, 1966. Poissons tétrao- nieuwe soorten van zoetwatervisschen. Natuurk. Tijd-

dontiformes du Cambodge. Bull. Mus. natn. Hist. nat. schr. Ned.-Indië, 2: 57—70.

(2) 38 (5): 1—4. 185 ld. Nieuwe tot de kennis der Paris, 554—561, figs. —, bijdrage ichthyolo-

AXELROD, H. R., C. W. EMMENS, D. SCULTHORPE, W. gische fauna van Borneo met beschrijving van eenige

1967. Exotic VORDERWINKLER & N. PRONEK, tropical nieuwe soorten van zoetwatervisschen. Natuurk. Tijd-

fishes: M 3.00—M 121.00, P 2.00— P 93.00, C 3.00— schr. Ned.-Indië, 1: 259—275.

C F 1.00—F 900 F. H. 1851e. Over drie nieuwe soorten Tetraödon 48.00, 608.00, ca. figs. (T. —, van van

Publications, Jersey City). den Indischen Archipel. Natuurk. Tijdschr. Ned.-Indië,

BEAN, B. A. & A. C. WEED, 1912. Notes on a collection 1: 96—97.

of fishes from made Owen and 185If. Vierde tot de der Java, by Bryant —, bijdrage kennis ichthyolo-

William Palmer in 1909, with description of a new gische fauna van Borneo met beschrijving van eenige

natn. species. Proc. U.S. Mus., 42 (1919): 587—611, nieuwe soorten van zoetwatervisschen. Natuurk. Tijd-

figs. 1—3, pis. 73—75. schr. Ned.-Indië, 2: 193—208.

BEAUFORT, L. F. 1913. Fishes of the eastern of 1851g. Vijfde bijdrage tot de kennis der DE, part —, ichthyolo-

with on its the Indo-Australian Archipelago remarks gische fauna van Borneo met beschrijving van eenige

zoogeography. Bijdr. Dierk., 19: 95—164, figs. 1—8, nieuwe soorten van zoetwatervisschen. Natuurk. Tijd-

pi. 2. schr. Ned.-Indië, 2: 415—442.

1926. van den Indischen Archipel: 1852a. Bijdrage de kennis der vis- —, Zoogeographie —, tot blootkakige

1 schen den Soenda-Molukschen be- 1—202, figs. 1—8, map (Erven Bohn, Haarlem). van Archipel met

1939. On collection of freshwater fishes of the nieuwe Verh. batav. —, a schrijving van eenige soorten.

island of Billiton. Treubia, 17 (3): 189—198, 1 fig. Genoot. Kunst. Wet., 24 (10): 1—26.

1951. of the land and inland 1852b. de kennis der —, Zoogeography waters: —, Bijdrage tot ichthyologische

i—viii, 1—208, figs. 1—10 (Taylor & Francis, fauna van Singapore. Natuurk. Tijdschr. Ned.-Indië, 3:

London). 51—86.

1962. The fishes of the Indo-Australian 1852c. van nieuwe of —, Archipelago. —, Diagnostische beschrijvingen

XI. Scleroparei, Hypostomides, Pediculati, Plectogna- weinig bekende vischsoorten van Sumatra. Tiental

thi, Opisthomi, Discocephali, Xenopterygii: i—xi. I—IV. Natuurk. Tijdschr. Ned.-Indië, 3: 569—608.

1852d. Zesde tot de kennis der 1—481, figs. 1—100 (Brill, Leiden). —, bijdrage ichthyologi-

1964. Bleeker's collection in the Museum sche fauna van Borneo. Visschen van —. Zoological Pamangkat,

of Amsterdam. Beaufortia, 11 (131): 23—25. Bandjermassing, Praboekarta en Sampit. Natuurk.

BECK, P., 1950. Traité complet de la vie des animaux en Tijdschr. Ned.-Indië, 3: 407—442.

1—103 1853a. de ichthyologische fauna van aquarium: 1—269, figs. (Payot, Paris). —, Nalezingen op

BENL, G., 1956. Siisswasser-Kugelfische. Aquar. - u. Bengalen en Hindostan. Verh. batav. Genoot. Kunst.

Terrar.-Z., 9 (6, 7, 8): 141—147, 172—175. 202— Wet., 25 (8): 1—164, tabul. 1—6.

1853b. de fauna van 205, Abb. 1—14. —, Nalezingen op ichthyologische

1957a. Tetraodon leiurus brevirostris, nova. het eiland Banka. Natuurk. Tijdschr. Ned.-Indië, 5: —, subsp.

Aquar.- u. Terrar.-Z., 9: 63—65, Abb. 1—4. 175—194.

1957b. Carinotetraodon 1853c. Vierde tot de kennis der —, chlupatyi nov. gen., nov. —, bijdrage ichthyologi-

ein Kiel sche fauna Amboina. Natuurk. Ned.- spec., Kugelfisch mit Kamm und (Pisces, van Tijdschr.

Fam. Tetraodontidae). Opusc. zool. Münch., 5: 1—4, Indië, 5: 317—352.

1 fig. 1853d. Vierde bijdrage tot de kennis der ichthyologi- —, Celebes. Natuurk. 1959. Zur Nomenklatur Siamesischer sche fauna van Tijdschr. Ned.-Indië, —, Kugelfischfor-

men. Aquar.- u. Terrar.-Z., 12 (2): 42—44. 5: 153—174.

G. & P. 1957. eine 1853e. Zevende tot de kennis der Benl, Chlupaty, Carinotetraodon, —, bijdrage ichthyologi-

sche fauna Borneo. Zoetwatervisschen Sam- neue Kugelfischgattung. Aquar.- u. Terrar.-Z., 10 van van

Ned.- (9): 227—229, Abb. 1—2. bas, Pontianak en Pangaron. Natuurk. Tijdschr.

Bleeker, P., 1849a. Bijdrage tot de kennis der ichthyolo- Indië, 5: 427—462.

fauna van het eiland 1854a. waarnemingen, gedaan ver- gische Madura, met beschrijving —, Ichthyologische op in de residentie Natuurk. van eenige nieuwe soorten. Verh. batav. Genoot. schillende reizen Bantam.

Kunst. Wet., 22 (7): 1—16. Tijdschr. Ned.-Indië, 7: 309—326.

Downloaded from Brill.com10/11/2021 05:35:07AM via free access BIJDRAGEN TOT DE DIERKUNDE, 45 (1) - 1975 137

1854b. der fauna Suma- Indico habitationi- —, Overzigt ichthyologische van Archipelago observatarum, adjectis

met nieuwe soorten. bus ubi recentiores tra, beschrijving van eenige Na- citationibusque, descriptiones earum

tuurk. Tijdschr. Ned.-Indië, 7: 49—108. reperiuntur, nee non speciebus musei Bleekeriani Ben-

1854c. de kennis der i- —, Vijfde bijdrage tot ichthyologi- galensibus, Japonicis, Capensibus Tasmanicisque:

sche fauna van Amboina. Natuurk. Tijdschr. Ned.- xxxvi, 1—276 (Langii & Soc., Batavia (= Djakarta)).

tot kennis der visch- Indië, 6: 455—508. —, 1859b. Negende bijdrage de

tot de kennis der fauna van Banka. Natuurk. 18: —, 1854d. Vijfde bijdrage ichthyologi- Tijdschr. Ned.-Indië,

sche fauna van Celebes. Natuurk. Tijdschr. Ned.- 359—378.

Indië, 7: 225—260. 1859c. Twaalfde bijdrage tot de kennis der vischfauna —,

1855a. Achtste de kennis der van Borneo. Visschen van Sinkawang. Verh. natuurk. —, bijdrage tot ichthyologi-

sche fauna van Borneo. Zoetwatervisschen van Ver. Ned. Ind., 5 (7): 1—10.

Natuurk. 8: 151— 1859d. eene visschen van Bandjermasin. Tijdschr. Ned.-Indië, —, Verslag van verzameling

168. westelijk Borneo, aangeboden door J. H. A. Sonneman

Rebentisch. Natuurk. Ned.-Indië, 16: 433— 1855b. Achtste tot de kennis der Tijdschr. —, bijdrage ichthyologi-

Natuurk. 436. sche fauna van Celebes. Tijdschr. Ned.-

Indië, 9: 281—314. 1859e. Vischsoorten, van Banjoewangi —, toegezonden

1855c. de vischfauna van Sumatra. door den heer andere, verzameld —-, Nalezingen op Zollinger, eenige te

Visschen van Lahat en Sibogha. Natuurk. Tijdschr. Buitenzorg door den heer H. Von Rosenberg. Natuurk.

Ned.-Indië, 9: 257—280. Tijdschr. Ned.-Indië, 16: 47—48.

1855d. bijdrage tot de kennis der 1859f. Vischsoorten van Palembang, verzameld door —, Negende ichthyolo- —,

gische fauna van Borneo. Zoetwatervisschen van Pon- E. A. Lange en F. J. P. Storm van 's-Gravensande.

tianak Natuurk. Ned.- 263—266. en Bandjermasin. Tijdschr. Natuurk. Tijdschr. Ned.-Indië, 16:

415—430. Vischsoorten verzameld door Indië, 9: —, 1859g. van Palembang,

tot de kennis der 16: —, 1855e. Zesde bijdrage ichthyologi- R. V. Heyliger. Natuurk. Tijdschr. Ned.-Indië,

sche fauna van Amboina. Natuurk. Tijdschr. Ned.- 338—341.

tot vischfauna Indië, 8: 391—434. —, 1859h. Zevende bijdrage de kennis der

1856a. Beschrijvingen van nieuwe en bekende van Sumatra. Visschen van Verh. natuurk. —, weinig Palembang.

vischsoorten van Amboina, verzameld op eene reis Ver. Ned. Ind., 5 (6): 1—12.

door den Molukschen in het 1860a. Achtste tot de kennis der vischfauna Archipel gedaan gevolg —, bijdrage

van den Gouverneur-Generaal Duymaer van Twist, van Sumatra. (Visschen van Benkoelen, Priaman,

in September en Oktober 1855. Verh. natuurk. Ver. Tandjong, Palembang en Djambi). Verh. natuurk. Ver.

Ned.-Indië, 1 (5): 1—76. Ned.-Indië, 8 (2): 1—88.

1856b. Beschrijvingen van nieuwe of weinig bekende —, 1860b. Derde tot de kennis der vischfauna —, bijdrage

vischsoorten van Manado en Makassar, grootendeels 20: van Singapoera. Natuurk. Tijdschr. Ned.-Indië,

verzameld eene reis naar den Molukschen op Archipel 446—456.

in het den Gouverneur-Generaal gevolg van Duymaer 1860c. Dertiende tot de kennis der visch- —, bijdrage

van Twist. Verh. natuurk. Ver. Ned.-Indië, 1 (6): fauna van Borneo. Verh. natuurk. Ver. Ned.-Indië, 8

1—80. (4): 1—64.

1856c. Verslag van eenige verzamelingen van zee- en —, 1860d. Dertiende bijdrage tot de kennis der visch- —,

zoetwatervisschen van het eiland Banka. Natuurk. fauna van Celebes. (Visschen van Bonthain, Badjoa, Tijdschr. Ned.-Indië, 11: 415—420. Sindjai, Lagoesi en Pompenoea). Verh. natuurk. Ver. 1857a. Achtste de kennis der vischfauna —, bijdrage tot Ned.-Indië, 8 (7): 1—60.

Amboina. Verh. natuurk. Ver. Ned. 2 van Ind., (7): 1860e. tot de kennis der vischfauna —, Negende bijdrage 1—102. van Sumatra. (Visschen uit de Lematang-Enim en van

1857b. moluccensium Verh. Ver. 8 —, Conspectus specierum piscium Benkoelen). natuurk. Ned.-Indië, (3):

hucusque cognitarum. Verh. natuurk. Ver. Ned. Ind., 2 1—12.

1—23. 1860f. Nieuwe vischsoorten van verzameld (4): —, Singapore,

1857c. Tiende tot de kennis der door Fr. Graaf de Castelnau. Natuurk. Ned.- —, bijdrage ichthyologi- Tijdschr.

sche fauna van Borneo. Visschen van de rivieren Indië, 20: 236—239.

Barito, Kahajan en Verh. natuurk. Ver. Ned. Zoetwatervisschen van Sintang, verzameld door Kapoeas. —, 1860g. Ind., 2 (6): 1—21. den heer A. Thepass. Natuurk. Tijdschr. Ned.-Indië,

1858a. Zesde bijdrage tot de kennis der vischfauna 20: 199—200. —,

van Sumatra. Visschen van Padang, Troessan, Pria- 1861a. Iets over de vischfauna van het eiland Pinang. —, Akad. 12: man, Sibogha en Palembang. Verh. natuurk. Ver. Versl. Meded. K. Wet. Amst., 64—80.

Ned. Ind., 3 (9): 1—50. 1861b. omtrent vischsoorten, nieuw voor —, Mededeeling

de kennis der fauna Versl. Meded. K. 1858b. Elfde tot de kennis der van Singapoera. —, bijdrage ichthyologi- Akad. Wet. Amst., 12: 28—63. sche fauna van Borneo. Visschen van Sinkawang.

Verh. natuurk. Ver. Ned. 3 1—4. 1861c. nieuw de kennis der fauna Ind., (13): —, Vischsoorten, voor

1858c. Enumeratio specierum piscium van Singapoera, verzameld door Fr. Graaf de —, javanensium

hucusque cognitarum. Natuurk. Tijdschr. Ned.-Indië, Castelnau. Natuurk. Tijdschr. Ned.-Indië, 22: 101—

15: 359—456. 102.

1858d. Tweede de kennis der 1863. Over vischsoorten van het eiland —■, bijdrage tot vischfauna —, eenige Bangka,

van Singapore. Natuurk. Tijdschr. Ned.-Indië, 15: het Rijks Museum van Natuurlijke Historie te Leiden

door 241—254. aangeboden, den heer J. F. R. S. van den Bos-

1859a. Enumeratio specierum in sche, Resident. Ned. Dierk., 1: 72—73. —, piscium hucusque Tijdschr.

Downloaded from Brill.com10/11/2021 05:35:07AM via free access 138 W. J. DEKKERS - REVIEW OF TETRAODON

1864. Notice faune de Siam. of State for India in Council. Edited —, sur la ichthyologique Secretary by

Fishes. — Vol. Versl. Meded. K. Akad. Wet. Amst., 16: 352—358. W. T. BLANFORD. II: i—xiv, 1—509,

1865a. Atlas des Indes Orientales 1—177. (Taylor & London, etc.). —, ichthyologique figs. Francis,

sous les auspices du Néêrlandaises, publié gouverne- DELSMAN, H. C. & J. D. F. HARDENBERG, 1934. De Indi- ment colonial Néêrlandais. Tome V. Baudroies, Ostra- sche zeevisschen en zeevisscherij. Bibliotheek van de

cions, Gymnodontes, Balistes: 1—152, pis. 194—231 Nederlandsch-Indische Natuurhistorische Vereeniging,

(Frédéric Muller, Amsterdam). 6: i-viii, 1—388, figs. 1—273, 1 pi. (Visser & Co., 1865b. Énumération des espèces de poissons actuelle- —, Batavia).

ment de l'île d'Amboine. Ned. travail inédit de connues Tijdschr. DUMÉRIL, A. H. A., 1855. Note sur un Dierk., 270—297. 2: Bibron relatif aux poissons Plectognathes Gymnodon- la faune 1865c. Nouvelle notice sur ichthyologique tes et Revue 7: —, (Diodons Tétrodons). Mag. Zool., (2) Siam. Ned. 2: 33—37. de Tijdschr. Dierk., 274—282.

1865d. Sixième notice sur la faune ichthyologique de 1904. Die Fische der Halb- —, DUNCKER, G., Malayischen Ned. 2: 171—176. Siam. Tijdschr. Dierk., insel. Mitt. naturh. Mus. Hamb., 21 (2): 133—207,

1866. Synonyma Balistidorum, Ostracionidorum Gym- 1—2. —, fig. 1, Taf. ob- nodontidorumque Indo-archipelagicorum hucusque 1912. Die Mitt. naturh. —, Süsswasserfische Ceylons. servatorum revisa, habitationibus citationibus- adjectis Mus. Hamb., 29 (2): 241—272, 1 map, 1 pl.

ubi descriptiones figuraeque eorum recentiores que DUNCKER, G. & E. MOHR, 1929. Die Fische der Südsee-

Ned. 3: 20—40. reperiuntur. Tijdschr. Dierk., Expedition der Hamburgischen Wissenschaftlichen

1868a. Notice sur la faune ichthyologique de l'île de Stiftung 1908—1909. Mitt. zool. St. Inst. Hamb., 44: —, Guébé. Versl. Meded. K. Akad. Wet. Amst., (2) 2: 57—84, figs. 1—9.

Fresh-water 271—272. FEIGS, G., 1955. puffers spawn. Aquarium,

sur la faune de —, 1868b. Sixième notice ichthyologique Philad., 24 (22): 373—374, 2 figs.

l'île de Bintang. Versl. Meded. K. Akad. Wet. Amst., FOWLER, H. W., 1905. Some fishes from Borneo. Proc.

(2) 2: 289—294. Acad. nat. Sei. Philad., 57: 455—523, figs. 1—16.

1860. on some fishes recieved chiefly 1928. The fishes of Oceania. Mem. Bernice P. BLYTH, E., Report —, Bishop from the Sitang River and its tributary streams, Mus., 10: i-iii, 1—540, figs. 1—82, pis. 1—49.

Tenasserim Provinces. J. Asiat. Soc. Beng., 29: 138— 1934a. results of the Third De Schauensee —, Zoological

174. Siamese Expedition. Part 1. Fishes. Proc. Acad. nat.

BOESEMAN, M., 1957. On a collection of East Asian Sei. Philad., 86: 67—163, figs. 1—127, pi. 12.

fishes. Zoöl. Meded. 35 69—79. Leiden, (7): 1934b. results of the Third De Schauensee —, Zoological 1972. Preface. In: W. H. LAMME ed., Pieter Bleeker. —, Siamese Expedition. Part V. Additional fishes. Proc.

tot de van de Indonesische Bijdragen ichthyologie Acad. nat. Sei. Philad., 86: 335 —352, figs. 1—13. verzameld bewerkt uit het Natuurkundig archipel, en 1935. results of the Third De Schauensee —, Zoological

Nederlandsch-Indië I-XXII, 1851— Tijdschrift voor Siamese Expedition. Part VI. Fishes obtained in 1934. 1861. I: v-viii & Co, Vaals/Amsterdam). (Asher Proc. Acad. nat. Sei. Philad., 87: 89—163, figs. 1—

BREDER, C. H. & E. CLARK, 1947. A contribution to the 132.

and of 1937. results visceral anatomy, development relationships —, Zoological of the Third De Schauensee

the Plectognathi. Bull. Am. Mus. nat. Hist., 88 (5): Siamese Expedition. Part VIII. Fishes obtained in

287—320, figs. 1—8, pis. 11—14. 1936. Proc. Acad. nat. Sei. Philad., 89: 125—264,

CANTOR, T., 1849. Catalogue of Malayan fishes. J. Asiat. figs. 1—300.

Soc. 18: 983 1—14. 1938. A list of the fishes known from Fish. Beng., i-xii, —1443, pis. —, Malaya.

CHABANAUD, P., 1923. Description d'un Tetrodon Bull. Singapore, 1: 1—268, i-lvi.

nouveau du Cambodge. Bull. Mus. natn. Hist. nat. FRASER-BRUNNER, A., 1940. Notes on the Plectognath

Paris, 29 (2): 137—140. Fishes. IV. Sexual dimorphism in the family Ostracio-

CHEVEY, P., 1932. Inventaire de la faune ichthyologique dontidae. Ann. Mag. nat. Hist., (11) 6: 390—392,

de l'Indochine. Deuxième liste. Note Inst. océanogr. 1 fig.

19: 1—31. 1943. Notes the fishes. VIII. The Indoch., —, on Plectognath

1936. Le Grand Lac du les causes classification of the suborder Tetraodontoidea, with a —, Cambodge: pro-

fondes de Note Inst. océano- of the Ann. sa richesse ichthyologique. synopsis genera. Mag. nat. Hist., (11)

gr. Indoch., 29: 39—49, figs. 1—3. 10: 1—18, figs. 1—4.

CHEVEY, P. & F. LE POULAIN, 1940. La pêche dans les GEISER, W., 1958. Aufzucht von Tetraodon leiurus brevi-

eaux douces du Cambodge. Trav. Inst. océanogr. rostris. Aquar.- u. Terrar.-Z., 11 (4): 100—101, 1 fig.

Tetraodontoidea. U.S. Indoch., S: 1—78, pis. 1—68, maps 1—7, figs. 1—119. GILL, T., 1892. Notes on the Proc.

CHLUPATY, P., 1955. Haltung und Pflege von Kugel- Natn. Mus., 14: 705—720, pl. 34.

fischen. Aquar.- u. Terrar.-Z., 8 (8): 197—200. GROBE, J., 1956. Tetraodon palembangensis. Aquar.- u.

DAY, F., 1865. On the fishes of Cochin, on the Malabar Terrar.-Z., 9 (4): 111.

Coast of India. Part. II. Anacanthini. Proc. zool. Soc. the material for GUDGER, E. W., 1924. The sources of

Lond., 1865: 286—318, 8 figs. Hamilton-Buchanan's Fishes of the Ganges, the fate

1878—88. The fishes of natural of his and and the use —, India; being a history collections, drawings notes,

fishes known to and fresh 19 of the inhabit the seas made of his data. J. Proc. Asiat. Soc. Beng., (4):

waters of India, Burma, and Ceylon: i-xx, 1—778, pis. 121—136.

1—198 (privately printed, London). GÜNTHER, A., 1870. Catalogue of the fishes in the British

1889. The fauna of British Museum. Vol. 8. of the con- —, India, including Ceylon Catalogue Physostomi, Published and Burma. under the authority of the taining the families Gymnotidae, Symbranchidae,

Downloaded from Brill.com10/11/2021 05:35:07AM via free access BIJDRAGEN TOT DE DIERKUNDE, 45 (1) - 1975 139

Muraenidae, Pegasidae, and of the Lophobranchii, drainage of North Borneo. Fieldiana Zool., 37: 47—

Plectognathi, Dipnoi, Ganoidei, Chondropterygii, Cy- 90, figs. 10—13.

clostomata, Leptocardii, in the British Museum: i-xxv, INGER, R. F. & P. K. CHIN, 1962. The fresh-water fishes

1—549 (Trustees Brit. Mus., London). of North Borneo. Fieldiana Zool., 45: 3—268, figs.

1—120. HAMILTON, F., 1822. An account of the fishes found in the

W. 1935. Exotic a work of river Ganges and its branches: i-vii, 1—405, pis. 1— INNÉS, T., aquarium fishes, general reference, edited by G. S. MYERS (19th ed.): 39 (A. Constable & Co, Edinburgh/Hurst, Robinson 1—541, ca. 450 1 pl. Publ. Philadel- & Co, London). figs., (Innés Co., phia). HARDENBERG, J. D. F., 1931. The fishfauna of the Rökan JOHNSEN, P., 1964. Notes on fishes the river Kwae mouth. Treubia, 13 (1): 81—168, fig. 1, 1 map. along Noi in Western Thailand. Pt. 2. Nat. Hist. Bull. Siam 1934. Additional "The fishfauna of —, notes to my paper Soc., 20: 257—263. the Rökan mouth". Treubia, 14 (3): 299 —312, figs. JORDAN, D. S. & R. E. 1909. Check-list of 1—4. RICHARDSON,

the of fishes known from the 1935. On a collection of fishes from Samarinda. Na- species Philippine —, Archipelago. Pubis Philipp. Bur. Sei., 1: 1—78. tuurk. Tijdschr. Ned.-Indië, 95 (3): 231—239.

1936. On collection of fishes from the and J., 1882. Prodromus piscium Asiae Orientalis a —, a estuary KÂROLI,

the lower and middle course of the river Kapuas (W. domine Joanne Xantus annis 1868-70 collectorum.

Borneo). Treubia, 15 (3): 225—254. Természetr. Filz., 5: 147—187.

1937. Hydrological and ichthyological observations in KLAUSEWITZ, W., 1957a. Neue Süsswasserfische aus —,

the mouth of the Kumai-River (S. W. Borneo). Treu- Thailand. Senckenberg. biol., 38 (3/4): 193—204, Abb.

bia, 16 (1): 1—14, fig. 1. 1—5, Taf. 17—18.

1957b. Tetraodon somphongsi n. ein weiterer HERRE, A. W. C. T., 1924. Poisonous and worthless —, sp.,

neuer Kugelfisch aus Thailand (Pisces, Tetraodonti- fishes; an account of the Philippine plectognaths. 38 Abb. Philipp. J. Sei., 25 (4): 415—510, pis. 1—2. dae). Senckenberg. biol., (3/4): 205—208, 1—2. 1933. A check list of fishes from Sandakan, British —, 1957c. Eine Siam: Tetraodon North Borneo. J. pan-Pacif. Res. Instn, 8 (4): 2—5. —, neueKugelfisch-Art aus

somphongsi. Aquar.- u. Terrar.-Z., 10: 115—116, 1940. On a collection of littoral and freshwater fishes —, 2 from the Andaman Islands: supplement. Ree. Indian figs.

KNER, 1867. Reise der österreichischen No- Mus., 42: 1—8. R., Fregatte die Erde in den Jahren 1859 1941. A list the known from Andaman vara um 1857, 1858, —, of fishes the

unter den Befehlen des Commodore B. von Wüllers- Islands. Mem. Indian Mus., 13 (3): 331—403. torf-Urbair. Theil. Erster Band. 1953. Check list of fishes. Res. U.S. Zoologischer Fische, —, Philippine Rep. dritte Taf. XII-XVI Hof- Fish Wildl. Ser., 20: 1—977. Abtheilung:273—433, (K.-K. und 1958. Marine fishes in rivers and lakes. Staatsdruckerei, Wien). —, Philippine

Philipp. J. Sei., 87 (1): 65—88. KOUMANS, F. P., 1947. Zoological notes from Port Dick-

A. W. C. T. & G. S. 1937. A contribution II. A small collection of fishes from HERRE, MYERS, son, Malaya.

to the ichthyology of the Malay Peninsula. Bull. Zoöl. Meded. Leiden, 27 (3—4): 309—311.

Raffles Mus., 13: 5—75, pis. 1—7. LADIGES, W., 1954. Tropische Fische — Tropical fish:

HERVEY, G. F. & J. HEMS, 1963. Freshwater tropical 1—207, 284 figs., 43 pis. (G. Wenzel & Sohn, Braun-

aquarium fishes: i-xiv, 1—432, figs. 1—89, pis. 1—8, schweig).

A-X (Spring books, London). LE DANOIS, Y., 1959. Étude ostéologique, myologique

et systématique des poissons du sous-ordre des Orbi- HORA, S. L., 1923a. Fauna of the Chilka Lake. Fish. Part culates. Annls Inst. Monaco, (N.S.) 36: V. Mem. Indian Mus., 5: 739 —769, figs. 22—35. océanogr., 1—221. 1923b. On collection of fish from Siam. J. Hist. 1—274, figs. —, a nat. des de orbiculates du —, 1961. Catalogue poissons Soc. Siam, 6 (2): 143—184, pis. 10—12. types Muséum National d'Histoire Naturelle II. Familles 1924. results of a in the Far East. —, Zoological tour des Tetraodontidae, Lagocephalidae, Colomesidae, Fish of the Talé Sap, Peninsular Siam. (Part II). Diodontidae et Triodontidae. Bull. Mus. natn. Hist, Mem. Asiat. Soc. Beng., 6: 479—501, figs. 4—10. nat. (2) 33 (5): 462—478. 1929. An aid to the of Hamilton Buchanan's Paris, —, study 1962. Transitions et "Gangetic Fishes". Mem. Indian Mus., 9 (4): 169— —, morphologiques ostéologiques entre les espèces du Dichotomycterus (Tetrao- 192, 1 fig., pis. 13—23. genre Bull. Soc. zool. 86: 1933. Buchanan's entitled dontidae). Fr., 694—700, figs. —, ichthyological manuscript 1—11. "Piscium Bengalae Inferioris Delineationes". J. Proc.

LÜLING, K. H., 1958. Zufallsbeobachtungenan Tetraodon Asiat. Soc. Beng., (N.S.) 27 (1931) (1): 123—135. somphongsi und Gyrinocheilus aymonieri (Tirant). HORA, S. L. & K. K. NAIR, 1941. Notes on fishes in the 11 (1): 11—15, Abb. 1—4. Indian Museum XLI. New records of freshwater fish Aquar.- u. Terrar.-Z., MCCULLOCH, A. R., 1922. Check list of the fish and from Travancore. Ree. Indian Mus., 43: 387—393,

fish-like animals of New South Wales. Part. iii. Aust. figs. 1—4. Zool., 2 (3): 86—130, pis. xxv-xliii. HUBRECHT, A. A. W., 1879. Catalogue des collections

formées laissées M.- P. MARION DE 1822. Sur nouvelles et par Bleeker...i-iv, PROCÉ, plusieurs espèces

1—71 Breuk & de de M. Marion (De Smits, Leiden). poissons et crustacés observées par

INGER, R. F., 1953. A new fish from North Borneo. de Procé, D. M. P., membre correspondant de la

Genus Tetraodon. Fieldiana Zool., 34 (11): 149—152, Société Philomatique. Bull. Soc. philom. Paris, 1822:

fig. 27. 129—134.

notes 1956. —, 1955. Ecological on the fish fauna of a coastal MEINKEN, H., Mitteilungen der Fischbestimmungs-

Downloaded from Brill.com10/11/2021 05:35:07AM via free access 140 W. J. DEKKERS - REVIEW OF TETRAODON

stelle des VDA. XXI. Tetraodon palembangensis ROXAS, H. A. & C. MARTIN, 1937. A check list of Philip-

Bleeker, 1852, der Palembang-Kugelfisch. Aquar.- u. pine fishes. Tech. Bull. Commonw. Philipp. Dep. Agr.

Terrar.-Z., 9 (3): 61 —63, 1 fig. Commerce Manila, 6: 1—314.

Verzeichniss der in dem Museum der MENON, A. G. K., 1961a. On a collection of fish from RÜPPELL, E., 1852.

Lake Chilka, Orissa. Ree. Indian Mus., 59: 41—69, Senckenbergischen Naturforschenden Gesellschaft auf-

fig. 1. gestellten Sammlungen. Vierte Abtheilung: Fische und

1961b. On collection of fish from the Coromandel deren Skelette: i-iv, 1—40 (J. D. Sauerländer, Frankfurt —, a

Coast of India including Pondicherry and Karaikkal am Main).

H. 1883. Sur collection de areas. Ree. Indian Mus., 59 (4): 369—404. SAUVAGE, E., une poissons

Harmand. recueillie dans le Mé-Nam (Siam) par M. MENON, A. G. K. & G. M. YAZDANI, 1963. Catalogue of Bull. Soc. philom. Paris, (7) 7: 150—155. type-specimens in the Zoological Survey of India.

Schwimmsweise Part 2. Fishes. Ree. zool. Surv. India, 61: 91—190. SCHNEIDER, H., 1964. Untersuchungen zur

MERCKENS, P. J., 1958. Zoetwater-kogelvissen 1. Aqua- der Kugelfische. 1. Die Flossenmuskulatur des Fluss-

rium, Den Haag, 29 (3): 50—54, 8 figs. kugelfisches (Tetraodon fluviatilis)in Vergleich zu der

3. Den Schleie Z. Ökol. 54: —, 1959. Zoetwater-kogelvissen Aquarium, Haag, (Tinea tinea). Morph. Tiere,

29 (8): 170—172, 2 figs. 414—435, 15 figs.

of MUNRO, I. S. R., 1955. The marine and fresh water fishes SEALE, A., 1910. Fishes of Borneo, with descriptions

of Ceylon: i-xvi, 1—351, figs. 1—19, pis. 1—56 (Dept. four new species. Philipp. J. Sei., 5 (4): 263—289,

external affairs, Canberra). pis. 1 —4.

1958. The fishes of the New Guinea A check G. E. & E. O. 1938. The fishes of —, Region. SHAW, SHEBBEARE,

list of the fishes in New Guinea incorporating new northern Bengal. J. R. Asiat. Soc. Beng., 3: 1—137,

records of species collected by the Fisheries Survey figs. 1—130, pis. 1—6.

Vessel Fairwind the 1948 1950. H. 1945. The fresh-water fishes of during years to Papua SMITH, M., Siam, or

New Guin. agric. J., 10 (4): 97—369, fig. 1, maps Thailand. Bull. U.S. natn. Mus., 188: i-xi, 1—622, figs.

1—3. 1—107, pis. 1—9.

1967. The fishes of New Guinea: S. MONKOLPRASIT & P. 1971. —, i-xxxvii, 1—651, SONTIRAT, S., WONGRAT,

Agriculture, Klausewitz, 1957, a figs. 1—23, col. pis. 1—6, pis. 1—78 (Dep. Notes on Tetraodon Somphongsi,

Stock and Fisheries, Port Moresby). puffer fish from Thailand. Notes Fac. Fish. Kasetsart

OGILBY, J. D., 1886. Catalogue of the fishes of New Univ., 6: 5—6.

their 1—67 Fishes of Uttar Pradesh: South Wales, with principle synonyms: SRIVASTAVA, G. J., 1968. eastern

(Th. Richards, Sydney). i-xxii, 1—163, figs. 1—86 (Vishwavidyalaya Prakashan,

OSTERMÖLLER, W., 1957. Einige Beobachtungen über Varanasi).

Tetraodon somphongsi. Aquar.- u. Terrar.-Z., 10 (11): STEINDACHNER, F., 1870. Ichthyologische Notizen (X).

288—290, 1 fig. Sber. Akad. Wiss. Wien, 60 (1): 623—642, Tab. 1—5.

PELLEGRIN, J. & P. W. FANG, 1940. Poissons du Laos STERBA, G., 1959. Süsswasserfische aus aller Welt. Teil 2:

recueillis MM. Ed. Blanc. Abb. 360—620. Taf. 145—280 par Delacour, Greenway, 357—688, (Urania-

Description d'un genre, de cinq espèces et d'une Verlag, Leipzig/Jena/Berlin). variété. Bull. Soc. zool. Fr., 65: 111 —123, figs. 1—6. SUVATTI, C., 1936. Index to fishes of Siam: A-F, 1—226,

J. & P. 1940. Poissons nouveauxou 1 1 of PELLEGRIN, CHEVEY, pl., map (Bureau Fisheries, Bangkok).

rares de Cochinchine. Descriptions de deux espèces et SWAINSON, W., 1839. The natural history of fishes,

de deux variétés. Bull. Soc. zool. Fr., 65: 153—158, amphibians, & reptiles, or monocardian animals. II:

figs. 1—2. i-vi, 1—452, figs. 1—135 (Longman, Brown, Green Sc

POPTA, C. M. L., 1905. Suite des descriptions préliminai- Longmans, London).

recueillies res des nouvelles espèces de poissons au TIRANT, G., 1885. Notes sur les poissons de la Basse-

Bornéo Central le A. par M. Dr. W. Nieuwenhuis en Cochinchine et du Cambodge. Excurs. Reconn., 9:

1898 et en 1900. Notes Leyden Mus., 25 (4): 171 —186. 413—438; 10: 91—198. Id., réimprimé en 1929: Notes

1906. Résultats des scientifi- Stn marit. 6: 43—163. —, ichthyologiques voyages Cauda,

de Monsieur le Professeur Dr. A. W. Nieuwen- ques TWEEDIE, M. W. F., 1940. Additions to the collection of huis dans le de Bornéo et Notes centre (1898 1900). fishes in the Raffles Museum. Bull. Raffles Mus., 16:

27: pis. I-X. Leyden Mus., 1—304, 68—82.

B. & D. D. 1929. The fish of the PRASHAD, MUKERJI, 1952. Notes fresh-water fishes. Bull. —, on Malayan Indawgyi Lake and the streams of the Myitkyina Raffles Mus., 24: 63—95, figs. 1—4. District (Upper Burma). Ree. Indian Mus., 31 (3): TYLER, J. C., 1963. A critique of Y. Le Danois' work on VII-X. 161—224,pis. the classification of the fishes of the order Plectogna-

1934. Tetrodon cutcutia und Tetrodon RANDOW, H., thi. Copeia, 1963 (1): 203—206.

fluviatilis Hamilton-Buchanan. Wschr. u. Aquar.- 1964. A diagnosis of the two of South —, species Terrarienk., 31: Abb. 1. 561—563, American puffer fishes (Tetraodontidae, Plectognathi) C. 1902. On the classification of the fishes REGAN, T., of the Colomesus. genus Proc. Acad. nat. Sei. Philad., of the suborder with notes and Plectognathi; descrip- 116 (3): 119—148, figs. 1—16.

tions of new species from in the British specimens VAILLANT, L., 1893a. Contribution à l'étude de la faune

Museum collection. Proc. zool. Soc. Lond., 1902 (II): ichthyologique de Bornéo. Nouv. Archs Mus. Hist, 284—303, text-figs. XXIV-XXV. 56—59, pis. nat. Paris, (3) 5: 23—112, pis. 1 —2.

R. Where Pieter Bleeker's East- 1893b. Sur collection de recueillie ROFEN, R., unpubl. are —, une poissons par Indian fishes? (Undated and unpublished manuscript M. Chaper à Bornéo. Bull. Soc. zool. Fr., 18: 55 —62.

in the Institute of Taxonomie Zoology, Amsterdam). 1902. Résultats de scientifi- —, zoologiques l'expédition

Downloaded from Brill.com10/11/2021 05:35:07AM via free access BIJDRAGEN TOT DE DIERKUNDE, 45 (1) - 1975 141

Néerlandaise Bornéo Central. Poissons. Notes nebst über den der que au Archipels, Bemerkungen Ursprung

Leyden Mus., 24 (1—3): 1—166, figs. 1—47, pis. Fauna von Celebes. In: M. WEBER ed., Zoologische 1—2. Ergebnisse einer Reise in Niederländisch Ost-Indien.

VINCIGUERRA, D., 1883. Appunti ittiologici sulle collezioni III: 405—476 (Brill, Leiden).

del Museo Civico di Genova. V. Enumerazione di 1895. Fische von dem —, Ambon, Java, Thursday Island,

alcuni pesci raccolti a Minhla suil' Irrawaddi dal Burnett-Fluss und von der Südküste von Neu-Guinea.

capitano cav. G. B. Comotto. Annali Mus. civ. Stor. Denkschr. med.-naturw. Ges. Jena, 8: 259—276, 1

nat. Genova, 18: 651—660. Textfig.

1890. Viaggio di Leonardo Fea in Birmania e 1911. Die Fisohe der Aru- und Kei-Inseln. Ein —, regioni —, Beitrag

nat. vicine. XXIV. Pesci. Annali Mus. civ. Stor. zur Zoogeographie dieser Inseln. Abh. Senckenb.

Genova, (2a) 9: 129—367, tav. VII-XI. naturforsch. Ges., 34: 1—49, figs. 1—11, pis. 1 —2.

1926. dei raccolti a Borneo dai Sigg. 1913. Süsswasserfische aus Niederländisch Süd- und —, Catalogo pesci —,

Marchese G. Doria e Dott. O. Beccari negli anni Nord-Neu-Guinea. Nova Guinea (Zool.), 9 (4): 513—-

1865 —67. Annali Mus. civ. Stor. nat. Giacomo Doria, 613, figs. 1—36, pis. 12—14.

(3a) 10: 532—628, tav. I. WEBER, M. & L. F. DE BEAUFORT, 1912. Fische. In: A.

VLUGT, P. J. VAN DER, 1968. Carinotetraodonsomphongsi. MAASS ed., Durch Zentral-Sumatra. II: 1—20 (C.

Aquarium, Den Haag, 38 (10): 218—224, 7 figs. Marschner, Berlin).

VOLZ, W., 1903. Fische von Sumatra (Reise von Dr. WHITEHEAD, P. J. P., M. BOESEMAN & A. C. WHEELER,

Walter Volz). Zool. Jb. (Syst.), 19 (4): 1 1966. The types of Bleeker's Indo-Pacific elopoid and

Abb., Taf. 25—26. clupeoid fishes. Zool. Verh. Leiden, 84: 1—159, pis.

1904. Fische von Sumatra von Herrn G. —, gesammelt 1—19.

Schneider. Revue suisse zool., 12 (2): 451—493. WHITLEY, G. P., 1957. List of type-specimens of recent

1906. Catalogue of the fishes of Sumatra. Natuurk. fishes in the Australian Sydney: i-iii, 1—40 —, Museum, Tijdschr. Ned.-Indië, 66 (1 —2): 35—250. (mimeograph, Australian Museum, Sydney).

WAITE, E. R., 1904. A synopsis of the fishes of New WILLEM, V., 1947. Contributions à l'étude des organes

South Wales. Mem. N. S. W. Nat. Club, 2: 3—59, respiratoires chez les Téléostéens Plectognathes (1).

i-iv. 5e Partie: Tétrodontes et Diodon. Bull. Mus. r. Hist,

WEBER, M., 1894. Die Siisswasser-Fische des Indischen nat. Belg., 23 (17): 1—17, figs. 1—10.

INDEX TO THE TAXONOMIC PART

Main references are printed in bold type bergii, Tetraodon 103, 109, 111 erythrotaenia, Tetraodon

Bergii, Tetrodon 108 92, 93, 114, 115-117, 120; figs. 24-25 biocellatus, Tetraodon 119-120 erythrotaenia, Tetraödon 115 borneensis, Tetraodon 94, 97, 99 erythrotaenia, Tetrodon 115 borneensis, Tetrodon 97 fangi, Tetraodon 92, 93,

cambodgiensis, Tetraodon 95, 102, 103, 104, 107-108, 112-114; figs. 13, 21-23

103, 104, 108, 109, 110; figs. 13, 18-20 fangi, Tetrodon 112

cambodgiensis, Tetrodon 108 fluviatilis-group 92, 93, 102, 104, 112, 117-120

caria, Tetraodon 93-94 fluviatilis, Chelanodon 127

caria, Tetrodon 95 fluviatilis, Chelonodon 127

caria, Monotreta 94, 97 fluviatilis, Crayracion 120, 124, 127 chlupatyi, Carinotetraodon 93, 94, 97, 99 fluviatilis, Dichotomycter 121, 124

chlupatyi, Tetraodon 94 fluviatilis, Dichotomyctère 123

cutcutia-group 92, 93-95, 100, 102, 114 fluviatilis, Dichotomycterus 124, 127, 129

cutcutia, Arothron? 95 fluviatilis, Tetraodon 108, 117, 118,

cutcutia, Leiodon 95 119, 120-121, 123, 124, 127, 130, 134, 135; figs. 32-37

cutcutia, Leiosomus 95 fluviatilis, Tetraödon 127

cutcutia, Monotrète 95 fluviatilis, Tetrodon 120-121, 124, 127, 135

cutcutia, Monotretus 95 fluviatilis fluviatilis, Tetraodon

cutcutia, Tetraodon 92, 93, 117, 119, 127-130; figs. 32-34

92, 93, 94, 95-97, 100, 101, 112; figs. 3-5 fluviatilis var. A, Tetrodon 119, 127

Tetrodon cutcutia, Tetraodon 95 fluviatilis var. B, 127

95 var. cutcutia, Tetrodon fluviatilis ocellata, Crayracion .... 119, 132, 134

cutcutia, cutcutia, Monotreta 108 fluviatilis sabahensis, Tetraodon cutcutia palembangensis, Monotreta 102, 105 92, 93,117, 120, 130-132; figs. 35-37

dorsovittatus, Arothron 119, 127 gernaerti, Epipedorhynchus 115

dumerili, Chelonodon 94, 97 Gernaerti, Epipedorhynchus 114-115

Dumerili, Chelonodon 97 gularis, Monotreta 94, 95 erijthrotaenia, Tetraödon 115 gularis, Tetraodon 93-94 erythrotaenia-group 92, 93, 102, 114-115, 120 gularis, Tetrodon 95

Arothron 115 hilgendorfii, Tetraodon 13 erythrotaenia, .... 103, 104, 109, 111; fig. erythrotaenia, Crayracion 114, 115 Hilgendorfii, Tetrodon 108 erythrotaenia, Monotretus 115 honckenii, Geneion 115

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javanicus, Dichotomy ctère 124 palembangensis, Tetrodon 105, 108, 132

kappa, Tetraodon 108 palembangensis Variété, Tetrodon 105

kretamensis, Tetraodon 92, 93, patoca, Chelonodon 92

92 114, 117, 118, 120, 121-123, 127, 130, figs. 26-28 patoca, Monotreta

leiurus-group 92, 93, 102-104, 114; fig. 13 patoca, Tetraodon 92, 103, 112, 134

leiurus, Arothron 107, 108 pelambangensis, Tetraodon 105

leiurus, Crayracion 108 pinguis, Tetraodon 102, 105, 106

leiurus, Tetraödon 108 pinguis, Tetrodon 108

leiurus, Tetraodon 92, 93, 94, 97, 102, 103, pleurogramma, Sphoeroides 114

123 104, 105, 107-108, 108-112, 121, 124; figs. 13, 16-20 potamophilus, Arothron

leiurus, Tetrodon 108, 109, 112 potamophilus, Tetraodon 118, 124

leiurus brevirostris, Tetraodon . . 95, 103, 109, 112, 113 potamophilus, Tetraodon 123

lineatus, Tetraodon 92 rangoonensis, Dichotomycterus 124, 127, 129

104 127 lineatus, Tetraödon rangoonensis dorsovittatus, Dichotomycterus .. 119,

liurus, Tetraodon 109 rangoonensis, Tetraodon 119

liurus, Tetrodon 108 reticulatus, Tetraödon 104

lorteti, Tetraodon 92, 93, 94, 97-101; figs. 6-9 sans tache, Tetrodon 108

lorleti, Tetrodon 97 semimaculatus, Arothron 119,120

Lorteti, Tetrodon 97 simulans, Arothron 123-124 marmoratus, Leiodon 94 simulans, Tetraodon 118, 124, 125

95 123 marmoratus, Leiosomus 94, simulans, Tetrodon modestus, Chonerinus 92 somphongsi, Carinotetraodon 97 naritus, Chonerinus 92 somphongsi, Tetraodon 94, 97 nigroviridis, Tetraodon 92, 93, 115, steindachneri, Tetraodon 92,

117, 118, 119, 120, 121, 123-127, 134; figs. 29-31 93, 112, 114, 117, 118, 120, 132-134; figs. 38-40 nigroviridis, Tetrodon 123 tiranti, Monotreta 94, 97, 105 oblongus, Sphoeroides 134 travancorius, Tetraodon ocellaris, Tetraodon 103, 112, 113, 134 92, 93, 94, 100, 101-102, 114; figs. 10-12 ocellatus, Sphoeroides 92 viridipunctatus, Tetraodon 119, 127 ocellatus, Tetraodon 119-120, 134 waandersii-group 92, 93, 134, 135 palembagensis, Tetraodon 105 Waandersii, Arothron 134-135 palembangensis, Arothron 102, 105 Waandersii, Leiodon 135 palembangensis, Arothron? 105 Waandersii, Rohita 134 palembangensis, Crayracion 105 waandersii, Tetraodon 92, 93, 134-135; figs. 41-42 palembangensis, Tetraodon 92, 93, 94, 97, 102, Waandersii, Tetraödon 134

104-107, 108, 109, 111, 112, 119, 132, 134; figs. 14-15 waandersii, Tetrodon 135 palembangensis, Tetraödon 104-105 werneri, Tetraodon 94, 97

Received: 9 October 1974

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