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Fossil Bovidae from the Malay Archipelago and the Punjab

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Fossil Bovidae from the Malay Archipelago and the Punjab FOSSIL BOVIDAE FROM THE MALAY ARCHIPELAGO AND THE PUNJAB by Dr. D. A. HOOIJER (Rijksmuseum van Natuurlijke Historie, Leiden) with pls. I-IX CONTENTS Introduction 1 Order Artiodactyla Owen 8 Family Bovidae Gray 8 Subfamily Bovinae Gill 8 Duboisia santeng (Dubois) 8 Epileptobos groeneveldtii (Dubois) 19 Hemibos triquetricornis Rütimeyer 60 Hemibos acuticornis (Falconer et Cautley) 61 Bubalus palaeokerabau Dubois 62 Bubalus bubalis (L.) subsp 77 Bibos palaesondaicus Dubois 78 Bibos javanicus (d'Alton) subsp 98 Subfamily Caprinae Gill 99 Capricornis sumatraensis (Bechstein) subsp 99 Literature cited 106 Explanation of the plates 11o INTRODUCTION The Bovidae make up a very large portion of the Dubois collection of fossil vertebrates from Java, second only to the Proboscidea in bulk. Before Dubois began his explorations in Java in 1890 we knew very little about the fossil bovids of that island. Martin (1887, p. 61, pl. VII fig. 2) described a horn core as Bison sivalensis Falconer (?); Bison sivalensis Martin has al• ready been placed in the synonymy of Bibos palaesondaicus Dubois by Von Koenigswald (1933, p. 93), which is evidently correct. Pilgrim (in Bron- gersma, 1936, p. 246) considered the horn core in question to belong to a Bibos species closely related to the banteng. Two further horn cores from Java described by Martin (1887, p. 63, pl. VI fig. 4; 1888, p. 114, pl. XII fig. 4) are not sufficiently well preserved to allow of a specific determination, although they probably belong to Bibos palaesondaicus Dubois as well. In a preliminary faunal list Dubois (1891) mentions four bovid species as occurring in the Pleistocene of Java, viz., two living species (the banteng and the water buffalo) and two extinct forms, Anoa spec. nov., and Bos 1 2 D. A. HOOIJER (or Bison?) spec. nov. The alleged Anoa was named Anoa santeng Dubois (in Jentink, 1891), but later renamed Tetraceros kroesenii Dubois (1907). The Bos or Bison was found to be a Leptobos (Anonymus, 1892). In his 1908 survey of the fossil vertebrate fauna of Java Dubois enumerates six species of Bovidae, all but the first of them new to science, as follows : Tetracerus kroesenii, Leptobos groeneveldtii, Leptobos dependicornus, Bibos palaesondaicus, Bibos protocavifrons, and Bubalus palaeokerabau. Stremme (1911) raised "Tetracerus" kroesenii to distinct generic rank, Duboisia, and added very materially to our knowledge of some of the forms named by his descriptions and figures of skull and dentition of Duboisia santeng, Bibos palaesondaicus, and Bubalus palaeokerabau. The collection of the Selenka expedition to Trinil studied by Stremme does not contain any material of Leptobos, and none of Bibos protocavifrons either. Soergel (1913) records a skull from Kedoeng Loemboe (Elbert collection) answering the description of Bibos protocavifrons) however, he does not provide any de• scription or figure. In his work on the Limbangan fauna Stehlin (1925) gives a description of a horn core most probably belonging to Dubois's Leptobos groeneveldtii, and further mentions fragmentary remains of large bovids as well as of an antelope. Descriptions and figures of fossil skulls of banteng and water buffalo from various localities in Java have been given by Van der Maarel (1932). Van der Maarel considers his fossil material to be conspecific with the living forms but is careful in stating that in the absence of any figures of Dubois's speci• mens he is unable to determine whether Dubois's species of Bibos and Buba• lus are valid and distinct from the recent species. In the opinion of Von Koenigswald (1933, 1934) Bibos palaesondaicus and Bubalus palaeokerabau are only subspecifically distinct from the extant species. Three of Dubois's new species, viz., Leptobos groeneveldtii, L. depen• dicornus, and Bibos protocavifrons, are declared "nomina nuda". Five names are added to the list of Pleistocene Java Bovidae, viz., "Antilope" gracilicornis Von Koenigswald (1933, p. 82), Antilope modjokertensis, Antilope saätensis, and Leptobos cosijni Von Koenigswald (1934, pp. 192 and 193), and, finally, Probibos djulangensis Von Koenigswald (1950, p. 92). The first and the last of these should be kept on the paleontological suspense account : the bat• tered skull on which "Antilope" gracilicornis is based may be cervid rather FOSSIL BOVIDAE 3 than bovid in characters (the poor figure does not permit of a decision one way or the other), while the only diagnosis of Probibos djulangensis available to date is : "The jaw is about two third the size of a Bibos jaw; the dentition suggests a relation with that group". Both of Antilope modjokertensis and of Antilope saätensis generic alloca• tion is admittedly provisional; in the absence of horn cores the genus cannot be determined (Von Koenigswald, 1934, p. 192). Hence, there is no justifica• tion for placing Antilope, a genus of Pallas, 1766, today confined to a single Indian species, Antilope cervicapra (Linnaeus), on the list of fossil genera of Java as done by Von Koenigswald (1940, p. 71). It would have been less confusing if Von Koenigswald had introduced his supposedly new species as "Genus indet. (cf. Antilope) modjokertensis nov. spec", and "Genus indet. (cf. Antilope) saätensis nov. spec"; it seems quite legitimate to describe new species the genus of which is indeterminate (see, e.g., Pilgrim, 1939, pp. 36, 40, 42, 57, 86, 107, and 114). It is Von Koenigswald's opinion that Duboisia santeng occurs only in the Trinil beds, whereas his Modjokerto antelope would be confined to the under• lying Djetis beds. Likewise, "Bos (Bibos) banteng palaeosondaicus Dubois" and "Bos (Bubalus) bubalis palaeokerabau Dubois" are found in the Trinil beds, and "Leptobos cosijni" in the Djetis beds. The stratigraphical position of "Antilope saätensis" is uncertain (Von Koenigswald, 1934, p. 192/193). However, a single referred specimen was found at Sangiran I (Djetis beds) (Antilope cf. saätensis: Von Koenigswald, 1934, p. 193, pl. HI fig. 8; re- figured as Antilope saätensis in Von Koenigswald, 1940, pl. II fig. 5). Thus, Von Koenigswald would have us believe that "Antilope" and Leptobos are from the Djetis, and Duboisia, Bibos, and Bubalus from the Trinil beds (see Von Koenigswald, 1940, p. 61). There is a great deal to say respecting this matter, not only as regards the correct identification of the bovid types of Dubois and of Von Koenigswald as to genus and species and the synonymies involved, but also as regards the stratigraphical side of the question. Let us first consider systematically how many of Dubois's new species are valid, then go on to Von Koenigswald's new species, and finally look into the stratigraphie evidence adduced by Von Koenigswald. The fact that Dubois (1908) gave only brief diagnoses of his new species and no figures at all is no reason to consider these names invalidly published. Stremme (1911), whose excellent report on the Selenka Trinil collection is still the best work on the mammalian fauna of the Trinil beds as a whole, found Dubois's species often strikingly well characterized. "So skeptisch ich 4 D. A. HOOIJER anfangs den Bestimmungen der Säugetiere bei Dubois gegenüberstand, so musste ich doch auf die Dauer fast in allen Fällen seinen Ansichten zustim• men" (I.e., p. 82/83). As far as Duboisia is concerned there can be no doubt that the material described by Stremme is identical with Dubois's Tetraceros kroesenii. Moreover, this has been confirmed by Dubois when visiting the Selenka collection at Berlin (Stremme, I.e., p. 123). The skulls found by Dubois and first described and figured in the present paper resemble that dealt with by Stremme in every detail. Since Dubois explicitly states that his Tetracerus kroesenii is the same species as that originally named Anoa san• teng (Dubois, 1908, p. 1260, footnote 1), and since it is clearly stated in the paper by Jentink (1891) that the description of Anoa santeng was prepared by Dubois, the correct name of this, the only known species of Duboisia Stremme, is Duboisia santeng (Dubois). The water buffalo, a fine skull of which has been described by Stremme (1911) as "Buffelus palaeokerabau Dubois" doubtless is identical with Bubalus palaeokerabau of Dubois's (Stremme, I.e., p. 135), and should be so named; Bubalus Smith, 1827, has 38 years' priority over Buffelus Rütimeyer (Pilgrim, 1939, p. 254). In his paper of 1908 Dubois named two species of Bibos with the new specific names palaesondaicus and protocavifrons respectively. In the hand• written catalogue of specimens of his collection preserved in the Leiden Mu• seum Dubois even added a third new specific name, for a Bibos skull collected at Trinil in 1900 (Coll. Dub. no. 2774), but this name has never been publish• ed by Dubois and will also be omitted by me in order to keep the synonymy as simple as possible. As it happens, the synonymy of the fossil banteng of Java is complicated enough already for Dubois thrice gave a name to one and the same species. Not only do Bibos palaesondaicus Dubois (1908, p. 1262), type skull from Trinil (Coll. Dub. no. 2798) found in 1900, and Bibos protocavifrons Dubois (1908, p. 1262), type skull found at Tegoean in March, 1891 (Coll. Dub. no. 2797) refer to the same species, but Dubois's second species of "Leptobos", L. dependicornus Dubois (1908, p. 1261/1262), type skull from Trinil, 1900 (Coll. Dub. no. 2812), likewise is a Bibos; it repre• sents the female of Bibos palaesondaicus Dubois. Although Leptobos dependi• cornus has line precedence over Bibos palaesondaicus as well as over Bibos protocavifrons the three names were published in the same paper and on the same date. I select Bibos palaesondaicus, to the exclusion of the others, as the name to be used for the fossil banteng of Java because this name has been in use ever since it was first published, while the status of Leptobos dependi• cornus and of Bibos protocavifrons has never been well understood.
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