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From Madagascar On the specific identification of subfossil Cryptoprocta (Mammalia, Carnivora) from Madagascar Steven M. GOODMAN Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605 (USA) and WWF, B.P. 738, Antananarivo (101) (Madagascar) [email protected] [email protected] Rodin M. RASOLOARISON Département de Paléontologie et d’Anthropologie biologique, B.P. 906, Université d’Antananarivo, Antananarivo (101) (Madagascar) and Deutsches Primatenzentrum, Kellnerweg 4, D-37007 Göttingen (Germany) Jörg U. GANZHORN Zoologisches Institut und Zoologisches Museum, Martin-Luther-King-Platz 3, D-20146 Hamburg (Germany) [email protected] Goodman S. M., Rasoloarison R. M. & Ganzhorn J. U. 2004. — On the specific identifica- tion of subfossil Cryptoprocta (Mammalia, Carnivora) from Madagascar. Zoosystema 26 (1): 129-143. ABSTRACT Grandidier (1902) described a large form of Cryptoprocta ferox Bennett, 1833, a genus of Carnivora endemic to Madagascar, from subfossil material dating from the presumed Holocene that he referred to the variety spelea. Subsequently there has been varying opinion on the validity of this taxon. In this paper 159 subfossil and 32 modern osteological specimens of Cryptoprocta are examined and analyzed to determine if indeed two separate KEY WORDS Mammalia, forms can be morphologically distinguished within the sample. On the basis Carnivora, of these tests we conclude that C. spelea Grandidier, 1902 is a valid species Cryptoprocta, that existed on the island in the recent geological past. It was presumably a subfossil, Holocene, formidable predator that may well have feed on a variety of large lemur Madagascar. species that are now extinct. ZOOSYSTEMA • 2004 • 26 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 129 Goodman S. M. et al. RÉSUMÉ Identification spécifique de Cryptocarpa subfossiles (Mammalia, Carnivora) de Madagascar. À partir de matériel subfossile datant probablement de l’Holocène, Grandidier (1902) a décrit une forme de Cryptoprocta ferox Bennett, 1833 de grande taille, carnivore endémique de Madagascar, qu’il a nommé « variété » spelea. Par la suite, il y eut des opinions variées quant à la validité de ce taxon. Dans le présent article, des spécimens ostéologiques subfossiles (n = 159) et modernes (n = 32) de Cryptoprocta sont examinés et analysés afin de détermi- ner si deux formes séparées peuvent vraiment être distinguées à partir de MOTS CLÉS Mammalia, l’échantillon. Sur la base de ces tests, la conclusion est que C. spelea Carnivora, Grandidier, 1902 est une espèce valide ayant existé sur l’île au cours d’un Cryptoprocta, passé géologique récent. C’était probablement un redoutable prédateur qui se subfossile, Holocène, nourrissait d’une variété d’espèces de lémuriens de grande taille, éteintes Madagascar. actuellement. INTRODUCTION Bones referable to the genus Cryptoprocta have been recovered from a wide variety of Holocene The modern native Carnivora fauna of Mada- paleontological sites on Madagascar, particularly gascar is composed of eight species, the largest of in the west and extreme south; none of these which is Cryptoprocta ferox Bennett, 1833. This specimens show signs of mineralization and are animal shows sexual dimorphism in body mass hence referred to as “subfossils”. Grandidier with adult males reaching between 6.2-8.6 kg (1902, 1905) studied osteological material of and adult females between 5.5-6.7 kg; average Cryptoprocta from the subfossil sites of body mass across adults is 6.8 kg (Hawkins 1998, Ambolisatra (23°03’S, 43°24’E) to the north 2003). C. ferox occurs in a variety of natural and of Toliara in the southwest and the cave of human modified habitats. Its diet is largely Andrahomana (25°50’S, 46°40’E) to the west of composed of vertebrates, particularly mammals, Tolagnaro in the extreme southeast and conclud- although a wide assortment of other items are ed that the material represented a form larger taken (Albignac 1973; Rasolonandrasana 1994; that extant C. ferox and he proposed to name it Rasoloarison et al. 1995; Goodman et al. 1997; as a distinct “variété” C. ferox var. spelea. He did Rasamison 1997; Goodman 2003). It is a very not designate a holotype specimen. Petit (1935) proficient hunter, being to pursue prey both on subsequently concluded that specimens housed the ground and in trees (Laborde 1986). The sys- in the Muséum national d’Histoire naturelle, tematic position of Cryptoprocta Bennett, 1833 Paris, and presumably part of the Grandidier (e.g., Veron & Catzeflis 1993; Veron 1995) and collection, and redescribed the form spelea as a Malagasy Carnivora in general have been the sub- distinct species. This latter author mentioned no ject of numerous studies that have reached very holotype. different conclusions about the familial and inter- Lamberton (1939) conducted a detailed osteolo- familial relationships of these animals. However, gical and morphometric analysis of Cryptoprocta recent molecular studies indicate that all native bones collected at a variety of paleontological Malagasy Carnivora form a monophyletic group sites on the island, all presumably dating from (Yoder et al. 2003). the Holocene. He concluded that indeed subfos- 130 ZOOSYSTEMA • 2004 • 26 (1) Subfossil Cryptoprocta (Mammalia, Carnivora) from Madagascar sil Cryptoprocta were for the most part larger than in 1658 (reprinted edition 1995: 221): « C’est the living species of this genus and concurred une bête grande comme un grand chien qui la with Petit (1935) in considering Grandidier’s tête ronde et au rapport des Nègres, elle a la res- form spelea to be a full species. Material referred semblance d’un léopard, elle dévore les hommes to C. spelea by Lamberton was excavated from the et les veaux. Elle est rare et ne demeure que sites of Beavoha (25°04’S, 44°19’E), grotte dans les lieux des montagnes les moins fré- d’Ankazoabo (north of Itampolo), Beloha quentés. » (25°10’S, 45°03’E), Belo-sur-Mer (20°44’S, Since Lamberton’s study of Cryptoprocta more 44°00’E), Bemafandry (25°07’S, 44°16’E), and modern and subfossil osteological specimens have Tsiandroina (24°59’S, 44°07’E). Lamberton also become available. The purpose of this paper is to noted that at some sites certain subfossils were reevaluate these subfossil specimens and specifi- closer in measurements to extant C. ferox than cally to assess morphological variation in extant the extinct C. spelea, particularly material from C. ferox, then to compare this range of variation Antsirabe, Beloha, and Beavoha – indicating the to subfossil material, in order to examine if at last possibility that at some sites the two species lived some of these bones can be separated from extant in temporal sympatry. However, given the lack of members of this genus. stratigraphic control associated with the material studied by Lamberton and that no corroborative radiocarbon dates are available, it cannot be sub- SPECIMENS AND MEASUREMENTS stantiated if the two species actually lived at the same time and in the same area. In 1989 a substantial portion of the paleontologi- The work on Malagasy subfossils conducted by cal collections from the Académie malgache were Lamberton has been of paramount importance in transferred to the Laboratoire de Primatologie et subsequent work in Quaternary paleontology Paléontologie des Vertébrés, Université d’Anta- (e.g., Godfrey & Jungers 2003). However, one of nanarivo. This included a considerable number the weak points of his monograph on Crypto- of the Cryptoprocta specimens described by procta was the lack of modern comparative mate- Lamberton (1939) in the collections of the rial to assess intraspecific variation in C. ferox. On Académie malgache. A number of recent paleon- the basis of comparisons he made, it appears that tological field studies have resulted in new mate- a maximum of three skeletons of recent C. ferox rial of subfossil Cryptoprocta and we have had were available to him. Given the range of sexual access to these collections (e.g., the sites of dimorphism in this species (Hawkins 1998) and Ankarana and Lakaton’ny akanga). Further, we the possibility of geographic variation associated have been able to measure subfossil and modern with the island’s dramatic bioclimatic zonation material of Cryptoprocta in several other museums (Ljungquist 1930), this number of specimens was (see below). insufficient to capture the possible range of varia- A series of measurements were taken by the first tion in this species. Subsequently certain authors author with a dial calipers to an accuracy of have considered C. spelea to be a synonym of 0.05 mm. The measurements include: C. ferox (Savage 1978; Köhncke & Leonhardt 1986). Cranial measurements Finally, amongst the subfossils studied by Condylobasal length: from the anterior edge of the premaxillae to the posteriormost projection of the Lamberton was a mandible from Tsiandroina occipital condyles. that had a form distinctly different from other Skull width: greatest skull width perpendicular to the species of Cryptoprocta, which he named as a greatest skull length, above bullae. distinct species, C. antamba. The name antam- Interorbital width: minimum width between the orbits. ba was derived from a reputed animal living in Nasal length: greatest length of nasal bone (rostral end southern Madagascar and described by Flacourt to fusion with frontal). ZOOSYSTEMA • 2004 • 26 (1) 131 Goodman S. M. et al. Zygomatic breadth: greatest breadth across the zygo- Modern and subfossil material was examined from matic process, perpendicular to skull length at the the following collections (Appendices 1 and 2) junction of the zygomatic-orbital suture. AMNH American Museum of Natural History, Palatal length: from anterior edge of premaxillae to New York; anteriormost point on posterior edge of palate. FMNH Field Museum of Natural History, Chicago; Incisive foramen: the inside length of one of the fora- MCZ Museum of Comparative Zoology, Harvard men. University; Upper molar row: length across the occlusal surface of MNHN Muséum national d’Histoire naturelle, the maxillary molar series.
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