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La position systématique de Cryptoprocta ferox (Carnivora). Analyse cladistique des caractères morphologiques de carnivores Aeluroidea actuels et fossiles

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Afin de determiner la position phylogenetique de l'espece malgache Cryptoprocta ferox, nous avons etudie l'anatomie de differents genres de carnivores Aeluroidea et de deux taxons fossiles, Proailurus et Palaeopnonodon, auxquels il avait ete compare. L'analyse cladistique de 77 caracteres morpho-anatomiques place Cryptoprocta en groupe-frere des Felidae sur la base de caracteres dentaires essentiellement. Proailurus est le groupe-frere de l'ensemble Felidae + Cryptoprocta. Cette etude demontre surtout que Cryptoprocta doit etre exclu des Viverridae au sein desquels il est generalement classe.
... le cryptoprocte ne partage des caractères dérivés qu'avec les Herpestidae (VÉRON, 1995). Tous les autres carnivores autochtones de Madagascar sont, comme le cryptoprocte, endémiques de l'île. ...
... Les hypothèse de relations phylogénétiques au sein des Viverridae ainsi nouvellement délimités restaient à étudier ; les premières analyses cladistiques montraient surtout la nature homoplasique de nombreux caractères morphologiques (VÉRON, 1995 ;GAUBERT et al., 2002). Les premiers résultats moléculaires (VÉRON & HEARD, 2000) suggèrent la monophylie des civettes palmistes asiatiques, les Hemigalinae et les Paradoxurinae. ...
... Les linsangs africains et asiatiques furent rassemblés dans la tribu des Prionodontini (GRAY, 1864), mais par la suite les auteurs ont considéré Poiana comme une genette primitive, qui a acquis un morphotype similaire à celui des linsangs asiatiques par convergence du fait de leur mode de vie similaire dans les forêts tropicales (GREGORY & HELLMAN, 1939 ;SIMPSON, 1945 ;CRAWFORD-CABRAL, 1993). Prionodon présente des ressemblances aussi bien avec les Felidae qu'avec les Viverridae et la base du crâne est très proche de celles de féliformes fossiles de l'Oligocène, Paleoprionodon et Proailurus (HORSFIELD, 1824 ;TEILHARD DE CHARDIN, 1915 ;GREGORY & HELLMAN, 1939 ;VÉRON, 1995 ;HUNT, 2001). Des analyses moléculaires (GAUBERT & VÉRON, 2003) placent pour la première fois les linsangs asiatiques en position de groupe-frère des Felidae. ...
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Recent molecular studies have shed some light on the phylogeny of the feliform carnivores and have clarified the systematic position of some debated taxa that were previously placed within the Viverridae. The African palm civet, the Malagasy viverrids and the Asian linsangs are no longer included in the Viverridae, which now consists of four subfamilies: the Hemigalinae (banded palm civets and otter civet); the Paradoxurinae (palm civets and binturong); the Genettinae (genets and oyans); and the Viverrinae (terrestrial civets). The African palm civet (Nandiniidae) is the sister-group of all other feliforms, and the Asian linsangs (Prionodontidae) are the sister-group of the cat family (Felidae). The Malagasy civets (Euplerinae) and Malagasy mongooses (Galidiinae) form a monophyletic group (Eupleridae), which is the sister-group of the mongooses (Herpestidae). The Herpestidae family is itself split into two subfamilies, the Herpestinae, including the solitary species, and the Mungotinae, grouping the social species. These studies have also clarified which morphological features are reliable for the phylogeny of the feliform carnivores.
... This family includes digitigrade terrestrial species (civets), semi-digitigrade semi-arboreal species (genets), and plantigrade arboreal species (palm civets) (Ewer, 1973;Wozencraft, 1984;Taylor, 1988;Veron, 1994Veron, , 1999Nowak, 2005;Jennings and Veron, 2009). Their systematics was based mainly on the morphology of the basicranium (size and shape of the anterior and posterior chambers of the auditory bullae, position of the foramens at the base of the skull), the dentition, the feet, and the perineal gland (a scent pouch that lies between the genitals and the anus) (Gray, 1864;Flower, 1869;Mivart, 1882;Pocock, 1915aPocock, ,b,c,d, 1916bPocock, ,c, 1929Pocock, , 1933aPocock, ,b,c, 1934a1939;Gregory and Hellman, 1939;Hunt, 1974Hunt, , 1987Hunt, , 1989Hunt, , 1991Hunt, , 2001Wozencraft, 1984Wozencraft, , 1989aVeron, 1994Veron, , 1995. The perineal gland is specific to the Viverridae, and its product, called civet, is used in scent marking (see Jennings and Veron, 2009). ...
... The relationships within the Viverridae have been long debated, largely due to their large diversity of forms and also because this family was a dumping ground for all 'viverrid-like' species of feliforms (Pocock, 1916c; and see reviews in Wozencraft, 1984;Veron, 1994Veron, , 1995Bininda-Emonds et al., 1999;Veron and Heard, 2000;Hunt, 2001;Gaubert et al., 2002;Gaubert and Veron, 2003;Jennings and Veron, 2009). Until recently, the Viverridae was divided into six subfamilies (Wozencraft, 1993): two endemic to Madagascar, the Cryptoproctinae Gray, 1864 and the Euplerinae Chenu, 1852; two Asian subfamilies, the Hemigalinae Gray, 1864 and the Paradoxurinae Gray, 1864, one monospecific African subfamily, the Nandiniinae Pocock, 1929, and one subfamily, the Viverrinae Gray, 1821, with both Asian and African representatives. ...
... Until recently, the Viverridae was divided into six subfamilies (Wozencraft, 1993): two endemic to Madagascar, the Cryptoproctinae Gray, 1864 and the Euplerinae Chenu, 1852; two Asian subfamilies, the Hemigalinae Gray, 1864 and the Paradoxurinae Gray, 1864, one monospecific African subfamily, the Nandiniinae Pocock, 1929, and one subfamily, the Viverrinae Gray, 1821, with both Asian and African representatives. However, the systematic position of several viverrid species were questioned, in particular the African palm civet (Nandinia binotata), the Malagasy viverrids (the fossa Cryptoprocta ferox, falanouc Eupleres goudotii, and Malagasy civet Fossa fossana), and the Asian linsangs (the spotted linsang Prionodon pardicolor and banded linsang Prionodon linsang) (see Gray, 1864;Milne-Edwards and Grandidier, 1867;Flower, 1869;Filhol, 1879Filhol, , 1894Mivart, 1882;Carlsson, 1911;Gregory and Hellman, 1939;Pocock, 1940;Beaumont, 1964;Petter, 1974;Hunt, 1987Hunt, , 2001Flynn et al., 1988;Wozencraft, 1984Wozencraft, , 1989aHunt and Tedford, 1993;Wyss and Flynn, 1993;Veron, 1994Veron, , 1995Gaubert, 2003a). The first molecular phylogenies revealed a conflict with the traditional taxonomy (e.g. ...
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The phylogenetic relationships of the extant feliform carnivores, Felidae (cats), Herpestidae (mongooses), Hyaenidae (hyenas and aardwolf), and Viverridae (civets, genets, and oyans), have been debated for a long time, with several proposed hypotheses for the relationships of these families (Flower, 1869; Gregory and Hellman, 1939, see Figure 3.1; Simpson, 1945; Hunt, 1987; Flynn et al., 1988; Wayne et al., 1989; Wozencraft, 1989a; Hunt and Tedford, 1993; Wyss and Flynn, 1993; Veron, 1994). The position of the Viverridae family is still unresolved (see e.g. Gaubert and Veron, 2003; Flynn et al., 2005; Koepfli et al., 2006; Holliday, 2007). The mongooses were initially included within the Viverridae (Flower, 1869; Mivart, 1882) until Pocock (1916a, 1919) advocated for a family rank, to which he gave the name Mungotidae. Gregory and Hellman (1939) also placed them in a separate family, the Herpestidae Bonaparte, 1845. This separation was not followed by Simpson (1945) and several other authors (e.g. Albignac, 1973; Ewer, 1973; Petter, 1974; Rosevear, 1974; Coetzee, 1977; Kingdon, 1977; Payne et al., 1985; Stains, 1987; Taylor, 1988; Schreiber et al., 1989; Dargel, 1990; Skinner and Smithers, 1990). However, this split has been supported by further studies, based on morphology, chromosomes and molecular data (e.g. Wurster, 1969; Fredga, 1972; Radinsky, 1975; Bugge, 1978; Neff, 1983; Hunt, 1987; Wozencraft, 1984; Hunt and Tedford, 1993; Veron and Catzeflis, 1993; Wyss and Flynn, 1993; Veron, 1994, 1995; Flynn and Nedbal, 1998; Veron and Heard, 2000; Gaubert and Veron, 2003; Veron et al., 2004a; Flynn et al., 2005), and it is now generally accepted that the mongooses should be placed in a separate family, the Herpestidae (see Honacki et al., 1982; Wozencraft, 1989b, 1993, 2005; Gilchrist et al., 2009).
... To date, the systematics of the Hemigalinae and Paradoxurinae has been mainly assessed by morphological data (Pocock, 1933;Gregory and Hellman, 1939;Wozencraft, 1989;Veron, 1994Veron, , 1995. Molecular studies of feliform carnivorans have included very few representatives of these subfamilies due to the difficulties in obtaining biological material (see Veron and Heard, 2000;Gaubert 2. Materials and methods ...
... Some taxa were placed among the Viverridae on the basis of shared morphological traits, but some of these characters are now deemed plesi-omorphic (Gaubert et al., 2005). Several studies led to the exclusion of some 'viverrid-like' taxa and helped redefine the Viverridae sensu stricto as a monophyletic group (Veron and Catzeflis, 1993;Veron, 1995;Veron and Heard, 2000;Gaubert and Veron, 2003;Yoder et al., 2003;Gaubert et al., 2004aGaubert et al., , 2005, but little attention was paid to the Hemigalinae and Paradoxurinae. This study confirms the definition of the Viverridae as containing the Genettinae, Hemigalinae, Paradoxurinae, and Viverrinae. ...
... Following Carbone et al. (2007), we classified Bsmall-prey specialists^as species targeting prey smaller than themselves, Blarge-prey specialists^as those targeting prey larger than themselves, and Bmixed-prey specialists^as opportunists that regularly target either. Despite being distantly related to cryptoprocts (family Eupleridae), felids make an excellent reference population for cryptoprocts because, like (Yoder et al. 2003;Eizirik and Murphy 2009), and morphological research confirms that Cryptoprocta is cat-like in cranial as well as postcranial traits (e.g., Legendre and Roth 1988;Véron 1995). ...
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The extent to which Madagascar’s Holocene extinct lemurs fell victim to nonhuman predators is poorly understood. Madagascar’s Holocene predator guild included several now-extinct species, i.e., crocodiles, carnivorans, and raptors. Here we focus on mammalian carnivory, specifically the roles of Cryptoprocta spelea and its still-extant but smaller-bodied sister taxon, C. ferox, the fosa. Cryptoprocta spelea was the largest carnivoran on Madagascar during the Quaternary. We ask whether some extinct lemurs exceeded the upper prey-size limits of C. spelea. We use univariate and multivariate phylogenetic generalized least squares regression models to re-evaluate the likely body mass of C. spelea. Next, we compare characteristics of the forelimb bones of C. ferox and C. spelea to those of other stealth predators specializing on small, mixed, and large-bodied prey. Finally, we examine humeri, femora, crania, and mandibles of extinct lemurs from six sites in four ecoregions of Madagascar to identify damage likely made by predators. We test the relative prevalence of carnivory by mammals, raptors, and crocodiles at different sites and ecoregions. Our data reveal that crocodiles, raptors, and the largest of Madagascar’s mammalian predators, C. spelea, all preyed on large lemurs. Cryptoprocta opportunistically consumed lemurs weighing up to ~85 kg. Its forelimb anatomy would have facilitated predation on large-bodied prey. Social hunting may have also enhanced the ability of C. spelea to capture large, arboreal primates. Cryptoprocta carnivory is well represented at cave and riverine sites and less prevalent at lake and marsh sites, where crocodylian predation dominates.
... Despite these threats, little molecular systematics work has been done on this group; knowledge of their phylogenetic diversity, intraspecific genetic diversity and structure, and population differentiation is crucial for defining conservation priorities and for the identification of evolutionarily significant units (ESUs, Ryder 1986;Moritz 1994;Haig 1998;Isaac et al. 2007). The systematics of the Hemigalinae had previously only been assessed using morphological data (Pocock 1933;Gregory & Hellman 1939;Wozencraft 1989;Veron 1994Veron , 1995 until Patou et al. (2008) proposed a molecular phylogeny of the Hemigalinae and Paradoxurinae. Patou et al. (2008) found that the banded civet and Owston's civet were sister species, with the otter civet as the sister taxon to these two (as also obtained later by Agnarsson et al. 2010). ...
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Due to the difficulty in obtaining samples, the systematics of the Hemigalinae civets has not been fully resolved. The aim of this study was to clarify the relationships of the species and the intraspecific diversity within this subfamily, and to explore the environmental factors that might have affected its evolution. Using two mitochondrial and two nuclear markers, we confirmed that the Hemigalinae comprises Owston’s civet, the otter civet, Hose’s civet and the banded civet, but also the Sulawesi palm civet (formerly included in the Paradoxurinae). Our study showed that the banded and Owston’s civets are sister species, and suggested that Hose’s civet is sister to these two. Within the banded civet, we observed a high divergence between individuals from the Mentawai Islands and those from Sumatra and Borneo (while the latter two were not strongly divergent), likely due to the deep sea channel between the Mentawai Islands and Sumatra. Unexpectedly, the Sumatran and Peninsular Malaysian individuals were not closely related, despite the fact that these two regions have repeatedly been connected during the last glaciations. No high polymorphism was found within Owston’s civet, although three groups were obtained: southern China, northern Vietnam and central Vietnam, which might be related to Pleistocene climatic fluctuations.
... With respect to the phylogenetic relationships among carnivoran families, early small feliforms such as Stenoplesictis and Palaeoprionodon are placed at a sister taxon position to the extant Feliformia excluding the Nandiniidae (Véron 1995;Hunt 1998), while the Nimravidae is treated as the first divergence within the Feliformia or as a sister taxon to the crown Carnivora (i.e. Feliformia þ Caniformia; Flynn et al. 2010). ...
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This study reports occurrences of feliform carnivorans from the Upper Eocene Ergilin Dzo Formation and Alag Tsav locality of southeastern Mongolia. Nimravus mongoliensis (Nimravidae) is distinguished from other species in having a deep mandible, a longer p/1–2 and m/2 relative to p/4, and relatively wider P3/. Eofelis sp. (Nimravidae), a genus previously known only from the Oligocene of France is found in the Ergilin Dzo Formation. Alagtsavbaatar indigenus comb. nov. (Alagtsavbaatar gen. nov.; Feliformia) is established for new specimens from the Ergilin Dzo Formation and the previously known Stenoplesictis specimen from the Alag Tsav locality based on its characteristics such as moderately developed buccal cingulid and cingular and accessory cuspids on p/3–4, wide m/1 trigonid and double-rooted m/2 with a trenchant talonid. Stenoplesictis simplex from the Ergilin Dzo Formation is revised to Asiavorator gracilis, extending its chronological range back to the Late Eocene. Geographical and chronological distributions and morphological comparisons suggest that the Nimravidae originated by the Middle Eocene in southern East Asia and migrated northward in the Late Eocene, whereas the early small feliforms immigrated to northern East Asia in the Late Eocene and stayed within the middle-to-high-latitude area.
... Hunt (1989Hunt ( , 1998Hunt ( , 2001 reinforced this position with further studies of various extinct and extant feliforms. Subsequently, phylogenetic analyses of morphological (e.g., Véron 1995;Spaulding and Flynn 2012) and DNA data (e.g., Gaubert et al. 2005; have supported Hunt's position for N. binotata as sister to other extant feliform, which as noted above is widely held today (see also , Wozencraft 2005;Gaubert 2009). ...
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The external and endocranial surfaces of the skull of the African palm civet, Nandinia binotata (Gray, 1830), are described and illustrated in detail based on 30 specimens (from Carnegie Museum of Natural History and American Museum of Natural History). With the inclusion of a newborn and six juveniles with deciduous dentitions, a reasonable ontogenetic series is represented. The bone-by-bone descriptions are primarily based on the condition in an adult female and the newborn with consideration of variation across the sample. The principal cranial foramina are treated in a glossary, and the hyoid apparatus and larynx are described from a single specimen. The sample exhibits a remarkable degree of variability in cranial features that are often used as different states of characters in phylogenetic analysis (e.g., number and position of palatal foramina, the orbital mosaic, and composition of the lacrimal foramen). Nandinia binotata, the only taxon in the Nandiniidae, has been identified as the most basal extant feliform in recent phylogenetic analyses of both molecular and morphological data. It has long been recognized that its ear region with its uninflated auditory bulla exhibits a primitive level of organization. To assess the primitive nature of the skull of N. binotata, comparisons are made with three extant carnivorans, the felid Felis catus Linnaeus, 1758, the viverrid Genetta genetta (Linnaeus, 1758), and the canid Canis lupus Linnaeus, 1758. Of the three, N. binotata shares numerous resemblances across the skull with G. genetta, which accounts for its historical inclusion in the Viverridae. Whereas aspects of the ear region of N. binotata are clearly unique among extant carnivorans, the rest of its skull is not similarly so.
... The Viverridae form a heterogeneous taxon, which is probably not monophyletic (Hunt 1987, 1989, Hunt and Tedford 1993, Veron and Catzeflis 1993, Flynn and Nedbal 1998 and with uncertain intra-family relationships (Veron 1994(Veron , 1995. ...
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A survey of the morphology of pads within the Viverridae has been related to locomotion, ecology, ethology and to other morphological features. Species with digitigrade hind foot are lacking metatarsal pads, have an elongated pes, a short tail, and non retractile daws. They are terrestrial and their fur is spotted. Species with plantigrade hind foot, have broad and long metatarsal pads, which form a bare sole. They have retractile claws, a long tail, and fur with few markings or are one-coloured. They are arboreal and most of them are omnivorous or fruit-eaters. The genets and Poiana are intermediate forms because they have digitigrade feet, but possess long metatarsal pads, retractile claws, a long tail and spotted coat. However, some of them are close to the plantigrade state and have a plantigrade manus. The form of the pes of the genets is an adaptation for jumping. Prionodon is arboreal but have no metatarsal pads and is digitigrade. The ancestral condition is assumed to be plantigrady and arboreality and the presence of all the pads, which could be approach by some genets or Poiana, while the morphology of the foot of the Paradoxurinae, Nandinia and Cryptoprocta seems to be a secondary condition.
... Felidae, Hyaenidae, Mustelidae) may have an alisphenoid canal, while more derived and/or extant relatives may not (see, e.g. Veron, 1995;. The presence of two upper and lower molars, a character state that may also be defined as the loss of m3, is primitive for Carnivora. ...
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The red panda, Ailurus fulgens, is a peculiar recent carnivoran whose systematic relationships have been disputed since its first description in 1825 (Figure 4.1). While occurring with a single species today, Ailuridae was represented by several genera in the past. Flynn and Nedbal (1998) were the first to place Ailurus in its own family based on molecular evidence. While subsequent analyses of molecular data have confirmed this, paleontological findings of the last 10 years shed much more light on the natural history of Ailuridae and the morphology of its members. Before 1997, Ailuridae consisted of 5 genera with about 16 species. Only two genera belonged to Ailurinae (Ailurus and Parailurus). Today, the family contains 9 genera (5 of which are considered to be ailurines) with about 26 species. It now is agreed that the earliest ailurid is Amphictis, the earliest species of which are known from the Late Oligocene (MP 28, about 25 Mya) of Europe. Even though Ailurus today is restricted to Southeast Asia, it is apparent that ailurids were once present in all Northern continents and were most diverse in Europe throughout most of their history. In this chapter, we examine the recent and fossil evidence supporting the assignment of Ailurus fulgens and its ancestors (including Amphictis) to a distinct family, Ailuridae. We find such an assignment fully justified and corroborated by morphological data. We review all molecular and morphological studies in which Ailurus has been included. While new molecular studies continue to support a family Ailuridae, a review of the morphology of A. fulgens led to the identification of a number of characters which could be traced within the fossil relatives of Ailurus. This survey led to the development of emended diagnoses of all included genera. We also include a list of all ailurid species and resolve some outstanding issues concerning the taxonomy or systematic relationships of several taxa. Finally, we briefly recapitulate the natural history of Ailuridae. We follow Smith and Dodson (2003) for anatomical notation and orientation of dentitions.
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