Middle Miocene Chalicotheriinae (Mammalia ... - Wiley Online Library

Blackwell Publishing LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082© The Linnean Society of London? 2007

2007

1513 577608 Original Article MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCEJ. ANQUETIN ET AL.

Zoological Journal of the Linnean Society, 2007, 151, 577–608. With 11 figures

Middle Miocene Chalicotheriinae (Mammalia, Perissodactyla) from France, with a discussion on chalicotheriine phylogeny JÉRÉMY ANQUETIN1*, PIERRE-OLIVIER ANTOINE2 and PASCAL TASSY3 1

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK Équipe de Géodynamique, LMTG, Institut des Sciences de la Terre, 14 avenue Édouard Belin, 31400 Toulouse, France 3 Muséum National d’Histoire Naturelle, Département Histoire de la Terre, USM 203/UMR 5143 Paléobiodiversité, 57 rue Cuvier, 75231, Paris cedex 5, France 2

Received March 2006; accepted for publication April 2007

Middle Miocene chalicotheriine remains from France have played an important part in the palaeontological history of chalicotheres. For example, both the first known skull and first complete individual come from Sansan (MN 6, Gers, France). Nevertheless, these remains have never been described in detail until now. Firstly, cranial remains of Anisodon grande from Sansan are described. A review of nomenclatural problems supports Anisodon Lartet, 1851 as the appropriate genus name for this species and a new lectotype is designated. Secondly, the description of new remains of Chalicotherium goldfussi from Saint-Gaudens (MN 8, Haute-Garonne, France) considerably increases knowledge of this taxon. Thirdly, a skull from La Grive Saint-Alban (MN 7–8, Isère, France) is reassessed. A new cladistic analysis of the Chalicotheriinae is proposed incorporating these new data. This analysis, including 12 taxa and 51 cranio-mandibular and dental characters, supports the division of Middle and Late Miocene chalicotheriines into two clades. A novel pattern of relationships is proposed and the systematics is revised. However, chalicotheriine relationships remain labile because of the need for both a revision of pre-Middle Miocene taxa and the addition of postcranial data. © 2007 The Natural History Museum, London. Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608.

ADDITIONAL KEYWORDS: Anisodon – Chalicotherium – cladistics – Macrotherium – nomenclature – systematics.

INTRODUCTION Chalicotheres are perissodactyls with a very peculiar morphology. The most conspicuous feature is that they do not have hooves, but instead have large bifid claws that look like those of pangolins or sloths. The Chalicotherioidea Gill, 1872 are known from Middle Eocene to Pleistocene deposits. In current classifications (e.g. McKenna & Bell, 1997), there are two families within chalicotherioids: the Eomoropidae Matthew, 1929 and Chalicotheriidae Gill, 1872. The first is paraphyletic with respect to the second, and recently Hooker & Dashzeveg (2004) included ‘eomoropids’ within the Chalicotheriidae. However, until the whole phylogeny of chalicotheres s.l. is better established we follow *Corresponding author. E-mail: [email protected]

McKenna & Bell’s classification here. During the Neogene, two subfamilies are recognized within Chalicotheriidae (sensu McKenna & Bell, 1997): the Schizotheriinae Holland & Peterson, 1914 and Chalicotheriinae Gill, 1872. The chalicotheriines are known from Africa and Eurasia (e.g. Butler, 1965; Coombs, 1989), and persist until the Pleistocene in Asia (e.g. Falconer, 1868). They exhibit the most derived morphology among chalicotherioids and have been referred to as ‘chevauxgorilles’ (gorilla-horses) in popular accounts (Tassy, 1978). In fact, the forelimbs are very elongated whereas the hindlimbs are quite shortened, resulting in a clinograde posture (like gorillas). Moreover, according to Zapfe (1979), chalicotheriines practised a kind of knuckle-walking resembling in some respects the stance of gorillas.

577 © 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608

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Since the work of von Koenigswald (1932), it has been thought that one chalicotheriine genus was present during the Miocene, i.e. Chalicotherium Kaup, 1833a. More recently, de Bonis et al. (1995) described a new chalicotheriine species from the Late Miocene (MN 13) of Dytiko 3 (Macedonia, Greece) and produced the first cladistic analysis of chalicotheriines (based exclusively on cranial and dental characters). This study identified two clades among Middle and Late Miocene chalicotheriines: one including Chalicotherium goldfussi Kaup, 1833a (the type species of Chalicotherium), and the other including the new species from Greece (Late Miocene) and the species from Sansan, France [Middle Miocene; assigned to Chalicotherium grande (de Blainville, 1849) until this time]. According to de Bonis et al. (1995) the two clades are easily distinguished by skull anatomy, with the most obvious feature being the much reduced snout length of the Dytiko 3 and Sansan species. They proposed to resurrect the genus name Macrotherium, which was originally proposed by Lartet (1837b), for this second clade and named the new species from Greece Macrotherium macedonicum , whereas the species from Sansan became Macrotherium grande. In this paper, we present a detailed study of historic specimens from Sansan and La Grive Saint-Alban and a description of new remains from Saint-Gaudens; this new information allows us to test de Bonis et al.’s (1995) results. First, we reassess the validity of the name Macrotherium, and reject this name in favour of Anisodon Lartet, 1851, as proposed but not demonstrated by Geraads, Spassov & Kovachev (2001). A new lectotype for the species Anisodon grande (de Blainville, 1849) is designated. Anatomical descriptions are divided into three parts: (1) reassessment of the cranial remains from Sansan, which have never been accurately described and which were not included in de Bonis et al. (1995); (2) description of new remains (a right hemimandible and a symphysis) from SaintGaudens, France, which are assigned to C. goldfussi; (3) comparison of a skull from La Grive Saint-Alban, France, to other known chalicotheriine cranial remains. New additional characters (around 34%) were then included in a data matrix derived from that of de Bonis et al. (1995), and a cladistic analysis was performed. The result confirmed that there were two chalicotheriine clades during the Middle and Late Miocene, but a new, somewhat different pattern of relationships is found. As a consequence, we also revise the systematics of Chalicotheriinae.

HISTORICAL FRAMEWORK AND NOMENCLATURE From the initial discovery of the first chalicothere remains at the beginning of the 19th century until the

description of a complete individual from Sansan (Filhol, 1890), chalicothere skulls and postcrania were assigned to completely different taxa. The first chalicothere remains to be found were ungual phalanges from the Late Miocene of Eppelsheim, Germany. Kaup (1833b) assigned them to Deinotherium (which is now referred to the Proboscidea but was an enigmatic taxon at that time), whereas Cuvier (1822) interpreted them as the remains of a ‘pangolin gigantesque’ (gigantic pangolin). Kaup (1833a) described teeth of a new perissodactyl from Eppelsheim and named it C. goldfussi, without realizing that the teeth and claws pertained to the same taxon. During the 1830s, in France, Édouard Lartet began excavations in the Sansan deposits. Lartet (1837a) pointed out the presence of teeth of a large Anoplotherium and announced the discovery of postcranial remains of an ‘Edentata’ similar to Cuvier’s ‘pangolin gigantesque’; later he named the latter Macrotherium (Lartet, 1837b). After several years of excavation, the Sansan deposits yielded numerous chalicothere fossils. In his Ostéographie, de Blainville (1849) published a description of chalicothere cranial remains under the new species Anoplotherium grande . De Blainville died before he was able to describe postcranial remains, but plates and captions were published posthumously under the name Macrotherium (de Blainville, 1855). Until 1890, Macrotherium was regarded as an ‘edentate’ known only by its postcranium, whereas Chalicotherium and Anoplotherium grande were thought to be ungulates known only from their dentition. The discovery of a complete, articulated specimen (MNHN Sa 15671) at Sansan (Filhol, 1890) demonstrated that these different elements belonged to the same animal. This specimen is still on display in the Exhibition Hall of Palaeontology at the Muséum National d’Histoire Naturelle (MNHN), Paris, with the label: ‘Macrotherium sansaniense Lartet, 1851’. After Filhol (1890), the name Macrotherium was often used for Middle Miocene chalicotheres (mostly remains from France), whereas Chalicotherium was applied to Late Miocene forms (especially remains from Germany and Eastern Europe). Von Koenigswald (1932) argued that it was not useful to distinguish two genera during the Miocene, leading him to synonymize Macrotherium with Chalicotherium (subjective synonyms). This idea prevailed when de Bonis et al. (1995) proposed their reevaluation of Macrotherium. In addition to these disagreements concerning the nature and diversity of chalicotheres, there are nomenclatural problems. Butler (1965) provided an exhaustive synonymy list for each chalicotheriine species known at that time. Geraads et al. (2001) revealed that Macrotherium is a junior synonym of Chalicotherium Kaup, 1833a. The following is a historical review of names relevant to the problem surrounding the

© 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608

MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE availability of the name Macrotherium and to the application of the valid name for the species from Sansan. Detailed explanations regarding their current status under the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1999), abbreviated as ICZN below, are given below (such crucial explanations are missing in Geraads et al. 2001). 1837a Anoplotherium – Lartet, p. 88: Lartet announced dental remains of a large anoplothere from the Sansan deposits (note that the authorship of the name Anoplotherium is Cuvier, 1804). Lartet’s comment was just an announcement without assignment to an existing species, or a description, definition or indication of a new species (see Article 12 in ICZN, 1999: 16). 1837b Macrotherium – Lartet, p. 424 Nomen nudum: Lartet proposed the name Macrotherium for postcranial remains of the ‘edentate’ from Sansan (see above). There is a reference (Lartet, 1837b: 418) to the description of these remains in Lartet (1837a). However, Lartet (1837b) is not consistent with the Principle of Binominal Nomenclature and fails to satisfy the Article 11.4.1 (ICZN, 1999: 10). Consequently, Macrotherium Lartet, 1837b is unavailable. 1839 Anoplotherium magnum – Lartet, p. 26 Nomen nudum: The binomen applies to dental remains mentioned by Lartet (1837a). Butler (1965) designated this name as a nomen nudum because it is not associated with a description of the teeth concerned. Following Article 12 (ICZN, 1999: 16), the absence of description is not sufficient to designate a name as a nomen nudum and Butler’s argument fails. However, we were unable to find the publication of Lartet (1839) and to find out whether there is a definition or indication for this species, so we cannot say whether the species epithet magnum is available or not. This is a very delicate issue because, according to the Principle of Priority, if the name magnum is available, it would become the valid epithet for the species from Sansan. Since 1839, the epithet magnum was used by only three authors other than Lartet himself: Filhol (1890), Abel (1920, 1922) and Bohlin (1936) (see Butler, 1965). Consequently, until the publication of Lartet (1839) is found, it is preferable to grant the benefit of the doubt to Butler (1965) and continue to consider A. magnum Lartet, 1839 as a nomen nudum in order to avoid doubtful change in the already complex chalicotheriine nomenclature. 1844 Macrotherium giganteum – Pictet, pp. 232–233: Pictet proposed the first valid binomen with respect to ICZN (1999). Unfortunately, this is a hybrid species based on an ungual phalanx from Eppelsheim (the type locality of C. goldfussi) and postcranial remains

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from Sansan (the type locality of Anisodon grande). The species epithet giganteum has priority over grande despite the prevailing usage. But Butler (1965) saved the latter by choosing the ungual phalanx from Eppelsheim as the lectotype of M. giganteum. Actually, M. giganteum becomes a junior synonym of C. goldfussi Kaup, 1833a, and giganteum loses its priority over grande. Consequently Macrotherium is a junior synonym of Chalicotherium (as M. giganteum is the type species of Macrotherium). Hence, following the lectotypification of Butler (1965), Macrotherium is available and invalid. 1848 Anisodon (Choelichotherium) – Pomel, p. 687 Nomen nudum: Pomel cited the name Anisodon in a list of genera without nomenclatural value (no indications either of a species or of remains). As a matter of fact, this name appears in Lartet’s hand-written sale catalogue [which is not an official publication, as it does not satisfy Article 8.1.3 (ICZN, 1999: 6)], and refers to cranial and dental remains from Sansan. Pomel doubtless took this name from this catalogue. So, Anisodon is unavailable at this time. Lartet’s catalogue is dated from 1847 and is currently housed at the MNHN. 1849 Anoplotherium grande – de Blainville, p. 66: de Blainville described and named cranial and dental remains from Sansan. The species name grande is available (it becomes valid after the nomenclatural act of Butler 1965). De Blainville (1849: 68) mentioned that these specimens were temporarily called Anisodon in Lartet’s catalogue. But, Article 11.5.2 of the ICZN (1999: 11) specifies that ‘the status of a previously unavailable name is not changed by its mere citation’. So, Anisodon was still unavailable in 1849. Butler (1965) made a mistake in saying that de Blainville (1849) described another species from Sansan based on a mandible displaying a pathological position of P2. According to Butler (1965) this other species was named Chalicotherium anisodon . Indeed, de Blainville (1849) said that remains from Sansan were named Anisodon in Lartet’s catalogue and that they are quite similar to remains from Eppelsheim described by Kaup (1833a) as a new genus, Chalicotherium. De Blainville did not think that a new genus name (i.e. Chalicotherium) was needed for the remains from Sansan and for those from Eppelsheim. The valid name for him was clearly Anoplotherium grande, but he also proposed two junior synonyms: Chalicotherium europaeum and C. anisodon (this can be seen on legends of plate 8 in de Blainville, 1849). Butler (1965) was probably misled by plate 9 (de Blainville, 1849), which represents the hemimandible MNHN Sa 9376 with its peculiar position of P 2 in labial view. The illustration is labelled with both A. grande and C. anisodon. But MNHN Sa 9376 is


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also represented on plate 8 as A. grande, demonstrating that de Blainville saw only one species among remains from Sansan. Following Article 11.6 (ICZN, 1999: 11), C. europaeum is unavailable. This is not the case for C. anisodon because it was treated before 1961 as a valid name, and so satisfies Article 11.6.1 (ICZN, 1999: 11) (see also Butler, 1965). 1850 Chalicotherium grande – Gervais, p. 91: In his Zoologie et Paléontologie Françaises, Gervais assigned cranial and dental remains from Sansan to Chalicotherium (note that Chalicotherium was still considered to be an anoplothere at this time). He noted that these remains had not yet been compared with the other Chalicotherium species from Eppelsheim described by Kaup (1833a). This is a subjective synonymy. 1850 Macrotherium giganteum – Gervais, p. 136: Again in the Zoologie et Paléontologie Françaises, Gervais noted this ‘edentate’ from Sansan and Eppelsheim, and used the name proposed by Pictet (1844). Note that Gervais (1850: 136) incorrectly attributed the name M. giganteum to Lartet (1837a). 1851 Macrotherium sansaniense – Lartet, p. 22: Lartet gave this name to postcranial remains from Sansan. As the postcranial remains in question belong to the same taxon as cranial remains described as Anoplotherium grande by de Blainville (1849), the species name sansaniense is a junior synonym of grande. 1851 Anisodon magnum – Lartet, p. 30: Lartet published the generic name he had coined in his catalogue for cranial remains from Sansan [the same described by de Blainville (1849) as Anoplotherium grande]. For Lartet (1851: 30), the species epithet is that of Anoplotherium magnum Lartet (1839) (see discussion above). Anisodon becomes available as it is associated both with a species epithet and a description of remains. We follow Butler (1965) in considering Anoplotherium magnum Lartet (1839) as a nomen nudum (see above). In the context of this study, the species epithet magnum has been available since 1851, and so is a junior synonym of the epithet grande. The foregoing discussion shows the complexity of chalicotheriine nomenclature, and why it has to be studied thoroughly. The nomenclatural act of Butler (1965) plays an essential role. In choosing to save the species name grande, he condemned the use of the generic name Macrotherium Pictet, 1844. So, as argued by Geraads et al. (2001), Macrotherium is a junior synonym of Chalicotherium. The senior available generic name for the species from Sansan is Anisodon Lartet, 1851, and the senior available species name is Anoplotherium grande de Blainville (1849). Some doubt remains for the species epithet, as Anoplotherium magnum Lartet (1839) might be a senior synonym of grande (see discussion above).

Terminology: Dental terminology follows that of Coombs (1978) and Butler (1965). Institutional abbreviations: AMNH, American Museum of Natural History, New York, USA; BMNH, The Natural History Museum, London, UK; CCECM, Centre de Conservation et d’Étude des Collections du Muséum, Lyon, France; HLM, Hessischen Landesmuseum, Darmstadt, Germany; HMS, Heimatmuseum Stegersbach, Burgenland, Austria; LNK, Landessammlungen für Naturkunde, Karlsruhe, Germany; MHNT, Muséum d’Histoire Naturelle, Toulouse, France; MNHN, Muséum National d’Histoire Naturelle, Paris, France; MPA, Museum of Palaeontology (Branch of the National Museum of Natural History, Sofia), Assenovgrad, Bulgaria; PMM, Prirodonaucen Muzej na Makedonija, Skopje, Republic of Macedonia; UT, University of Thessaloniki, Thessaloniki, Greece; XBDD, North-west University Geological Department, Xian, Shaanxi, China.

SYSTEMATIC PALAEONTOLOGY ORDER PERISSODACTYLA OWEN, 1848 SUBORDER ANCYLOPODA COPE, 1889 SUPERFAMILY CHALICOTHERIOIDEA GILL, 1872 FAMILY CHALICOTHERIIDAE GILL, 1872 SUBFAMILY CHALICOTHERIINAE GILL, 1872 ANISODON LARTET, 1851 (NON POMEL, 1848) Nestoritherium Kaup, 1859 Diagnosis: The following synapomorphic characters (see cladistic analysis below) diagnose Anisodon as a member of the Chalicotheriinae: the height of the mandibular corpus increases posteriorly; the protoloph of P3 and P4 is lost; the protoloph does not reach the protocone on molars; upper molars are subsquare. Anisodon differs from other chalicotheriines (including Chalicotherium) mainly by its brevirostry. Anisodon displays a quite derived cranial anatomy and can be diagnosed by 16 unambiguous synapomorphies (see cladistic analysis below): upper tooth rows convex in lateral view; lacrimal tubercle present; shortened maxilla; presence of a groove between the ventral border of the zygomatic arch and M3; postorbital process of the jugal present; swollen braincase; laterally orientated paroccipital process; vertical occiput; upper tooth rows converging strongly anteriorly in ventral view; loss of the retromolar space on the maxilla; anterior wall of the tympanic bulla extending more anteriorly than the postglenoid process; angulus mandibulae expanded ventrally; metacone more labially placed than the paracone on M 1 and M2; external walls of the metacone and metastyle subparallel to the mesiodistal lengthening on M 3; weak ‘metastylid’ on lower molars.


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE Anisodon retains some plesiomorphic traits (small occipital condyles, and postfossette of upper molars not more pinched on M3 than on other molars), and differs regarding these characters from Chalicotherium. Type species: Anoplotherium grande de Blainville, 1849. Included species: Anisodon macedonicus, Anisodon wuduensis, Anisodon sivalense.

ANISODON

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under the mandibular symphysis, retaining a tubercle on the ventral border of the external auditory meatus and a protocone relatively more anterior in position on molars, having a less swollen braincase and a foramen caroticum that perforates the tympanic bulla; and from A. sivalense in retaining the paraconule on upper molars. Lectotype: Right maxilla (MNHN Sa 9339, formerly A.C. 4232) with M1–M3, P4 alveoli, and distal alveolus of P3 (Fig. 1). The tooth row was illustrated by de Blainville (1849: plate 8; see Fig. 2B) in association with a P4 that could not be identified in the MNHN collection

Anoplotherium grande de Blainville, 1849 Chalicotherium anisodon de Blainville, 1849 Macrotherium sansaniense Lartet, 1851 Anisodon magnum Lartet, 1851 Diagnosis: Anisodon grande displays two autapomorphies: absence of the canalis alisphenoideus; weak entoconid on P4. The species also retains, unchanged, the 16 synapomorphies of Anisodon (see above). Anisodon grande is distinguishable from other species of the genus by several plesiomorphic characters: short symphysis extending until the level of P 2; distal cingular crest of M3 stopping at the base of the hypocone. Additionally, A. grande differs from A. macedonicus by lacking the pterygoid lacuna and tuberosity

Figure 1. Lectotype of Anisodon grande (de Blainville, 1849); right maxilla MNHN Sa 9339 (formerly A.C. 4232) with M1–M3. Scale bar: 1 cm.

Figure 2. Plate 8 (genus Anoplotherium) of de Blainville (1849) showing: the cranium MNHN Sa 15670; A, the right hemimandible MNHN Sa 9376 with the problematic symphysis; B, the tooth row of the maxilla MNHN Sa 9339 (the new lectotype). MNHN Sa 9339 was confused with MNHN Sa 9340 by Schaefer & Zapfe (1971). Orientations are reversed from the actual specimens (see text). © 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608

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(as de Blainville’s illustration is not accurate enough). Another lectotype, previously selected by Schaefer & Zapfe (1971), is not valid under ICZN regulations (see below). Paralectotypes: The other remains described by de Blainville (1849): a fragment of a left hemimandible (MNHN Sa 9374); the cranium MNHN Sa 15670 (illustrated: see Fig. 2 this paper); a complete mandible (left side MNHN Sa 9341 and right side MNHN Sa 9376 – the latter was illustrated: see Fig. 2A this paper); a fragment of a right juvenile hemimandible (MNHN Sa 9377). Type locality: Sansan, Gers, France. Age: Middle Miocene, Astaracian (MN 6). Why a new lectotypification? Schaefer & Zapfe (1971) selected a lectotype for Anisodon grande among remains from Sansan. This specimen was a left maxilla (MNHN Sa 9340, formerly A.C. 4233) with P4–M3. Unfortunately, this specimen is not part of the type series as it was not described by de Blainville (1849), so it could not be chosen as the lectotype (Article 74.1, ICZN, 1999: 82). Moreover, according to Article 74.2 (ICZN, 1999: 82), ‘If it is demonstrated that a specimen designated as a lectotype was not a syntype, it loses its status of lectotype’. Schaefer & Zapfe (1971) stated that the left maxilla MNHN Sa 9340 was illustrated on plate 8 of the Ostéographie (de Blainville, 1849). In fact, although there is a tooth row on plate 8, it is that of the right maxilla MNHN Sa 9339. This error is the result of two factors. Firstly, the illustrations published by de Blainville are reversed, as a result of the engraving techniques used at that time. Secondly, Schaefer & Zapfe (1971) were confused by the presence of the P4 on the illustration of the tooth row (Fig. 2B), as this tooth is present in MNHN Sa 9340 but not in MNHN Sa 9339. Conversely, the illustration of MNHN Sa 9339 is accurate in that the M 2 is unworn, and that the distal alveolus and the distal part of the mesial alveolus of P3 are present (compare Figs 1 and 2B). Hence, the P4 was probably added to the illustration, maximizing the information provided. As there are other isolated P4s in the collection, we do not know whether de Blainville used the premolar of MNHN Sa 9340 when adding the missing P 4 to the illustration of the tooth row of MNHN Sa 9339. Then, among the syntypes, the right maxilla MNHN Sa 9339 appears to be the best choice for the lectotype. It is well preserved and was illustrated by de Blainville (1849), thus following ICZN Recommendation 74B (ICZN, 1999: 83). Moreover, it clearly has some relation to Schaefer & Zapfe’s (1971) reasoning. In the following section, we provide a detailed description of the cranium (MNHN Sa 15670) found by Lartet

and described briefly by de Blainville (1849). We also present the first description of the skull that belongs to the articulated skeleton found by Filhol (1890) at Sansan (MNHN Sa 15671).

Cranium MNHN Sa 15670 This specimen represents the first ever known chalicothere skull. It is obvious that Filhol (1890) examined the specimen, as he gave measurements missing from the original description by de Blainville (1849). Since that time, however, there has been no published evidence that any subsequent authors have studied this skull. De Bonis et al. (1995) were unable to locate it. Subsequently, this cranium was identified in the collection of the MNHN by one of us (PT), and Geraads et al. (2001) subsequently published a brief comment, but did not provide a detailed description. Furthermore, their comment is surprising with respect to the specimen (see discussion below). Consequently, a complete description of this specimen is provided with as much detail as possible. Comparing the illustrations by de Blainville (1849: plate 8; see Fig. 2) with the cranium today (Fig. 3), it is obvious that it was subsequently damaged, probably after 1890, as Filhol (1890: 296) provided measurements of the condyles, which are now lost. The two M3s were originally complete, but are now broken. De Blainville’s illustrations show the anterior part of the left zygomatic arch, which is absent today. Finally, there is a large vertical fracture on the posterior part of the cranium showing that the occipital region was dissociated from the remaining skull (the condyles were probably lost at that time).

Dorsal view: Sutures between the different bones are not apparent. The frontal crests are low, especially anteriorly. They converge posteriorly, but do not fuse to form a sagittal crest. Following this, the crests diverge to define the ‘interparietal triangle’ (Barone, 1966; see also de Bonis et al., 1995), but because the superior part of the occiput is broken we cannot determine its orientation. The braincase is tubular. Lateral view: The anterior part of the snout is broken, especially on the right side. On the left side, the orbital area is distorted: the inferior part of the maxilla and the anterior part of the orbit are separated and joined to the remainder of the skull in a more posterior and lower position than would have occurred in life. This probably explains de Blainville’s (1849: 68) error when he described the orbit as ‘very small, round and quite low’ (our translation from the French original). Indeed, the anterodorsal margin of the orbit is present on the skull roof clearly anterior to its suggested position. The foramen infraorbitale lies above the distal border of M1. The alveolar border is convex in lateral


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Figure 3. Anisodon grande. Cranium MNHN Sa 15670: A, left lateral view; B, ventral view with an enlarged and labelled view of the basicranium; C, dorsal view. Anatomical abbreviations: bo, basioccipital; bs, basisphenoid; eam, external auditory meatus; fc, foramen caroticum; fo, foramen ovale; gu, guttural fossa; ma, matrix; pog, postglenoid process; shf, stylohyoid fossa; tb, tympanic bulla; vc, vaginal crest. Scale bars: 1 cm. © 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608

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view. There is an outgrowth of the maxilla just above the mesial root of M2. Such an outgrowth is unknown in other chalicotheriines. Sutures between the maxilla and jugal, and between the maxilla and lacrimal, are marked by a shallow vertical groove. The jugal extends anteriorly to a point situated above the parastyle of M2. The orbitosphenoidal area is badly damaged, but it is still possible to observe the deep groove accommodating the optic nerve on the orbit floor. The postglenoid process is strong and flattened anteroposteriorly. It is separated from the external auditory meatus by a narrow furrow. Above the meatus, there is a strong tuber that develops from the lambdoid crest. The meatus is bordered posteriorly by the posttympanic process of the squamosal. The ventral margin of the meatus displays a blunt, rounded process. The squamosal is slightly domed, so the braincase is swollen in transverse section. The occipital face is vertical. Occipital view: Superficially, the occipital face seems to be low, but this results from the lack of its lower part (including the condyles, foramen magnum, and paroccipital processes). Its upper edge is rounded. The occipital tubercle is high on the occiput and continues ventrally as a vertical ridge. The paroccipital process continues dorsally on the occipital face as a vertical ridge. Ventral view: The skull is strongly distorted and the anterior part is translated laterally. Moreover, the anteroposterior axis of the basicranium is not exactly the same as that of the snout. A large part of the guttural area is missing. The snout is broken in front of the P 3, but the distal end of the left P2 alveolus is preserved. The tooth rows converge strongly anteriorly. The zygomatic arch extends as far forward as the distal part of M 2. The anterior part of the ventral edge of the zygomatic arch displays a strong elongate tuberosity. Between this tuberosity and M3 there is a deep anteroposterior groove. There is no retromolar space. The palate is broken displaying numerous fractures, and the choanal opening is missing. What is preserved of the guttural fossa shows that it was divided sagittally by a high crest that issued from the basisphenoid, anterior to the basioccipital crest. The postglenoid processes are anteroposteriorly flattened and their anterior faces display a subvertical, ovoid articular facet. The tympanic bulla is elbowshaped and flattened ventrally. On both sides, the anterior borders of the bullae are broken, but it is obvious that they extended more anteriorly than the postglenoid processes. The auditory duct is pressed against the posterior face of the postglenoid process. The vaginal crest and stylohyoid fossa are only partially visible, but it is still obvious that they were positioned as far posteriorly as the external auditory

meatus. Although the paroccipital processes are broken at their base, it is clear that they were orientated ventrally and externally. The foramen ovale opens in front of the anterior wall of the bulla. The foramen lacerum medium cannot be located. Anterior to the foramen ovale, there is no trace of a canalis alisphenoideus. Although its absence cannot be confirmed in MNHN Sa 15670 because the sphenoid area is badly damaged, specimen MNHN Sa 9398 (see below) displays no canalis alisphenoideus. The foramen caroticum perforates the bulla on its posterior ventral margin, whereas the foramen metoticum and canalis nervi hypoglossi cannot be observed. Teeth are badly preserved on this specimen, and yield no useful new information. What is visible does not differ from the other remains from Sansan.

Skull of the complete individual discovered by Filhol (MNHN Sa 15671) When Filhol (1890) announced the discovery of a complete, articulated skeleton, he emphasized the association between a ‘Macrotherium’ postcranium (which was thought to be an ‘edentate’ at this time) and a skull with a Chalicotherium-type dentition, but he did not describe the cranial anatomy. In fact, nobody has done this to date. This omission may be because this skull is quite crushed and fragmented, and taking its distortion into account is therefore difficult. The specimen (Fig. 4A) lies on its left side and is embedded in a large slab of lacustrine limestone from the lower levels of the Sansan deposits (Filhol, 1890). A cast of the cranium was made to facilitate study. Although badly crushed, this skull yields a number of important observations.

Cranium: Viewed as a whole (Fig. 4B), the cranium seems very high throughout its length. The upper profile of the skull roof seems to be horizontal and the occiput is exceptionally bulged posteriorly and ventrally. Finally, the lengthening of the skull appears to be too great for what is traditionally considered as a ‘macrothere’. However, these first impressions do not reflect the original morphology. In fact, the degree of distortion misrepresents the original morphology and only a few features can be confidently identified on this specimen. Anteriorly, the two maxillae are closely pressed against one another. The right maxilla is represented only by the M 3 and a part of its associated alveolar margin. The rest of the right maxilla consists of small bony fragments lying inside the nasal cavity. The left maxilla is better preserved: the three molars are present and it is possible to follow the alveolar margin up to the alveolus of P 2. The anterior border of the left maxilla seems to indicate the position of the nasal opening, but it is difficult to be sure whether it is damaged or not, so the shape



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Figure 4. Anisodon grande. A, complete skeleton MNHN Sa 15671 found by Filhol (1890) at Sansan (currently on display in the Exhibition Hall of Palaeontology at the MNHN, Paris) (copyright MNHN); B, cast of the skull of MNHN Sa 15671 (the white arrow indicates the isolated canine). Anatomical abbreviations: anm, angulus mandibulae; lc, lambdoid crest; lt, lacrimal tubercle; or, orbit; por, postorbital process of the jugal; ptm, post-tympanic process; zy, zygomatic arch. Scale bars (black or white bar): A, 30 cm; B, 1 cm.

of the nares remains conjectural. Dorsally, fragments of the nasal(s) and frontal(s) are present. This area is so badly crushed and distorted that it cannot be concluded that the nasals are as far anteriorly extended

as they appear to be, in a way that somehow resembles extant tapirs. Posteriorly, the skull roof is better preserved. However, deformation of the originally horizontal skull roof into a two-dimensional, vertical plane


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gives the misleading impression that the skull is very deep. There is no sagittal crest. The lower part of the lambdoid crest can be observed as can the posttympanic process of the squamosal, which lies slightly posterior to the lambdoid crest. The preserved part of the lambdoid crest indicates that the occiput did not project posteriorly. All of the occiput, and what is visible of the basicranium in lateral view, is crushed, giving the misleading impression that the skull bulges posteriorly and ventrally. The principal features of interest in this skull are the preservation of the zygomatic arch and anterior border of the orbit. Despite its bad condition, this specimen is the only one from Sansan that displays these areas. The rounded anterior border of the orbit shows a strong lacrimal tubercle. The zygomatic arch is shattered, but the remaining fragments are easily distinguishable. The jugal bears a moderate, but obvious, postorbital process just anterior to the jugalsquamosal suture. The zygomatic arch is slightly bowed posterodorsally, but in any case is not as high as the superior border of the orbit. Mandible: The two hemimandibles are pressed against one another (Fig. 4B). Only one observation can be made concerning the symphysis: on the left hemimandible, a dorsal part of the symphysis is present and extends posteriorly at least up to the P 2, but the right hemimandible hides its complete extent. The corpus increases in height posteriorly. Its ventral border is straight. The angulus mandibulae expands ventrally. In lateral view, the tooth rows are slightly concave. The slope of the anterior border of the ramus is orientated at about 90 degrees relative to the corpus. There is no retromolar space. The coronoid process is partly broken and the crest present on its anterolabial border is low. Upper dentition: Only the left M1–M3 and right M3 are present. This specimen was probably a young adult when it died as tooth wear is minimal. On each molar the protoloph only reaches the paraconule, and continues as a low crest until the protocone summit. The protocone is receded but its position is still relatively anterior, leaving a large, U-shaped distal valley behind it. A vertical crest descends from the summit on both the labial and lingual walls of the paracone. The mesial cingulum is strong and not reduced in front of the protocone. On M1 and M2 the metacone is more labial than the paracone. The metaloph is shorter on M 3 than on M2 (because of the contraction of the distal part of M 3). On M3 the metacone is labially displaced. The external wall of the metacone and metastyle is nearly parallel to the mesiodistal length. The mesiolingual face of the hypocone of M3 is very abrupt and the distal cingular crest does not reach the summit of this cusp. Finally, the postfossette (sensu Butler, 1965) is not more pinched on

M3 than on the two other molars (see below). The lingual cingulum is weakly developed. Lower dentition: Right M1–M3 are undamaged, whereas left M2–M3 are broken. The left hemimandible displays the alveoli of P2–M1, whereas the right is broken anterior to the alveoli of P 4. On the molars, the paraconid is low and the metaconid is the highest cusp. The ‘metastylid’ [perissodactyls lost the metastylid early in their evolution, this pseudometastylid is the distal cusp of a twinned metaconid (Hooker, 1994)] is observable, although it is not particularly distinct from the metaconid. It shows a well-developed metacristid. The talonid is slightly more elongated than the trigonid and both are V-shaped. The mesial wall of the hypolophid and the distal walls of the protolophid and entolophid show a weak vertical, medial incisure (see description of MHNT VAL-3, below). The cingulum is thickened on the distal face and weak elsewhere. The other teeth are represented by their alveoli, and it is notable that P2 has two alveoli. An isolated tooth (Fig. 4B) is embedded in the sediment several centimetres in front of the mandible. This tooth is complete and has only one root. It is either a lower canine or a lower incisor. Unfortunately, little is known of such teeth in Anisodon grande. Zapfe (1979) tentatively assigned some isolated teeth from Neudorf (MN 6, Slovak Republic) to incisors of A. grande, but in the absence of any association. These teeth are slightly flattened labiolingually. The labial face is flat, whereas the lingual one shows two enamel crests descending toward the lateral border of the tooth and joining a basal, lingual point. They are either sharp or rounded, probably as a result of differential attrition by the tongue and food. The incisors of the holotype of Anisodon macedonicus (UT DKO 234) are much reduced, rounded and button-shaped (de Bonis et al., 1995). The isolated tooth associated with MNHN Sa 15671 does not correspond to any of these descriptions, and is more compatible with lower canine morphology: a triangular crown with a distal crest showing a weak lingual, vertical groove (see description of the symphysis MHNT VAL-4 from Saint-Gaudens, below). For the moment, this tooth is provisionally identified as a right lower canine. Additional observations Other cranial remains from Sansan (Appendix 4) provide some additional character information that is not seen in the two skulls described above. MNHN Sa 9398 includes several cranial fragments that belong to the cerebral region. The occipital condyle is relatively small, oval, low, and narrow. The ‘interparietal triangle’ is displaced on the dorsal face of the skull. A fragment of the left basicranium confirms the absence of the canalis alisphenoideus.


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE The crowns of P2 and P3 are unknown at Sansan. On P4, the protoloph does not reach the protocone and ends at the paraconule. On very worn teeth it only reaches the base of the protocone. On molars, the crest extending the protoloph continues until either the protocone summit or the paraconule base (intraspecific variation). The development of the crest on the lingual wall of the paracone is also variable. Sometimes, the distal part of M3 is less reduced and the metacone is less displaced labially (e.g. MNHN Sa 9373). The symphysis is never completely preserved in any specimen found at Sansan, but several remains suggest that it extends posteriorly up to the level of the P2–P3 boundary. There is no retromolar space on the mandible. The angle between the anterior border of the ramus and the corpus is variable. When it is preserved, the angulus area always expands ventrally. The P2 is similar to that of UT DKO 234. Only one complete P4 is known from Sansan. It is molar-shaped, but the talonid is lower than the trigonid. The entoconid is weak.

The awkward symphysis described by de Blainville De Blainville (1849: 68–69, plates 8 and 9) described and illustrated the right hemimandible MNHN Sa 9376. According to him, three incisors were present in a rather long symphysis that expands laterally forming a palette-shaped structure (Fig. 2A). There is no doubt that de Blainville illustrated the specimen MNHN Sa 9376 – which, nevertheless, now lacks the symphysis. With the exception of the symphysis, the outline of the specimen is exactly the same and P2 is drawn in its conspicuous pathological position (note that only M2 and M3 are present in MNHN Sa 9376, but de Blainville completed the dentition on plate 8 probably for the needs of illustration as he did for MNHN Sa 9339). The symphysis illustrated by de Blainville is somewhat reminiscent of that of a hippopotamus, because of its great dilatation. Yet, neither MNHN Sa 9376 nor any other remains from Sansan [nor mandibles from Neudorf described by Zapfe (1979)] display evidence for such a morphology. From his description, it is obvious that de Blainville observed a symphysis. Was he misled in his identification, assembling two isolated pieces erroneously, or has this material been misinterpreted by de Blainville and lost since? We tentatively interpret this specimen somewhat differently. Comparison with other known chalicotheriine symphyses (especially MHNT VAL-4 from Saint-Gaudens, see below) reveals that the original remains observed by de Blainville would have been distorted. The anteroposterior axis of the symphysis seems to sit at an angle of 20–30° to that of the corpus. If this interpretation is correct, the symphysis would not be more expanded than in other chalicotheres.

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More importantly, if this symphysis is genuine, there would have been two incisors instead of three as de Blainville thought (see below). Such a hypothesis can only be confirmed by study of this symphysis, but no such specimen could be located in the MNHN collection. Note concerning remains from Neudorf Zapfe (1979) described chalicotheriine remains from Neudorf an der March, now Devínska Nová Ves (Slovak Republic), and assigned them to Chalicotherium grande. These remains were found in fissure fills referred to the Middle Miocene (MN 6). The chalicotheriines from Neudorf are represented by at least 60 individuals, including both cranial and postcranial remains (Zapfe, 1979). Geraads et al. (2001: 600) raised doubts concerning the conspecific status of the Neudorf and Sansan remains. Our study confirms this suspicion as some specimens from Neudorf differ from their equivalents at Sansan. This means that perhaps two taxa are associated at Neudorf (probably Anisodon grande and a Chalicotherium). The Neudorf remains need to be reviewed to establish their taxonomic status.

CHALICOTHERIUM KAUP, 1833A Macrotherium Pictet, 1844 Diagnosis: Amoung Chalicotheriinae (height of the mandibular corpus increasing posteriorly; protoloph of P3 and P4 lost; protoloph not reaching the protocone on molars; upper molars subsquare), Chalicotherium is diagnosed by two unambiguous synapomorphies (see cladistic analysis): occipital condyles elongated vertically; postfossette more pinched on M3 than on other molars. The cranial anatomy of Chalicotherium is far less derived than that of Anisodon and retains numerous plesiomorphic traits, such as: a rather long snout and skull; parallel (in ventral view) and straight (in lateral view) upper tooth rows; lack of a lacrimal tubercle; no groove between the ventral border of the zygomatic arch and M3; postorbital process on the jugal absent; tent-shaped braincase (not swollen); ventrally orientated paroccipital process; upper part of the occiput projected posteriorly; retromolar space present on both the maxilla and mandible; tympanic bulla not extended anterior to the postglenoid process; angulus mandibulae straight (not expanded ventrally); metacone at least as lingual as the paracone on M 1 and M2; external walls of the metacone and metastyle subperpendicular to the mesiodistal lengthening on M 3; strong ‘metastylid’ on lower molars. Type species: Chalicotherium goldfussi Kaup, 1833a. Included species: Chalicotherium brevirostris.


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CHALICOTHERIUM

GOLDFUSSI

KAUP, 1833A

Macrotherium giganteum Pictet, 1844 Diagnosis: In the present paper, C. goldfussi is not diagnosed by any autapomorphy. This is the consequence of our choice not to include the specimen CCECM Lgr 1065 from La Grive Saint-Alban within C. goldfussi for the time being, despite both anatomical evidence and the result of our cladistic analysis (see below). Chalicotherium goldfussi displays the two synapomorphies of Chalicotherium, and almost all plesiomorphic characters mentioned in the diagnosis of the latter (except those of the skull, as no skull is known for this species). Both C. goldfussi and CCECM Lgr 1065 have a complete protoloph on P3 [unambiguous synapomorphy of the clade (C. goldfussi, CCECM Lgr 1065)]. Lectotype: Right M3 (HLM Din. 3167) described and illustrated by Kaup (1833a). This lectotype was designated by Schaefer & Zapfe (1971), who reviewed and illustrated the specimen. Paralectotype: Right lower molar described by Kaup (1833a) as an M3 (HLM unknown number). Type locality: Eppelsheim (Rheinhessen, Germany). Age: Late Miocene (Vallesian), Deinotherium sands (MN 9).

Remains from Saint-Gaudens De Bonis et al. (1995) described chalicothere remains from Saint-Gaudens (Haute-Garonne, France) in the MHNT collection: a left maxilla and a right M 3. These remains come from the Valentine Quarry, which is attributed to the late Middle Miocene MN 8 biozone (Antoine, Duranthon & Tassy, 1997). Since then, one of us (POA) found a right hemimandible and a complete symphysis in the MHNT Valentine collection, both undescribed. In fact, this chalicothere from SaintGaudens was mentioned (without description or illustration) by Harlé (1898: 382) consisting of two half-jaws (‘demi-mâchoires’). These remains correspond to the maxilla described by de Bonis et al. (1995) and the hemimandible described herein. There is also an unnumbered M1. These five specimens belong to the same individual although they bear different catalogue numbers. The maxilla and the M 3 are numbered MHNT VAL1 and MHNT VAL-2, respectively [both unnumbered when de Bonis et al. (1995) described them], whereas the hemimandible and the symphysis are MHNT VAL-3 and MHNT VAL-4, respectively. Chalicothere remains from Saint-Gaudens are important because they can be confidently assigned to

Figure 5. Chalicotherium goldfussi. Specimen from SaintGaudens (Valentine Quarry). Symphysis MHNT VAL-4: A, right lateral view; B, frontal view, note the three alveoli per side for the incisors (right alveoli are indicated); C, dorsal view. Right hemimandible MHNT VAL-3 with M1–M3 (same individual as MHNT VAL-4): D, labial view; E, occlusal view of the tooth row; F, distolabial view of M3 and M2 illustrating the presence of the medial incisures (see text). Scale bars: 1 cm.

C. goldfussi (sensu this study) and include the first mandibular symphysis known for this species.

Right hemimandible MHNT VAL-3 The section anterior to the distal root of P 4 is missing, but the rest of the mandible is complete (Fig. 5D). The three molars are preserved. The corpus increases in height posteriorly. In transverse section, the inferior margin of the corpus is slightly swollen on its labial face. The ventral border of the corpus is straight and there is no ventral expansion of the angular area. There is a large retromolar space. The anterior border of the ramus is nearly perpendicular to the ventral border of the corpus. The ramus is elongated anteroposteriorly. The foramen mandibulare opens slightly


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE ventral to the tooth row, beneath the coronoid process. The latter is flattened and curved posteriorly, and displays a strong mesiolabial crest. The area between the coronoid process and the condyle is more strongly concave in a vertical plane than in UT DKO 234, and is similar to the state in Moropus (Schizotheriinae). The condyle is flat. Ventral and lingual to it, there is an oval articular surface for the postglenoid process. Just below, a small tubercle is present. The molars (Fig. 5E) bear a talonid longer than the trigonid. In occlusal view the former is V-shaped, whereas the latter is U-shaped. The paraconid is low, but well defined and underlined by the mesial cingulum. The ‘metastylid’ is strong and distinct from the metaconid: consequently, the hypolophid does not reach the metaconid summit, even on worn molars. The metacristid is strong. The cingulum is absent lingually and enlarged distally. There are well-developed vertical medial incisures on the distal walls of the metalophid and entolophid, and on the mesial wall of the hypolophid (Fig. 5F). Although these structures have not been described before, they can also be observed on remains from Sansan and on the specimen from La Grive Saint-Alban (CCECM Lgr 1065).

Symphysis MHNT VAL-4 The complete symphysis MHNT VAL-4 includes the beginning of the two corpuses (Fig. 5C). Only the right canine is preserved, but other teeth are represented by their alveoli. There are three incisors on each side (Fig. 5B). Alveolus size increases from I 1 to I3. The canine is just behind the third incisor, and its lateral outline is triangular with a subvertical mesial face. There is a vertical crest running from the summit to the distolingual border of the tooth, which defines a large lingual groove. In dorsal view (Fig. 5C) the incisor border is slightly convex anteriorly. Posterior to the canine, the symphysis constricts before broadening suddenly at the level of the corpuses. Unfortunately, there is no trace of the P2 alveolus, so it is impossible to determine the posterior extent of the symphysis. Yet, the diastema between the canine and P 2 was long. The dorsal surface of the symphysis is concave. In lateral view (Fig. 5A), the symphysis is very slender anteriorly and reaches the minimum thickness necessary for incisor insertion. The foramen mentale opens just posterior to the symphysis constriction. In frontal view (Fig. 5B) the arrangement of canines and incisors is arched ventrally. The canine is anterodorsally implanted. The incisor border forms a plateau and the incisors are implanted strictly forward (as in Moropus). The ventral face of the symphysis is smooth and no tuberosity is present.

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Comparison with remains from the Late Miocene Deinotherium sands, Germany Additional remains from Eppelsheim (type locality of C. goldfussi) include a right maxilla (HLM Din. 3168) with P2–M3 (Zapfe, 1979: fig. 7). Zapfe (1989) also reported the presence of C. goldfussi at Höwenegg (Hegau, Germany), represented by a left M 2 (LNK te pli 3) and by a right P4 (LNK te pli 2), M1, and M3 (LNK te pli 1). Comparison between these remains and those from Saint-Gaudens supports assignment of the latter to C. goldfussi. A left mandible with M 1–M3 found in the Late Miocene (MN 12–13) of Henndorf (HMS unknown number), Austria, was also assigned to this species (Schaefer & Zapfe, 1971: fig. 3a, b; Zapfe, 1979: fig. 40). However, several features, i.e. the absence of retromolar space, a symphysis extending posterior to the mesial part of P4, a short diastema, and a V-shaped trigonid on the molars, contradict this assignment. These characteristics correspond to that of Anisodon. The Henndorf mandible, assigned to C. goldfussi since its first description, is the specimen that prevented authors recognizing the existence of two different chalicotheriine lineages. The mandible from Titov Veles (Garevski & Zapfe, 1983) and those of Chalicotherium wuduensis Xue & Coombs (1985) from China were referred to Chalicotherium because they resemble the mandible from Henndorf. Now, with the description of new remains of C. goldfussi from SaintGaudens, we can confidently exclude the Henndorf mandible from this genus.

THE CHALICOTHERE FROM LA GRIVE SAINT-ALBAN (MN 7–8) PREVIOUS

WORK

Chalicothere remains from La Grive Saint-Alban (Isère, France) were described for the first time by Depéret (1892). This material consists of a partially damaged cranium (CCECM Lgr 1065; Fig. 6), with a fragment of each hemimandible (same number) associated with postcranial remains. Depéret (1892) attempted to emphasize differences between these remains and those from Sansan by erecting a race: Macrotherium grande race rhodanicum. According to Depéret (1892), these differences demonstrated a transition between Macrotherium grande and C. goldfussi. De Bonis et al. (1995) presented a brief review of the cranial remains. Based on their phylogenetic analysis, they assigned this specimen to C. goldfussi race rhodanicum. Here, the cranial remains are compared with other equivalent remains so that the striking difference between the cranial anatomy of Anisodon and that of


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Figure 6. Skull CCECM Lgr 1065 (Chalicotherium ?goldfussi) from La Grive Saint-Alban: A, right lateral view; B, ventral view; C, dorsal view. Scale bars: 1 cm.

Chalicotherium can be illustrated easily. Comparative specimens include: MNHN Sa 15670 from Sansan (Anisodon grande), MHNT VAL-1, -2, -3, -4 from Saint-Gaudens (C. goldfussi), UT DKO 234 from Dytiko 3 (Anisodon macedonicus), and also the schizotheriine Moropus and AMNH 26518 from Tung Gur, Mongolia [Chalicotherium brevirostris (Colbert, 1934)].

CRANIUM Dorsal view The cranium is large and quite elongate. The nasals are slightly abbreviated, a condition correlated with the regression of the upper border of the nasal opening, but they are still longer than wide (Fig. 5C). The above features separate this skull from that of


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE A. grande and A. macedonicus (MNHN Sa 15670 and UT DKO 234, respectively). After fusing to form a sagittal crest, the temporal crests split and fuse with the lambdoid crests. They therefore define what Depéret (1892) called the ‘méplat triangulaire’ (triangular plane), corresponding to the ‘interparietal triangle’ sensu Barone (1966). This structure is displaced onto the dorsal surface of the skull, as in A. grande, but differs from the condition seen in A. macedonicus. Lateral view The maxilla is high and notably elongated (Fig. 6A). As in MHNT VAL-1, C. brevirostris and Moropus, the tooth row is straight in lateral view. The zygomatic arch seems narrow because the upper portion of the squamosal part is damaged. This distortion also gives the impression that the postorbital process of the jugal is well developed, whereas this was probably not the case. The following features distinguish CCECM Lgr 1065 from MNHN Sa 15670 and UT DKO 234: there is a strong pillar-shaped postorbital process on the frontal that is distinct from the supraorbital process (see discussion about character 6 in Appendix 2); the stocky postglenoid process is separated from the external auditory meatus by a wide groove; the upper part of the occiput greatly projects posteriorly to overhang the occipital condyles; the braincase is tentshaped (not swollen) in transverse section. Occipital view The occipital face is high as in anisodonts, but is unlike the condition seen in Moropus (the occipital face is unknown in both C. goldfussi and C. brevirostris). Its outline is trapezoidal and less rounded than in anisodonts (Fig. 7J, K). The occipital tubercle is high. The preserved part of the paroccipital processes shows that they were orientated only ventrally (ventrally and laterally in anisodonts). The occipital condyles are more expanded vertically than in anisodonts. A vertical crest runs dorsally from the paroccipital process and borders medially the suture between the occipital and petrosal bones. The occipital part of the petrosal bone is oblique with respect to the occiput and is underlined by a subvertical wide and shallow gutter. However, these features cannot be observed in detail in other chalicotheriines because their occipital regions are less well-preserved (see description of MNHN Sa 15670). Ventral view The tooth rows do not converge anteriorly. There is a retromolar space and the choanae open far behind M 3,

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as in Moropus and AMNH 26518. The anterior part of the ventral border of the zygomatic arch displays a low crest, but no strong tuberosity. There is no anteroposterior groove between this crest and M 3 (Fig. 6B). All the above features separate CCECM Lgr 1065 from A. grande and A. macedonicus, and are also found in Chalicotherium and Moropus. The postglenoid process is globular, a unique feature of this specimen. The tympanic bulla does not extend anteriorly to the anterior face of the postglenoid process, as it does in specimens from Sansan and UT DKO 234. The foramen caroticum perforates the bulla in its posterior part, i.e. it is included in the ventral wall of the bulla, as in A. grande.

MANDIBLE A fragment of each hemimandible was found in association with the cranium. The right one is the most complete, with a corpus including P3–M3 and a fragmented ramus. The left hemimandible is reduced to a partial corpus with P4–M3. The retromolar space is large, unlike the condition in anisodonts. The angular area (observed on the right hemimandible) is quite fragmented. Although Depéret (1892) and de Bonis et al. (1995) noted that the angulus mandibulae is slightly expanded ventrally, it is very difficult to interpret this with confidence because of fragmentation. If there is an expansion it is quite weak and not as strong as the condition seen in A. grande and A. macedonicus. It is more comparable with the very slight expansion seen in MHNT VAL-3 (Fig. 5D) and Moropus. The symphysis extends posteriorly up to P2.

UPPER

DENTITION

Depéret (1892) described the P2. This tooth is now missing but its root indicates that it was longer than wide, the reverse of the state seen in A. macedonicus (UT DKO 234). P3 has a complete protoloph, although it is represented by a relatively thin crest. There is no protoloph on P4. On both P3 and P4, the paraconule is partly fused to the paracone lingual wall, whereas it is medially placed with respect to the crown mesial border in anisodonts. On the molars, the mesial cingulum is not lowered in front of the protocone, in contrast with the state seen in A. macedonicus. The following features separate CCECM Lgr 1065 from Anisodon: on M2, the metacone is as lingually positioned as the paracone; the metacone and metastyle external walls are subperpendicular to the anteroposterior lengthening; and on M3, the postfossette is more pinched than in other molars.


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Figure 7. Cranial character states. A, ventral view of the skull of Anisodon macedonicus (MNHN SLQ 1054a – cast of the holotype UT DKO 234); B, ventral view of the skull of Chalicotherium brevirostris (cast of AMNH 26518); C, ventral view of the skull of Moropus elatus (modified from Holland & Peterson, 1914); D, right lateral view of the skull of A. macedonicus (MNHN SLQ 1054a – cast of the holotype UT DKO 234); E, right lateral view of CCECM Lgr 1065; F, left lateral view of the skull of C. brevirostris (cast of AMNH 26518) (the drawing is reversed for comparative purposes); G, left lateral view of the skull of M. elatus (modified from Holland & Peterson, 1914) (drawing reversed); H, dorsal view of the skull of A. macedonicus (MNHN SLQ 1054a – cast of the holotype UT DKO 234); I, dorsal view of the skull of M. elatus (modified from Holland & Peterson, 1914); J, occipital view of the skull of A. macedonicus (MNHN SLQ 1054a – cast of the holotype UT DKO 234); K, occipital view of CCECM Lgr 1065. Not to scale. © 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608


LOWER

DENTITION

The P2 is now missing from the specimen, but was described by previous authors (Depéret, 1892; de Bonis et al., 1995). P4 is molar-shaped and shows a strong entoconid. On the molars, the trigonid is Ushaped as in C. goldfussi and unlike the condition in other chalicotheriines. This pattern is characterized by a more elongated hypolophid that terminates before reaching the metaconid summit, and by a protolophid perpendicular to the mesiodistal lengthening.

TAXONOMIC

ATTRIBUTION

Cranial and dental remains from La Grive SaintAlban are indistinguishable from those attributed to C. goldfussi (note that the skull of this species is known only by maxillae and mandibles). Such a referral is critical as CCECM Lgr 1065 would represent by far the most complete cranial remains known for this species. For the time being, we refrain from making a formal referral [and thus from discussing the assignment of CCECM Lgr 1065 to C. goldfussi rhodanicum by de Bonis et al. (1995)] in order to avoid further complications in chalicotheriine systematics. We therefore refer this specimen to Chalicotherium ?goldfussi. Future studies, especially on postcranial remains, might confirm the specific attribution to C. goldfussi (and deal with the legitimacy of a subspecies level for this specimen). We included CCECM Lgr 1065 as a separate terminal taxon in the cladistic analysis (see below).

PHYLOGENETIC RELATIONSHIPS OF CHALICOTHERIINES: CLADISTIC ANALYSIS PREVIOUS

WORK

The phylogenetic analysis proposed by de Bonis et al. (1995) is the only one available for chalicotheriines. It did not include either MNHN Sa 15670 or MNHN Sa 15671. It did include the skull from La Grive Saint-Alban (CCECM Lgr 1065), but did not update its description. Finally, de Bonis et al. were unaware of the mandible and symphysis from SaintGaudens (MHNT VAL-3 and MHNT VAL-4), which convey crucial information on C. goldfussi.

TAXON

SAMPLING

All taxa used by de Bonis et al. (1995) in their cladistic analysis are included here, except Chalicotherium styriacum. This species was described by Bach (1912) and is known only from a partial left mandible from Eggersdorf. De Bonis et al. (1995) examined a cast of this mandible at the BMNH. They expressed doubts concerning the identity of the teeth present in this specimen and concluded that it would be necessary to

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X-ray the original to ensure that the M 3 is present within the corpus. Eomoropus and Moropus are used as outgroups. Eomoropus Osborn, 1913 is one of the earliest representatives of the Chalicotherioidea (sensu McKenna & Bell, 1997), and is known from the Middle Eocene. This terminal taxon was coded mainly on the basis of the type species Eomoropus amarorum (Cope, 1881), following Osborn (1913) and Lucas & Schoch (1989). Moropus Marsh (1877) is a member of the Schizotheriinae. This terminal taxon was coded using its bestknown species, Moropus elatus Marsh, 1877 from the Early Miocene of North America, following Holland & Peterson (1914) and Coombs (1978). Anisodon macedonicus from the Late Miocene (MN 13) of Greece was coded following de Bonis et al. (1995), with supplementary observations made on a cast of the holotype (UT DKO 234) housed in the MNHN as SLQ 1054a, b and c, for the skull, left hemimandible and right hemimandible, respectively. The coding of Macrotherium brevirostris Colbert (1934) was problematic for de Bonis et al. (1995). This species is known only on the basis of one skull (AMNH 26518) from Tung Gur, Mongolia (MN 7–8). Only outline drawings of the left side and ventral view of this skull were given by Colbert (1934). De Bonis et al. (1995) mainly based their observations on poor photographs of the specimen. Subsequently, Louis de Bonis acquired a good quality cast of AMNH 26518, which we were allowed to study (this cast is now housed at the MNHN). Anisodon grande was coded on the basis of remains from Sansan (MN 6), the type locality (Appendix 4 lists cranial remains from Sansan available at the MNHN and used in the present study). In contrast to de Bonis et al. (1995), remains from Neudorf are excluded from this terminal (see above). Chalicotherium goldfussi was coded using remains from Eppelsheim and Höwenegg (MN 9), and also remains from Saint-Gaudens (MN 8). De Bonis et al. (1995) did not use the latter, but unlike us, did include the mandible from Henndorf (HMS unknown number) (Schaefer & Zapfe, 1971: figs 3a, b; Zapfe, 1979: fig. 40). However, according to us, this mandible does not belong to C. goldfussi but rather pertains to an anisodont (see above). The specimen (CCECM Lgr 1065) from La Grive Saint-Alban (MN 7–8) was included as a separate terminal taxon in order to test its relationships with C. goldfussi. Remains from Vathylakkos described by Arambourg & Piveteau (1929) (i.e. a partial skull, a hemimandible, three isolated upper teeth, an ungual phalanx, and a metatarsus; MNHN SLQ 65, 66, 67, 68, and 69, respectively) were also included as a terminal taxon.


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This taxon was coded following the original description and from direct observations on remains. Vathylakkos is a set of localities of the Late Miocene (MN 11) of Salonica (Greece). This terminal taxon is labelled ‘Vathylakkos’ in the following analysis. Chalicotherium wuduensis was coded following the original description by Xue & Coombs (1985). This taxon is known only from two specimens: two mandibles pertaining to an adult (XBDD 48Wd0079) and a juvenile (XBDD 48Wd0080) from the Late Miocene (Turolian?) of the Longjiagou Valley, Wudu Country, Gansu Province, China. Butleria rusingensis (Butler, 1965), from the Early Miocene of East Africa, is commonly known as the sister group of Eurasiatic chalicotheriines (although this is probably just an artefact resulting from the incomplete taxonomic assemblage of the present study). It was coded on the basis of the original description by Butler (1965). This description is informative, especially concerning the tooth morphology, but lacks photographs and detailed illustrations that would permit coding of new characters. A better understanding of chalicotheriine relationships requires the revision of this taxon (especially concerning nondental characters). Garevski & Zapfe (1983) described a mandible (PMM unknown number) from the Late Miocene (Turolian?) of Titov Veles (now Veles, Republic of Macedonia) and assigned it to C. goldfussi. De Bonis et al. (1995) pointed out that this mandible resembles that of A. macedonicus and chose to introduce it as a separate terminal taxon. We follow these authors and use

it as a terminal taxon (called ‘Titov Veles’ below) to test its position. We coded it following the original description by Garevski & Zapfe (1983). The genus Nestoritherium Kaup, 1859 is coded using its type species Nestoritherium sivalense (Falconer & Cautley, 1843) (see also, Colbert, 1935). It is one of the most recent chalicotheriines, known from the Plio-Pleistocene of the Siwalik Hills (Pakistan). The coding follows the description and plates of Falconer (1868), augmented by direct observations on remains available at the BMNH. The latter include a left hemimandible with P 4–M3 (BMNH M36374), and two maxillae that are thought to be parts of the palate of the same individual (BMNH M15366 with right P4– M3, and BMNH M15367 with left P3–M3). The BMNH catalogue listed this palate as the holotype, but this might not be the case according to M. C. Coombs (pers. comm., 2005). There is also a cast (BMNH M2710) of the anterior part of the skull and jaw illustrated by de Blainville (1849: plate 9) and Falconer (1868). The original specimen has probably been lost (Holland & Peterson, 1914). Table 1 lists taxa used in the present analysis with their specimen numbers (or references) and localities. Note that the systematics correspond to the results presented below.

CHARACTERS The analysis was based on a matrix of 51 characters (Appendix 1) comprising 29 cranial, 7 mandibular, and 15 dental (11 upper dentition and 4 lower) characters,

Table 1. List of terminal taxa used in the phylogenetic analysis with specimens and/or references used and localities Terminal taxa

Specimens/references used

Localities

Anisodon grande

Collections from Sansan at the MNHN (see Appendix 4) UT DKO 234 MNHN SLQ 65; 66; 67 PMM unknown number XBDD 48Wd0079; 48Wd0080 BMNH M36374; M15366; M15367; M2710 (cast) AMNH 26518 HLM Din. 3167; 3168/LNK te pli 1; 2; 3/MHNT VAL-1; 2; 3; 4

Sansan, France

Anisodon macedonicus Vathylakkos (Anisodon sp.) Titov Veles (Anisodon sp.) Anisodon wuduensis Anisodon sivalense Chalicotherium brevirostris Chalicotherium goldfussi

CCECM Lgr 1065 (Chalicotherium ?goldfussi) Butleria rusingensis

CCECM Lgr 1065

Eomoropus Moropus

see Osborn (1913), Lucas & Schoch (1989) see Holland & Peterson (1914), Coombs (1978)

see Butler (1965)

Dytiko 3, Greece Vathylakkos, Greece Veles, Macedonia Gansu Province, China Siwalik Hills, Pakistan Tung Gur, Mongolia Eppelsheim and Höwenegg, Germany/Saint-Gaudens, France La Grive Saint-Alban, France Various localities including Rusinga, Kenya North America, Eurasia North America

Note that the nomenclature of terminal taxa follows the new proposed systematics. © 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608

MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE which were coded for the 12 taxa mentioned above and listed in Table 1. Three characters (2, 29 and 35) have multiple states and are treated as ordered. The character list and discussions regarding coding are given in Appendix 2. The majority of character states are illustrated in Figs 7 (cranial), 8 (mandibular), and 9 (dental). From now on, references to characters used by de Bonis et al. (1995) are flagged with a number sign (e.g. #5). Six characters (#2, #3, #11, #16, #20, and #23) were not retained. Character #2 concerns the length of the nasal, but the data are insufficient and this character is redundant with #1. Similarly, there are not enough data to code character #3 (perinasal border with or without maxillar depression) accurately. Character #11 concerns the development of the ‘pterygoid tubercle’ (see de Bonis et al., 1995). In fact, the pterygoid blade is almost always broken and the ‘pterygoid tubercle’ is thereby unobservable. Character #16 con-

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cerns the shape of the vaginal crest (flattened or rounded). The presentation of this trait makes it ambiguous. According to de Bonis et al. (1995), the vaginal crest is rounded only on AMNH 26518 and CCECM Lgr 1065, but in reality the crest is broken in both specimens. The same applies to character #20 [paroccipital process: massive (0); or flattened (1)] as state 0 corresponds to a broken paroccipital process. Character #23 [temporal crests: not merged at the level of the occipital crest (forming an ‘interparietal triangle’) (0); or merged (1)] is not retained because there is indeed an ‘interparietal triangle’ in CCECM Lgr 1065, the only taxon possessing state 1 according to de Bonis et al. (1995). Other characters from de Bonis et al. (1995) have been used, checked, and recoded when necessary, with many multistate characters being coded as binary characters here (Appendix 2). Eighteen new characters are proposed.

Figure 8. Mandibular character states. A, dorsal view of the symphysis of Chalicotherium ?goldfussi (MHNT VAL-4); B, dorsal view of the mandible from Titov Veles (modified from Garevski & Zapfe, 1983); C, ventral view of the symphysis of C. ?goldfussi (MHNT VAL-4); D, ventral view of the symphysis of the specimen from Titov Veles (modified from Garevski & Zapfe, 1983); E, lingual view of the right hemimandible of Anisodon macedonicus (MNHN SLQ 1054c – cast of the holotype UT DKO 234) (note that UT DKO 234 has a supernumerary lower molar on both sides, but that this does not change the coding of character 30); F, lingual view of the right hemimandible of C. ?goldfussi (MHNT VAL-3); G, labial view of the left hemimandible of Moropus elatus (modified from Holland & Peterson, 1914). Not to scale. © 2007 The Natural History Museum, London Journal compilation © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 577–608

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Figure 9. Dental character states. A, M2–M3 of Anisodon grande (lectotype MNHN Sa 9339) (drawing reversed); B, P2–M3 of Moropus elatus (modified from Holland & Peterson, 1914); C, P3–M3 of Anisodon macedonicus (MNHN SLQ 1054a – cast of the holotype UT DKO 234); D, P3–M3 of Chalicotherium ?goldfussi (left maxilla, MHNT VAL-1); E, M1–M3 of Chalicotherium ?goldfussi (right hemimandible, MHNT VAL-3) (drawing reversed); F, M2–M3 of A. grande (left hemimandible, MNHN Sa 9341). Not to scale.

RESULTS The parsimony analysis was carried out using PAUP 3.1.1 (Swofford, 1993). A Branch-and-Bound search generated 14 most parsimonious trees (MPTs) with a length of 62 steps (consistency index, CI = 0.871; CIexcluding uninformative characters = 0.840; retention index, RI = 0.877). The strict consensus tree (65 steps; CI = 0.831; CIeuc = 0.792; RI = 0.831) is presented (Fig. 10). Branch support values (Bremer, 1994) show that both Chalicotheriinae and Anisodon are the strongest clades. The retention indices are high, indicating a strong phylogenetic signal. The phylogeny is well-resolved: only the relationships within the group (A. macedonicus, Vathylakkos, ‘C.’ wuduensis, Titov Veles, N. sivalense) are unresolved. The hypothesis of de Bonis et al. (1995) is supported, with two main chalicotheriine lineages during the Middle and Late Miocene. The first includes A. grande, A. macedonicus and the Vathylakkos specimen, i.e. taxa previously called macrotheres (now anisodonts). The second incorporates Chalicotherium s.s.. The branch supporting the clade Anisodon is the strongest, in terms of both its length and Bremer support value (Fig. 10). The most obvious result of the present analysis is the novel pattern of relationships that is proposed. Three terminal taxa – i.e. ‘C.’ wuduensis, Titov Veles, and N. sivalense – formerly close to C. goldfussi (de Bonis et al., 1995: fig. 9), become closer to both

A. macedonicus and the Vathylakkos form. These three terminals are still poorly known, but the increased data provided herein shows that they share several derived features with other anisodonts (see below). Relationships of ‘M.’ brevirostris (AMNH 26518) were unresolved in de Bonis et al. (1995) (lying either as a basal chalicotheriine, a basal Anisodon, or a basal Chalicotherium), but a more complete coding has been possible as a result of observations on the cast of AMNH 26518. ‘M.’ brevirostris is the sister group of the clade (C. goldfussi, CCECM Lgr 1065) and is included in the clade Chalicotherium. C. goldfussi and CCECM Lgr 1065 are indistinguishable based on the characters used in this analysis (both are supported by a zero-length branch): they are likely to be the same taxon. Butleria keeps its status of sister group to post-Early Miocene chalicotheriines, representing the ancestral morphotype of the latter for the coded characters (no autapomorphy). According to the present analysis, several modifications to chalicotheriine systematics are required. Table 2 summarizes the proposed systematics of the Chalicotheriinae and compares it with that presented in major recent papers: Coombs (1989), de Bonis et al. (1995), and Geraads et al. (2001). Chalicotherium wuduensis becomes Anisodon wuduensis (Xue & Coombs, 1985). Some authors (Coombs, 1989; de Bonis et al., 1995) had indicated that a generic distinction was probably inappropriate for Nestoritherium. In a way, this point of view is confirmed here: Nestorithe-



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Figure 10. Phylogeny of the Chalicotheriinae based on 51 cranio-mandibular and dental characters: strict consensus of the 14 most parsimonious trees (MPTs: 62 steps; CI = 0.871; RI = 0.877) (strict consensus: 65 steps; CI = 0.831; RI = 0.831). Nodes are labelled with letters. Numbers above branches are Bremer branch support values. Numbers under branches are the number of nonambiguous synapomorphies. ‘Vathylakkos’ and ‘Lgr 1065’ are only supported by ambiguous characters with DELTRAN optimization (see Appendix 3): they are represented as zero-length branches.

Table 2. Systematics of the Chalicotheriinae – a comparison between present results and major recent papers Assignment in Coombs (1989)

Assignment in de Bonis et al. (1995)

Assignment in Geraads et al. (2001)

Present work

C. grande – – C. goldfussi C. wuduensis N. sivalense C. brevirostris C. goldfussi – C. rusingense

M. grande M. macedonicum M. macedonicum Chalicotherium C. wuduensis N. sivalense ‘M.’ brevirostris C. goldfussi C. goldfussi race rhodanicum B. rusingensis

A. grande ‘M.’ macedonicum – – – – ?A. brevirostris C. goldfussi C. goldfussi –

Anisodon grande Anisodon macedonicus Vathylakkos (Anisodon sp.) Titov Veles (Anisodon sp.) Anisodon wuduensis Anisodon sivalense Chalicotherium brevirostris Chalicotherium goldfussi CCECM Lgr 1065 (C. ?goldfussi) Butleria rusingensis

Abbreviations: A., Anisodon; B., Butleria; C., Chalicotherium; M., Macrotherium; N., Nestoritherium.

rium Kaup, 1859 becomes a junior synonym of Anisodon Lartet, 1851 (subjective synonymy). Phylogenetic relationships of the specimens from Vathylakkos and Titov Veles are still unresolved, so they are assigned to Anisodon sp., awaiting further investigation. AMNH 26518 is attributed to C. brevirostris (Colbert, 1934). Finally, the generic name Butleria de Bonis et al., 1995 is conserved here.

In the following sections, we describe the distribution of unambiguous synapomorphies along the strict consensus tree (Fig. 10). Our results are compared with those of de Bonis et al. (1995): for each node, common characters and conflicts are listed. The latter are discussed further in Appendix 2. Monophyly of Chalicotheriinae (Fig. 10, node B) is supported by four unambiguous synapomorphies (two


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are homoplastic): the height of the mandibular corpus increases posteriorly (30; RI = 0.5); loss of the protoloph on P3 (38; RI = 0.667); loss of the protoloph on P 4 (39; RI = 1); the protoloph does not reach the protocone on molars (42; RI = 1). The only character in common with de Bonis et al.’s (1995) analysis is 30 (#25). In de Bonis et al. (1995), this node was also supported by characters #26 (31) and #36 (51). The first is recoded from a three-state to a binary character, whereas the second was unsatisfactorily observed by de Bonis et al. (1995) (for discussions, see Appendix 2). Monophyly of post-Early Miocene Chalicotheriinae (Fig. 10, node C) is supported by one nonhomoplastic, new character: the upper border of the nasal opening above P3–P4 (2; RI = 1). This node shows how poorly known the basal chalicotheriine phylogeny is. This illustrates the necessity of reviewing remains of Butleria and integrating information from both Old World schizotheriines and Early Miocene chalicotheriines from Asia. In de Bonis et al. (1995), this node was supported by characters #1 (5), #3 (only in trees 0 and 2) and #27 (37). Character #3 is not retained in the present analysis. Characters #1 and #27 are recoded from a three-state to a binary coding with suppression of the intermediate state (see Appendix 2). Monophyly of the Anisodon clade (Fig. 10, node D) is supported by 16 unambiguous synapomorphies (14 being nonhomoplastic): convex upper tooth row in lateral view (1; RI = 1); lacrimal tubercle present (4; RI = 1); shortened maxilla (5; RI = 1); presence of a groove between the ventral border of the zygomatic arch and the M3 (7; RI = 1); postorbital process of the jugal present (8; RI = 1); short zygomatic arch (10; RI = 1); swollen braincase (13; RI = 1); laterally orientated paroccipital process (15; RI = 1); vertical occiput (18; RI = 1); upper tooth rows converging anteriorly (21; RI = 1); no retromolar space on the maxilla (22; RI = 0.75); anterior wall of the tympanic bulla extending anterior to the postglenoid process (27; RI = 1); angulus mandibulae quite expanded ventrally (31; RI = 1); metacone more labial than the paracone on M 2 (44; RI = 1); external wall of the metacone and metastyle subparallel to the anteroposterior lengthening on M3 (45; RI = 1); weak ‘metastylid’ on lower molars (51; RI = 1). Characters 4, 7, 18, 21, and 44 are new. In de Bonis et al. (1995), this node was supported by characters #0, #1, #2, #4, #6, #7, #9, #15, #22, #28, and characters #19 (trees 0 and 1), #26 (trees 0 and 2), and #14 (trees 1 and 2). Seven of these characters are also found in the present analysis, i.e. #1 (5), #4 (22), #6 (10), #9 (13), #22 (1), #19 (15), and #26 (31). The absence of character #0 (12) is the result of a coding mistake in de Bonis et al. (1995) concerning AMNH 26518. Character #2 is not retained in the present study. Character #7 (11) has an ambiguous distribution here, mainly because of the state

observed in Eomoropus. Character #15 (26) is recoded from a three-state to a binary coding. Character #14 (16) is completely reviewed here (Appendix 2). Monophyly of node E (A. macedonicus, Vathylakkos, N. sivalense, Titov Veles, ‘C.’ wuduensis) (Fig. 10) is supported by two nonhomoplastic, unambiguous synapomorphies: elongated symphysis reaching the level of P3–P4 boundary (33; RI = 1); distal cingular crest reaching the hypocone summit on M 3 (47; RI = 1). Neither of these is found in de Bonis et al.’s (1995) phylogeny. In the previous study, the clade (A. macedonicus, Vathylakkos) was supported by characters #29 (39), #30 (42), and #31 (41) (only the third in tree 0). The latter has an ambiguous distribution here, as the derived state (i.e. state 1) is only present in A. macedonicus and the specimen from Vathylakkos. The first two characters are recoded from a threestate to a binary coding and now support node B only (see above). In de Bonis et al. (1995), relationships between ‘M.’ brevirostris and Chalicotherium are supported by one tree among the three MPTs (i.e. tree 2). This clade was supported by character #21 (19). This is also the case in the present study. Moreover, the new character 46 also supports this clade. Both characters (19 and 46) are nonhomoplastic (RI = 1) and unambiguous. CCECM Lgr 1065 and C. goldfussi did not form a clade in de Bonis et al. (1995), mainly because of the inclusion of the problematic Henndorf mandible (HMS unknown number) within C. goldfussi (see discussion above). Here, the monophyly of the clade (C. goldfussi, Lgr 1065) is supported by the homoplastic, unambiguous character 38 (RI = 0.667).

EVOLUTION

OF THE SYMPHYSIS AND LOWER INCISORS

According to de Bonis et al. (1995: 162), ‘la réduction et la diminution du nombre de ces dents (. . . incisives inférieures) semblent donc être des tendances propres à toute la sous-famille des Chalicotheriinae’ [i.e. the reduction and diminution in the number of these teeth (. . . lower incisors) seem to be a trend in all Chalicotheriinae]. Indeed, their results proposed a convergence between derived anisodonts like A. macedonicus (with two very reduced lower incisors) and members of the clade Chalicotherium (with the loss of lower incisors). The loss of lower incisors was optimized in C. goldfussi and CCECM Lgr 1065. This hypothesis is refuted here: in reality, C. goldfussi had three well-developed lower incisors (see description of MHNT VAL-4). On the other hand, de Bonis et al. (1995: 162) proposed that the reduction then loss of the lower incisors was followed by a lengthening of the posterior extent of the symphysis. According to their results, acquisi-


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE tion of a symphysis extending to the P 3–P4 boundary is also convergent between the two above-mentioned lineages. This conclusion is again a result of the inclusion of the Henndorf mandible (HMS unknown number) within C. goldfussi (Schaefer & Zapfe, 1971: fig. 3a, b; Zapfe, 1979: fig. 40), which blurred the distinction between Anisodon and Chalicotherium. However, comparison with the mandible from Saint-Gaudens (MHNT VAL-3) demonstrates that the specimen from Henndorf can no longer be assigned to this genus. Thus, the new remains from Saint-Gaudens, coupled with the first accurate description of cranial remains from Sansan, places N. sivalense, the Titov Veles form and ‘C.’ wuduensis (all of which possess an extended symphysis and lack lower incisors) within the Anisodon clade. However, the distribution of character 35 (number of lower incisors) is ambiguous in the present study. With ACCTRAN optimization, the loss of lower incisors is optimized in A. grande and the specimen from Vathylakkos. With DELTRAN optimization, the loss is only optimized in the Vathylakkos specimen. In both cases there is a reversal (states 0–1) in A. macedonicus. If the symphysis described by de Blainville (1849) actually exists (see above), A. grande has lower incisors (three according to de Blainville, but perhaps only two if our interpretation is correct) and the first optimization would be contradicted. However, we guess that none of these two scenarios (ACCTRAN and DELTRAN) is correct. Our (speculative) point of view is that UT DKO 234 and the specimen from Vathylakkos probably form a clade, as they share two synapomorphies: subcentral position of the protocone on molars [41 (1)]; and presence of a notch on the mesial cingulum of upper molars [43 (1)]. Moreover, we surmise that the terminal taxa Vathylakkos and A. macedonicus are probably the same species, or at least very close species. If true, the Vathylakkos specimen would probably be optimized to have two incisors, and lower incisors would be lost only in the Titov Veles form, A. wuduensis, and A. sivalense. More data are necessary to improve our understanding of this part of the tree.

WHEN

IS A CLADE A GENUS?

Some authors could use the phylogeny presented here (Fig. 10) to propose another classification. Obviously, a single cladogram can yield several different phylogenetic classifications (e.g. Tassy, 1988). The eternal problem between taxonomic ‘splitters’ and ‘lumpers’, so well summarized by Simpson (1945: 23), has not been solved by Hennigian principles. The choice between subordination or sequencing (Nelson, 1972), the use of plesions for fossils following Patterson & Rosen (1977), the abandonment of nomenclatural cat-

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egories following Hennig (1969), or the emergence of the PhyloCode (Cantino & de Queiroz, 2004), all witness attempts to adapt classifications to phylogenetic methods. Plurality exists; consensus does not, even among cladistic classifications. A major part of this problem doubtless arises from the accommodation of ranks within a cladogram. Indeed, ranks are subjective categories and there are no rules to govern whether a given clade would be either a genus or a family. Adoption of ranks (and consequently of names) above the species level is almost always an arbitrary choice. But, the case of the genus is probably one of the most problematic as it has a great impact on species designation. Concerning the present cladogram, some might consider the Chalicotheriinae as a single genus, Chalicotherium, whereas others might choose to conserve the subfamily divided into two genera expressing the basal sister groups, Butleria and Chalicotherium. Yet, the salient result of the present study is the existence of two main clades during the Middle and Late Miocene. The divergence between the two clades can be seen in the 18 nonambiguous derived characters that separate them (16 for Anisodon and two for Chalicotherium). Hence, chalicotheriine diversity is somehow reflected by the proposed classification. Nevertheless, systematists’ preferences regarding rank choices are trivial: what is important is the pattern of relationships.

NOTE

CONCERNING THE

KALIMANTSI

CHALICOTHERE

The most recent contribution to chalicotheriine knowledge is an important new skull (MPA K631) recovered from the Late Miocene (MN 11–12?) of the Kalimantsi Formation (Bulgaria), which was described as a new genus, Kalimantsia, by Geraads et al. (2001). However, there are numerous problems with both the skull description and the authors’ hypotheses concerning chalicotheriine phylogeny. These problems are discussed below.

Cranium The skull of Kalimantsia shows several plesiomorphic features that suggest it should be excluded from the clade Anisodon (as defined here). These are: (1) subparallel upper tooth rows, (2) no lacrimal tubercle, (3) long zygomatic arch, (4) no groove between the anterior part of the zygomatic arch and M3, (5) upper retromolar space present, (6) choanal opening situated far behind M3, (7) metacone as lingual as the paracone on M2, (8) external wall of the metacone and metastyle subperpendicular to the mesiodistal lengthening. On the other hand, the only two characters supporting the clade Chalicotherium in the present study are not observable on MPA K631: the occipital condyles


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(character 19) are missing and the degree of development of the postfossette of M3 (character 46) is not clearly observable in Geraads et al.’s (2001) illustrations. Resulting phylogeny including Kalimantsia Only 15 characters among our set of 51 could be coded on the basis of the description provided by Geraads et al. (2001). A search using the Branch-and-Bound exact algorithm generated 98 MPTs with a length of 62 steps (CI = 0.871; CIeuc = 0.840; RI = 0.877). The strict consensus tree (68 steps; CI = 0.794; CIeuc = 0.750; RI = 0.785) reflects the impact of the aforementioned plesiomorphic characters on the position of Kalimantsia (Fig. 11). As we did not have the opportunity to study MPA K631, we leave the question open. For the moment, the only peculiarities (see discussion below) of this specimen are: P3 and P4 differing more in length than in width and having a lingual cingulum slightly thicker than in CCECM Lgr 1065 and the specimen from Saint-Gaudens, and molars more elongated than those of other chalicotheriines. Given the current state of knowledge, these peculiarities could either be autapomorphic or plesiomorphic. Discussion Geraads et al. (2001) criticized the phylogenetic contribution of de Bonis et al. (1995) in several respects. The hypotheses of relationships proposed by Geraads

Figure 11. Phylogeny after the inclusion of Kalimantsia Geraads et al., 2001 in the analysis. Strict consensus of the 98 most parsimonious trees (MPTs: 62 steps; CI = 0.871; RI = 0.877) (strict consensus: 68 steps; CI = 0.794; RI = 0.785).

et al. are quite different from those of previous authors and from those presented here. According to Geraads et al. (2001: 601), Kalimantsia differs from Chalicotherium for the following reasons: (1) P3 and P4 differ more in length than in width (Geraads et al., 2001: fig. 4); (2) their lingual cingulum is thicker; (3) the molars are elongated. In fact, as we state above, MPA K631 differs from all other known chalicotheriines with respect to these features. Also, according to these authors, Kalimantsia cannot be included in Anisodon (which corresponds only to A. grande for them) because in the latter (Geraads et al., 2001: 600): ‘(1) the occipital is low and broad; (2) the temporal fossa is short, at least at Sansan (it is unknown at Neudorf); (3) the sagittal crest is short or absent (a difference of doubtful taxonomic significance); (4) the face is long, the muzzle is deep, with the tooth-row diverging forwards from the upper skull profile; (5) the lingual cingulum of P 3 and P4 is reduced; the latter tooth is much broader than the former, although not much longer; (6) molars have a long protoloph; (7) molars are wide in respect to their length’. We agree with reasons (2), (5), and (7); the first is character 10 of our analysis, the last two correspond to the aforementioned peculiarities of MPA K631 (see above). Reason (1) is weakened by the fact that on MNHN Sa 15670 the lower part of the occipital face (including the occipital condyles) is missing. Reason (3), as stated, is doubtful; the presence of the sagittal crest is probably due to sexual dimorphism (Coombs, 1975). Finally, based on the new description of MNHN Sa 15670, reasons (4) and (6) appear unjustified (Figs 3, 9A). Geraads et al. (2001: 601) argued that almost all the features in which Anisodon differs from Kalimantsia are also found in AMNH 26518, except that the temporal fossa [reason (2)] of AMNH 26518 is long. We agree partly with reason (4); AMNH 26518 does have a long snout, but Anisodon grande surely does not. Reasons (5) and (7) again correspond to peculiarities of MPA K631. Reason (6) is incorrect (Colbert, 1934; J. Anquetin, pers. observ. on the cast). Finally, features (1) and (3) are not observable on AMNH 26518. According to Geraads et al. (2001: 601), Kalimantsia shares several similarities with A. macedonicus from Dytiko 3 (UT DKO 234): ‘the muzzle is shortened (more so at Dytiko 3), the orbit is square, the cerebral skull is rather long (more so at Kalimantsi), there is a sagittal crest, the protoloph is short on the molars, P4 is not much broader than P3’. Although the snout of MPA K631 seems shortened, it is quite difficult to determine its true shape as it is very damaged. Uniting UT DKO 234 and MPA K631 on the basis of a long cerebral skull is surprising with respect to other chalicotheriine skulls (e.g. CCECM Lgr 1065, see Fig. 6). The protoloph is short on the molars of both specimens, but this is also the case on those of all other


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE chalicotheriines (except in A. wuduensis and the specimen from Titov Veles, where this feature cannot be observed as both are known only from the mandible). The observation that ‘P4 is not much broader than P3’ in UT DKO 234 and MPA K631 (Geraads et al., 2001: 601) is unclear because it is contradicted by the authors’ own data (Geraads et al., 2001: fig. 4). According to Geraads et al. (2001), an Anisodon clade containing the species from Sansan and Dytiko 3 can no longer stand. They proposed five arguments to support this conclusion (Geraads et al., 2001: 601): ‘(1) the cerebral skull is in fact much lower in A. grande of Sansan (MNHN) and Neudorf (Zapfe, 1979: fig. 33) than at Dytiko 3; (2) the face is in fact very long in A. grande, both at Sansan (MNHN) and Neudorf (Zapfe, 1979: fig. 34); (3) the nasal is unknown at Neudorf, and not distinct from the frontal on the Sansan skull, but most unlikely to have been much reduced; (4) the fragment of zygomatic arch figured by Blainville (1849) is now missing, but it is low at Neudorf (Zapfe, 1979: fig. 34); and (5) the braincase is in fact quite narrow at Sansan’. Obviously, argument (3) is rather unsound. Argument (4) should be considered with prudence as the taxonomic assignment of part of the Neudorf remains is doubtful (see above). Argument (2) is erroneous: following the present study, one cannot argue that the face is ‘very long’ in MNHN Sa 15670 (Fig. 3). Finally, arguments (1) and (5) remain, but these only support the hypothesis that the cranial morphology of A. grande (MN 6) is less derived than that of A. macedonicus (MN 13) for the concerned features. Apart from the three initially cited characters concerning P3, P4, and the molar length, which are specific for MPA K631, no other features help to solve its relationships. The foregoing discussion shows that almost all of the features used by Geraads et al. (2001) to support their views are plesiomorphic. Geraads et al. (2001: 601) ‘do not believe that the relationships of Miocene Chalicotheriinae can be satisfactorily solved by phylogenetic parsimonious analysis at the present time, because too few reasonably complete skulls are known [. . .]’. Yet, they did recognize two groups within Miocene chalicotheriines: one associating Anisodon from Sansan and Neudorf, and perhaps C. brevirostris; and one including Kalimantsia, A. macedonicum, and perhaps C. goldfussi. The present study, based on shared derived characters, disagrees with these propositions: as far as the 15 characters we were able to code are meaningful, Figure 11 shows a basal position for Kalimantsia. MPA K631 needs to be reviewed and included in a phylogenetic analysis. Only after that can its phylogenetic relationships be inferred and an appropriate name applied. Indeed, within the frame of a classification reflecting the phylogeny, the establishment of a

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new genus should be based on a phylogenetic study of its characters.

CONCLUSION Despite their great historical importance, chalicotheriine remains from France have never been accurately described. Here, a detailed study of the most complete cranial remains is given alongside a description of new material from Saint-Gaudens. Both considerably increase knowledge of chalicotheriine cranial anatomy. Based on these new data, a cladistic analysis was performed which supports the hypothesis that there are two main chalicotheriine lineages during the Middle and Late Miocene. These two lineages display divergent evolutionary trends. Anisodon has a strongly derived cranial anatomy, whereas Chalicotherium retains numerous plesiomorphic traits. Another result of this study is the novel proposed pattern of relationships. Previous authors suggested that N. sivalense, ‘C.’ wuduensis and the specimen from Titov Veles were close to C. goldfussi. The present analysis shows that they belong to the Anisodon lineage, and that consequently a generic distinction for N. sivalense is unnecessary [note that this was also implied by de Bonis et al.’s (1995) results]. The phylogeny of Chalicotheriinae remains labile and we hope this preliminary study will be a first step followed by many others. First, it would be useful to review available cranial remains from the Early Miocene (especially Butleria and Asian forms), and those from Titov Veles and Kalimantsi. The present study is based only on cranio-mandibular and dental characters, so the inclusion of postcranial remains will be required to improve future studies. It is also crucial to extend the analysis to include schizotheriines and eomoropids, as both are known from cranial and postcranial remains. Schizotheriine remains are generally more common than those of chalicotheriines, and relationships of Eurasiatic schizotheriines are little studied. For the moment, our knowledge of chalicothere phylogeny is mostly speculative. This study demonstrates the progress that can be made by coupling detailed, comparative anatomical study with critical cladistic analysis. Not only is the phylogeny of chalicotheriines poorly known, but so is their dental anatomy, even though it has been the basis for most of the published taxonomic studies since the 19th century. Teeth from La Grive Saint-Alban (MN 7–8) described by Depéret (1892) as deciduous teeth of M. grande race rhodanicum in fact belong to a schizotheriine [referred to Metaschizotherium fraasi by von Koenigswald (1932), although this is unlikely (M.C. Coombs, pers. comm., 2006)]. We also identified a schizotheriine in Le Fousseret (MN 7),


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Haute-Garonne, France (unprepared, unnumbered upper and lower teeth from Noulet’s collection at the MHNT). Thus, there were three different, contemporaneous chalicothere lineages (i.e. Anisodon, Chalicotherium, and a schizotheriine) during the Middle and Late Miocene, at least in France. This diversity was unexpected.

ACKNOWLEDGEMENTS This paper presents the results of JA’s Master thesis (MSc) at the MNHN, which was supervised by PT and POA. We thank: Francis Duranthon, Pierre Dalous, and Guillaume Fleury, curators at the MHNT, who allowed us to study remains from Saint-Gaudens and Noulet’s collection; Didier Berthet (CCECM) for access to specimens from La Grive Saint-Alban; and Jerry J. Hooker (BMNH) for access to specimens of A. sivalense (‘Nestoritherium’). Louis de Bonis (University of Poitiers, France) was especially helpful in providing casts of UT DKO 234 (A. macedonicus) and AMNH 26518 (C. brevirostris). Moreover, the donation of these casts to the MNHN, Paris, was deeply appreciated by PT. We also thank Margery C. Coombs (University of Massachusetts, MA, USA) for providing useful information regarding ‘Nestoritherium’ and schizotheriines, and for reviewing the manuscript. Claire Sagne’s (MNHN) goodwill fruitfully helped JA in Paris. Paul M. Barrett and Jerry J. Hooker (BMNH) reviewed an earlier version of the manuscript and their suggestions were very valuable. Philippe Loubry (MNHN) made the photographs of MNHN Sa 9339, 15670, and 15671 (Figs 1, 3, 4B, respectively). Drawings and other photographs were made by JA. The first author also deeply thanks Guillaume Billet (MNHN) for really enriching discussions and for his precious help with nomenclatural questions.

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de Blainville HMD. 1849. Ostéographie ou description iconographique comparée du squelette et du système dentaire des Mammifères récents et fossiles – Genus Anoplotherium. 4 BB, Paris: J.B. Baillère, 66–70. de Blainville HMD. 1855. Ostéographie ou description iconographique comparée du squelette et du système dentaire des Mammifères récents et fossiles – Genus Macrotherium. 4 EE, Paris: J.B. Baillère, 37–38. Bohlin B. 1936. Notes on some remains of fossil mammals from China and Mongolia. Bulletin of the Geological Society of China 15: 321–330. de Bonis L, Bouvrain G, Koufos G, Tassy P. 1995. Un crâne de chalicothère (Mammalia, Perissodactyla) du Miocène supérieur de Macédoine (Grèce): remarques sur la phylogénie des Chalicotheriinae. Palaeovertebrata 24: 135–176. Bremer K. 1994. Branch support and tree stability. Cladistics 10: 295–304. Butler PM. 1965. East African Miocene and Pleistocene chalicotheres. Bulletin of the British Museum (of Natural History) 10: 163–237. Cantino PD, de Queiroz K. 2004. Phylocode: a phylogenetic code of biological nomenclature, Version 2b. Available at http://www.ohiou.edu/phylocode/preface.html Colbert EH. 1934. Chalicotheres from Mongolia and China in the American Museum. Bulletin of the American Museum of Natural History 67: 353–387. Colbert EH. 1935. The proper use of the generic name Nestoritherium. Journal of Mammalogy 16: 233–234. Coombs MC. 1975. Sexual dimorphism in chalicotheres (Mammalia, Perissodactyla). Systematic Zoology 24: 55–62. Coombs MC. 1978. Reevaluation of early Miocene North American Moropus (Perissodactyla, Chalicotheriidae, Schizotheriinae). Bulletin of the Carnegie Museum of Natural History 4: 1–62. Coombs MC. 1989. Interrelationships and diversity in the Chalicotheriidae. In: Prothero DR, Schoch RM, eds. The evolution of perissodactyls. New York: Oxford University Press, 438–457. Cope ED. 1881. The systematic arrangement of the order Perissodactyla. Proceedings of the American Philosophical Society 19: 377–403. Cope ED. 1889. The Vertebrata of the Swift Current River, II. American Naturalist 23: 151–155. Cuvier G. 1804. Des animaux qui diffèrent du Palaeotherium pour le genre, mais qui sont de même ordre, et particulièrement du genre Anoplotherium, et de ses espèces. Annales du Muséum National d’Histoire Naturelle, Paris 3: 370–382. Cuvier G. 1822. Recherches sur les Ossemens Fossiles, 2nd edn, Vol. 5. Paris. Depéret C. 1892. La faune de mammifères miocènes de La Grive-Saint-Alban (Isère) et quelques autres localités du bassin du Rhône. Archives du Muséum d’Histoire Naturelle de Lyon 5: 1–93. Falconer H. 1868. On Chalicotherium sivalense. In: Murchison CA, ed. Palaeontological memoirs and notes of the late hugh falconer. London: Robert Hardwicke, 208–226. Falconer H, Cautley. 1843. On some fossil remains of Anoplotherium and giraffe, from the Siwalik Hills, in the North


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE of India. Proceedings of the Geological Society of London 4: 235–249. Filhol H. 1890. Études sur les mammifères fossiles de Sansan. Bibliothèque de l’École des Hautes Études, section Sciences Naturelles 37: 1–319. Garevski R, Zapfe H. 1983. Weitere Chalicotheriiden-Funde aus der Pikermi Fauna von Titov Veles (Mazedonien, Jugoslawien). Acta Musei Macedonici Scientiarum Naturalium 17: 1–20. Geraads D, Spassov N, Kovachev D. 2001. New Chalicotheriidae (Mammalia) from Bulgaria. Journal of Vertebrate Paleontology 21: 596–606. Gervais FLP. 1850. Zoologie et Paléontologie française (animaux vertébrés), ou nouvelles recherches sur les animaux fossiles de la France, Vol. 2. Paris. Gill TN. 1872. Arrangement of the families of mammals with analytical tables. Smithsonian Miscellaneous Collections 11: 1–98. Harlé É. 1898. Une mâchoire de dryopithèque. Bulletin de la Société Géologique de France 26: 377–383. Hennig W. 1969. Dies Stammesgeschichte der Insecten. Frankfurt: W. Kramer, Senckenberg-Buch n°49. Holland WJ, Peterson OA. 1914. The osteology of the Chalicotherioidea with special reference to a mounted skeleton of Moropus elatus Marsh, now installed with Carnegie Museum. Memoirs of the Carnegie Museum 3: 189–406. Hooker JJ. 1994. The beginning of the equoid radiation. Zoological Journal of the Linnean Society 112: 29–63. Hooker JJ, Dashzeveg D. 2004. The origin of chalicotheres (Perissodactyla, Mammalia). Palaeontology 47: 1363–1386. International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, 4th edn. London: The International Trust for Zoological Nomenclature (Natural History Museum). Kaup JJ. 1833a. Description d’ossements fossiles de Mammifères inconnus jusqu’à-présent, qui se trouvent au Muséum grand-ducal de Darmstadt. Second Cahier. Darmstadt: J. G. Heyer. Kaup JJ. 1833b. Der Krallen-Phalanx von Eppelsheim, nach welchem Herr von Cuvier seinen Riesen-Pangolin, Manis gigantea, aufstellte, gehört zu Dinotherium. Neues Jahrbuch für Mineralogie, Geologie und Palaontologie (1833): 172–176. Kaup JJ. 1859. Beitraege zur naeheren kenntniss der Urweltlichen Saeugethiere. Viertes Heft. Darmstadt: Eduard Zernin. von Koenigswald GHR. 1932. Metaschizotherium fraasi, n. g. n. sp., ein neuer Chalicotheriide aus dem Obermiocän von Steinheim a. Albuch. Palaeontographica 8: 1–24. Lartet É. 1837a. Note sur les ossements fossiles des terrains tertiaires de Simorre, de Sansan, etc., dans le département du Gers, et sur la découverte récente d’une mâchoire de singe fossile. Comptes Rendus de l’Académie des Sciences 4: 85–93. Lartet É. 1837b. In: de Blainville HMD. Rapport sur un nouvel envoi de fossiles provenant du dépôt de Sansan. Comptes Rendus de l’Académie des Sciences 5: 417–427. Lartet É. 1839. Notice géologique. Extrait de l’Annuaire du Département du Gers, année 1839. Lartet É. 1851. Notice sur la colline de Sansan. Extrait de l’Annuaire du Département du Gers, année 1851. Auch: J.-A. Portes.

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0 0 0 ? 0 1 1 0 0 0 0 0 ?

CHARACTER

1 1 1 1 1 1 1 1 ? 1 ? 0 1 1 1 0 ? 0 0 0 0 0 0 1 0 ? 1 1 0 0 0 0 ? 0 ? ? ? ? ? 0 0 1 0 ? ? ? ? ? ? ? ? ? ? 1 0 1 0 1 ? ? ? ? ? ? 1 – = nonapplicable.

Eomoropus Moropus A. macedonicus ‘M.’ brevirostris A. grande Lgr 1065 C. goldfussi Vathylakkos ‘C.’ wuduensis Butleria Titov Veles Nestoritherium Kalimantsia

? 0 1 0 1 0 0 1 ? 0 ? 1 ?

? 0 2 1 ? 1 ? ? ? 0 ? ? ?

0 1 0 0 ? ? ? ? ? ? ? ? ?

0 0 1 0 1 ? ? ? ? ? ? ? 0

? 0 1 0 1 0 ? 1 ? 0 ? ? ?

1 1 0 ? ? 1 ? 0 ? ? ? ? ?

? 0 1 0 1 0 0 ? ? ? ? 1 0

? 0 1 0 1 ? ? ? ? ? ? ? ?

? 0 1 0 ? ? ? ? ? ? ? ? ?

10

1 0 1 0 1 ? ? ? ? ? ? ? ?

1 0 1 ? 1 1 ? ? ? ? ? ? 1

0 0 1 ? 1 0 ? 1 ? ? ? ? ?

0 0 1 1 ? 1 ? ? ? ? ? ? ?

? 0 1 0 1 0 ? ? ? ? ? ? ?

1 0 1 1 1 0 ? ? ? ? ? ? ?

1 1 0 1 1 1 ? ? ? ? ? ? ?

1 1 0 ? 0 1 ? ? ? ? ? ? ?

1 1 1 0 1 0 ? ? ? ? ? ? ?

20

? 1 0 1 0 1 ? 0 ? 1 ? ? 1

0 1 0 1 0 1 1 0 ? 1 ? 0 1

? 0 1 0 ? 0 ? 1 ? ? ? ? 0

1 1 1 1 0 ? ? 1 ? ? ? ? ?

MATRIX

APPENDIX 1

1 0 1 1 1 1 ? ? ? ? ? ? ?

1 1 0 1 0 1 ? ? ? ? ? ? ?

0 0 0 1 0 0 ? ? ? ? ? ? ?

? 0 2 2 1 1 ? ? ? ? ? ? ?

30

1 1 0 ? 0 1 1 0 ? ? ? ? ?

0 1 0 ? 0 1 1 0 0 ? 0 ? ?

0 0 1 ? 0 0 ? 1 1 0 1 1 ?

? ? 1 ? ? 0 ? ? 0 0 1 0 ?

2 2 1 ? ? 2 ? ? 0 2 0 0 ?

1 0 1 ? ? ? 1 ? 1 1 1 1 ?

0 0 1 0 ? 0 ? ? ? 0 ? 1 ?

0 0 1 1 ? 0 0 1 ? 1 ? 1 1

0 0 1 1 1 1 1 1 ? 1 ? 1 1

40

0 0 1 0 0 0 0 1 ? 0 ? 0 0

0 0 1 1 1 1 1 1 ? 1 ? 1 1

0 0 1 0 0 0 0 1 ? ? ? 0 0

– 1 0 1 0 1 1 0 ? ? ? – 1

– 1 0 1 0 1 1 ? ? 1 ? 0 1

1 1 1 0 1 0 0 ? ? 1 ? 1 ?

1 ? 0 1 1 1 1 ? ? ? ? 0 ?

1 1 1 ? 1 1 ? ? 1 1 0 1 ?

1 1 1 ? 0 1 1 ? 1 1 1 1 ?

50

1 1 0 ? 0 1 ? 0 0 1 0 ? ?

APPENDIX 2 LISTING AND COMMENTS

Cranium 1. Lateral outline of the upper tooth row: straight (0); convex (1). This corresponds partly to character #22 of de Bonis et al. (1995), which separated straight tooth rows from those that are externally convex. It is reformulated here in order to agree with our observations (see character 21). 2. Upper border of the nasal opening: above P2–P3 (0); P3–P4 (1); P4–M1 (2). New character. This character is treated as ordered. 3. Nasal-lacrimal contact: absent (0); present (1). New character. In the limited context of the present study (focused on chalicotheriines), it only supports the Moropus branch. 4. Lacrimal tubercle: absent (0); present (1). New character. 5. Maxilla: elongated (0); shortened (1). This corresponds to character #1 of de Bonis et al. (1995), which was coded in three states: elongated, short, and very short. 6. Postorbital process of the frontal: absent (0); present (1). New character. De Bonis et al. (1995) described a postorbital process perforated by a large supraorbital foramen in UT DKO 234. Such a process is usually described in all other Chalicotheriidae. In fact, there are two distinct processes on the frontal (see description of CCECM Lgr 1065 in the text). Behind the orbit there is a true postorbital process, and above the orbit there is a flattened supraorbital process perforated by the supraorbital foramen. These observations are confirmed in Eomoropus and Moropus elatus. These two processes can either be separated (as in CCECM Lgr 1065), or almost fused (as in Moropus elatus). 7. Groove between the ventral border of the zygomatic arch and M3: absent (0); present (1). New character. Although observed by de Bonis et al. (1995), this character was not retained in their analysis. State 1 corresponds to a real wide and deep groove, not to a slightly curved area. 8. Postorbital process of the jugal: absent (0); present (1). This corresponds to character #5 of de Bonis et al. (1995), which was coded in three states: absent, weak, and strong. Such a distinction is abandoned. In the analysis of de Bonis et al. (1995), the only terminal taxon with state ‘weak’ is CCECM Lgr 1065. Determining the development or absence of this process in this specimen is difficult (see above). We code only the presence or absence of this process; CCECM Lgr 1065 is coded as missing data. 9. Orientation of the jugal-squamosal suture with respect to the horizontal plane: parallel (0); oblique (1). This nearly corresponds to character #8 of de Bonis


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE et al. (1995). The coding of these authors was: vertical or oblique. No vertical suture was observed by us. 10. Lengthening of the zygomatic arch: short (0); long (1). This is character #6 of de Bonis et al. (1995) used without modifications, although the coding was reversed [to have a homogeneous strategy of coding throughout analysis, the coding was reversed for some characters issued from de Bonis et al. (1995)]. 11. Posterior part of the zygomatic arch: reaching the same level as the upper border of the orbit (0); always lower than the level of the upper border of the orbit (1). This corresponds to character #7 of de Bonis et al. (1995), but the coding was reversed. 12. Braincase: low (0); high (1). This is character #0 of de Bonis et al. (1995). These authors coded ‘low’ for Eomoropus, Moropus, AMNH 26518 and CCECM Lgr 1065. In fact, only Moropus has a low braincase. Moreover, coding AMNH 26518 is impossible as nearly all of the skull roof is missing. 13. Transverse section of the braincase: tent-shaped (0); swollen (1). This corresponds to character #9 of de Bonis et al. (1995). 14. Occipital condyles in lateral view: projected posteriorly to the skull (0); not projected behind the lower part of the occiput (1). New character. De Blainville (1849) and Filhol (1890) described projected condyles in MNHN Sa 15670, but as the condyles are now destroyed we coded Anisodon grande as missing data. This character has no relation with character 18. 15. Orientation of the paroccipital process: ventral (0), ventral and lateral (1). This is character #19 of de Bonis et al. (1995). 16. Furrow separating the postglenoid process from the external auditory meatus: wide (0); thin (1). This corresponds to character #14 of de Bonis et al. (1995), but was recoded. There is a furrow between the postglenoid process and the meatus in every specimen. This furrow is wide only in Moropus elatus and CCECM Lgr 1065 [according to de Bonis et al. (1995), there is a furrow only in Anisodon macedonicus and Anisodon grande; this is not verified by our observation). 17. Ventral border of the external auditory meatus: smooth (0); displaying a tubercule (1). New character. State 0 is only present in UT DKO 234. 18. Occiput: vertical (0); with the upper part projected posteriorly (1). New character. 19. Occipital condyle: elongated vertically (0); not high and almost rounded (1). This is character #21 of de Bonis et al. (1995), but the coding was reversed. 20. Pterygoid lacuna: absent (0); present (1). This is character #10 of de Bonis et al. (1995), but the coding was reversed. A lacuna is only present in UT DKO 234. 21. Upper tooth rows: converging anteriorly (0); subparallel (1). This character was mentioned by de Bonis

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et al. (1995) but was not used in their analysis (or in part with character #22). 22. Retromolar space on the maxilla: absent (0); present (1). This is character #4 of de Bonis et al. (1995). 23. Choanal opening: far behind M3 (0); at the level of M3 (1). New character. 24. Canalis alisphenoideus: absent (0); present (1). This is character #12 of de Bonis et al. (1995), but the coding was reversed. 25. Basioccipital: with a medial crest (0); smooth (1). This is character #17 of de Bonis et al. (1995). Each occipital condyle gives rise to a low crest anteriorly on the basioccipital. These crests can either fuse medially and form a single medial crest (state 0), or continue anteriorly in parallel and progressively disappear (state 1). 26. Shape of the tympanic bulla: oval (0); elbowshaped (1). This corresponds to character #15 of de Bonis et al. (1995), which was coded in four states: globular (0), anteroposteriorly elongated (1), narrow anteriorly (2), and very narrow anteriorly (3). These four states obviously code two distinct types of observation: the general outline of the bulla (states 0 and 1), and the anterior width of the bulla (states 2 and 3). It is true that there is a width variation of the anterior part of the bulla. But given the current state of knowledge, it seems better to code a binary character for the shape of the bulla, without coding its anterior width. 27. Anterior wall of the tympanic bulla: extending anterior to the postglenoid process (0); at the level of or a little behind the postglenoid process (1). This is character #13 of de Bonis et al. (1995). 28. Stylo-hyoid fossa: at the same level as the external auditory meatus (0); displaced posteriorly with respect to the meatus level (1). New character. In both cases, the fossa perforates the ventral wall of the bulla. The relative position of the fossa is stable (state 0) in both long- and short-snouted chalicotheriines, except in AMNH 26518. 29. Foramen caroticum: between the basioccipital and tympanic bulla (0); perforating the posterior margin of the bulla (1); posterior to the bulla (2). This corresponds to character #18 of de Bonis et al. (1995). States have been recoded according to the new data presented here. This character is treated as ordered. Note that in UT DKO 234 the foramen caroticum is fused to the foramen metoticum (de Bonis et al., 1995), whereas in AMNH 26518 these foramina are close but still separated. In the present analysis both specimens are coded as state 2, but may represent two nonhomologous states. Mandible 30. Height of the mandibular corpus: increasing posteriorly (0); constant (1). This is character #25 of de Bonis et al. (1995), but the coding was reversed.


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31. Angulus mandibulae: quite expanded ventrally (0); not expanded (1). It corresponds to character #26 of de Bonis et al. (1995), which was coded in three states: angular inflection absent (0), present (1), and accentuated (2). See description of CCECM Lgr 1065 for our interpretation of the angular area. 32. Retromolar space on the mandible: absent (0); present (1). New character. One can argue that it is redundant with character 22, but state distributions are not the same, so we decided to keep the two characters. Here, deletion of one of them does not change the resulting phylogeny. Future authors will determine whether these two are redundant or not. 33. Posterior extent of the symphysis: at the level of P2 (0); at the level of the boundary of P 3–P4 (1). This is character #24 of de Bonis et al. (1995). 34. Tuberosity under the symphysis: absent (0); present (1). New character. Such a tuberosity is observed in both UT DKO 234 and the specimen from Titov Veles. 35. Lower incisors: absent (0); two (1); three (2). This corresponds to character #37 of de Bonis et al. (1995), but for these authors lower incisors are either absent or present (binary character). This character is treated as ordered. 36. Lower canine: absent (0); present (1). New character.

Upper dentition 37. P 2: longer than wide (0); wider than long (1). This corresponds to character #27 of de Bonis et al. (1995). A binary coding is preferred to the states ‘shortened’ (1) and ‘very short’ (2) proposed by these authors. 38. P 3: protoloph reaching the protocone (0); protoloph stopping at the paraconule (1). This is character #28 of de Bonis et al. (1995). The observations of these authors are incorrect [the protoloph of P3 is neither complete in AMNH 26518 nor in Butleria (Butler, 1965: 182; fig. 3)]. 39. P 4: protoloph reaching the protocone (0); protoloph stopping at the paraconule (1). This character is not redundant with the previous (see matrix). It is character #29 of de Bonis et al. (1995). These authors incorrectly coded a complete protoloph (state 0) in Butleria (Butler, 1965: 182; fig. 3) and in Anisodon grande. So, state 1 is no longer a parallelism between (Anisodon macedonicus, Vathylakkos) and the Chalicotherium clade as proposed by de Bonis et al.’s results, but appears to be a synapomorphy of Chalicotheriinae (see Appendix 3). 40. Paraconule on molars: absent (0); present (1). This is a subdivision (see character 42) of character #30 of de Bonis et al. (1995). 41. Position of the protocone on molars: slightly posterior to the paracone and paraconule (0); subcentral

with respect to the lingual side of the tooth (1). Among chalicotheres, the protocone is never at the same level as the paracone and paraconule, but state 1 is only known from UT DKO 234 and the specimen from Vathylakkos. This state is illustrated by a narrower and V-shaped distal valley. 42. Protoloph on molars: reaching the protocone (0); not reaching the protocone (1). This corresponds to character #30 of de Bonis et al. (1995), which was coded in three states: reaching the protocone (0), stopping at the paraconule (1), and loss of the paraconule (2). Herein, the loss of the paraconule is coded in a separate character (40), and occurs only in Anisodon sivalense. Indeed, there is no suggestion that this loss is part of a hypothetical trend to protoloph reduction within chalicotheriine, as de Bonis et al.’s (1995) coding implied. It is also important to note that de Bonis et al. considered the presence of a crest continuing the protoloph as the same state as the protoloph physically reaching the protocone. We disagree with this conclusion (moreover, remains from Sansan show that the development of this crest is intraspecifically variable). 43. Mesial cingulum on upper molars: regularly lowered until the lingual border (0); displaying a notch in front of the protocone (1). New character. State 1 is observed only in UT DKO 234 and the specimen from Vathylakkos. 44. M2: metacone more labial than paracone (0); metacone at least as lingual as paracone (1). New character. In fact, it is not restricted to M2: M1 can be used if unworn. This character is inapplicable for Eomoropus and Anisodon sivalense (‘Nestoritherium’). In Eomoropus, the molar pattern is relatively primitive: i.e. the paracone and the metacone are on the labial border of the molars (the ectoloph is not yet W-shaped). In A. sivalense the metacone has migrated mesiolingually, obstructing the central valley. The metacone is as lingual as the paracone, but it is impossible to say from which state this peculiar position has evolved. 45. External wall of the metacone and metastyle on M3: subparallel to the mesiodistal lengthening (0); subperpendicular to the mesiodistal lengthening (1). This corresponds to character #32 of de Bonis et al. (1995), which was coded in three states: metacone rather labial in position (0), metacone displaced lingually (1), and external wall of the metacone subperpendicular to the sagittal plane (2). The character was treated as ordered. Here, we only code the orientation of the external wall of the metacone and metastyle, not the position of the metacone, which is intraspecifically variable (see description of remains from Sansan and Saint-Gaudens). 46. Postfossette on M3: more pinched than in other molars (0); as wide as in other molars (1). New character. ‘Postfossette’ is used sensu Butler (1965).


MIDDLE MIOCENE CHALICOTHERIINAE FROM FRANCE 47. Distal cingular crest on M 3: reaching the hypocone summit (0); stopping at the base of the hypocone (1). This corresponds to character #33 of de Bonis et al. (1995), but it was recoded. State 0 is present only in UT DKO 234 and Anisodon sivalense. It should be noted that the M3 are missing in the specimen from Vathylakkos.

Lower dentition 48. P2 : absent (0); present (1). This is character #35 of de Bonis et al. (1995), but the coding was reversed. The absence of P2 in Titov Veles may be an artefact (de Bonis et al., 1995: 170). 49. Entoconid of P4 : weak (0); conspicuous (1). This is character #34 of de Bonis et al. (1995), but the coding was reversed. 50. Trigonid of lower molars: V-shaped (0); U-shaped (1). New character. The U-shape is characterized by the elongation of the hypolophid, which fails to reach the metaconid, and by a protolophid perpendicular to the anteroposterior lengthening. 51. ‘Metastylid’ of lower molars: weak (0); strong (1). This corresponds to character #36 of de Bonis et al. (1995), but was recoded. Contrarily to de Bonis et al.’s analysis, Anisodon grande is coded as having a weak ‘metastylid’ (see Fig. 9F), whereas Eomoropus and Moropus are coded as having strong one.

APPENDIX 3 TREE

DESCRIPTION

(FIG. 10)

Node B: Chalicotheriinae Unambiguous: 30(0), 38(1), 39(1), 42(1). ACCTRAN: 14(1), 25(0), 29(1). Node C: post-Early Miocene chalicotheriines Unambiguous: 2(1). DELTRAN: 14(1), 25(0), 29(1).

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Node D: Anisodon Unambiguous: 1(1), 4(1), 5(1), 7(1), 8(1), 10(0), 13(1), 15(1), 18(0), 21(0), 22(0), 27(0), 31(0), 44(0), 45(0), 51(0). DELTRAN: 11(1). ACCTRAN: 2(2), 6(0), 9(1), 11(1), 20(1), 23(1), 35(state 2–0), 37(1). Node D to A. grande: Unambiguous: 24(0), 49(0). Node E: (A. macedonicus, Vathylakkos, ‘C.’ wuduensis, Titov Veles, N. sivalense) Unambiguous: 33(1), 47(0). DELTRAN: 6(0), 23(1), 35(state 2–0), 37(1). ACCTRAN: 17(0), 29(2), 41(1), 43(1). Node E to A. macedonicus: Unambiguous: 34(1), 35(1). DELTRAN: 2(2), 9(1), 17(0), 20(1), 29(2), 41(1), 43(1). Node E to Vathylakkos: DELTRAN: 41(1), 43(1). Node E to Titov Veles: Unambiguous: 30(1), 34(1), 48(0). Node E to N. sivalense: Unambiguous: 40(0). ACCTRAN: 41(0), 43(0). Node F: Chalicotherium Unambiguous: 19(0), 46(0). ACCTRAN: 32(1). Node F to ‘M.’ brevirostris: Unambiguous: 28(1), 29(2). Node G: (C. goldfussi, Lgr 1065) Unambiguous: 38(0). DELTRAN: 32(1), 50(1). ACCTRAN: 16(0). Node G to Lgr 1065: DELTRAN: 16(0).


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APPENDIX 4 CRANIO-MANDIBULAR MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN MNHN

Sa 9339 Sa 9340 Sa 9341 Sa 9342 Sa 9373 Sa 9374 Sa 9375 Sa 9376 Sa 9377 Sa 9378 Sa 9379 Sa 9380 Sa 9381 Sa 9382 Sa 9383 Sa 9384 Sa 9385 Sa 9386 Sa 9387 Sa 9388 Sa 9389 Sa 9390 Sa 9391 Sa 9392 Sa 9393 Sa 9394 Sa 9395 Sa 9396 Sa 9397 Sa 9398 Sa 9568 Sa 9649 Sa 9650 Sa 9651 Sa 9652 Sa 9870 Sa 9960a Sa 9960b Sa 9960c Sa 9960d Sa 9961 Sa 9962 Sa 10305 Sa 10306 Sa 10741 Sa 10944 Sa 15642 Sa 15670 Sa 15671 unnumbered

AND DENTAL REMAINS FROM

SANSAN

AT THE

MNHN, PARIS

(lectotype) right maxilla with M 1–M3 (formerly A.C. 4232) left maxilla with P4–M3 (formerly A.C. 4233) (lectotype chosen by Schaefer & Zapfe (1971)) left hemimandible with M1–M3, part of the complete mandible described by de Blainville (1849) left juvenile hemimandible with the symphysis and D 2–M1 (Filhol’s collection) upper dentition with left P4–M3, and right P4 and M2–M3 left hemimandible with P4–M2 right juvenile hemimandible with D 2–M1 (Filhol’s collection) right hemimandible with M2–M3, part of the complete mandible described by de Blainville (1849) fragment of right hemimandible with M 3 (or M2?) fragment of right maxilla with D 3–M2 (formerly A.C. 4234) partial left maxilla with D4–M2 (formerly A.C. 4235), probably associated with MNHN Sa 9378 left hemimandible with P4 and a fragment of M1 right hemimandible without teeth left hemimandible without teeth right hemimandible without teeth fragment of left hemimandible with alveoli of M 1–M3 ectoloph of right upper molar mesiolingual fragment of right upper molar lingual half of left upper molar mesial border of left upper molar crown of right P4 fragment of left maxilla with M1 right M2 (formerly A.C. 4229) fragment of right juvenile hemimandible with D 2 and D3 left P3 right P2 or P3 vestibular border of right lower molar left P3 upper D? (missing specimen) skull fragments (maxilla, condyles, tympanic bulla, occiput, and basicranium fragments) distal fragment of right corpus condyle and coronoid process of a juvenile left hemimandible fragment of left maxilla with teeth in section fragment of maxilla talonid of left lower molar right upper molar (M3?) juvenile mandible with right D2–D4, symphysis and left D2 left D4 left D3 left lower molar (probably M1) juvenile lower incisor? juvenile lower incisor? D2 (missing specimen) canine? (missing specimen) fragment of upper molar ectoloph left D3 distolingual half of upper molar (M 2?) skull discovered by Lartet (de Blainville, 1849) complete specimen discovered by Filhol (1890) mesial fragment of right ramus
