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On Buno/Istríodon (=Euro/Istriodon) and Kubanochoeres

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On Buno/Istríodon (=Euro/Istriodon) and Kubanochoeres ¡! Proc. Kon. Ned. Akad. v. Wetensch. 100(1-2), 141-160 June 23, 1997 . - On Buno/istríodon (=Euro/istriodon) and Kubanochoeres ¡ by J. van der Made ¡ Consejo Superior de Investigaciones Científicas. Museo Nacional de Ciencias Na/urale.v, c. José Guliérrez Abascal 2, 28006 Madrid Communicated by Pror. J.E. van Hinte at tbe meeting of June 23. 1997 ABSTRACT The name Eurolislriodon Pickford and Moya-Sola, 1995was meant to replace Bunolislriodon, for a wide-spread group of Miocene pigs. However, Bunolislriodon Arambourg 1963is a valid name, and thus Eurolislriodon is a junior synonym. Three groups of derived characters are shared by kubanochoeresand listriodonts: 1) meso-distal diameter of the incisors and related characters such as anterior mandible shape,2) decreasein the heigbt of the crown of the incisors, and 3) the morphology and the orientation of the first upper incisor. These characters show that the kubanochoeresbelong to the Listriodontinae, though most of the characters tend to be more developed in tbe Listriodontini than in the Kubanochoerini. INTRODUCTION Unfortunately the classification ofthe Listriodontinae sufferedfrom instability ~ during the last twenty years. Wi1kinson (1976) placed a number of lis- triodontine suids in the genus Buno/istriodon, Wilkinson (1978) placed these ., species in Kubanochoerusand Pickford (1986) placed them in Libycochoerus. Savage (1990, p. 593), reviewing African faunas, comp1ained:'However, the taxonomic identity of African suids (and anthracotheres) is very shaky; every paper published in recent years has produced yet another horrendous changeof namesand the synonomy now surpassesthat of any other mammalian stock on the continent. A careful and responsibletaxonomic revision for both suids and anthracotheres is sorely needed.'A classification of the Suidae at the genuslevel is presentedin figure 1 and figure 2 gives the stratigraphical position of the type ,I;~,. 141 fM1I1ySuMIM: Subfamily Tribe Geous SeIecIed lYIMIIIYmI C8inochosiD8e KM)8rMI .4II.-.,u~. ..wnlll,... LiiOi'~=-~=.,,:~ po " li ~ockxIIiJIi. N-'-:-:..~~ N..- ~-- ~~7"'" .."-:.-;~--~-_8IK K~_-,.. ~nu \... ~ .. ~~ ~ u.w. CIIIJ,-"úJLf. TapirotherilUll, '. Lophjochoerus HyodICIÜDae -- ... A8r - .;& x.-.,.. ::-jOIfo.,-'-:ChlellG.JlOChoerus Tc~ ~ ~ ~ ~nu lAPIIocI-rMS S~ T.:.-~ B8ru88 ~ ~..4 ::':.'_G.'N_-:-.¡r ~ 1'tI~.-::c;r~~ lfiRII*yini SI-.,., ~ Mff~ ~ro4/ju ~ I-.-i;;~::':~-~~-'"' . '-'~~~"---'-- «-~..,., HI~ ~--j% E.-iocMerM.I Koipochoerus MU«6OInu H-~.. s.- s... Pwr* Fil. 1. A classification of the Suidae. material of each species (dots) and the phylogenetic relationships of tbe Lis- triodontinae (modiefied after Van der Made, 1996). In a recent paper, Pickford and Moya-Sola (1995), introduced the new name 142 ~ .. Fig. 2. The evolution and stratigraphical position of the type materia! (dots) the Listriodontinae. Stratigraphy modified after Alvarez Sierra et al. (1997) and Van der Made (1996). Sets are Fauna! Sets of Pickford (1981) for Africa, Ramblian and Aragonian biozones from various papcrs by Daams. Alvarez Sierra and Freudenthal, and MN Neogene Mammal units from Mein (1975, 1990) and De Bruijn et al. (1992). Namachoerini are known from Africa, K. minheensisfrom China, K. robustusfrom China, Russia and the Indian Subcontinent, K. mancharensisfrom Turkey and the Indian Subcontincnt, remaining Kubanochoerus spp. from Africa, L. splendens and possibly B. lockharti (= ? B. intermedius)froro Europe and China, B. adelJi- meidamon lineage froro Europe and Turkey, B. guptai - L. pentapotamiae from the Indian Subcontinent, the B. affinis - jeanneli lineage froro Africa and the Indian Subcontinent and B. akatikuha.r from Africa. Euro/istriodon to replace the widely used llame Buno/istriodon. They state that Buno/istriodon is a nomen nudum and include B. /ockharti, type speciesof Bu- no/istriodon in Euro/istriodon. They further introduced a new species,E. ade//i on the basis of a comparison of the material from EIs Casots with that from Can Canals, but omit a comparison with the types of B. /ockharti and B. be/- siacus.The absenceof p2 and P2 in the type of B. ade//i is probably pathological and B. ade//i might be identical to Bunolistriodon aff. /atidens ofVan der Made (1996) and be ancestral to B. /atidens. Pickford (1986, 1988, 1995) and Pickford and Moya-Sola (1995) included Kubanochoerusand Libycochoerusin the Kubanochoerinae, and believed them to be different from the Listriodontinae. Van der Made (1996) considered the genera synonymous and belonging to the Kubanochoerini of the Lis- triodontinae. In the following, the informal 'kubanochoeres' and 'listriodonts' will be used whenever the taxonomic level is under discussion. 143 ~ This paper discussesthe status of Eurolistriodon and the characters that have a bearing on the status of the kubanochoeresin arder to answer the question: 'should the kubanochoeresbe included in the Listriodontinae or do they con- situte a subfamily of their own?' ABREVIATIONS OF MEASUREMENTS AND COLLECTIONS DMD Meso-distal diameter in mm, taken as indicated by Van der Made (1996). DMD' DMD of an incisors, expressedas a percentage of the width of tbe MI (MI for upper incisors, MI for lowers). DLL Labio-lingual diameter in rom, taken as indicated by Van der . Made (1996). 1 The meso-distal diameter (DMD) expressedas a percentage of the labio-lingual diameter (DLL): 1 = 100DMD/DLL i". BNHM Beijing Natural History Museum. BSPHGM Bayerische Staatssammlung fúr Palaontologie und Historische Geologie, München. GSP Geological Survey of Pakistan, Islamabad. HLD HessischesLandesmuseum Darmstadt. 1M lndian Museum. Calcutta. IPS lnstitut de Paleontologia. Sabadell. IVAU Instituut voor Aardwetenschappen,Utrecht. IVPP lnstitute for Vertebrate Palaeontology and Palaeoanthropology, Beijing. KNM Kenya National Museums. Nairobi. MNCN Museo Nacional de Ciencias Naturales. Madrid. MNHNP Muséum National d'Histoire Naturelle. Paris. MPZ Palaeontological Museum of the University of Zaragoza. MTA Maden Tetkik ve Arama. Ankara. NMB Naturhistorisches Museum, Basel. NNML Nationaal Natuurhistorisch Museum. Leiden. PDTFAU Paleontropoloii. Dil ve Tarih Cografya Facultesi, Ankara Uni- versitesi. SLJG SteiermarkischesLandesmuseum Joanneum. Graz. SMNS Staatliche Museum für Naturkunde in Stuttgart. UCBL Université Claude Bernard, Lyon. ULAR Universitá La Sapiensa,Rome. ZMA Zoologisch Museum, Amterdam. THE STATUS OF BUNOLISTRIODON Arambourg (1933) introduced the name Bunolistriodon conditionally for a group of bunodont and sublophodont suids, that at that time were included in Listriodon. He did not designatea type species.After 1930,the introduction of 144 a genus should be accompanied by the indication of a type species(Interna- tional Code of Zoological Nomenclature, article 13b; Ride et al. 1985),also the conditional introduction of a genus.Thus Bunolistriodon Arambourg, 1933is a nomen nudum. A nomen nudum is a not available name that may be made available later, taking authorship and date of that later publication (ICZN). Arambourg (1961) described a mandible from Gebel Zelten and based the new generic and specific names Libycochoerusmassai on it. At that moment he considered the genus to be related to Propotamochoerusand Conohyus.A skuil ofthe samespecies was found later at Gebel Zelten, as well as a large collection of bones and teeth. This made Arambourg (1963) aware of the similarity be- tween Libycochoerus and bunodont listriodonts and he considered Liby- cochoerusArambourg, 1961as ajunior synonym of Bunolistriodon Arambourg, 1933.At this stage Arambourg (1963)designated 'Sus lockharti Pomel, 1843'as type speciesof Bunolistriodon. It is evident that Arambourg (1963) meant Sus lockharti Pomel, 1848, since Arambourg (1933), cited by Arambourg (1963), referred to Pomel (1848).The fact that in 1933no type specieswas indicated is thus no reason, why Bunolistriodon Arambourg, 1963should not be available. Pickford and Moya-Sola (1995, p. 352) state that Arambourg (1961) de- scribed a skull from Libya, which gives the false impression that Arambourg (1961)based Libycochoeruson good and abundant material, whereasin reality this additional evidence convinced Arambourg (1963) that the name Liby- cochoerusis a synonyrn of Bunolistriodon and should not be used anymore. Leinders (1976),Pickford (1995) and Pickford and Moya Sola (1995)were of the opinion that the name Bunolistriodon is not available, for at least one of two reasons: problems with the diagnosis or with the type species.Pickford and Moya Sola (1995)considered the genus as a nomen nudum and introduced the name Eurolistriodon to replace it. The type speciesof Eurolistriodon is E. adelli. Two other speciesare included in the new genus: B. lockharti, type speciesof Bunolistriodon, and B. latidens. E. adelli might be identical with B. aff. latidens of Van der Made (1996) and we agree that both species (B. lockharti and B. adelll) belong to the samegenus. Arambourg's (1963) diagnosis of Bunolistriodon has been criticized (Lein- ders, 1975;Pickford, 1993;Pickford and Moya Sola, 1995).Pickford and Moya Sola (1995, p. 353) cite Arambourg's (1963) diagnosis comple,tely,underlining characters that they could not observe in B, lockharti and underlining double characters that they consider to be primitive. Though it is common to give a diagnosis with the introduction of a taxon, the word diagnosis is not used in the ICZN. The ICZN (article 13ai) puts it in this way: 'accompanied by a description or definition that statesin words characters that are purported to differentiate
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