Bol/ettinodel/4 Societ4 Paleontologictl ltalia1l4 1 pl. Modena,Ottobre 1989
European Suinae (Artiodactyla) &om the Late Miocene onwards
Jan VAN DER MADE Salvador MoYA-SoLA InstitUutvoor Aardwetenschappen Institut de Paleontologia "M. Crusafont" RijksuIÚversiteitte Utrecht Sabadell
KEY WORDS- Mammalfa,Art1odactyia, Sutnae, Neogene, Eu~.
ABS1RACf - Eumpean Suldae tbat haLlea time dlslrlbutton nMr tbe Mto-Pliocene boundary haLle~ studied. 1bey beiong lo /UU trlbes: Dtcot):1Jhocboerlnt(Korynochoerus, Mi~onyx and Eumaiochoerus) and Sutni (oniy tbe genus Sus). 1be ftrst hibe was wel/ ~ted tn lbe Late Mlocene and tbe otber tribe was ~.from tbe Eariy PlIocene onwa~. ]be tmmtgrants K. palaeochoerus, Mi~onyx and K. provincial~ enten1din tbe /ate5tMtddJe Miocene and Late Miocene. ~onyx Lsa lineage of tb~ forms; lbe endemic island form Eumalochoerus Lsan offsboot of tbLsItne. In tbe PHoceneSus arvemensis and S. strozzü tmmigrated. Susscrofa ente1-edlate tn tbe Eariy Pletstocene.]be endemtc SarrJtntan Sus nanus LsbeIteIKNi lo be a descendant o/S. arvemens~. A range cbartfor tbe Suinae Lsg~. Eu~n Mlcrostonyx was a ItnWJgecbaractenza:I by a si:zedecrease o/ lbe denIttton and m o/ pnmIola1S:M. antlquus, M. major major and M. major erymandllus. "Propotamochoerus" provinclalls tsfound lo be related lo KorynochoenJs¡Yd!aeochoerus and Lstncluded in Ibis genus. 1Me /UU spectesare not re/ated lo ProP
RIASSUNro- [Suinae (Artiodactyla) europei a partire dal Miocene superiore) - Sanostatt studtattt sutdt europetche banno una dtstrlbuztone tempomle vicina al Itmtte Mio-Plíocene. Em ~no a due trlbll.. DicOr)pbochoerint (KOIyfiochoerus. Microstonyx e Eumaiochoerus) e Sutni (ti solo genen1Sus). La prima trlbtJ era '--m ~ alla ftne del Mtocene mentre la seconda co~re a pantre da! primo Pliocene. Korynochoerus palaeochoeros. K. provincialis e Microstonyx giutlsero come tmmigrantt alla ftne del Mtocene medio e nel Miocene superiore. Microstonyx costtJUtsce una Itnea ftlettca di #refi>nne; la fonna insufare endemica Eumaiochoerus trova la sua origine tn ~ linea. Ne/ Pliocene tmm~nmo Sus arvemensis eS. strozzii. S. scrofa gtunse nel tardo PleisIOCenein/more. Si pensa cbe la fonna endemica Sus nanus deDa Sardegna dtscenda da S. arvemensis. VIene stt/ata una carta di dtstribuztone del Sutnae. La línea eoo/ub"meu~ di ~onyx.fu camttenzzata da/Ja dtmtnuztone dt tagHa della dentíztone e daDaperrJita de; premoiari: Mi~onyx antiquus, M. major major e M. majar eryrnandúus. VIene ~ cbe "Propotamochoerus" provincialis e coliegato a Korynochoerus palaeochoerus e pe11antoincluso tn quesi'ultimo genere. Questedue spec;ienon sono tn relazione con Propotamochoerus. 1 resti~lí riferiti al sottogenereP~oerus vengono considemti victni a Sus e non a Potamochoerus. Vengono/omUt i ciadogmmmi del/e trlbtJ Dicorypbochoerlni e Sutni.
INrRODucnON mE TAXA AND mElR CHARAcrERISI1CS
The African Pliocene and PleistoceneSuidae are Thic;section has two purposes.One purpose is to consideredto be of great value in sttatlgraphy.White & give the chamcters of the teeth that are frequendy Harris (1977) provided an altemative sttatigraphy on found, so that a taxon can be recognized in a collec- fue basis of suid evolution in cases where absolute tion. Another purpose is to give charactersthat can be methods were controversial. used in phylogenYi these chamcters are usually only It is doubtless under fue influence of that and seen in rare fossils such as complete skulls. similar papers that interest in EurasianSuidae is grow- ing. The workshop on continental faunas at fue Mío- cene/Pliocene boundaIY is a good opportunity for Dicoryphochoerini Schmidt-Kittler, 1971 drawing up an inventoIY of fue data available on fue European Suidae of that time. The Dicoryphochoerini share several characters, Apart from a suid from Samos little attention is such as fue morphology of P4 and fue morphology of paid to Asian suids in this paper. The Samossuid might fue upper incisors. also prove to occur in Europe. The P. has a rnain cusp positioned labially to fue 330 J. VANDERMADE, S.MOYA-SOU
midline. Frorn this cusp an antero-lingually directed "ProjJotamocboetUS"provinctaJís (Gervais, 1859) ridge descends to the anterior CingulUffi ending so- mewhat linguaUyof the middle of the tooth. This ridge "P." provtncíalís has dentition that is a little larger is not smooth, but mar have swellings,which look like than in K. paiaeochoetUS.The size range of fue teeth is cusps.A second main cusp is placed a little behind the not well known but it is known that the two species other and much more linguany. Posteriorly these two overlap in size. The molars have somewhat more pro- cusps are connected by a ridge or cusp to the muro nounced accessorycusps and fue labial wall of fue lower posterior cusp of the talonid. On both sides a labial CUSpsof fue upper molars is more convexoThe 13 cingulum runs up to this cusp and fuses with it. The resemblesthat of K. pa/aeochoerus. anterior and posterior ends of the tooth are about the In a female skull from PtolemaisKardia fue occiput same height and cleariy lower than the maincusps. is not elevated as it is in fue male K. pa/aeochoerus A detailed study of the incisors of the Suoidea is skull from Johnsdorf (see Mottl, 1966 and Thenius, being prepared by one of us Q. v. d. M.); it is sufficient 1972). In Sus Mrix.u'us and Metrldiochoerus andrewsi to deal only with the 12and 13in this paper. The second both males and females have elevated occiputs. Alth- upper incisorsare long and narrow. They have a poste- ough fue skull structure is different, this indicates that ro-lingual cingulum. fue difference in fue shape of fue occiput of fue male AI1 Dicoryphochoerini have more elongated mo- K. pa/aeocboenJSand of fue female "P." provincíalís Iars than primitive Suidae.The males of K. pa/aeochoe- skulls is not due to sex but to specific characters.The ros, "P." provinctalts and Mlcrostonyx and prot:y.-bly Johnsdorf skull has inflated zygomatic arches,whereas also Eumaioch0en4Shave weU-developed alveolar in fue skull from Ptolemais fuese arches are not in- crests above the canines. The alveolar crests of 'P." flated. This is a feature that is related to mastication;in provinctalts are more slender.The femalesof K. paJae- Potamochoerusfue shape of fue zygomatic arches in och0en4S,"P." provinctalts and probably also of Mi- femalesand malesdoes not differ much, although their crostonyxand EumaiochoentSlack such crests.(This is zygomatic arches differ greatly from those of a prim- also the casein modero suids such as Potamoch0en4S, itive type suchas those of S. scmja. In Metrldiochoerus s. ~, S. batfutUS and S. celebensts). however fue degree of inflation is surprisingly related to sexo Korynoch0en4Spaiaeoch0en4S (Kaup, 1833) We expect that "P."provincialís does not have fue zygomatic arches inflated nor fue occiput elevated. In K. pa/aeochoentSis about the size of a recent wild fue crushed Ptolemaisskull fue anterior rim of fue orbit boar. The males have much inflated zygomatic arches seemsto be somewhere above fue M3.The individual that depart nearly at right angiesfrom the snout. In the is fully adulto It should be borne in mind that this femalesthis angie is less acute, it is not known if their characteris ase dependant.The "P." protltncíalís from zygomatic arches are also inflated. The occiput was Montpellier has lost its 13.The skull from Ptolemais elevatedin the males,whereas in the femalesthis is not Kardia has lost both r, fue alveoli are still open but known. The anterior end of the orbit is behind the M3' were in fue process of closing when fue animal died. The 13is large but not elongated; it does not always The loss of r and P during life seems to have been have a postero-lingual cingulum. common.
EXPIANA'nON OP PLATE 1
- K. provtnctaJls.Right p., Venta del Moro, VM ~, MNCN (Museo Nadonal de Clendas Naturales,Madrid). Fig. 1 2 - Microstonyx major erymantbius. Right p., KerassIa,Ke 72, IVAV (Instltuut voor Aardwetenschappen,Vtrec!1t). Fig. - EumaiochoenLSetn¿scus. Right p., Pavolona (Bacclnello V2), Museum of Natural History, Basle. Fig. 3 Fig. 4 Koryn0cb0en4SpaJoeocboeru.s. I.eft P. Can Ponsic 1, IPS 1001. Instituto de Paieontologia,Sabadell. 5 - Sus'scrofa, Right p., recen! from Benaban'e(Huesca), coll. JvdM. Fig. - Sus nanus. Left p', Capo Figari, Ty 5353, Museum of Natural History, Basle. Fig. 6 7 - Potamocb0en4S.Right p., recent, RMNm 1686, Rijksmuseumvoor Natuurlijke Historie, Leiden. Fig. - K. provtncialts. Right 1', Venta del Moro, VM 115, MNCN. Fig. 8 - K. paiae!Xb0en4S. 9) Left P, Venta del Moro, VM 613, MNCN; 10) 1', Castell de BarlJed, IPS -; 11) 1', HOStalets, IPS 1077. Fig.'i. 9-11 12-13 MicrostonJIX 12) Right 1', IPS Ta78; 13) Right 12 and 1', Sarnos, Sam. 27, Museum of Natural History, BasIe. Figs. - Potam0cb0en4S.!tIght12 and 1', RMNHL. Fig. 14 Fig. 15 Sus scrofa. Left 12and 1', recen! BenabaIre (Huesca), roIl. JudM. J. VANDER MADE, S. MOYA-SDf.)., EUROPEAN SUINAE FRON L MIOCENE Pl.l J. VAN DERM.-WE; S. MOYA-soL1.
placed in MN 12. Maybe Tudurovo should be correlat- ed to MN 12 if this is not contradictory to evidence Mtcrostonyx is a iarge suid with an elevated occi- from other taxa. pUl, lengthenedsnout, relatively small canines,relative- Since fue srna11estfonn has not been found in Iy shallow mandibles, r' that are greatiy elongated and westem Europe one of us (S. M.) think¡¡ that it is a that have postero-lingual cingula, inflated zygomatic geographical subspecies. But as fue size decrea.-.e archesithe parietal ridges are wide apart (at least in the seemsto occur in many places, fue smaller fonn may male skulls that we know), The anterior rim of the orbit a1sobe a younger subspecies.The iarge fonn has been is well behind the W, found in France, Gerrnany and Turkey; fue intenne- A lower Vallesian forro has a very large dentition diate fonn in Spain, France, Russiaetc. and fue srna1l and retains its p1 and P1'An upper Vallesianto middle fOlm in Greece and Russia. Turolian forro has a smaller dentition and also retains its f1fSt premolars (the PI is probably lost early in life), EumatochoenJSetnJSCus (Michelotti, 1861) There is also an upper middle Turolian forro from Pikermi and Kerasia that has a sti1l smaller dentition, Eumatoch0en4SHürzeler, 1982 is an endernic is- but the skull is just as long as it is in upper Vallesian land fOlm. It has relativeIy small canines like Mícrosto- and middle Turolian forros, it has no P1 andoften lacks nyx and greatiy elongated 13,but is srnaller. Its denti- its p1 (which is probably lost during the life of the tion is of about fue sarnesize as in K. palaeochoerus. individual), Gaudry (1862-1867)reports alveoli for the The anterior edge of fue orbit is above fue anterior end pl on two of six skulls from Pikermi. He did not find of fue M3 (in one adult animal) and fue orbit was any indication of the existenceof a Pl' It seernsthat the probabIy higher than in most other Suidae.These cha- size of the canine and the depth of the mandible racters are probably related. The talan and talonid of decreasefrom the ftrst to the last forro (these characters fue M3 and M3 tend to be wider than in K. palaeochoe- are probably related). ros and in "P." províncíalis (although overIap occurs). Other differences between forros of Mtcrostonyx The talonid of fue M3has two larger posterior cusps,as that have been mentioned (for instance by Thenius, occur frequendy in Microstonyx, to a lesser degree in 1972, pp, 555-557)are: 'P. n províncialis and not at a11in K. palaeoch0en4S. 1) The degree of complication of p., We failed to Compared to K. paiaeochoenJSfue mandible of Eu- find significant differences: this apparent difference matochoernsis relatively shallow in view of fue size of was probably an artefact of individual variation or of fue dentition. The depth measuredlingually in a man- wear. roble from Can Uobateres is about 49 rnm below M¡ 2) The lenght of the snout (indicated by the dis- and M2; in a rnandible with only slighdy smaller teeth lance Cx - P3 and by the lenght and position of the fue depth is about 32 mrn in E. etroscus.Eumaíochoe- symphysis), It should be realized that both characters ros may have a P11 but this tooth may also be lacking. are related to sex and prüOOblyto ontogenetic age. We do not know whether Eumaíoch0en4Shada pl, but 3) The length of the part of the skull anterior to the it is likely since a P¡ usua11ydisappears flrst during orbit relative to the lenght of the posterior part of the dental reduction. skull. This ratio is infIuenced by the different rates of development of the different parts of the skulli fuese Suini developments are ontogenetic ase related. 4) The structure of the M3' In both lower Vallesian The Suini share fue shape of the P4' and probably and the middle Turolian forros the third lobe generaily the shape of their indsors. consists of a srnall cusp on the axis of the tooth and The P4 is one high meso-dista11ydirected ridge. two larger cusps behind this srnall CUSPithe morphol- There is one main cuspj a second equally large cusp ogy of the M3 is probably not of much utility, mar be placed irnmedjateIypostero-lingually to it. After TroflffiOV (1954) gives measurementsfor Russian only a little wear fue two cusps can no longer be Microstonyx. Measurementsfrom Eldar, Berislave and differentiated. From the main cusp a ridge is directed Grossulovo (MN 11) and Grebeniki and Taraklia (MN anterolinguallY.This is a homologous ridge to the one 12) are similar to those of contempornry Mtcrostonyx seen in the Dicoryphochoerini, but it does not slope from westem Europe.The Microstonyx from Tudurovo downwards. is comparable in size to Mtcrostonyx from Pikermi, Its What is called the anterior cingulum in that group mean size is greater than that of Mtcrostonyx from is either absent or is directed nearly vertically and Kerassia,indicating a greaterase (?).Tudurovo is dated forms lingual and labial Stylids.The cusp of the talonid MN 13 (Mein, 1975), Kerassiasti1l has Deínotberlum is nearly as high as the main cusp. What is homologous and Dorcatberium: Kerassia probably should be to the posterior cingulum is absent in Suini or it has EUROPEAN SUlNAE FROM L. MIOCENE 333 been converted into a lingual and a labial, nearly verti- Susnanus van der Made,1988 cal stylid. The 12 is very low and elongated,a postero-linguai Sus nanus is a dwarf fonn from Sardinia. It is cingulum is lacking (this mar be due to reduction). We 75-800/0of the size of Susa~ensis. Its p4 and molars do not know this tooth in S. a~ hut suppose are slightly more hypsodont than fue sanle teeth in that it has the samemorphology as in fue other species. most speciesof Sus.The enamel surfaceof the molars The 13is small and short in S. scrofa; we suppose it to looks very smooth and simple. The talan of the M3 be like fue 13in fue other species, since short 13are consistsof one cusp and in the M3 only a small cusp primitive and cornmon in Suidae. and a larger cusp behind it follow the second pair of The genus StlS is currently divided into two cusps.Its P2 and P3 are reducedin size ¡md a P1 is groups: fue spedes with a "verrucosic" male lower absent. The diastema between P2 and the canine is canine and those with fue "scrofic" male lower canine. shortj the snout is shortened. The section of fue lower canine is verrucosic if fue length of the lingual side is about fue same as fue Susstrozzii Meneghini length of fue antero-labial side (measured horizontal- Vety iarge species.The prenlolar row is shortened Ir). If fue lingual side is much wider than the antero- COffiparedto fue statein S. a~ensis. Usually the last labial side fue canine is scrofic. SirK:efemale lower molar has a simple talon(id). The M3 from Valdeganga canines have a verrucosic section this is often thought 11and Tegelen are much more elongated; other teeth to be fue primitive state. But primitive pigs like Cono- from Tegelenseem to be large as well. Thesedifferenc- byu.s,Hyotberlum, peccaries like Ba~yu.s, Alba- es cannot be explained by evolution towards more nob.yus,Taucanamo have scrofic canines. This indi- elongated molars since fuese localities are among 01- cates the contrary. In Listriodon spiendensboth types dest at which S. strozzii has been found. It is not of canines occur. The chamcter is not sufficiently un- impossible that this material represents ) 'et another derstood for great phylogenetic or taxonomic weight to speciesof suid. The wider molars of fue regular type be attached to it. can be differentiated from those of S. scrofa by their The verrucosic graup (spedes with a canine like relatively great width. Azzaroli (1954) repol1sthat one that of S. wrrucosus, which for this reason are thought cranium from Senezestill had its 13and that the youn- to be related to this species)is characterizedby severa! ger crania of Valdamo had lost the 13. other features.There are well developed alveolar crests above fue canines in fue males. These are higher and Sus scrofa Linnaeus, 1758 more slender than in K. paiaeocboerusand Microsto- nyx. In most forros fue zygomaticarches depart abrupt- The premoIar row is shorter than in S. aroernensis. ly fram the skull. In all verrucosicspecies fue stylesand In fue European fonns fue talan and talonid of the last stylids of the premolars, especially of fue P4, are more molars always seem to be complicated; a tl1ird pair of pronounced and fue basesof fue premolars are much cuspsand a tenninal cusp are cornmon in the M3in fue lower than the bases of fue molars compared with S. oldest popuIations (Mosbach). These teedl are more scrofa. The last chamcter is not a constant one, being elongated than those of fue Dicoryphochoerini, S. ar- dependent on age and being difficult to measure.Ho- t;ernenS~and S. nanus. wever it seemsto be useful within certain limits. The skull structure and canine shape could not be checked PHYLOGENY AND TAXONOMY in S. nanus. In fue scrofic group fue zygomatic arches are more slender,fue alveolar crestsabove fue canines Suinae Van Zittel, 1893 in fue males are less developed and fue base of fue premolars is higher. In recent literature all taxa described here are in- cluded in fue Suinae. Mtcrostonyx has traditionally Sus arvernenstsCroizet & Jobert, 1828 been consideredto belong to fue Suínae(Many Suídae were ftrst described as a species of Sus. Later some Susaroernensis is fue smallestSus known fram fue authors attached more ímportance to fue dífferences mainland of Europe. The M;s have a simple talonid and made new genera; others created a tribe or sulr with one terminal cusp. The talan of fue M3 is also family. For some reason, in the minds of paleontol- simple. The anterior and posterior ends of fue P3 are ogists Mtcrostonyxremained connected with Sus.)On- much lower than in S.strozzii and S. scrofa.The length Iy Thenius (1972) placed this genus in fue Hyotherií- of premolar raw of S. a~is relative to fue molar nae.He did this becausehe recognizedfue relantionship raw is large for S~ the relative length is probably fue betWeen "Hyotberlum" paiaeochoerus and Mtcrosto- nyx. "Hyotherlumn paiaeochoeruswas thought (O be same as in the Dicoryphochoerini. 334 J. VANIBMADE, S.MOYA-SDU descendantof Hyotherium soemmerlngtuntil Schmídt- Potamocb0en4Shas a P. with one main cusp. A P. Kittler (1971) proved this idea to be false; he created with only one main cusp is probably prirnitive in Suoi- the genus Korynochoerusfor the Spedes "H." paJae- dea. lbe Dicoryphochoerini P. is farther from fue sup- ocboerusand created fue tribe Dicoryphochoerini for posed ancestralfonn than fue Potamocb0en4SP4' Di<;orypbocbQen4S(including only D. I#an with <;er- The M3 of Potamochoernsdoes not llave a third tainty). Mtcrostonyxand Korynocboe1US.This tribe was pair of cusps. The talonid consists of severa! srnaller characterizedby the morphology of the P. and placed cusps and one 1argercusp. The latter mar be divided in the Suinae.Sclmúdt-Kittler probably placed the Di- by a cIeft. In severa!M, of "P." provtnctalts this cusp coryphochoerini in the Suinaebecause Sus is trndition- can already be seen to be divided. The M3 of this ally thought to be derived from Dtcorypbochoerus(in speciesis a1somore eIongated(lower DT/DAP ratio). If this case Dtcorypbochoerusincludes all spedes that this is an evolving feature it seems strange that fue were included by Pilgrim. 1926). He derived the SU-'- specimensfrom Montpellier are equa1lyprogressive or type P4 from the Dtcorypbocboerustype by supposing even more progressivethan recent POtamoch0en4SM,. that the position of the linguai cusp had Changedand "P." provtnctalts possibIy starts to tose its e and l' that the anterior part of the tooth had been raised. early in life, there is no sire reduction of fuese teeth. However the S~ P4 can be derived aIso from the Potamochoenlsretains its (small) ¡2 into old ase. Only PotamocboerusP 4 by supposing that the anterior end one Potamocb0en4Sin fue collection of fue Rijksmu- of this tooth has been raised. Biologists believe that seum V. Natuurlijke Historie in Leyden-(RMNHL)had Potamocboerusand Sus are closely related; they mar not replaced its De. It is unlikely that Potamocb0en4S have shareda long part of their history. The Suinaeare or its forerunners ever had an elongatedIl, since short a group with many more different forms than other ¡2 are primitive. subfamiliesof the Suidaeand we were unable to find any synapomorphy uniting all Suinae. Consequently The relationship between Korynocb0en4Spaiaeocboe- we do not know if this group is really monophyletic; ros and "P." provtnctalts however we follow current taxonomy. since at the moment we cannot offer a better alternative. It has been suggestedthat K. paiaeoch0en4Smight be fue ancestorof "P."provtnctalts (for instanceGins- The relationship between Potamochoerusand "Pro/)()- burg 1980). Both species resemble each other very tamocboerus"provincíali5 much in fue morphology of their dentition. K. palae- ocb0en.L5has an eIevated occiput. In mis respect it Severalauthors considerexplicitly or implicitly that resembIesMicrostonyx. ProbabIy "P."provtnctalts and "P." provinctali5 is closely related to Potamocboerus.or K. palaeocb0en4Sare closely related, but are not an even to be its immediate ancestor(for instanceGuérin evolutionary lineage. Moreover their ranges overlap. & Faure, 1985). This is unlikely for several reasoDs. Stehlin (1899-1900) derived Potamocboen¿sfrom K. PropoIamochoenlS palaeochoenlS.This is also not likely to be con'ect for ahnost fue same reasons. The type Spedes of PropotamochoerusPilgrirn, Stehlin (1899-190())already noted fue trend in Sui- 1926is P. salínus Pilgrirn, 1926.The morphology of fue fonnes to acquire more elongated molars. This hap- P4 was considered important by Pilgrim (1926, p. 22): pened parallel in many branches,even within fue Sui- "The fact that fue height of fue main cusp of P-1 much dae. Potamocboerusis more conservative in this res- exceeds that of fue anterior and posterior cusps dis- pect than severa!odler Suidae.We take the M2 as an tinguishes it from fue Sus - Híppohyus series. The example. cone -like characterof fue main cusp of P.¡distinguish- es it from fue corresponding tooth of fue Dícorypbo- cb0en4Sbranch, ...". The morphology of fue P-1 of fue D'nJ8x/DAP holotype of P. salínus colTespondswell to this descrip- Mz n-.nnD.-. tion (pl. 7, fig. 1). "Propotamochoerus"províncíalís was included in this genus by Pilgrim (pl. 1), although fue Potamocboerus(specimen in stroctureof its P4 is more like that of Dícorypbochoerus. RMNHL) 8 0.77 0.70 O~ K. paiaeocboe1US(Deinothe- From fue definition of Dícorypbochoerus (Pilgrim, riensande, data from Hüner- 1926,p. 37): "The main cusp of fue last lower premolar, mann, 1968) 39 0074 0066 0.79 instead of being single as in fue Conobyusand Pota- "P." provindalis (div. loco) 6 0072 0069 0076 mochoenJSlines, is divided into two of approximately Mícrostonyx (from Pikenni, equal value, which may be known as the principal data from Pearson, 1928) 15 0.73 0.64 O.~ cusp and fue inner cusp.The latter stand" generally a EUROPEANSUlNAE FROM L. MIOCENE 335
little behind fue otherj in some speciesit is almost on a level with it, but it never standsdirectly behind it, as in many species of Sus.". "P." provincialis has a P4 with two main cusps arranged in fue same way as in fue P4 of Dicoryphochoerus tIran, fue type species of thio; genus. However in other charactersit differs from D. titan. '~." provincialis resembles Korynochoeruspa- Iaeochoerusin many features,as we have seen above. The most ilnportant differences are fue shape of fue ocdput and a slight size differenceof fue dentition. The differences are like those between Sus scrofa vittatus and Sus barbatus, which are sympatric. It would be better to place "P." provincialis in Korynocboerus Schmidt-Kittler, 1971.
Microstonyx Text-flg. 1 - The Dicoryphod1oerini cladogram Currendy two speciesare recognized:Mícrostonyx antiquus Kaup, 1833Chololype from Eppelsheim) and M. majorGervais, 1848/1852Chololype from Cucuron). arches are shared by K. palaeochoenJSand Mtcrosto- M. erymantbtusRO{h & Wagner, 1854(from Pikenni) is nyx and probably also by EumaíochoenJSbut probably thought to be a synonym of M. major. According to not by K. provincialis. In K. palaechoenJSand Mícros- Thenius 1972fue two speciesare contemporaneusbut tonyx the anterior rim of fue orbit is placed behind M3. lived in different habitats. Ginsburg 1980 ~ they In fue Ptolernais-Kardiaskull it is difficult to seewheth- fonn one lineage: M. antiquus in MN 9 and 10 and M. er this was already fue case in K. provincialis, but fue majorin MN 11 and 12. As we have seen, three fonns molars were probably in a less fOfW'clfdposition. In can be recognized. We failed to fmd any reliable in- Eumaíochoon.isfue orbit is placed further forward, but dication that two fonns existed at fue same moment this is related to other changes in the skull. and we think that fue three fonn are a lineage. Nade 3 - Elongated 13,canines of reduced size Pearson(1928) gives measurementsfrom Pikenni, and, related to fuese, shallow mandibles are shared by fue M3 range in length from 38.1 to 43.2 mm (n - 12). Mícrostonyxand EumaiochoenJS.A wide occipital and The holotype of M. major is an M3 wrnch is 45 mm parietal zone are probably also shared, but evidence long (Gervais, 1848-1852).The M3 of fue Eppelsheim concerning Eumaíochoenl.Sis lacking. This is fue be- fonn is not known. Judging from fue M3' a mean length ginning of fue Mícrostonyx lineage: Mícrostonyx antí- of 50 to 52 mm can be expected.The fonn of Kerassia quus. is even smaller than that of Pikermi. The size of fue Nade 4 - In EumQÍQChoen.l.5the anterior rim of type of M. major does not indicate that this animal is fue orbit lies above fue anterior end of M3. This is identical to one of fue two other "species",but there is probably due to size reduction of fue part of fue skull certainly a size overIap between fue majar and ery- that is used for mastication. The orbit lies relatively mantbius fonns. The differences between fue three high above fue dentition; this mar be related to fue fonns are very small: presenceor absenceof a premo- relatively high position of the glenoid combined with a lar and small size differences of fue dentition. Diffe- oockward shift of fue dentition. (The eres, brain-case, rences between recent subspeciesare larger. In arder glenoid and occipital condyles are considered to be not to change fue accepted nomenclature too much stationary, but fue dentition is considered to move and in arder to reflect in taxonomy fue differenceswe relative to this group of elements.)In K. paiaeochoerus see,we recognizeM. antiquus, M. majar majorand M. and Mícrostonyx fue anterior rim of fue orbit was major erymantbtus. behind fue W but in K. provincialis it is not quite clear. Nade 5 - Further size decreaseof dentition, com- t The Dicotyphochoerini cladogram plete loss of P1and loss of pl early in life. Nade 1 - All share fue "Dicoryphochoerini-rype The Suini claOOgmm p,,", fue elongated f with postero-lingual cingulurn, molars that are somewhat elongated and some rype of Node 1 - A1l share: "Sus-typep." and probably alveolar crest above fue male canines. also elongated ¡2 widtout postero-lingual cingulum, Node 2 - Elevatedocciput and inflated zygomatic aldtough this is not known for Sus arvernensisand S. 336 J. VANDERMADE,S.MOYA-s0L4
"Postpotamocboorus"from Sarnos
Thenius (1950) described a suid frorn Sarnasas PotamochoenlS (PostpotamocboenJSnov. subgen.)
byotberioides. But its P4 has fue typical morphology of a SusP 4. The talonid and fue anterior part of the tooth are high. The anterior part of the P4 of PotamocboenJS is low. The anterior and posterior ends of the P3 are higher relative to the main cusp than in PotamocboenJS (and S. aroernensts).In addition the rnolars are elon- gated. PotamocboenJSmolars are conservative in this respecto The animal from Sarnasresernbles Sus strozzii in the rnorphology of its individual teeth. Apart frorn this the base of its p4 is rnuch lower than the MI (in both the upper and lower dentition). However the molars are smaller than in S. strozzii trorn Seneze and the Text-fig.2 - The Suini cIadograrn. prernolars are larger. Sus arvernensis has a prernolar row that is relatively longer than that of Sus strozzii. nanus. Recent Sus species such as S. ver7UCOSUSand The premolar row of S. strozzii is probably reduced in others have such an r and are consideredto be closely length. In Suidae a long premolar row is generally related to S. arvernensisand S. strozzii. primitive. It is possible that the animal frorn Sarnasis Nade 2 - Elevation of anterior and posterior end an ancestralforro of Susstrozzii. The description of the of P3 and P2, reduction in the length of premolar row skull given by Thenius does not exclude this possibil- compared to molar row and, at least in fue European ity. forms, a strong elongation of M3. All fuese charncters The figures and data presented by Thenius in- have dcvcloped parallel in at least some speciesof fue dicate that the Sarnos animal is Sus and not Pota- venucosus-grou p. mocb0en4S.The sub-genus is ba.c;edon dIe specics Node 3 - Strong alveolar crests above C' of fue created by Schlos.c;er,1903: Sus byotberioides.It was males,C. of fue males "verrucosic",stronger stylesand Pearson (1928) who related this species to Potamo- stylids in p' and probably wider zygomatic arches in Cb0en4S.The syntypes of S. byotberioide.,do not in- both sexes.(This ís fue "ver7UCOSus-groUp".)Primitive clude a P4. We do not wish to speculatehere about the pigs lack alveolar cresl". ~ charncter is given more specific identity of the Sarnasmaterial, nor about the weight here than fue canines. generic identity of Sus?byotberioides. Nade 4 - The known common charnctersof fuese speciesare all primitive or difficult to interpret Oike fue mE STRAllGRAPHIC RANGE-'; OF mE TAXA morphology of P3' which is in course of reduction in S. nanus but resemblesfue morphology of P3 in S. ar- vernensiS).S. nanus must derive from a European Sus. Text-fig. 3 gives fue stratigrdpruc ranges of fue Thc primitive morphology of its last molars make fue taxa. The f1fStand last Iocalities where a species has decrescentof S. nanus from S. anX!rnensÍSmost prob- been found are given. The biozonations of Mein (1975) able; that i" why fue two are placed together in fue and Augusti et al. (1987) have been used in this dia- cladogram. gr.¡m. Nade 5 - Increase of hypsodonty, loss of P17 Information on fue tempoml mngesof Susstrozzií, reduction of function of P2 and p 3 (neither of fue two Sus anX!rnenSisand KorynocboenlSprottíncialís has ocdudes, P2 has only one roou, shorteningof fue snout been taken and adapted from Guérin & Faure (1985) and 20-25%reduction of absolute linear size. and Faure & Guérin (1984). Somethingshould be said Nade 6 No typical specializationsof this species here about fue stratigmpruc position of fue Tegelen are known. It differs from fue other speciesof ver7UCD- fauna. Guérin 0981) pIaces fue macro-mamrnalfauna sus- group by having charactersthat are all primitive. from Tegelen much higher than fue micro-mammal Nade 7 - Elevation of fue anterior and posterior fauna. The small manunalswere collected from a chan- end" of P3 and P2 and reduction of fue length of fiel wruch had cut into ciar and it was in this ciar that premolar row comparedto molar row. ~ last charnc- fue macro-marnmalswere found. The channel is imme- ter has a parallel in S. scrofa; in this character S. diately overlain by another formativo. For these rea- arvernensisis like the Oícoryphochoerini. sons fue macro-marnmalscannot be younger than fue EUROPEANSlnNAE FROM L. MIOCENE 337
micro-marnmals; dIey must be slightly older (V. d. (without EumatochoenJS)is con-elatedto MN 11 or MN Meulen pers. com.). 12. Sedimentsin fue V2 level have been dated radio- K. palaeocboenlShas been cited from the Bacci- metrically at 8 MY (Hürzeler & Engesser,1976). The nello V3 level (Hünern1ann,1969; Hürzeler & Engesser, M:N 12 Sarnosand Marngheh faunas have also been 1976) which dIe last audIors placed in MN 13 or 14. dated radiometricalIy (Weidmann et al., 1984; Camp- Some flrst upper incisors from this level lack a distal belLet al., 198(). Their agesare 7.35 MY (mean age for cusplet which is normally present in K. paiaeochoerus. theiMain Bone Bem) and 7.6 MY (mean for fue highest Some molars of this level are closer in morphology to level with Microstonyx) respectively. Berggren el al. K. provincialis. Measurementsof dIe dentition of dIe (1985) dated Crevillente VI (MN 13; with fue oldest two species averiar. A more detailed study might SpanishApodemus) 6.5 MY and they date ~ Text-flg. 3 - $tmtigrnflc mnge of the Suinae. 338 J. VANDBRMADE,S.MOYA-s0L4 9 M3 10 9 M3 9 1 ~ 5 Q" . 5 3 . 1 c~ 8~ ti . 2 6 . . 6 . ~ 32 3~ ~2~2 2 #. :;.2 a a . Z 2 2 8 . a . . a B . . c,1fI . ~ 8 . D QD 006 D . p D D. o o Q D~ 0.0\ D 0.0 D Oo. .. ~ ,a " c ..'. O c%c Ób . ~ ~ a a a a a 15 20 25 30 DT Text-fig. 4 - Greatest length (DAP) versus greatest width (OT) of M' and M,. . KOrynocboenlSpa/QeocbCN!1us from Gau Weinhelm (data from HOnennann 1968, excl. no. 1933/833 wl1ich IDÍght be Conobyus).. KQryTI(x;bCN!1usprovInctaIts from various k>ca1ities. Mfcrostony.x ant1qu~ 10) Montrigaud, 9) Eppelshelm (data frorn Hünennann 1968). Mfcrostony.x major maja1: 8) Stratzing (measurernentstaken from figures gIven by Thenius 1972; comparison of dJfferent views indicate that me drnwings are not exacdy to scale), 7) Tenasa, 6) Montredon, 5) Piera, 4) Cucuron (holotype; data frorn GeJVais1848/52), 3) Concud Mtcmstony.x majar erymantbtUor;2) Pikem1i (hokJtype; data from ~ & Wagner 1854), 2) Pikem1i (data from Pea~ 1928), 1) Kerassia. O Eumatochoerusetruscus ff{XrI various localities. . SusanJemensis from various Iocalities. . Susstrozztt from various localities. o Sus nanus ff{XrI Capo Figari. O Sus scrofa, recent and f~il from various Iocalities, including f~iI domestic pigs. EUROPEANSUINAE FROM L. MIOCENE 339 fue level. Even M. majar erymantbius and Apodemus -, 1975,Sus scrofa priscus GoI~ im Pleistozanvon Süssen- did not coexist in Europe. bom bei Weimar: Palaeont Abh., Berlin, A, 3 (3/4): 611-616, 3 JanOssy(1986) reported S. arvernensisfrorn Süro, figs. 1 tab. which is correlated to MN 17. The animal is certainly HÜBZELBR,J. &: ENGESSER,B., 1976, Les faunes de Marnmif'eres Neogenes du Bassin de Baccinello (G~o, ltalle): C. R. present in Etouaires and Hajnacka (MN 16b). Acad Sc. Paris, 283 D: 333-336. It should be noted that Susscrofa arrived in North JANOssY, D., 1986, Plelstocene vertebrate faunas of Hungary, Deve- Africa at about fue same time that it arrived in Europe lopments in Paleontoi