Quaternaire, 18, (1), 2007, p. 65-71

THE PLIOCENE-PLEISTOCENE BOUNDARY: WHICH SIGNIFICANCE FOR THE SO CALLED “WOLF EVENT”? EVIDENCES FROM WESTERN EUROPE Ⅲ

Raffaele SARDELLA1 & Maria Rita PALOMBO1

ABSTRACT

The radiation of canids referable to Canis ex gr. etruscus in Western Europe, the so-called “Wolf Event” by Azzaroli (1983), is an important part of a faunal renewal that occurred approximately at Pliocene-Pleistocene transition, strongly related to climatic and environmental changes. Actually, taking into account the recently available data, the meaning of the so called “Wolf-Event” must be reconsidered in a wider sense, and it has to be considered as different dispersal events occurred in a quite short time span to be placed at the end of Pliocene. The reconsideration of the available fossil record suggests that: 1) The “Wolf Event”, in its original acception, is just one of the different bioevents (dispersal of Canis, Lycaon, but also Pachycrocuta and Panthera) part of the important faunal renewal that affected the carnivore guilds in Western Europe; 2) New hunting strategies (patch hunters) took place in Western Europe at this moment; 3) The dispersal of different lineages of canids has to be related to the diffusion of open habitats; 4) The “Wolf Event”, in a new and wider acception, occurred during latest Pliocene, thus it cannot be considered as a “marker” of the begin- ning of Pleistocene.

Key words: Canids, Carnivora, Richness, Turnovers, Pliocene, Pleistocene, North Western Mediterranean.

RÉSUMÉ

QUELLE IMPORTANCE A LE « WOLF EVENT » POUR LA DÉFINITION DE LA LIMITE ENTRE PLIOCÈNE ET PLÉISTOCÈNE ? LES ÉVIDENCES DONNÉES PAR LES FAUNES DE L’EUROPE SUD OCCIDENTALE La diffusion des espèces du genre Canis est un « bioevent » dont l’importance a été reconnue depuis longtemps. L’apparition de Canis etrus- cus (« Wolf Event ») avait été vue comme le signal du début du Pléistocène. Canis etruscus a été reconnu pour la première fois dans la faune locale de Olivola (Valdarno supérieur, Italie) dont l’âge était considéré comme Pléistocène inférieur. Des découvertes plus récentes nous obligent à placer le « Wolf Event » dans le Pliocène: Canis cf. C. etruscus est présent en Italie à Costa S. Giacomo (MN17) et les trois canidés Villafranchiens (C. etruscus, Canis arnensis et Lycaon falconeri) sont déjà présents dans le Pliocène final à Fonelas. Par ailleurs, dans le Bassin du Valdarno, C. etruscus et C. arnensis ont été trouvés en association avec Chasmaporthethes lunensis et Pachycrocuta brevirostris dans des dépôts d’âge Pliocène final, corrélés avec l’événement paléomagnétique d’Olduvai. Le « Wolf Event » va donc se produire en Méditerranée Nord occidentale au cours du Pliocène supérieur, bien que de façon différente dans chaque région, comme le montre l’observation des richesses spécifiques et des taux d’apparition et d’extinction. De plus, ce bio-évènement n’inclut pas seulement la dispersion de Canis et Lycaon, mais également de Pachycrocuta et, au moins pour l’Italie, de Panthera, produisant un renouvelle- ment important de la guilde des carnivores en Europe occidentale. En conclusion, puisque ce renouvellement va se développer en grande partie durant le Pliocène final, le « Wolf Event », dans sa nouvelle acception plus étendue, ne peut pas être considérer comme un “marqueur” du début du Pléistocène.

Mots clés : Canidés, Carnivores, Richesse spécifique, renouvellement faunique, Pliocène, Pléistocène, Méditerranée Nord occidentale.

1 - INTRODUCTION an important part of a faunal renewal that occurred ap- proximately at Pliocene-Pleistocene transition. The Since the Late Pliocene, a serie of climatic cycles term “WE” initially defined the arrival in Western Eu- have caused latitudinal displacements of vegetational rope of the first representative of the genus Canis cover and biomes in Europe. Terrestrial mammals re- “sensu stricto” and the beginning of the late acted to such climate changes by expanding their diffu- Villafranchian (Olivola Faunal Unit), at the time re- sion area or migrating. The radiation of canids garded as roughly coincident with the beginning of the referable to Canis ex gr. C. etruscus in Western Europe, Early Pleistocene (as formally defined by Aguirre & the so-called “Wolf Event” (WE) by Azzaroli (1983), is Pasini, 1985; see Pillans 2004 and Clague 2006 for a

1 Dipartimento di Scienze della Terra, Università “La Sapienza, CNR - Istituto di Geologia Ambientale e Geoingegneria, Piazzale A. Moro, 5 - 00185 ROMA, Italy. E-mail: [email protected]

Manuscrit reçu le 08/07/2006, accepté le 09/10/2006 66 discussion). In this paper a concise overview of the falconeri. The large sized Lycaon falconeri characte- most recent discoveries is presented together with a re- rised the late Villafranchian assemblages referred to consideration of the biochronological value of impor- Olivola and Tasso FUs, being replaced by the more tant bioevents as suggested by the analyses of the derived Lycaon lycaonoides, recorded in Italy at Pliocene and Pleistocene carnivore guilds in the Medi- Villafranchian-Galerian transition (Pirro & Colle Curti terranean region (Palombo, in press; Palombo et al., in FUs). As stated by Werdelin & Lewis (2005) such a press). phyletic trend seems plausible, although it must be noted that “leaving Lycaon with full generic status ren- ders Canis paraphyletic” and for this reason they prefer 2 - CANIS DISPERSAL IN THE WESTERN to refer these species, as well as the ancestral species of EUROPE AND RELATED ISSUES the lineage “Lycaon” falconeri,toCanis. Only the revi- sion of the genus Canis will enable the researchers to The origin of the genus Canis is matter of debate. clarify such a topic. Many authors agree in suggesting an Asian origin for In France, some dental remains belonging to a me- the genus. An Early Pliocene occurrence of Canis in dium-sized dog from Senèze had been already referred Europe has to be considered with caution, being pre- to Canis cf. C. arnensis by Schaub (1943). 30 years sent in coeval deposits taxa very similar in morphology later, Martin (1973) referred these fossils to the new and size to true dogs, but that can be referred to the ge- species Canis senesensis because of its primitive tooth nus Eucyon (see Spassov & Rook, 2006 for a discus- features. Fossil remains showing great affinities with sion). this species have been recorded even in the late Actually, wolf-like canids dispersal bioevent oc- Pliocene mammal faunas from the early middle curred during the Late Pliocene when such animals, al- Villafranchian Liventsovka and Khapry LFAs (South- ready reported in North China around 3.0 Ma (Mazegu ern Russia) (Sotnikova et al., 2002). At Liventsovka Formation), dispersed from Asia into Europe (Flynn et this canid has been recorded together with the two al., 1991). Turner (1995) pointed out that the genus hyaenas Pachycrocuta brevirostris and Pliocrocuta Canis, represented by several species of jackals, first perrieri, while at Khapry Canis cf. C. senezensis co- appeared in Africa approximately at 2.5 Ma, after exists with P. perrieri. Thus the dog from Senèze could dispersion from Eurasia. be considered as a primitive step of the Canis arnensis Recent discoveries in central Italy (Anagni and lineage (Palombo, in press). Valdarno Basins) provide data for updating the Also recent discoveries in Spain puts in evidence the biochronological framework of such bio-event. early appearance of the three Villafranchian canids in At Costa S. Giacomo, Canis cf. C. etruscus has been Europe. A rich vertebrate fauna with “middle retrieved from a faunal assemblage including several Villafranchian taxa”, such as Gazella borbonica, co- middle Villafranchian taxa (Cassoli & Segre Naldini, mes from Fonelas P1 (Southern Spain) (Arribas et al., 1984, 1993). Size and morphology of those specimens 2004). In this site Canis aff. C. etruscus, Canis aff. (a fragmentary right lower carnassial and two premo- C. arnensis and Lycaon falconeri have been recorded lars) suggest close relationships to the Villafranchian together with three different hyaenids such as wolf-like canids Canis ex gr. C. etruscus (Rook & Pachycrocuta brevirostris, Chasmaporthetes lunensis Torre, 1996). On one hand, new magnetostratigraphic and Parahyaena brunnea. The occurrence of a primi- calibrations have demonstrated a Late Pliocene age for tive form of the living brown hyaena, perhaps a syno- faunas belonging to Olivola FU such as the “late nym of Pliocrocuta perrieri, is currently an open issue. Villafranchian” Poggio Rosso local fauna (LFA), in The appearance of different canids in Europe during which Canis arnensis marks its lower occurrence to- the late Pliocene is testified by an increasing number of gether with Pachycrocuta brevirostris,butwhere fossil evidences and it is worth nothing that not only Chasmaporthetes lunensis is still present (Mazza et al., Canis etruscus but also the primitive wild dog could be 2004). Poggio Rosso is one of the richest and most in- appeared in correspondence of the middle teresting sites in the Upper Valdarno basin, whose Villafranchian faunas (MN17). Indeed at Puebla de latest Pliocene age has been proved by Valverde, some dental remains belonging to a large magnetostratigraphical evidences (upper part of the canid have been referred to Canis cf. C. falconeri Olduvai chron; Napoleone et al., 2001, 2003). Taking (Kurtèn & Crusafont Pairò, 1977). Thus, evidences for into account the recently available data the so called an early occurrence in the Late Pliocene of Europe of a “WE” has to be considered as different dispersal events primitive wild dog are becoming more and more signifi- occurred in a quite short time span to be placed at the cant and numerous even if problematic as well as their end of Pliocene. phylogeography. In the early late Villafranchian faunal assemblages It is worth noting that in Fonelas P1 LFA Panthera (Olivola + Tasso FUs, sensu Palombo, 2005) related to gombazsoegensis has not been recorded yet. Actually, the end of the Olduvai paleomagnetic event, three the origin and dispersal of the Eurasian jaguar canids have been recorded: the wolf-like Canis Panthera gombaszoegensis is still unclear. A very early etruscus, the smaller coyote-like dog Canis arnensis, occurrence of a closed related felid is recorded in and the large sized hypercarnivorous wild dog Lycaon Africa at Laetoli (Upper Unit Laetoli I beds) (Tanzania) 67

(Leakey & Harris, 1987) even if an Asian origin cannot rates, data have been analysed using the methodology be ruled out, taking into account the quite large spread developed by Foote (2000). of P. gombaszoegensis in Near East Asia during the Actually, species richness patterns strictly relate on Pleistocene (Hemmer et al., 2001). Moreover, in East- origination and extinction rates, thus richness changes ern Europe this felid has been recorded at in the Late and turnover patterns are strictly connected. Increases Pliocene site of Slivnitza (Bulgaria) together with or decreases in richness indicate respectively dispersal Canis etruscus. or extinction events, whereas a change in taxonomical Consequently, uncertainty about the migration composition between two successive biochrons can be routes followed by canids, hyaenas and felids still per- regarded as a true faunal turnover when changes in spe- sist. The wolf like and coyote like dogs seems to have cies composition result from the concurrent extinction anAsianoriginaswitnessedbytheoccurrenceof of existing species and replacement by the immigra- closely related taxa in the Pliocene locality of Kuruksai tion/origination of new species. To compute origina- (Tajikistan) (Sotnikova, 1989), even if some uncer- tion and extinction rates, a taxon is assumed to have tainty in the taxonomic position is pointed out by originated within the biochron where it is first ob- Spassov & Rook (2006), who suggest a coyote-like ad- served, while a taxon is assumed to have become aptation for late forms of Eucyon. extinct within its last observed biochron (Foote, 2000). An African origin is possible for the giant hyaena Pachycrocuta brevirostris and for Lycaon, even if wild 3.2 - RESULTS AND DISCUSSION dogs remains have been recorded in Asian localities too (Martinez Navarro & Rook, 2004). Martinez Navarro Trends in richness (fig. 2), and origination/extinction & Palmqvist (1995, 1996) suggested that in the Early rates (fig. 3) are quite different in the examined territo- Pleistocene the African dirk-toothed cat Megantereon ries that are characterized by differences in physiogra- whitei dispersed into Europe, replacing the Pliocene phyaswellasinclime. Megantereon cultridens. The taxonomic position of the Indeed, even during the recent past, environmental Early Pleistocene Megantereon is matter of debate conditions affected in different ways migration and dif- (Pons Moya, 1987; Turner, 1987; Sardella, 1998; fusion of large mammal species into the North-western Hemmer, 2001). A possible African origin for the Early Mediterranean region. Pleistocene Megantereon is a crucial point in valuing the turnover dynamic but not as regard the “turnover” Therefore, new occurrences in the Iberian or Italian per se. peninsulas (if not related to endemic speciation) re- sulted from a previous route through France and central As a matter of fact, during the Late Pliocene different Europe. For this reason, French mammalian faunal species of canids, hyaenids and felids dispersed into complexes should been more affected by immigration the Mediterranean region, greatly changing the compo- waves from central Europe, as well as by taxa sition and the structure of the guild of the carnivores. latitudinal migrations. Such a framework seems to fit quite well the main re- sult of our analysis: both richness and origin/extinction 3 - CARNIVORA GUILD OF THE NORTH ratios have quite regular trends for Spain, articulated WESTERN MEDITERRANEAN: trends for Italy, and intermediate patterns for France. DIVERSITY AND TURNOVER Such trend differences seem to be independent by the FROM THE MIDDLE PLIOCENE quality of fossil record, even if it strongly affected any TO THE EARLY PLEISTOCENE analyses. In fact, although faunal complexes have been taken into account in the present paper instead of local faunal assemblages, fossil record gaps occurring in dif- 3.1 - METHODS ferent time spans and for different countries affect the detailed interpretation of data, but not the general Carnivora complexes detected according to the framework. biochronological scheme proposed by Palombo (in On the other hand, our results underline a moderate press) (fig. 1) have been analysed in terms of richness richness decrease at the Middle to Late Pliocene transi- as well as origination and extinction rates. tion (early to middle Villafranchian). A faunal renewal Richness can be measured by the total number of has been already noted for the whole faunal data and taxa actually or potentially occurring in each for herbivores. It has been related to the Pliocene cli- biochronological interval. Nevertheless, the richness mate worsening that followed the permanent Arctic of a single time interval may be overestimated when glacial cap onset (Alberdi et al., 1997; Azanza et al., first local appearances have been considered as occur- 1999, 2000; Palombo 2004, in press). As a matter of ring at the beginning of the interval and the taxa that fact, the renewal in the carnivore guild took place later disappeared as persistent up to the end of it, whereas than that of the herbivore one, and just in the Late Plio- they might not actually overlap in time. In order to re- cene such a guild is modified by appearances and duce this bias, and since taxonomic diversity relates to extinctions (Palombo, in press). It confirms the impor- origination/immigration and extinction/emigration tance of the structural changes that characterized the 68

Fig. 1: Biochronological range chart of selected Carnivora from North Western Mediterranean (biochronological framework from Palombo, 2005, in press) dating from the middle Pliocene to the early Pleistocene. Fig. 1 : Comparaison de la distribution chronologique durant le Pliocène moyen et supérieur et le Pléistocène inférieur des Carnivores d’Espagne, France et Italie (cadre biochronologique d’après Palombo, 2005, sous presse). 69

20 2

18 2

16 Spain 2 Spain

14 1

12 1

10 1

8 1

6 1

4 0

2 0

0 0 0,8 1,1 1,4 1,7 2 2,3 2,6 2,9 Ma 3,2 0,8 1,1 1,4 1,7 2 2,3 2,6 2,9Ma 3,2 Total Diversity Total Diversity minus singletons Per-taxon Origination rate Per-taxon Extinction rate Estimated mean standing diversity

20 2 18 France 2 France 16 2

14 1

12 1

10 1

8 1

6 1

4 0

2 0

0 0

0,8 1,1 1,4 1,7 2 2,3 2,6 2,9Ma 3,2 0,8 1,1 1,4 1,7 2 2,3 2,6 2,9Ma 3,2 Total Diversity Total Diversity minus singletons Per-taxon Origination rate Per-taxon Extinction rate Estimated mean standing diversity

20 2 18 Italy 2 Italy 16 2

14 1

12 1

10 1

8 1

6 1

4 0

2 0

0 0 0,8 1,1 1,4 1,7 2 2,3 2,6 2,9Ma 3,2 0,8 1,1 1,4 1,7 2 2,3 2,6 2,9Ma 3,2 Total Diversity Total Diversity minus singletons Per-taxon Origination rate Per-taxon Extinction rate Estimated mean standing diversity

Fig. 2 : “Total Diversity” and “Estimated mean standing diversity” Fig. 3: Per-taxon rates of origination and extinction of Spanish, of Spanish, French and Italian Carnivora calculated following French and Italian Carnivora calculated following Foote (2000). Foote (2000). Per-taxon origination rate = NFL + NFt)/Ntot/Δt

Total Diversity (Ntot)=NFL +NbL +NFt +Nbt; Estimated mean standing Per-taxon extinction rate = (NFL + NbL)/Ntot/Δt diversity (N ) = N – N /2 – N /2. esd tot o e NFL = taxa that exist only in the interval; NFt = taxa originate in the interval Number of originations (N )=N +N; Number of extinctions (N )= o FL Ft e and persisting beyond it; NbL = taxa that originate before the interval but NFL +NbL. go extinct within it; Δt = time interval. NFL = taxa that exist only in the interval; NbL = taxa that originate before Fig. 3 : Taux d’apparition et d’extinction «Per-taxon» des Carnivores the interval but go extinct within it; NFt = taxa that originate in the inter- d’Espagne, France et Italie calculés d’après Foote (2000). val and persisting beyond it; N = taxa that originate before the interval bt Taux d’apparition = NFL + NFt) /Ntot/ t ; taux d’extinction = (NFL + and persisting beyond it. NbL) /Ntot/ t. Fig. 2 : « Diversité totale » et « Diversité estimée » des Carnivores NFL = taxons qui existent seulement dans l’intervalle ; NFt = taxons qui d’Espagne, France et Italie calculées d’après Foote (2000). apparaissent durant l’intervalle et qui persistent durant l’intervalle sui- Diversité totale (Ntot) = NFL + NbL + NFt + Nbt ; Diversité estimée vant ; NbL = taxons qui apparaissent avant l’intervalle et disparaissent (Nesd) = Ntot – No/2 – Ne/2. durant l’intervalle, t = intervalle de temps. Nombre d’apparitions (No)=(No)=NFL +NFt ; Nombre d’extinctions

(Ne)= NFL +NbL. NFL = taxons qui existent seulement dans l’intervalle ; NbL = taxons qui apparaissent avant l’intervalle et disparaissent durant l’intervalle ; 1988; Palmqvist et al., 1999) seems to be more closely NFt = taxons qui apparaissent durant l’intervalle et qui persistent du- related to migrations affecting the guild of the rant l’intervalle suivant ; Nbt = taxons qui apparaissent avant l’inter- herbivores, than to environmental changes. valle et persistent durant l’intervalle suivant. The different temporal extension of decreasing in di- versity that affected the considered countries during whole North-western Mediterranean region at the be- the late Early Pleistocene relays on local disappear- ginning of Early Pleistocene, with Pachycrocuta ances of long survivor carnivores of Pliocene origin brevirostris and Canis etruscus as common elements. that occurred in the studied territories different times. Indeed, in France, Panthera gombaszoegensis is first For instance, the large felid Megantereon has been lat- recorded only in the latest Early Pleistocene LFAs (for est recorded at Pirro Nord in Italy and at Venta Micena instance Le Vallonnet) (Moullé, 1998), and Lycaon in Spain (Sardella, 1998) and disappeared in such re- falconeri and C. arnensis seems to be missing, even if gions before the Jaramillo event; Puma, that has been the latter has a closely related form (possibly co-spe- never recorded in Italy and was present in Spain and in cific) in Canis senezensis from Seneze (see above). As France during the Late Pliocene, survived in France a matter of fact, a new guild structure progressively (Le Vallonnet) (Hemmer et al., 2004) and central Eu- took place, with prevailing middle-sized and large car- rope (Untermassfeld) (Hemmer, 2001) in LFAs corre- nivores. The “WE” (Azzaroli, 1983; Azzaroli et al., lated to Jaramillo event. Actually, the occurrence at Le 70

Vallonnet of the puma is matter of debate, being the Upper Valdarno (Sardella, 1998). Accordingly, the re- fossils (2 lower carnassials) referred to Panthera newal guild occurred in a rather gradual way during the pardus by Moullé et al. (in press). In our opinion the latest Pliocene and the guild of large carnivores is al- limited recorded sample and its morphometrical ready definitely modified and in a new equilibrium at variability leave such taxonomical issue unsolved at the beginning of the Pleistocene. the moment. The results obtained support the argument that the During the late early Pleistocene, the reduction in “WE”, in a new and wider acception, actually includes biodiversity among canids bases more on the disap- different bioevents affecting the carnivore guilds in pearance of Lycaon, that survived until the end of the Western Europe (dispersal of Canis, Lycaon,butalso Early Pleistocene at Le Vallonnet and Colle Curti (Mar- Pachycrocuta and, at least in Italy, Panthera), thus it tinez-Navarro & Rook, 2003), than of Canis arnensis cannot be considered as a “marker” of the beginning of (latest recorded at Upper Valdarno) (Rook, 1993) be- Pleistocene, but, on the contrary, different quite cause canids of the group of Canis mosbachensis par- diachronic bioevents with the dispersal of different car- tially occupied the niche of the later species. The “end nivore taxa. Among them the most representative dis- Early Pleistocene” biodiversity decrease of Italian car- persion in Europe can be considered that of nivores (where it approximates the Jaramillo event) it Pachycrocuta brevirostris. possibly as consequence of the ongoing decrease of herbivores recorded at the beginning of Early Pleisto- cene (Palombo, 2005, unpublished data). In France, on ACKNOWLEDGEMENTS the other hand, in the LFAs related to Jaramillo, to- gether with the Villafranchian long survivor taxa (e.g. We wish to thank R.-D. Kahlke and P.-E. Moullé for the critical Puma, Acinonyx and Lycaon) both taxa found in France reading of the manuscript and the useful suggestions. Work supported for the first time (e.g. P. gombazsoegensis) and newly by MIUR 60% grants 2005 Prof. M.R. Palombo (“Evoluzione degli ecosistemi continentali dell’Italia peninsulare durante il Plio-Pleisto- appeared ones (possibly U. dolinensis, Garcia & cene: un approccio multidisciplinare”). 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