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The Pliocene-Pleistocene boundary : which significance for the so called “Wolf Event”? Evidences from Western Europe

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
Raffaele Sardella et Maria Rita Palombo
p. 65-71

Résumés

La diffusion des espèces du genre Canis est un « bioevent » dont l’importance a été reconnue depuis longtemps. L’apparition de Canis etruscus (« 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 renouvellement 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.

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Texte intégral

We wish to thank R.-D. Kahlke and P.-E. Moullé for the critical reading of the manuscript and the useful suggestions. Work supported by MIUR 60% grants 2005 Prof. M.R. Palombo (“Evoluzione degli ecosistemi continentali dell’Italia peninsulare durante il Plio-Pleistocene: un approccio multidisciplinare”).

1- Introduction

1Since the Late Pliocene, a serie of climatic cycles have caused latitudinal displacements of vegetational cover and biomes in Europe. Terrestrial mammals reacted to such climate changes by expanding their diffusion area or migrating. The radiation of canids referable to Canis ex gr. C. etruscus in Western Europe, the so-called “Wolf Event” (WE) by Azzaroli (1983), is an important part of a faunal renewal that occurred approximately at Pliocene-Pleistocene transition. The term “WE” initially defined the arrival in Western Europe of the first representative of the genus Canis sensu stricto” and the beginning of the late Villafranchian (Olivola Faunal Unit), at the time regarded as roughly coincident with the beginning of the Early Pleistocene (as formally defined by Aguirre & Pasini, 1985; see Pillans 2004 and Clague 2006 for a discussion). In this paper a concise overview of the most recent discoveries is presented together with a reconsideration of the biochronological value of important bioevents as suggested by the analyses of the Pliocene and Pleistocene carnivore guilds in the Mediterranean region (Palombo, in press; Palombo et al., in press).

2- Canis dispersal in the Western Europe and related issues

2The origin of the genus Canis is matter of debate. Many authors agree in suggesting an Asian origin for the genus. An Early Pliocene occurrence of Canis in Europe has to be considered with caution, being present in coeval deposits taxa very similar in morphology and size to true dogs, but that can be referred to the genus Eucyon (see Spassov & Rook, 2006 for a discussion).

3Actually, wolf-like canids dispersal bioevent occurred during the Late Pliocene when such animals, already reported in North China around 3.0 Ma (Mazegu Formation), dispersed from Asia into Europe (Flynn et al., 1991). Turner (1995) pointed out that the genus Canis, represented by several species of jackals, first appeared in Africa approximately at 2.5 Ma, after dispersion from Eurasia.

4Recent discoveries in central Italy (Anagni and Valdarno Basins) provide data for updating the biochronological framework of such bio-event.

5At Costa S. Giacomo, Canis cf. C. etruscus has been retrieved from a faunal assemblage including several middle Villafranchian taxa (Cassoli & Segre Naldini, 1984, 1993). Size and morphology of those specimens (a fragmentary right lower carnassial and two premolars) suggest close relationships to the Villafranchian wolf-like canids Canis ex gr. C. etruscus (Rook & Torre, 1996). On one hand, new magnetostratigraphic calibrations have demonstrated a Late Pliocene age for faunas belonging to Olivola FU such as the “late Villafranchian” Poggio Rosso local fauna (LFA), in which Canis arnensis marks its lower occurrence together with Pachycrocuta brevirostris, but where Chasmaporthetes lunensis is still present (Mazza et al., 2004). Poggio Rosso is one of the richest and most interesting sites in the Upper Valdarno basin, whose latest Pliocene age has been proved by magnetostratigraphical evidences (upper part of the Olduvai chron; Napoleone et al., 2001, 2003). Taking into account the recently available data the so called “WE” has to be considered as different dispersal events occurred in a quite short time span to be placed at the end of Pliocene.

6In the early late Villafranchian faunal assemblages (Olivola + Tasso FUs, sensu Palombo, 2005) related to the end of the Olduvai paleomagnetic event, three canids have been recorded: the wolf-like Canis etruscus, the smaller coyote-like dog Canis arnensis, and the large sized hypercarnivorous wild dog Lycaon falconeri. The large sized Lycaon falconeri characterised the late Villafranchian assemblages referred to Olivola and Tasso FUs, being replaced by the more derived Lycaon lycaonoides, recorded in Italy at Villafranchian-Galerian transition (Pirro & Colle Curti FUs). As stated by Werdelin & Lewis (2005) such a phyletic trend seems plausible, although it must be noted that “leaving Lycaon with full generic status renders Canis paraphyletic” and for this reason they prefer to refer these species, as well as the ancestral species of the lineage “Lycaonfalconeri, to Canis. Only the revision of the genus Canis will enable the researchers to clarify such a topic.

7In France, some dental remains belonging to a medium-sized dog from Senèze had been already referred to Canis cf. C. arnensis by Schaub (1943). 30 years later, Martin (1973) referred these fossils to the new species Canis senesensis because of its primitive tooth features. Fossil remains showing great affinities with this species have been recorded even in the late Pliocene mammal faunas from the early middle Villafranchian Liventsovka and Khapry LFAs (Southern Russia) (Sotnikova et al., 2002). At Liventsovka this canid has been recorded together with the two hyaenas Pachycrocuta brevirostris and Pliocrocuta perrieri, while at Khapry Canis cf. C. senezensis coexists with P. perrieri. Thus the dog from Senèze could be considered as a primitive step of the Canis arnensis lineage (Palombo, in press).

8Also recent discoveries in Spain puts in evidence the early appearance of the three Villafranchian canids in Europe. A rich vertebrate fauna with “middle Villafranchian taxa”, such as Gazella borbonica, comes from Fonelas P1 (Southern Spain) (Arribas et al., 2004). In this site Canis aff. C. etruscus, Canis aff. C. arnensis and Lycaon falconeri have been recorded together with three different hyaenids such as Pachycrocuta brevirostris, Chasmaporthetes lunensis and Parahyaena brunnea. The occurrence of a primitive form of the living brown hyaena, perhaps a synonym of Pliocrocuta perrieri, is currently an open issue. The appearance of different canids in Europe during the late Pliocene is testified by an increasing number of fossil evidences and it is worth nothing that not only Canis etruscus but also the primitive wild dog could be appeared in correspondence of the middle Villafranchian faunas (MN17). Indeed at Puebla de Valverde, some dental remains belonging to a large canid have been referred to Canis cf. C. falconeri (Kurtèn & Crusafont Pairò, 1977). Thus, evidences for an early occurrence in the Late Pliocene of Europe of a primitive wild dog are becoming more and more significant and numerous even if problematic as well as their phylogeography.

9It is worth noting that in Fonelas P1 LFA Panthera gombazsoegensis has not been recorded yet. Actually, the origin and dispersal of the Eurasian jaguar Panthera gombaszoegensis is still unclear. A very early occurrence of a closed related felid is recorded in Africa at Laetoli (Upper Unit Laetoli I beds) (Tanzania) (Leakey & Harris, 1987) even if an Asian origin cannot be ruled out, taking into account the quite large spread of P. gombaszoegensis in Near East Asia during the Pleistocene (Hemmer et al., 2001). Moreover, in Eastern Europe this felid has been recorded at in the Late Pliocene site of Slivnitza (Bulgaria) together with Canis etruscus.

10Consequently, uncertainty about the migration routes followed by canids, hyaenas and felids still persist. The wolf like and coyote like dogs seems to have an Asian origin as witnessed by the occurrence of closely related taxa in the Pliocene locality of Kuruksai (Tajikistan) (Sotnikova, 1989), even if some uncertainty in the taxonomic position is pointed out by Spassov & Rook (2006), who suggest a coyote-like adaptation for late forms of Eucyon.

11An African origin is possible for the giant hyaena Pachycrocuta brevirostris and for Lycaon, even if wild dogs remains have been recorded in Asian localities too (Martinez Navarro & Rook, 2004). Martinez Navarro & Palmqvist (1995, 1996) suggested that in the Early Pleistocene the African dirk-toothed cat Megantereon whitei dispersed into Europe, replacing the Pliocene Megantereon cultridens. The taxonomic position of the Early Pleistocene Megantereon is matter of debate (Pons Moya, 1987; Turner, 1987; Sardella, 1998; Hemmer, 2001). A possible African origin for the Early Pleistocene Megantereon is a crucial point in valuing the turnover dynamic but not as regard the “turnover” per se.

12As a matter of fact, during the Late Pliocene different species of canids, hyaenids and felids dispersed into the Mediterranean region, greatly changing the composition and the structure of the guild of the carnivores.

3- Carnivora guild of the northwestern mediterranean : diversity and turnover from the middle Pliocene to the early Pleistocene

3.1- Methods

13Carnivora complexes detected according to the biochronological scheme proposed by Palombo (in press) (fig. 1) have been analysed in terms of richness as well as origination and extinction rates.

14Richness can be measured by the total number of taxa actually or potentially occurring in each biochronological interval. Nevertheless, the richness of a single time interval may be overestimated when first local appearances have been considered as occurring at the beginning of the interval and the taxa that disappeared as persistent up to the end of it, whereas they might not actually overlap in time. In order to reduce this bias, and since taxonomic diversity relates to origination/immigration and extinction/emigration rates, data have been analysed using the methodology developed by Foote (2000).

15Actually, species richness patterns strictly relate on origination and extinction rates, thus richness changes and turnover patterns are strictly connected. Increases or decreases in richness indicate respectively dispersal or extinction events, whereas a change in taxonomical composition between two successive biochrons can be regarded as a true faunal turnover when changes in species composition result from the concurrent extinction of existing species and replacement by the immigration/origination of new species. To compute origination and extinction rates, a taxon is assumed to have originated within the biochron where it is first observed, while a taxon is assumed to have become extinct within its last observed biochron (Foote, 2000).

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: 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.

3.2 - Results and discussion

16Trends in richness (fig. 2), and origination/extinction rates (fig. 3) are quite different in the examined territories that are characterized by differences in physiography as well as in clime.

17Indeed, even during the recent past, environmental conditions affected in different ways migration and diffusion of large mammal species into the North-western Mediterranean region.

18Therefore, new occurrences in the Iberian or Italian peninsulas (if not related to endemic speciation) resulted from a previous route through France and central Europe. For this reason, French mammalian faunal complexes should been more affected by immigration waves from central Europe, as well as by taxa latitudinal migrations.

19Such a framework seems to fit quite well the main result of our analysis: both richness and origin/extinction ratios have quite regular trends for Spain, articulated trends for Italy, and intermediate patterns for France. Such trend differences seem to be independent by the quality of fossil record, even if it strongly affected any 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 different time spans and for different countries affect the detailed interpretation of data, but not the general framework.

20On the other hand, our results underline a moderate richness decrease at the Middle to Late Pliocene transition (early to middle Villafranchian). A faunal renewal has been already noted for the whole faunal data and for herbivores. It has been related to the Pliocene climate worsening that followed the permanent Arctic glacial cap onset (Alberdi et al., 1997; Azanza et al., 1999, 2000; Palombo 2004, in press). As a matter of fact, the renewal in the carnivore guild took place later than that of the herbivore one, and just in the Late Pliocene such a guild is modified by appearances and extinctions (Palombo, in press). It confirms the importance of the structural changes that characterized the whole North-western Mediterranean region at the beginning of Early Pleistocene, with Pachycrocuta brevirostris and Canis etruscus as common elements. Indeed, in France, Panthera gombaszoegensis is first recorded only in the latest Early Pleistocene LFAs (for instance Le Vallonnet) (Moullé, 1998), and Lycaon falconeri and C. arnensis seems to be missing, even if the latter has a closely related form (possibly co-specific) in Canis senezensis from Seneze (see above). As a matter of fact, a new guild structure progressively took place, with prevailing middle-sized and large carnivores. The “WE” (Azzaroli, 1983; Azzaroli et al., 1988; Palmqvist et al., 1999) seems to be more closely related to migrations affecting the guild of the herbivores, than to environmental changes.

21The different temporal extension of decreasing in diversity that affected the considered countries during the late Early Pleistocene relays on local disappearances of long survivor carnivores of Pliocene origin that occurred in the studied territories different times. For instance, the large felid Megantereon has been latest recorded at Pirro Nord in Italy and at Venta Micena in Spain (Sardella, 1998) and disappeared in such regions before the Jaramillo event; Puma, that has been never recorded in Italy and was present in Spain and in France during the Late Pliocene, survived in France (Le Vallonnet) (Hemmer et al., 2004) and central Europe (Untermassfeld) (Hemmer, 2001) in LFAs correlated to Jaramillo event. Actually, the occurrence at Le Vallonnet of the puma is matter of debate, being the fossils (2 lower carnassials) referred to Panthera pardus by Moullé et al. (in press). In our opinion the limited recorded sample and its morphometrical variability leave such taxonomical issue unsolved at the moment.

22During the late early Pleistocene, the reduction in biodiversity among canids bases more on the disappearance of Lycaon, that survived until the end of the Early Pleistocene at Le Vallonnet and Colle Curti (Martinez-Navarro & Rook, 2003), than of Canis arnensis (latest recorded at Upper Valdarno) (Rook, 1993) because canids of the group of Canis mosbachensis partially occupied the niche of the later species. The “end Early Pleistocene” biodiversity decrease of Italian carnivores (where it approximates the Jaramillo event) it possibly as consequence of the ongoing decrease of herbivores recorded at the beginning of Early Pleistocene (Palombo, 2005, unpublished data). In France, on the other hand, in the LFAs related to Jaramillo, together with the Villafranchian long survivor taxa (e.g. Puma, Acinonyx and Lycaon) both taxa found in France for the first time (e.g. P. gombazsoegensis) and newly appeared ones (possibly U. dolinensis, Garcia & Arsuaga, 2001) can be recorded (Moullé, 1998; Hemmer et al., 2004; Martinez Navarro & Rook, 2003; Palombo & Valli, 2004). In Spain too, the Eurasian jaguar and the primitive brown bear are recorded in slightly younger LFAs. Differences in trends can be due to the scantiness of well documented Early Pleistocene French LFAs. In Italy, LFAs referable to the Jaramillo event are quite rare, with an overestimation of uncommon species disappearances.

23Accordingly the major guild reorganisation occurred during the latest Pliocene and the latest Early Pleistocene.

Fig. 2 : “Total Diversity” and “Estimated mean standing diversity” of Spanish, French and Italian Carnivora calculated following Foote (2000).

Fig. 2 : “Total Diversity” and “Estimated mean standing diversity” of Spanish, French and Italian Carnivora calculated following Foote (2000).

Total Diversity (Ntot) = NFL + NbL+ NFt+ Nbt; Estimated mean standing diversity (Nesd) = Ntot – No/2 – Ne/2. Number of originations (No) = NFL + NFt; Number of extinctions (Ne) = NFL +NbL.NFL = taxa that exist only in the interval; NbL = taxa that originate before the interval but go extinct within it; NFt = taxa that originate in the interval and persisting beyond it; Nbt = taxa that originate before the interval and persisting beyond it.

Fig. 3: Per-taxon rates of origination and extinction of Spanish, French and Italian Carnivora calculated following Foote (2000).

Fig. 3: Per-taxon rates of origination and extinction of Spanish, French and Italian Carnivora calculated following Foote (2000).

Per-taxon origination rate = NFL + NFt)/Ntot/Δt Per-taxon extinction rate = (NFL + NbL)/Ntot/Δt NFL = taxa that exist only in the interval;NFt= taxa originate in the interval and persisting beyond it; NbL = taxa that originate before the interval but go extinct within it; Δt = time interval

4 - Conclusion: which biochronological significance for the already called “Wolf Event”?

24From the above mentioned issues a question arises: the renewal of the carnivore guild can be related to the climate change occurred at the Plio-Pleistocene transition? Is the “WE” a bioevent marking the Plio-Pleistocene transition?

25At the beginning of Pleistocene, the Carnivora’s guild is extensively reconstructed especially as regards to cooperative foraging canids and hyaenids. Pliocrocuta perrieri and Chamaporthetes lunensis disappeared whereas the powerful scavenger Pachycrocuta brevirostris became more and more frequent, the social canids are already present and spread in the grassland/savanna environments. On the other hand, among the large felids, Acinonyx pardinensis and possibly Puma pardoides persisted, while the new incomer Panthera gombaszoegensis inhabited more close environments together with Megantereon cultridens still present at least in the late Villafranchian faunas of Upper Valdarno (Sardella, 1998). Accordingly, the renewal guild occurred in a rather gradual way during the latest Pliocene and the guild of large carnivores is already definitely modified and in a new equilibrium at the beginning of the Pleistocene.

26The results obtained support the argument that the “WE”, in a new and wider acception, actually includes different bioevents affecting the carnivore guilds in Western Europe (dispersal of Canis, Lycaon, but also Pachycrocuta and, at least in Italy, Panthera), thus it cannot be considered as a “marker” of the beginning of Pleistocene, but, on the contrary, different quite diachronic bioevents with the dispersal of different carnivore taxa. Among them the most representative dispersion in Europe can be considered that of Pachycrocuta brevirostris.

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Table des illustrations

Titre 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.
URL http://journals.openedition.org/quaternaire/docannexe/image/969/img-1.jpg
Fichier image/jpeg, 110k
Titre Fig. 2 : “Total Diversity” and “Estimated mean standing diversity” of Spanish, French and Italian Carnivora calculated following Foote (2000).
Légende Total Diversity (Ntot) = NFL + NbL+ NFt+ Nbt; Estimated mean standing diversity (Nesd) = Ntot – No/2 – Ne/2. Number of originations (No) = NFL + NFt; Number of extinctions (Ne) = NFL +NbL.NFL = taxa that exist only in the interval; NbL = taxa that originate before the interval but go extinct within it; NFt = taxa that originate in the interval and persisting beyond it; Nbt = taxa that originate before the interval and persisting beyond it.
URL http://journals.openedition.org/quaternaire/docannexe/image/969/img-2.jpg
Fichier image/jpeg, 45k
Titre Fig. 3: Per-taxon rates of origination and extinction of Spanish, French and Italian Carnivora calculated following Foote (2000).
Légende Per-taxon origination rate = NFL + NFt)/Ntot/Δt Per-taxon extinction rate = (NFL + NbL)/Ntot/Δt NFL = taxa that exist only in the interval;NFt= taxa originate in the interval and persisting beyond it; NbL = taxa that originate before the interval but go extinct within it; Δt = time interval
URL http://journals.openedition.org/quaternaire/docannexe/image/969/img-3.jpg
Fichier image/jpeg, 41k
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Pour citer cet article

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Raffaele Sardella et Maria Rita Palombo, « The Pliocene-Pleistocene boundary : which significance for the so called “Wolf Event”? Evidences from Western Europe »Quaternaire, vol. 18/1 | 2007, 65-71.

Référence électronique

Raffaele Sardella et Maria Rita Palombo, « The Pliocene-Pleistocene boundary : which significance for the so called “Wolf Event”? Evidences from Western Europe »Quaternaire [En ligne], vol. 18/1 | 2007, mis en ligne le 01 mars 2007, consulté le 17 avril 2025. URL : http://journals.openedition.org/quaternaire/969 ; DOI : https://doi.org/10.4000/quaternaire.969

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Auteurs

Raffaele Sardella

Dipartimento di Scienze della Terra, Università “La Sapienza, CNR - Istituto di Geologia Ambientale e Geoingegneria, Piazzale A. Moro, 5 - 00185 ROMA, Italy. E-mail: raffaele.sardella@uniroma1.it

Maria Rita Palombo

Dipartimento di Scienze della Terra, Università “La Sapienza, CNR - Istituto di Geologia Ambientale e Geoingegneria, Piazzale A. Moro, 5 - 00185 ROMA, Italy. E-mail: mariarita.palombo@uniroma1.it

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