Palaeontographica, Abt. B: Palaeozoology – Stratigraphy Vol. 298, Issues 1–6: 169–181 Stuttgart, March 2013
Article
Axis eurygonos from Pirro Nord (Apricena, Southern Italy) by
Carmelo Petronio, Luca Bellucci and Giuseppe Di Stefano With 11 text-figures
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Zusammenfassung
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Häufige mittelgroße Cerviden Reste aus verschiedenen Karsthöhlen neben Pirro Nord (bei Apricena, Apulien Süditalien) werden hier beschrieben. Die morphologische und biometrische Analyse weist auf die Zugehörigkeit der Art Axis eurygonos (Azzaroli). Unsere Cerviden Reste unterscheiden sich von den mediterranen Populationen Europas nur durch ihre geringere Größe. Es werden dann auch die morphometrischen Unterschiede zwischen unserer Art und Rusa rhenanus aus Kontinentaleuropa markiert. Letzterer Art werden auch die vielen Reste aus Untermaßfeld zugeschrieben, welche vorher als “Cervus” nestii vallonetensis klassifiziert wurden.
Summary
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Schlüsselwörter: Oberes Villafranchian – Axis eurygonos – Rusa rhenanus
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The numerous medium-sized cervid remains collected in different times from some karst cavities of Pirro Nord (Apricena, Apulia, Southern Italy) are studied in this paper. The morphological and biometrical analyses suggest the reference of these remains to Axis eurygonos (Azzaroli), the only difference compared to the populations from Mediterranean Europe can be observed in the small size of these deer. The morphometrical differences between the bones of this species and those referable to Rusa rhenanus from Western Europe have been investigated, the numerous deer remains from Untermaßfeld, previuosly referred to “Cervus” nestii vallonetensis are attributed to this last species.
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Key words: Late Villafranchian – Axis eurygonos – Rusa rhenanus
Riassunto
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Sono analizzati in questo lavoro gli abbondanti resti di cervidi di media taglia rinvenuti in diversi tempi e in varie cavità carsiche di Pirro Nord, nell’area di Apricena (Puglia, Italia del Sud). L’analisi morfologica e biometrica indica l’appartenenza di questi resti ad Axis eurygonos (Azzaroli), le uniche differenze rispetto alle popolazioni dell’Europa mediterranea sembrano concentrarsi nelle modeste dimensioni di questi cervi. Vengono evidenziate anche le differenze morfometriche fra le ossa di questa specie e quelle di Rusa rhenanus dell’Europa continentale, cui vengono anche attribuiti i numerosi resti rinvenuti a Untermaßfeld, classificati precedentemente come “Cervus” nestii vallonetensis. Parole chiave: Villafranchiano superiore – Axis eurygonos – Rusa rhenanus
Table of Contents
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Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Short historical notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morphological analysis of the medium sized deer remains from Pirro Nord (Apricena, Southern Italy) . . . . . . . . . . . . . . Skull and antler remains . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
170 170 171 172
Postcranial skeleton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biometrical analysis of the remains from Pirro Nord . . . . Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
174 174 175 180 181
Address of the authors: Carmelo Petronio, Luca Bellucci and Giuseppe Di Stefano, Earth Science Department, Università “Sapienza” di Roma. P.le A .Moro 5, 00185 Roma, e-mail:
[email protected]
© 2013 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany
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Medium-sized cervid remains, similar to the living fallow deer are rather numerous during the last two millions years in the mammal faunal associations from Mediterranean Europe and in the Italian peninsula. On these forms, which grow into a considerable importance during time both in the biochonological and palaeobiogeographical point of views, a particular discussion has developed: this seems to be connected only apparently to a simple taxonomical problem but, on the contrary, it regards very important philogenetical and palaeobiogeographical questions. Therefore, the recognition of the synonymy between the genera Axis and Pseudodama (Di Stefano & Petronio 1998, 2002), as briefly exposed in the following historical notes, contributed to clarify a very precise biochronological sequence for the Mediterranean Europe, to simplify the various morphological features (above all these recognizable in the antlers), and to explain the philogenetical relationships among the living Asian genera and the Pleistocene European ones, also verifying the possible pathways of migrations of these cervids from Asia to the Mediterranean areas. The abundant medium sized deer remains collected in many cavities in the area of Apricena containing Villafranchian faunas have to be related with this picture and these problems. Besides to confirm the aspect previously mentioned, another purpose of this paper is to recognize the possible morphological and/or biometrical differences with the remains of deer populations collected from other Italian and European localities.
the antler morphologies for the position of the terminal fork, for the different length of the b span and, finally, for the angle between the brow tine and the beam. Moreover, the same author also suggested that the typical form of the subspecies eurygonos only occurred in the remains from Upper Valdarno and it was not connected to the subspecies nestii by any intermediate form. According to Azzaroli (1947, 1953), in the locality of Olivola the remains referable to the subspecies nestii dominates on those of the subspecies eurygonos which occurred in a not typical form. Finally, the author drew a philogenesis noting some affinities among these more evoluted forms and the primitive form of Cervavitus novorossiae (Khomenko 1913), he underlined some differences with the living fallow deer in the antlers and in the skull, also noting that the antlers of the subspecies eurygonos could “prelude” those of the living species. Heintz (1970) used the generical term “Cervus” s.l. to indicate the Villafranchian cervids from French and Spanish localities similar in size to the living fallow deer and he grouped them into the lineage C. pardinensis – C. philisi – C. perolensis. These species were considered by the author as subsequent stages of the same philetical lineage. Petronio (1979) analyzed the deer remains from Capena (Roma) and confirmed the philogenesis and the descriptions proposed by Azzaroli (1947), he also pointed out the occurrence of Dama nestii eurygonos in Late Villafranchian faunal assemblages. De Lumley et al. (1988) and Moullé (1990), referred the medium-sized cervid remains and their basal parts of antlers from Le Vallonet (Southern France) to “Cervus” (s.l.) nestii vallonetensis. Azzaroli (1992) recognized the synonymy between C. philisi and C. rhenanus Dubois, 1905, whose remains were described by Bernsen (1932) and by Kunst (1937). C. philisi was considered by the author as “junior synonym” of C. rhenanus and Dama nestii Azzaroli, 1947, could be included in this philetical lineage. The same author noted that Forsyth Mayor never described or figured Cervus nestii, therefore this is a nomen nuduum and, according to the rules of zoological taxonomy, the author of the species nestii must be Azzaroli 1947. In the same paper Azzaroli (1992) proposed the new genus Pseudodama, to make evident the different antler morphologies. The European species pardinensis, rhenanus, and perolensis, and the Italian species nestii were included into the genus Pseudodama
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Introduction
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Short historical notes
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The first remains referable to Dama-like cervids in the Italian peninsula have been described by Forsyth Mayor in 1879. While examining summarily the deer from Upper Valdarno the same author distinguished five or six species that cited in some faunal lists from Upper Valdarno (1885) and from Olivola (1890). One of these species, named Cervus nestii by the author in honour of Filippo Nesti who for the first time described the fossils from Valdarno, was subsequently included into the genus Dama by Schlosser (1924). Azzaroli (1947), reappraised these remains, confirmed the hypothesis of Schlosser and referred the forms from Valdarno to two new subspecies Dama nestii nestii and Dama nestii eurygonos. He underlined in his analysis that the two subspecies were different in
Axis eurygonos from Pirro Nord
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In the paleobiogeographical point of view, Di Stefano & Petronio (1998, 2002) indicate two possible pathways of migration from Central-Eastern Asia: the northern one, followed by the representatives of the genus Rusa and headed for the continental Europe, and the southern one, through Iran and Anatolia, followed by the representatives of the genus Axis. Petronio et al. (2007) confirm their opinion on the morphological links among Axis and the cervid forms from the Italian Early Pleistocene and among Rusa and the cervid forms from continental Europe, the authors hypothesize that the origin of the two genera Axis and Rusa could be recognized into the tribe of Chinese Pliocervini. A deer very close to Cervavitus novorossiae, that occurred above all in Late Miocene Chinese deposits, probably was the ancestor of the two mentioned genera. These deer spread from different climatic regions and adapted to the different ecological niches of continental Europe and of the Mediterranean areas. The two genera evoluted in these different geographical zones following different lineages also taking into account the climate changings in the Early Pleistocene and, above all, in the Middle Pleistocene. To complete the previous historical notes, the last works on taxonomy and on the relationships among the different cervid genera based on genetical and statistical data must be cited, because they confirm the general picture drew on the basis of the paleontological data (Meijaard & Groves 2004, Pitra et al. 2004).
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whose possible ancestor could be recognized in a deer close to Damacerus bessarabiae. The author also described two new species, P. lyra and P. farnetensis, in the Villafranchian remains from Valdarno. Azzaroli hypothesized two different philetical lineages: one comprising the species P. lyra, P. nestii and P. farnetensis and another one comprising the species P. pardinensis and P. rhenanus. Finally, the author suggested that the living fallow deer is not philogenetically related with the genus Pseudodama. According to Khalke (1997) the medium-sized deer from the Early Pleistocene localities of Le Vallonet, Untermaßfeld, Selvella, Casa Frata e Pirro Nord must be referred to “Cervus” nestii vallonetensis. Di Stefano & Petronio (1998) proposed a new genus, Euraxis, to explain better the close mophological and philogenetical relationships among the living genus Axis from Asia and the “Dama-like” forms from the Italian peninsula. The new genus would include three species (from the most archaic to the modern one): E. lyra, E. nestii and E. eurygonos. Moreover, according to the authors, cervids from continental Europe in the morphological point of view are different to those from the Mediterranean area above all for the antler characteristics that are more archaic, short, less bent and generally more massive in the first forms. Also the postcranial skeleton, according to the authors, is different in the biometrical point of view between the two groups and the cervids from continental Europe could be philetically linked with the Asian genus Rusa. However, the genus Pseudodama was already been instituted for the same cervid group and the type species, E. nestii, was already been considered as type species of the genus Pseudodama (Azzaroli 1992), therefore Di Stefano & Petronio (2002) considered the genus Euraxis as junior synonymous of Pseudodama and then as invalid (as also suggested by Van der Made 1999) and suggested to refer all Italian Dama-like forms to the genus Axis. Croitor (2001) recognized different philetical lineages for the medium sized deer from Italian Pleistocene. The first lineage is represented by Pseudodama nestii (Azzaroli 1947) that, according to the author, is closely related with the genera Axis and Cervus. P. lyra Azzaroli, 1992, is considered as junior synonymous of P. nestii. The second lineage includes the archaic fallow deer Dama eurygonos with two more advanced subspecies, D. eurygonos farnetensis Azzaroli, 1992, and D. vallonetensis (= Cervus s.l. nestii vallonetensis de Lumley et al. 1988).
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Morphological analysis of the medium sized deer remains from Pirro Nord (Apricena, Southern Italy) The fossil remains of the medium sized deer testify on the whole a homogeneous population with uniform morphological features, notwithstanding they were collected in different times by different researchers from the numerous karst cavities in the pit area of Apricena (Text-fig. 1). The state of preservation of the skeletal parts is good, few cases testify anatomical connections and only in very few cases the bones are broken and/or flowed: this may indicate that most of individuals died not far from the sites of discovery, sometimes almost complete parts of bodies were dragged before to decompose and seldom the skeletal parts were broken before they were buried and fallen into the karst cavities. Therefore, the taphonomy of the bone remains in the various karst cavities is differ-
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Text-fig. 1. Pirro Nord quarries (Apricena, Southern Italy).
rather primitive, brachiodont and with short interlobar styles, and the palaeomerycine fold lacks: these characteristics of the skull are typical and identifies the more advanced species of Axis from the Villafranchian faunas of the Italian peninsula (Di Stefano & Petronio 2002). However, the morphological features that, more than others, can clarify the systematical position of deer are the antlers. All the remains collected show, in their basal part, a very short pedicle (it can be observed of course in the antler remains connected with the frontal bone), the b span in practice lacks, the brow tine is long and very curved and forms an angle with the beam from scarcely obtuse (about 95°–100°) to very opened (more than 120°). The antler of the medium sized cervid from Apricena have three points and show an elongated and curved architecture (in the Italian peninsula antlers with the same architecture and a trez tine have been also found in many deposits), the beam has a subcircular section and it is curved upwards, where present, the terminal fork is parallel to the sagittal axis of the body, its size is variable but it is always well developed. These characteristics are present in all the populations of the same age as Pirro Nord and referable to Axis eurygonos from the various Italian deposits (Di Stefano & Petronio 2002), however, the remains from Apricena, even if maintaining the same structure, are slimmer and smaller than the latter ones. This can be observed in the antlers and, less evidently, in the skull remains. The
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ent: the sites that collected the carcasses of individuals dead nearby show numerous complete long bones sometimes in anatomical connections, while the few cavities containing many broken or flowed remains indicate they are not homogeneous in the chronological and territorial point of views. As it regards the number of the remains, the skull ones are scarce and represented above all by occipital fragments, frontal bones with basal parts of antlers and upper teeth, two skulls almost complete and stored in the collection of the University of Florence are exceptions. One of the skulls belongs to a subadult individual (Text-fig. 2) and shows both the antlers. Among the other skull remains three complete mandibles with premolars and molars, few mandibular fragments and many single lower teeth must be pointed out. Many fallen antlers almost always constituted in the rose, the brow tine and the basal part of the beam are represented, vertebrae, ribs and long bones are rather numerous and in many cases also complete (Text-fig. 2). Skull and antler remains
The two skulls show the same general features as in the skull remains from Valdarno (Azzaroli 1947, 1953, 1992) and from Capena (Petronio 1979), the splancnocranium is longer than the more archaic species of the genus Axis from Europe (Axis lyra and Axis nestii: Di Stefano & Petronio 2002), the teeth are
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Axis eurygonos from Pirro Nord
Text-fig. 2. Cervid remains from Pirro Nord: a1 – skull, a2 – particular of the upper teeth, b – part of an antler, c – mandible, d – radius, e – tibia, f – metacarpus, g – metatarsus. Metric scale 5 cm.
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Biometrical analysis of the remains from Pirro Nord
The biometrical analysis of the remains from Pirro Nord has been carried out comparing the cranial and post-cranial data of these deer with those of the medium sized cervids collected from many localities from the Italian peninsula (among them Valdarno, Capena, Venosa-Loreto but also other sites). The study suffered the scarcity and, in some cases, the lack of homogeneous data in some collections for the groups taken into account that, therefore, did not allow at this moment to make deeper statistical analyses. Notwithstanding the fossil remains, as already mentioned,
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The limb bones are, as already observed for the other bones, small sized but also showing morphological features similar to those recognized in the same species from other Italian localities. More particularly, the humerus shows in anterior view a very deep and marked coronoid fossa and, therefore, the condyles have a wide articulation surface in antero-posterior view, the same bone shows in posterior view a short and wide holecranic fossa. The most important characteristic in the radius of these cervids is the columnar diaphysis and the epiphyses that seem less developed in transversal view. This feature can be observed also in the metacarpus where the strong transversal development of the diaphysis is present. This bone does not show a well defined minimum transversal diameter, in posterior view the fossa between the two metacarpals is very wide and the internal and external edges are pronounced, the articular surfaces of condyles in the distal epiphysis are short. The only significant characteristic in the femur, in addition to the columnar appearance of the bone and already observed in the population of Axis eurygonos from Capena (Petronio
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Postcranial skeleton
1979), seems the particular height of the great trocanter in comparison with the head of the femur. The tibia shows a particular sinuous appearance but on the whole it seems massive if compared to the reduced size. A long tibial crest is present in the proximal epiphysis and, in the articular surface, the internal and external spinae are developed in the same way as in Axis eurygonos from other deposits. The metatarsus is on the whole small sized but also shows strong and columnar diaphysis and a remarkable slipping of the two metatarsals, the proximal articular surface is narrow in transversal view but very elongated in antero-posterior view, finally, the condyles are very short in anterior view. All these last characteristics are present in the populations of Axis eurygonos from the Italian peninsula.
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mandibles are not thick and seem rather short in correspondence of premolars and molars, the lower teeth are brachio-bunodont and the forth premolar is rather molarized: these latter features are also common with Axis eurygonos from other Italian deposits.
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Text-fig. 3. Comparison between the antler of Axis eurygonos (left) and Rusa rhenanus (right).
Axis eurygonos from Pirro Nord
than in other deer populations. Finally, the ratios between the length of the limb bones and their diameters show that the population from Apricena is a little more robust than the other ones considered: the points representing these ratios are in fact usually higher in the scattergrams and the regression lines are less sloping.
Discussion
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The morphological and biometrical analyses of the remains from Pirro Nord and other faunal deposits allowed recognizing the peculiar features of Axis eurygonos to take into account for the comparison with the remains of other populations. The occurrence of the genus Axis in Europe from Asian population is limited, on the basis of our knowledges, to the Mediterranean area and particularly to the Italian peninsula in faunal assemblages from Middle Villafranchian to Middle Galerian Faunal Units (Di Stefano & Petronio 2002). Western Europe,
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have been found in different karst cavities of the area, the differences observed in the statistical point of view among the collections (distribution, standard deviation, ANOVA, etc.), are too short to consider the population fully homogeneous. Even considering the scarcity of some groups, the biometrical analysis confirms the morphological one and the reference of the remains from Apricena to the species Axis eurygonos. A constant features observed in all the examined bones is the small size of this deer population. As an example, whose results can be also repeated by all limb bones, we propose the scattergram on the data of metacarpus in Text-fig. 3 that confirms this observation. In fact the points referable to Axis eurygonos from Apricena are in the left side of the diagram in comparison with the points of the same species from other localities. Another characteristic observed rather frequently is the presence of two separated groups into the population from Pirro Nord that can be referred to sexual dimorphism, more evident
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Text-fig.4. Scattergram on the length (L) and the transversal proximal diameter (TPD) of metacarpus in Axis eurygonos from Capena, Valdarno and Pirro Nord.
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on statistical data, referred the Dama-like cervid from Untermaßfeld to the subspecies Rusa rhenanus perolensis. To clarify definitively this question the writers compared the remains from Untermaßfeld with those of Axis eurygonos from Pirro and other localities to understand if the first deer, notwithstanding the systematic reference used, can be attributed to the same cervid which occurs in Italy and at Le Vallonet. The medium sized deer remains from Untermaßfeld are rather numerous but the skull and antler remains are poor and fragmented. However, the basal parts of antlers (Text-fig. 4) clearly show that the antler is more robust and shorter than those of Axis, the German form has short pedicle and also the b span is very short, the brow tine forms an angle of about 90° with the beam but it is strongly curved and seems subparallel with the same beam in its terminal part. The upper teeth from Untermaßfeld do not show peculiar features if compared with the deer from the Italian peninsula, on the contrary some important observations can be made on the lower teeth. In fact, the forth lower premolar is less molarized than the population attributed to Axis, the third lobe of the third lower molar is in the same line with the other two lobes in the deer from Untermaßfeld but it is in a oblique position in Axis, the same lobe is more rounded in the German deer and often shows an interlobar style which instead lacks in the Italian deer.
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beyond the Alpine range, is instead characterized by the presence of deer referable to the genus Rusa and by the lacking of deer referable to the genus Axis. This distribution, as already mentioned in the introduction, is far to be a simple systematic topic but shows important implications in the palaeobiogeographical and biochronological point of views. As already mentioned in the historical notes, the deer remains from Pirro and other Italian Late Villafranchian deposits have been referred to “Cervus” s.l. nestii vallonetensis (Kahlke 1997). This form, whose systematic attribution is still discussed (Croitor 2001, Di Stefano & Petronio 2002), surely belongs to the Dama-like cervids that characterize the Late Villafranchian faunas and has been identified for the first time at Le Vallonet (de Lumley et al. 1988). The morphological features of the remains collected from this locality are fully similar to those of the Italian populations and therefore they have been attributed to Axis eurygonos (Di Stefano & Petronio 2002). Also the remains from Untermaßfeld (Thuringien) have been referred to the subspecies “Cervus” s.l. nestii vallonetensis, however, this reference seems to contrast with the range of Axis known at the present day which seems to be limited to the Mediterranean area of Europe. Di Stefano & Petronio (2002) took into account this difference and, basing above all
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Text-fig. 5. Scattergram on the length of lower teeth and premolars in Axis eurygonos and in the medium sized deer from Untermaßfeld.
Axis eurygonos from Pirro Nord
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cervid to have the premolar line shorter than the other deer can be noticed. The tendency towards more slender limb bones in the remains from Untermaßfeld, already mentioned in the morphological observations, is testified, for example, also by Text-fig. 6 on the radius measurements. In fact, apart from the size, the points representing the ratios of the German remains is evidently in the lower part of the scattergram and well separated from the points representing the ratios of the Italian remains. The same considerations can be made from the various rations of measurement of the limb bones. Again for example, we propose two other scattergrams (Text-fig. 7 and Text-fig. 8): both confirm the previous observations, i.e., the limb bones of the Dama-like cervids from Untermaßfeld are always larger but also more slender than the population referred to Axis eurygonos from the Italian peninsula. The previous biometrical consideration and the morpholgical analysis, therefore, allow excluding the reference of the remains from Untermaßfeld to the same species to which the remains of the Dama-like populations occurring in the Italian peninsula in the Late Villafranchian faunas are attributed. To conclude the present analysis the attribution of the remains from Untermaßfeld is still to clarify. Some elements have been observed in the morphological analysis of the remains but more considerations
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The limb bones of the Dama-like cervid from Untermaßfeld are generally larger than those of the genus Axis, but also they show shorter transversal diameters and other features that sometimes allow separating the two forms. As an example, in the distal epiphysis of the humerus from Untermaßfeld the articular surface with the radius is less deep and more lengthened, therefore the coronoid fossa in less marked and the holecranic one is narrow and more lengthened to the diaphysis. The radius is longer and more slender and the great sigmoid fossa is less deep: this could testify an articulation with the humerus less suitable to run on uneven lands. There are few morphological differences in the metacarpus: we can observe a lower depth between the two metacarpals in the posterior view and, generally, a larger slenderness of the German population than those of Axis. Also the rear limbs do not show big differences between the German cervid and Axis: we most only point out the lower development of the epiphyses of tibia and metatarsus in comparison with the diaphyses and a lower sliding between the two metatarsals in the cervid from Untermaßfeld. As it regards the biometrical comparisons, the scattergram on the ratios of the measurements of lower premolars and molars, already shows some elements (Text-fig. 5). The difference in size among the remains from the Italian peninsula and those from Untermaßfeld is evident but also the trend in the German
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Text-fig. 6. Scattergram on the length (L) and the antero-posterior proximal diameter (APPD) of radius in Axis eurygonos and in the medium sized deer from Untermaßfeld.
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make easy and effective the observation of the data, the scattergrams are the same already showed (Text-fig 6, Text-fig. 7 e Text-fig. 8) with the addition of the data of Rusa rhenanus from the many Villafranchian
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can be added on the basis of the biometrical data. To this purpose three scattergrams on some ratios of radius, metacarpus and metatarsus measurements are proposed (Text-fig. 9, Text-fig. 10, Text-fig. 11). To
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Text-fig. 7. Scattergram on the length (L) and the transversal distal diameter (TDD) of metacarpus in Axis eurygonos and in the medium sized deer from Untermaßfeld.
Text-fig. 8. Scattergram on the length (L) and the transversal proximal diameter (TPD) of metatarsus in Axis eurygonos and in the medium sized deer from Untermaßfeld.
Axis eurygonos from Pirro Nord
last species are completely separated from those of Axis eurygonos. The other scattergrams show the same characteristic: the first one in a less evident way but still observ-
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deposits of Europe. The fist scattergram shows with evidence that the more slenderness already verified in the cervid from Untermassfeld is completely corresponding in Rusa rhenanus and that the points of this
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Text-fig. 9. Scattergram on the length (L) and the antero-posterior proximal diameter (APPD) of radius in Axis eurygonos, in the medium sized deer from Untermaßfeld and in Rusa rhenanus from continental Europe.
Text-fig. 10. Scattergram on the length (L) and the transversal distal diameter (TDD) of metacarpus in Axis eurygonos, in the medium sized deer from Untermaßfeld and in Rusa rhenanus from continental Europe.
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Text-fig. 11. Scattergram on the length (L) and the transversal proximal diameter (TPD) of metatarsus in Axis eurygonos, in the medium sized deer from Untermaßfeld and in Rusa rhenanus from continental Europe.
We can summarize the most important morphological features of the genus Rusa and the European species referred as follows: • R. pardinensis from the Early Villafranchian faunas of Central-Western Europe (Heintz 1970, Stefaniak 1995) is a medium sized deer characterized by primitive antlers with rather straight beam lacking of terminal fork, long b span, brow tine forming an acute angle with the beam, long pedicle and quite short splancnocranium. • R. rhenanus from the Early-Middle Villafranchian faunas of Central-Western Europe is larger than the previous species but the antlers features are very similar, three-pointed and lacking of the terminal fork, teeth seem more evoluted but some archaic features are still present as a weak palaeomerycine fold. • R. perolensis from Late Villafranchian faunas of Western Europe is represented only by few remains still discussed for their systematic rank (Azzaroli 1953, Heintz 1970, Geraads 1990, Di Stefano & Petronio 2002). In fact the antlers features are still very similer to those of R. rhenanus which is considered its ancestor, certainly the pedicle and the b spam are shorter than the last species and also the brow tine forms a wider angle with the beam, about 90°. As it regards the teeth and the limb bones there are no diagnostic elements allowing to separate R. rhenanus from R. perolensis. The high
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able, the second one is much more evident and confirms what already observed in the Text-fig. 9. In fact this scattergram show again that the ratios of measurements of R. rhenanus not only cover those of the cervid from Untermaßfeld but also that both these deer have more slender limb bones than Axis.
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As already pointed out, the occurrence of the genus Axis in Europe, migrated from Asia in the Late Pliocene, is referable only in the Mediterranean areas during the Plio-Pleistocene (Di Stefano & Petronio 2002, Petronio et al. 2007). According to the writers, the reference of the numerous remains from Untermaßfeld to the same specie which occurred in the Southern Europe cannot be considered as valid on the basis of the previous discussion. The middle sized deer remains from the German locality, basing on morphological and biometrical data, can be referred to one of the middle sized cervid that occurred in the Villafranchian faunas of continental Europe (Rusa pardinensis, Rusa rhenanus and Rusa perolensis). Moreover, the three species of Rusa, from the more archaic R. pardinensis to R. rhenanus and R. perolensis (Heintz 1970), seem to follow an evolutive trend very similar to the one followed by the genus Axis, where Axis lyra, with more archaic features, come before Axis nestii and the last and more evoluted Axis eurygonos.
Axis eurygonos from Pirro Nord
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De Lumley, H., Kahlke, H.-D., Moigne, A.M. & Moullé, P.-E. (1988): Les faunes de grands Mammiferes de la grotte du Vallonet Roquebrune-Cap Martin, Alpes-Maritimes. – L’Anthropologie (Paris) 92 (2): 465–496. Di Stefano, G. & Petronio, C. (1998): Origin of and relationships among the Dama-like cervids in Europe. – N. Jb. Geol. Palaont. Abh. 207 (1): 37–55. Di Stefano, G. & Petronio, C. (2002): Systematics and evolution of the eurasian Plio-Pleistocene cervini (Artiodactyla, Mammalia). – Geologica Romana 36: 311–334. Forsyth Mayor, C.I. (1879): Cervi Pliocenici del Val d’Arno superiore. – Atti Soc. Tosc. Sci. Nat. Proc. Verb. I: p. C. Forsyth Mayor, C.I. (1885): On the Mammalian Fauna of the Val d’Arno. – Quart. J. Geol. Soc. London 41: p. 1. Forsyth Mayor, C.I. (1890): L’ossario di Olivola in Val di Magra. – Atti Soc. Tosc. Sci. Nat. Proc. Verb. VII: p. 57. Geraads, D. (1990): Contribution des Cervidés à la chronologie des dèbuts de l’occupation humaine en Europe Occidentaile. – Quaternaire 3–4: 167–174. Heintz, E. (1970): Les cervides villafranchiens de France et d’Espagne. – Mem. Mus. Nat. Hist. C. 22: 1–206. Kahlke, H.-D. (1997): Die Cerviden- Reste aus dem Unterpleistozän von Untermaßfeld. – In: Kahlke, R.–D.: Das Pleistozän von Untermaßfeld bei Meiningen (Thüringen), Teil 1. – Monogr. Röm.-German. Zentralmus. Mainz 40 (1): 181–275. Khomenko, I. (1913): Meotian fauna of v. Tarakliya of Bendery district. – Annuaire geologique et mineralogique de la Russie. – 1. The ancestors of modern and fossil Cervinae. 2. Giraffidae and Cavicornia 15: 107–132 (in Russian). Kunst, C.E. (1937): Die Niederländischen Pleistozänen Hirsche. – Thesis, Drukkerij Lucto et Emergo, Leiden, 126 pp. Meijaard, E. & Groves, C.P. (2004): Morphometrical relationships between South-East Asian deer (Cervidae, tribe Cervini): evolutionary and biogeographic Implications. – J. Zool. 263: 179–196. Moullé, P.-E. (1990): Les Cervidés de la grotte du Vallonet (Roquebrune-Cap Martin, Alpes Maritimes). – Quaternaire 3–4: 193–196. Petronio, C. (1979): Dama nestii eurygonos Azz. di Capena (Roma). – Geologica Romana 18: 105–125. Petronio, C., Krakhmalnaya, T., Bellucci, L. & Di Stefano, G. (2007): Remarks on some Eurasian pliocervines: characteristics, evolution and relationships with the tribe Cervini. – Geobios 40: 113–130. Pitra, C., Fickel, J., Meijaard, E. & Groves, C.P. (2004): Evolution and phylogeny of old world deer. – Molecular Phylogenetic and Evolution 33: 880–895. Schlosser, M. (1924): Über die systematische Stellung jungtertiärer Cerviden. – Zentralbl. Mineral. Geol. Paläont: 640–654. Stefaniak, K. (1995): Late Pliocene cervids from Weze 2 in southern Poland. – Acta Palaeont. Pol. 40: 327–340. Van der Made, J. (1999): On Euraxis, on the rules of the ICZN, on methodology. – N. Jb. Geol. Paläont. 11: 676– 678.
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resemblance between the two forms produced the hypothesis to separate them only at a subspecific level (i.e., R. rhenanus perolensis) (Heintz 1970). As already said in a previous paper (Di Stefano & Petronio 2002), the cervid from Untermaßfeld completely fits with the characteristics of the more advanced European Rusa and, therefore, if the subspecific reference of the remains from Peyroles will be confirmed, it can be referred to Rusa rhenanus perolensis. The reference of the remains from Untermaßfeld to Rusa confirms the range of the genus Axis in Europe during the Plio-Pleistocene to the Mediterranean areas and, particularly, to the Italian peninsula. The morphological and biometrical features of the medium sized deer from Apricena, allow instead the attribution to Axis eurygonos. However, if compared with the typical form from Valdarno, the following differences have to be noticed: three-pointed antlers, very slender but also maintaining the typical architecture, the brachiodont teeth, with reduced interlobar styles in both upper and lower molars (in any case very similar to the typical Axis eurygonos), the post-cranial skeleton smaller than the other populations from Italy but also more robust than these latter. These differences can be referred to various elements, such as a light isolation of the population, that in the deer species often causes a fast size reduction, but also the geomorphological environment of the landscape and the trophic resources available by these arctiodactyls in an area that could be defined as marginal. Finally, it must be reminded that in any case these differences fit into the intraspecific range of Axis. References
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Azzaroli, A. (1947): I cervi fossili della Toscana con particolare riguardo alle specie villafranchiane. – Palaeontographia Italica 43: 1–45. Azzaroli, A. (1953): The deer of the Weyburn Crag and Forest Bed of Norfolk. – Bull. Brit. Mus. (Nat. Hist.), Geol. 2 (1): 3–96. Azzaroli, A. (1992): The cervid genus Pseudodama n.g. in the Villafranchian of Tuscany. – Palaeontographia Italica 79: 1–41. Bernsen, J.J.A. (1933/34): Eine Revision der fossilen Säugertierefauna aus den Tonen von Tegelen (IX) Cervidae. – Natuurhist. Maandblad 22 (11): 136–138, 23 (4, 6 & 7): 38–46, 71–77, 82–86, Maastricht. Croitor, R. (2001): Functional morphology of small-sized deer from the early and middle Pleistocene of Italy: implication to the paleolandscape reconstruction. – La Terra degli Elefanti, Proc. . First Internat. Congr., 97–102, Rome.
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