Deer Fauna from the Aulla Quarry (Val Di Magra, Northern Apennines)

Rivista Italiana di Paleontologia e Stratigrafia volume 101 numero 3 tavola 1 pagrne 341-348 Dicembre 1995

DEER FAUNA FROM THE AULLA QUARRY (VAL DI MAGRA, NORTHERN APENNINES). BIOCHRONOLOGICAL REMARKS

LATIRA ABBAZZI (,), GIOVANNI FICCARELLI (x) sc DANILO TORRE (.)

Key-uords: Mammalia, Cervidae, Early Villa{ranchian, Late rule out the possibility that the fossil remains belong to Pliocene, Italy. the Montopoli faunal assemblage. Indeed, the Montopo- li assemblage correlates with the Praetiglian pollen pha- Riassunto. Viene riponata la presenza nei sedimenti conrinen- tali di Cava Aulla, di cervidi tipici della fauna del Villafranchiano se, which coincides with a generalized cooling marked Inferiore, attribuiti a Procapreolus cusdnus e Pseudodarna ex gr. pardi- by the spread of steppe vegetation in Southern Europe nensis-lyra. rinvenimenti mettono in discussione Questi la correlazio- (cf. Bertoldi et a1.,1189). ne della successione sedimentaria al Pliocene Inferiore, effettuata sulla base del contenuto pollinico riconosciuto nei sedimenti. in Itaiy, fossils referable to the base of the Vilia- franchian (Triversa faunal unit) are relatively common. Abstract, The occurrence in the continental deposits of Aulla Various sites around Asti (S. Martino Unit, cf. Masini et quarry of deer characteristic of Early Villafranchian fauna, referred to aI., 1994) have yielded Pseudodarna pardimensis remains Procapreolus cusanus and Pseudodama ex gr. pard.inensis.lyra, is repor- associated masto d (A c e s - ted. These finds make questionable the correlation of the sedimenta- with o îts na n u s a ra rn en i s, 7, y go I o ry succession to the Early Pliocene, based on the pollen assemblage. phodon borsoni), Sus rninor, Dicerorbinus jeanaireti, and lcptobos stenometopon The lignites and clays at the base of the succession of the S. Barbara qu^rry (Upper Val- lntroduction. darno) contain remains referred to [Jrsus rninimus, Thpi- The discovery of the remains of medium ro small rus areernensis, Dicerorbinus sp., Icptobos sp. The Lower sized deer referable fo Procapreolus cusanus (Croizet & Vaidarno has yielded Tapirus arpernensis (Dominici et Jobert, 1828) and Pseudodama ex gr. pdrdinensis-lyra in al., in press), Pseudodarna lyra (Azzaroli, 1992), and the Aulla quarry allows the examination of the chrono- Nyctereutes *rgo*ottoidrs (Del Campana, 1,917). Finally logical position of the sediments exposed in this site. Anancus anternensis, Zygolopbodon borsoni, and Thpirus Bertoldi (1988) used the floral content of the Aulla de- arpernensis have been reported from Umbria-Sabina 1o- posits to define the Macriana pollen phase, which corre- calities in the Tiber basin (cf. Di Stefano et a1.,1994). lates with the Early Pliocene (foliowing the division of Paleontological, radiometric, and paleomagnetic ii- the Pliocene into Early and Late), and with the Rusci- nes of evidence show that the Villafranchian began du- nian Mammal Age. The association of Pseudodama par- ring the Gauss chron, between 3.0 and 3.6 Ma (cf. Al- dinensis-P lyra and Procapreolus cusdnus is characteristic, bianelli et al., in press). In the Spanish locality of in \festern Europe, of'the mammal assemblages of the Layna, the fossil remains, which represent a transitional Early Villafranchian (the Triversa, and possibly the phase between the Ruscinian and the Villafranchian, Montopoli faunal assemblages, referred respectively ro come from sediments with a positive magnetization that biozones MN16a and MN16b after Mein, 1990). The oc- correlates with the Gauss chron, thus excluding an age currence of these taxa in the sediments of the Aulla earlier than 3.6 Ma for the fossiliferous horizon (cf. quarry therefore contrasts with the attribution of the Baksi, 1993). At Vialette and Les Etouaires (French Cen- sedimentary succession to the Early Pliocene. The floral tral Massif), radiometric studies have yielded age be- assemblage from the Aulla succession contains elements tween 3.3 and 2.6 Ma (Bandet et al., 7978; Couthure Ee characteristic of a subtropical climate, which allow us to Pastre, 1983; Savage & Curtis, 1983) for the mammal

('f) Dipanimento di Scienze della Terra, Università degli Studi di Firenze, Via La Pìra 4,50121., Firenze. iro\ ni-.-i*.-'^,{; Scienze della Terra e Museo di Storia Naturale (Sezione Geo-Paleontologica), Università degli Studi di Firenze, Via LA rlra +- )u121- Flrenze- 342 L. Abbazzi, G. Ficcarelli G D. Torre

assemblages characteristic of the Triversa faunal unit. In Geological setting and chronology of the Aulla quarry. Italy, the sediments of the Fornace RDB quarry (Villa- The Aulla quarry, located north of the town of franca d'Asti, Northern Italy), bearing the type fauna of Auila, lies within the Aulla-Olivola basin (Fig. 2), the Triversa unit, show a positive magnetic interval in- which contains a 250 m succession of continentai depo- terlayered negative intervals that correlate between ts/o sits subdivided in 3 units (Federici et al., 1982). From with the Kaena and Mammoth events of the Gauss bottom to top, the first unit consists of lacustrine silts chron (Lindsay et ai., 19s0). The recent paleomagnetic and clays, the second of alternating fluviolacustrine silts analyses of the succession exposed in the S. Barbara and clays, and gravels, and the third of prevalently flu- qul.rry have yielded analogous results (Albianelli et al., vial pelitic sands and conglomerates ("Olivola Conglo- in press). Indeed, the part of the Gauss normal magneto- merates'), marking the final infilling of the basin. A chrone, with the reversed Kaena event dated as 3.10-3.17 Middle Villafranchian assembiage characterized by Pseu- Ma, has been identified in the fossiliferous lignites and dodama nestii, Sus strozzii, Stephanorhinus euuscus, Canis clays at the base of the deposit. etruscus (AzzaroIí, 1950; Azzaroli et al., 1988; Masini et In Fig. L a chronological scheme where radiome- al., in press) has been recovered in clay outcropping trical ages, paleomagnetical scale, geochronological units, near Quercia and Vaccareccia (Fig. 2) referable to the mammal ages, faunal units, and polien phase are correla- second unit (Federici et al., 1.982). ted, is shown. The Olivola Conglomerates yielded mammal re- mais upon which the Olivola faunal unit is defined, marking the beginning of the Late Villafranchian (Azza- 1.0 Galerian roli,1977). The sediments exposed in the Aulla quarry (Fig. 1.2 =o 3) are referable to the first unit, and have been the 1.4 object of a palynological study carried out by Bertoldi (1988), who showed that throughout the succession ar- 1.6 o boreal taxa predominated over non-arboreal ones. Pre- J 1.8 Tiberian phase dominant among the forest taxa are the so-called Tertia- I l ry elements, such as Tàxodiwrn, Sequoia, Magnolia, Sym- .o 2.0 plocos, Sapotaceae, and on the vrhole the pollen assem- p;-s9 2.2 blages indicate a temperate to warm climate. The predo- =-s minance of archaic plants at Aulla has chronological significance, in that in Europe these plants are generaily confined to the pre-Pleistocene. At Pietrafitta (Perugia, zo Central Italy), the sharp decrease in the Tertiary Taxo- o diaceae group that occurs at the base of the sedimeîtary 2.8 -g succession has in fact been used to locate the base of the 3.0 Tiberian, which correlates vzith the Plio-Pleistocene Kaena phase UJ Pretiberian boundary [-ona & Bertoldi, 1972). The presence of ele- 3.2 Mammoth 3.4

óó . o|-tVoLA

4.0 Macrian ohase

4.0 AÙ LELLA

Correlation scheme of Mammal ages, Faunal units, Pollen Fl" ) location map of the Aulla quarry (") in the Aulla-Olivola phases (according to I-ona 6c Benoldi, 1972; Benoldi, 1988). basin. The scale is 1 km. Deer fauna from tbe Northern Apennines J+J ments such as Syrnplocos and Magnoli4 makes the Aulla The pre and infra-Pliocene elements typical of the assemblage even more primitive than the assemblages re- Aulla succession are poorly represented in the sediments ferred to the Pretiberian phase. Indeed, in some Euro- of the RDB quarry, which are referable to the Late Plio- pean successions, some elements (e.g. Symplocos) appear cene. The analysis of the pollen content of the succes- to be limited to the Early Pliocene, and only reach the sion exposed in this locality (Bertoldi, 1988; Bertoldi, Pleistocene with a few forms. According to Bertoldi 1994) showing that it contains few elements that presen- (1988), the palynological sequence of Aulla rherefore re- tly have a sub-tropical distribution (e.g. Symplocos, cords a floral phase characterized, throughout its dura- Clethraceae/Cyrillaceae, and Sapotaceae), allows the pol- tion, by a pollen assemblage distinct from those of the len sequence to be attributed to the Pretiberian phase, Pretiberian and Tiberian phases. For this reason it was which was dominated by the Taxodiaceae group. chosen to define the Macriana pollen biozone (Early A floral assemblage indicative of a hot humid sub- Pliocene, Ruscinian Mammal Age). tropical climate, with Sapotaceae, Clerhraceae-Cyrilia- ceae, and Tàxodium, has recently been found in the lignites and basal clay of S. Barbara (Albianeili et a1., in press), bearing the Triversa mammal assemblage. This shows that the Tertiary elements whose substantial occurrence had been thought to be limited to the pre-Plio- cene or Early Pliocene, are also well represented in assemblages dated to subsequent intervals.

The Ruscinian and Villafranchian transition in the deer fauna.

The climatic and environmental variations that occurred during the Neogene had a profound influence on the Cervidae familir, favoring its radiation and diver- sification. One of the bioevents that affected this family occurred at the transition from the Ruscinian to the Vil- lafranchian, which seems to have been marked by a brief cold interval, and has been set near the limit between the Early and Late Pliocene. lVith the beginning of the Villafranchian. in addition ro rhe more derived species of genera present in the Ruscinian (Procapreolus, Croizetocero), medium to large sized deer such as Arper- noceros and "Cervus" spread into Europe (Vislobokova, 1992; Vislobokova et al., 1993).In \festern Europe there .was an assemblage characterized by the presence of Croi- zetoceros rAînosus, Procapreolus cusanî4s, Psewdodama pardinensis, Pseudodama lyra, Araernoceros ardei, and Cer- rLus perrieri. These taxa have also been found in Early Villafranchian localities of Central Europe, such as'W'eze 2 (Poland, cf. Kowalski, 1990). At the beginning of the Villafranchian there was, in addition to the dispersal of forms larger than those present in the Ruscinian assemblages, an increase in the complexity of the antlers, with the development of distal ramifications, and the appearance of a true basal Fig.3 - Stratigraphical section of Aulla quarry (from Bertoldi, tine, a few cm from the burr. The tendency towards an 1988, modified). On the right the zone where the deer increase in the morphological complexity of the distal remains have been recovered is shown. It is not possible to indicate on the section reponed by Benoldi the precise portion of the antler was, however, already observable provenance of the fossils, because at the present the in the preceding evolutionary phases (middleJate Turo- outcrop is somewhat modified by erosion. kgend: 1) gra- lian, Ruscinian), with the genera Procapreolus, Pliocer- vels andlor sands; 2) silty-sandy clays; 3) silty clays; 4) ows, and Croizetoceros (Azanza et al., 1989). In the Ru- clays with limonitic nodules; 5) lignitiferous clays. The scale is 1 m. The ondulating line in the upper part of the scinian Croizetoceros ralnosus the distal tines are esoecial- succession indicates the erosion of the lacustrine sediments. ly numerous (Heintz, 1970, 1974). 344 L. Abbazzi, G. Ficcarelli & D. Torre

The basal tine was also present from the beginning of the Early Pliocene. Azanza et al. (tlSf) report deer (Cervinae gen. and sp. indet.) with basal tines in the Spanish locality of Fonelas (early Ruscinian, biozone MN14). Czyzewska (tleS) descrlbes Ceraus aarthae, from 'Weze 1 (Poland, biozone MN15), underlining the presence of a basal tine. As occurred with the distal ramifications, this tine became common and widespread, in both simple small sized forms such as Pseudodama ,/ pardinensis and P. lyra, and more complex, larger sized forms such as Ceraus perrieri, only at the beginning of the Villafranchian. /r

The fossil remains from the Aulla quarry.

Provenance. The fossil deer found at Aulla comes from sandy sediments in the middle of the sedimentary succession (Fig. 2).

Systematics.

Procapreolus cusanus (Croizet Ec Jobert, 1828). At Aulla, this species is representedby a few proximal fragments of shed antlers (Pl. 1, fig. 1a, b), and by a specimen that lacks the basal section, and displays a marked flattening of the beam and a short ramification Fig. 4 - Comparison between one remain of Procapreolus cusanus Fig. +) A few fragments of isolated teeth can also be from Aulla quarry (on the right) and the neotype from l-es species. attributed to this Etouaires (from Heintz, 197A, redrawn). The scale is 10 cm. P cusanus was erected by Crorzet and Jobert (1828) on material from Les Etouaires. has also been It taxon (P1. 1, fig. 2a, b). Unfortunately, the specimen is found at (Heintz, 1970), and in Italy has been Vialette it heavily encrusted by well cemented sands (Pl. 1, fig. 3), (Masini a1., reported from the Montopoli locality et which alter its size, though not its morphology. 1.ee4). The beam presents a faínt concavity at the inser- The antlers attributed to this species display tion of the basal tine, which forms an angle of about at least the aduits, strong transversal flattening, and, in 50o with the beam and is located about 90 mm from from three to five tines oriented along the same ante- the burr. The proximal curvature of the beam indicates rior-posterior plane. that the antlers were fairly slanted backwards from the Procapreolws cusa.nus is typical of the Early Viila- skull. The beam is circuiar in cross section. These char- franchian of Europe (Heintz, L970; Triversa and 'ùTestern acters together with the size (taking into account the Montopoli faunal units), even though it was known encrustation that covers it completely) agree with the from the beginning of the Ruscinian (Vislobokova, attribution to Pseudodama. As a matter of fact the di- 1990). Representatives of the genus Procapreolus are stance separating the first tine from the burr and the known from throughout Eastern Europe beginning in angle that the first tine forms with the beam in the spe- the Late Miocene (Vislobokova, 1990), in faunal assem- cimen from Aulla are observed in the most archaic blages referable to the Vallesian (biozone MN10). forms of Pseudodama. Among of the major evolutionary The genus is closely related to Capreolus, whose changes undergone by this genus are the positioning and earliest known representatives referred by Vislobokova orientation of the basal tine, which, in the most derived et al. (1995) to C, costantini, have been found at Udun- forms (P nestii and P branches off closer to ga, an Early Villafranchian (biozone MN16) locality in farnetensis) the burr, and at an open angle with respect to the beam Transbaikalia (Vislobokova et al., 1993; Vislobokova et (Azzaroli, 1992; Cohcci et al., in progress). a1.,1995). On the contrary the Au11a specimen differs from Pseudodama ex gr. pardinensis-lyra. the European Ruscinian cervids such as Paraceruulus au- A right antler fragment with a.n incomplete basal stralis de Serres $4ontpeliier, biozone MN14) in that tine branching off an anterior position is attributed to this the latter has a laterally compressed shaft, and has the Deer fauna from tbe Northern Apennines 345

la 1b

PLATE 1 rrg. r Procapreolus cusanus (Crotzet Ec Joben). Right shed antler from Aulla quarry a) Lateral view; b) frontal vìew 1/2 natural size. Pseudodama ex gr. pardinensisJyra. Rrght shed antler from Aulla quarry. a) Frontal view; b) lateral view. 1,/2 natural size. Pseudodama ex gr. pardinensis'/yra. Cross section of the shed antler. The cemented sands envelopping the remain are shown. Natural size. first tine far from the burr. This second characteristic is ters are more derived, both in the anrlers (incipient also observable in the antlers of Croizetoceros ramosl.ts, third distal bifurcation and greater curvarure of the 'whose basal tine has a very wide insertion angle shaft), and in the dentition (lesser development of the (Heintz, 1970), wider than that observed in the Aulla cingulum in the upper teeth). Though the phylogeneric sPeclmen. relationships between the two species are not clear yet, The genus Pseudodama has been recently erected P lyra could be derived from P pardinensis, or rhey 6y Azzaroli (1992) ro group the medium sized deer wi- could be sister species (Azzarolì, 1992). despread in the Late Pliocene-Early Pleistocene, pre- The characters upon which the distinction be- viousiy referred ro Ddrna (Azzarolì, 1947) and to Ceraus tween the two species is made cannot be observed in (Heintz, 1970). According to Azzaroh (1,992), this genus the antler of Aulla because of its fragmenrary nature. includes six species that represent two distinct phyletic Therefore this specimen is referred to Pseudodama ex gr. lines. The former, including P. pardinensis, P rbenanus pardinensis-lyra. and P pereolensis (probably a synonym of P. nestii), seems to be widespread in more northern regions than Conclusions. the latter, which includes P lyra, P nestii and P. farneten- sls. This line is recorded in the mediterranean area. The presence of Procapreolus cusanus in the sedi- P lyra, which is the species of more recent erec- ments of the Aulla quarry has not a precise biochrono- tton (Azzarolí, 1992), is weii represented in the sedi- logic significance, in that though the species is typical ments of the Lower Valdarno (Early Pliocene), where it of the Early Villafranchian, it has also been found in occurs in association with mammals of the Triversa and Ruscinian localities. The identification of a primitive Montopoli faunal units. According to Azzaroli P lyra form of Pseudodama is on the other hand more impor- can be distinguished from P pardinensis found at Les tant. The occurrence of this taxon in fact seems to be Etouaires and Vialette, because its morphological charac- characteristic of Post-Ruscinian faunas. The association 346 L. Abbazzi, G. Ficcarelli & D. Torre

of P. cusanus and P ex gr. pardinensis-lyra would limit In conclusion in the Olivola-Aulla basin faunal as- the age of the Aulla deposit to the Late Pliocene, a con- semblages referred to Early Villafranchian (Triversa fau- clusion that contrasts with the results obtained from the nal unit), to Middle Villafranchian (Seneze faunal unit) pollen study, which indicates an Early Pliocene age and to Late Villafranchian (Olivola faunal unit) occur (Bertoldi, 1988). from W to NE. The lacking of the Montopoli faunal The recent identification o{ Tertíary subtropical unit could be attributabie to an erosive phase, recogniza- elements in the Late Pliocene sediments of the S. Barba- ble at the top of the Aulla succession. ra qirarry (Albianelli et al., in press) suggests however that the stratigraphic distribution of these taxa covers a time longer than that which had previously been Acknouled.gem.ents, thought. The abundance of elements whose present di- \Xle are gratefull to Prof. A. Azzarolì for the critical reading of stribution is tropical-subtropical in the sediments of the the text. \Ve thank Mr. I. Sirigu and Mr. A. Tanfi for having us indicated the presence of the cervid remains in the Aulla quarry. Aulla quarry could be due to the development, in that Many thanks also to Dr. V. Borselli for the help in the recovery of area, of a microclimate that supported a distinctive floral the fossils. Mr. F. Cozzini took care of the photos. assemblage. Vork supponed by M.U.R.S.T. and C.N.R. grants.

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