22 July 2021
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The vertebrate fauna from Cardamone (Apulia, Southern Italy): an example of Mediterranean mammoth fauna. / Rustioni
M.; Ferretti M.P.; Mazza P.; Pavia M.; Varola A.. - In: DEINSEA. - ISSN 0923-9308. - 9(2003), pp. 395-403. ((Intervento
presentato al convegno Second International Mammoth Conference tenutosi a Rotterdam nel 16-20 Maggio 1999.
Original Citation:
The vertebrate fauna from Cardamone (Apulia, Southern Italy): an example of Mediterranean mammoth fauna.
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M. Rustioni ',M.P Ferretti ',P Mazza ',M. Pavia '& A.Varola l I Museo di Stona Naturale, Universic.a degli Studi di Firenze
l Dipartimento di Scienze della Terra. Università. di Tonno
IOsservarorio di Fisica e Chimica deUa Terra e dell'Ambiente, Universìcà dì Lecce
The vertebrate fauna from Cardamone (Apulia,
southern Italy): an example of Mediterranean
mammoth fauna
RustÌoni, M., Fel'Tl:ni. M.P., Mazza, P., Pavia, M. & Varala, A., 2003 - The verrebrate fauna from Cardamone" (Apulia, southern ltaly): an examph:: of Mediterranean mammo:1h fauna -in: Reumer, l.W.F .• de Vos, J. & Mol, D. (eds .) . ADvANC1lS IN M.AA!MoTH REsPARCH (Proccedings or the SecoDd IDtemational Mammoth Conference. Ronerdam, May 16·20 1999) . DEINSEA 9: 395-403 (ISSN 0923-9308] Published 24 May 2003
Tb..: siLe of Cardamone .• a karstlc cavity in leeal Pljo-Pleì~toccnc calcarenitcs in th~ Olltsk.irts of Lecce (Apulia, SE ltaly), provided a wealthy amount of fossi l v<:rtt'br3tcs. Although dism~bered aod partly dispersed in severa} instirutions of Italy, thc fauna thar 5UTVi\'~d tht! vicis situdes of the years is srill enonnously rich, oount.ing 1886 mammalian Sp~ciLDcns and a la.rgt: a:mount of bird bone5. lt include.s rcmains of mSlD.IDoth (Mammuthus primigenius), woolly rllino (Coelodol1la
anriqulIaris), aurochs (Bos primigeniti3), rcd deer (Cervus dapnus), horsc (Equus fents), wolf (Canis IllpU.ç), red fox. (Vulpes vu/pes), spotLcd byena (Crocula crocuta), bedgl;!hog (Erinaceus
europaeus), bare (Lepus europaeus) md rabbit (OryclOlagus cunzcu/us). Thjs mammal comm.lln.i ty i:s suggeSTive of rclati\'t::ly open !aodscape;:s uDder [airly cold climatic conditions; a picrure also corroborated by thc avifaunal ~lements. Tbe fauna from Cardamone s~cms to be ao example of an ccotone wberein typical clements of the J1ammuthus-Coelodonta faunal complex (l"'{ primlgeni us , C. antiquilalis, E. ferus and C. crocuca) that probably immigrated from thc -east, art: mixed lO tbe more temperate, autochthonous fauna (characterized in particular by B. primigenius. C elap hus, L. europaeus and
o.
"mlculus). The extensive last Pleoiglacial se:1.·JevcJ !a.1l cambincd to intense late tectonics in tbe Adriatic area pn:sumably caused thc emersian of mOSI of thc: Adriatic sea-bottom. giving rise to a vaSi wide open plain which could" havt: acted as a corrido! b~twccn thc Balk.an peninsula andm
esouth-castem regions
ofItaly, permitting the arriva! of tht: eastcrn faun a! dements. This hypothesls would be coosiste nl with the abscnce of the Alammurhus-Coelodonla faunal complex ·elcmeDts in mast of centrai and sourhi:m lraly. AD alternative view is the classica! ODI.:, with thc pachyderrns foUowing a horseshoe coute from me north-cast soutbwards along the Adriatic shorcs of Italy. Finally, the abst'nce of E. hydruntinus and D. dama and the presencc of '\'fammwhus pnrnigenius and Cododonca may suggcst a dating of the assernblage ro the clima...;: of the last glac ial, that is at the sccond Wilimian Pleniglacial, which is dated some 22-18 k-y from pre sent.Comspondl.:ncc: ~larco Rustioni & Paul Mazza, MUSl:o di Storia ~aturaJe-Sezionc di Geologia e Paleonlologia, Università degli Studi di Firenze" Via La Ptra, 4·50121 Fircnz·e, Italy, \!-mail: pmazza@unifi.it; Marco P. Ferretti, Museo di Storia Naturale-Sezione di Geologia lo;} Paléontologia ..nd Dipartimento di Scienze: de.ua Terra, Università degli Studi di FirèDze, Via La Pira, 4-501 21 Firenze, ltaly, e-mai!: mferrett@geo.unifi.it; Marco Pavia, Università di Tor.ino, Dipan:iml!nto di Scienze della Terra, Via Accademia delle Scic:nzl;! 5, 10123 Torino, lt:lly, ~-mail: pavia@dst.uniro.it; Angelo Varola, Osservatorio di Fisica e Chimica della Terra I,,:: dcll'Ambientlt, Università di L~cce, Via Amesano, 73100 Lecce, ltaly
I
I
AoIf'N1J3 '" DEINSEA 9, 2003MAMMOTH REsEAACH
F~ I a-d: MOOlm'J!/Iu, pm'getlw a: Ircomplete jlMl\Ìle "13nd"," W1Ih mi in function,occlusa! vitm. 1/6 nat. size, b: detal cf ~sMwn In a: left m I. occlusal """",, I f2 not = c: ",complete juverile righI hemimandible with d4 In functIOf\ ocdusal vi... 1/6 nn
=
d: =ne .pecmeo .t.own IO '" lalerol VleW, 1/6 nat sile; e-m:Coelocbnto ortJqUIWtis e: .ncompk!te JlM'f'1e sIo"lI. leIì latenil ...""" 1/6 nat. 5IZe; f. incomplete juverule ,!<l,il.dorsaI vIew. 1/6 nat sile: g: Intanplet.l~ile sku!i. ocq,ital view. 1/6 nat.sile; h: iOCOmpk.'!e JLM:nI1. nghl nwdlary, v.ith M I and emerglnz M2, occIusal vi""" If2
'
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:i: inc~lete l""""e hemimandi bIe.left I,,~1/!eW, 1/6 nat Sile; I: iT-.:OfT4'Iete j"""",le hènwnandible. occ:lusal v",w. 1/6 nat >ile; m: incomplete j\M!nile n~t herri manclll*, (detaJl),1iI1gva1 view, 1/2 MI. size.RUSTlONI el 01: vertebrate fauna (rom Cardamone
INTROOUCTION
The rieh faunal assemblage fì-om Cardamone (Lecce, Apulia, southern Italy) \Vas found in a karslie cavity infilling in local Plio-PJeisto cene caJcarcnitcs. The cavity was fì.l!U1el-shapcd near the surface and passed to an e10ngated flue-shapcd vcrtical stm.cture. The latter widened abollt 15 m below thc surface and gave access to
a
subcircular underground cave about9
m in diameter. Thc cave was partly infilled with thc same sediment as thc vertical cavity, which piled up occluding the entrance to the cave. U. Botti, who is the author of the first report on th" fauna (Botti1890), collectcd rhe vertebrates. Subsèqucnt
shJdies were made in the 20" cen"tury by Vaufrey (1927), Guérin (1980) and Rustioni
(1998). The collèction was dismembered over
the years and partly disperscd to severa! insti tutions of Italy, while other specimens quoted in the literature now seem losl. Unfortunalcly, this prevented the reconsrmction of the ori ginai quantitative composition of thc fauna and therefore a taphonomic analysis was inhi bited. Retracing the bones stili preserved in the different institutions was the first concem of rhe authors. Allhough depleted, the fauna that survived the vicissitudes of the years is stili enormously rich and this Ijlakes it very significant from different viewpoints. MATERIAL ANO METHOOS The authors are aware that any possible observation is biased by the unavoidable sta
11/
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Figure 2 N""""" oi >l<eletal eItmeots of /VI. prit'ttige1IM (M P)
and C. GY'Ioqu;r,w, (CA).
tistical error represented by the incomplete nes> of the sample. Yet the impressive abul1 dance of thc specim.ens (1886 mammalian spccimens and 70 boxes each containing remains of single bird skt.:lctons) and the fact that practically allthe species rcported by Botti (1890) are stili reprcscnted (cxcluding
Felis, Arvicola and tvlm), convineed thc authors that thc fauna was worth eonsidering. The minimum number of individuals (lvINI) was thus caLculated and deemed to be t"idy reliabk The analytical approach adopted bere also included lhe construction oI dia grams of bOlle representarion of each specics and the observation or bone surface evidence. Most specimcns are parI of the natural histOly collections houscd by the Istiluto Tecnico O.G. Costa, at Lecce. Several others are pre served at the Museo Civico di Paleol1lologia c Paletnologia di Maglie, in rhe province of Lecce; a few at the Paleontological and Geological Seetion of thc Museo di Storia Naturale of Florence; an isolated Mammulhus primigenills toolh is at the Museo di Paleontologia G. Capellini, at Bologna. RESULTS
The bones do not show evidcnce of transpor tarion nor of butchering, scamification or of other human aClivity (Fig. I). Therefore they were not selected under natural circumstan ces. Diagrams of the bone representation of each species (Figs. 2 -5) confirm an expected artificial seleetion of the specimens, possibly
Figure 3 NLJ'!1l>er of >l<elet.ll ebnents of E.
r
'M
(EF). B.~(BF) ~C eJaph.Js (CE).r
I
Nwi'NCf!S IN MAMMOTH ReséAACHT,"* I Min;rrum rurber of ~als(jlM!o1les and Wb) and nurnb<!r of determ.-.ed >f"lC""""" per speàes.
5pecles juv adult NSP Idotonnlned)'
/,I pliiiiiOiiillus 3 D 971 C an tiqllilarJ.s 2 O 93 E. fe",s 9 29 542 a.tmmiaenius 8 13 321 C. .'aohUs 1 3 28 C.lupus 7 Q 103 V v!$O' 29 58 436 C.CrcCvt3 2 2. 2BJ Mustelidae E. europaeus
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1 43 Leuropaeu$ 6[ 3 73 O cunk:vlus ,atal 1 77 1 1 1191 5177
0
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due lO bOlh improper recovery (which proba bly explains the smali number of micromam mais and of other small vertebrates) and IO the following disl11cmbcring ofthe coliection. This is parlicularly evident in the anomaloLls ovcr- and under-reprcsentation of some post cranial clemenls wilh respect IO lhe number of boncs expected in condilions 'of normal accumulation in caves (Table l, Fig. 6). Notewol1hy is the extraordinary high number of rcd foxc. (87 individuals, 29 cubs and 58 adul1s) in the diagrams depicting the abun dance of the different species (Fìgs. 7 and 8); this may be the result of an attritional accu lTIulation of carcasses over a long time lapse. There are two possible explanalions for such an accumulation: (I) tbe cave was used as a shclter by these animals, which suggesls Ihat anòther entrance originaliy cxisted and was latcr obliterated, or (2) the cave acted as a
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(CC) and Mustd~ (Mu).DEiNSEA 9, 2003
natural trap. The former hypothcsis contrasls with the eco-ethology of Ihese carnivores. Red fox is not a gregarious animai and in nomlal conditions it does not form groups largcr than a female and its cub5. Thercfore the large amount of individllals encountered at Cardamone testifies the trapping of single specimcns over a long time lapse, although normally these animals are smart enough to avoid a natural trap, espccially ìf fdlow spe cimens fall into il. Assuming that the Wiirmian red [ox formcd family groups as large as those living today, the monaliry indexes would be deterrninant in assessing the lapse
01'
time possibly represented in the site. If one applies the mortality indcxes cal culatcd in extant red fox populations (Cavallini 1994) thc carcasses of these canids present in thc Cardamone tàuna would have accumulated in some 70 'lO 80 years. However, these mortality indexes, as those of ali European faunas, are heavi Iy biased by the statislical error of also including individllals killcd by modern man (specimens shot by hunters, killed by cars, etc.).lncidcntally, an interesting trait of thc red foxes presenUy living in tbe Mediterranean Basin is thcir genetic structure. Reccnt researches (Frati elal. 1998) have shown that red foxes can be
genetically divided inlo !wo distìnct groups: one includes the representatives frOtTI Spain, Sicily, Sardinia and Bulgaria, the other those from peninsular !taly and Austria. This has signitìcant paleogeographic implications
Figure 5 Number of sl<eletal c!encnts of
o.
ano.... (CC). l mopoeus (LE), E. ....-opoeus (EE)RUSTIONI et cl.: venebrrne fauna from Cardamone
I
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-F<gIR 6 Expected~. anualf, found abundonce or me bone elements!rom Carclamone.The CUNe of expec1.ed abund!InCe
depoct.s l!'te pattem of re!atM: ilbundances In an """"'8" skeleton. The C!'<01iun incll.ides alJ l!'te m.in par1S ,I IS composcd or (O(Cf p'llII. parietal, (rontal. e\c.).The pon:emages of specmens are reported on the __Ilcal ~,The diagram Is me&11 lO porttay the CNef'-or under-repre.c:nt3tion of bones
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,-espect to lhe 1~1sltw:tJon in wt.:h iIIJ bones tud been preserved and re<:o vered. Dari: grey t!l<pOOed NSP (%);ighl ~y: ~.,dNSP (%).which will be analysed more carefully later
ono
Besides foxes, tbe cave provided remains of mammoth (lv/!J/IIlnuthus primigenills),
woolly rhino (Coe/odonla unliqìlitalis),
aurochs (Bos primigenius), red deer (Cervus
elaphus), horse (EqIlLlsferus), wolves (Canis
lupus), spottcd hyena (Crocula crocula), hed
gcbog (Erillacells europaeus), hare (Lepus
europae/./s) and rabbit (Oryclolagus
cunicullls). Tbe perccntage of young indivi
duals is vCI")' high (Fig. 8); in particular the mammoth and thc woolly rhino are only
represented by juvenile specimens. The mam
malian community as a whole, and tbe high amotlnt of horse remains (Fìgs. 7 and 8), seem to be suggestive of fairly open forested stcppcs. This is apparen tly consistenr with note"'ortlty absences of some rypical repre sentalives of the Lat~ Pkistocene of !taly, such as Capreolu.\' capreolus, a characteristic woodland dwcllcr; Sus scrofa, a marshy woodland inhabitant; D. dama , which howe ver may be lacking for stratìgraphic reasons,
the Cardamone fauna likely postdating the last oc.:urrence of this species in Europe.
Caprine bovids are also lacking, perhaps excluded by the particularly flat physiograp hic characler oflhe area. At last, the absence
of E. hydruntinus and, on th.e other hand, the
presence of the \\'ooll'y rhino and of the mam motb, which substituted the more common
Elephas anli'flllls (=Elephas l1omadiclIs), are
suggestive of fairly cold climatic conditions, Tbc most significant occurrences from the stratigraphical viewpoint are those of the pachyderms, which scem to disperse in Italy dllring the last glacial (Palombo 1994;
Ferretti 1998). Tbe mammoth and, even more, the \\'oolly rhino are eXlremely rare in ltaly and many of the occurrences l'rom Ilaly reported in thc lilerature are due lO misatt'ri bution. Scanty remains of both were found in n0l1hem am! centrai Haly (a beautiful, but isolated skull of Coelodonla al1liq!litali~' was fOllnd at Monte Circeo promontory and described by Palmarell:i & Palombo 1981 )
400 200 100 O
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ÀDlANCI5 IN MAMMOTH~
T,t;je 1 Relat~ abundance in Ihe 3'M:
famiHy specimen Ardeidae Anatidae Accipitridae Falconidac Otidldae Scolopacidae I Columbidae 46 Cululidae Strigidae Corvidae other Pas5eriformes
ed in Apulia (Guérin 1980). The two species were never rcporLcd from any other part of southern !taly. This find l'rom Cardamone
thus confirms the occurrence of the woolly
rhino in Apulia and the sOlllhernmosl koown
occurrencc of Ihe mammoth in !Ialy, Cardamone cave also provided a weaithy
coll"ction of bird rcmains (Table 2). Ongoing sllidies on this material are performed by one of the authors (M, Pavia) and the results of a
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preliminary analysis can be given here, The avifauna is dorninated by Ihe Columbiformcs (pigeons), probably Co 111m ba livia, a bird Ihat I,ivcs in caves and on rock cliffs. Tbc presen
ce of water bircls, Anatidae (dllcks) and Ardeidac (herons), and some birds of prcy, Ha!iaelus albicilla and Circl/s sp" seem to indicale an extcnsion of fresh water or lagoons; Olidigae (buslards) also indicates open landscapes, The slighl amo'unt of small Passcriformes (songbirds) is probably due lo
improper colle.cting, The bird remains from
Cardamone are noi compklely rcpresentative of Ihe originai avifauna of the area; Ihis fact
inhibits taphonomic and detailed paleoecolo gical analyses. The prcl.iminary data on the
Cardamone avifauna suggest lemperate-cold
climatie conditions; the environm~nt of the area was probably charactcrised by an open
countryside with scattered wetlands or
lagoons not far from the cave.
436
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RUSTIONI et ul.: vertebrate fauna from Cardamone
I
MEDITERRANEAN MAMMOTH Saiga and Ovibos, on thc other hand, neverFAUNA reachcd lIaly.
Aithough e how et al.'s (1959) and Kahlke's The occurrencc 01' Mammllllllls and
(1994) Late Pleistocene MamnwlhusCoelodonra in limitcd arcas or northern Iraly,
Coelodol1la faunal complex in Italy is slightly in the centrai Tyrrhenian sidc of !taly and in
and incomplelely represented only in the the Salento peninsula, and their absence in
nOl1h, single components can also be found in most of the J\driatic belt, as \Vcll as the over other ltalian regions, RUl1g!ler tarandus is mentioned genetic similarity between rcd reportcd only from Liguria (Palma di Cesnola foxes presently living in southern Italy and 1983; Masseti el al. 1995); Alces alces from easlern Europe, may altogcther suggest that at
Liguria, Veneto and Trentino (Masseti et al. the time of the Ca rdamone fauna rhe paleo
1995); Megaloceros gigal1teus reached cc n geograpl1ic setting was quitc different from tralltal)' (Pasini 1969, 1970); Bisol1 priscus loday, The combinalion of an cxtensive sea disperscd to Sicily (Masseti el al. 1995); E. level fall and inte·nsl! late tectonics in the
fems is widcJy diffused in most of Italy; the Adriatic area may have caused the emersiOI1
cam ivores Calli.,· lupus, Ursus spelaeus, of 1110st of the Adriatic sea-bottom, giving
Panlhera leo,
P
pardus, I,ynx Iynx and rise to a vas t wide-opcn plain which actcd asCrocala crocuta are \Veli represenled a corridor l'or lhe pachydelllls of the Balkans
through-out Italy (C II/pus, P leo and C croto the sOllth-eastern regions of Ilaly and for red CI/la reached Sicily; Bonfiglio and Burgio foxes to southem Ilaly and Sicily, Sea-bottom
1992); Culo gulo was rare , Other steppe taxa maps of the Adriatiç Basin shol\' an abrupt
not mentioned by Kahlkc (1994) are thiàening of Ihe isoball1s ofr thc shorc-line
Dellolona and Sicisla, which are reported jllst north of Pescara and another less marked
from Veneto and from the Adriatic side of thickeniog ENE off the Gargano proll1onlory centrai Haly (Bartolomei 1980; Sala 1990). (Fig. 9). These two step-like structures are
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ADvANCES IN MAMMOTH RESEAROisouthwcst-northeast trending faull syste01s wh ich 10\Ver th~ sea-bonom from -150 m bsl to about -200 m and -500 m, respeclivcly. These structuml lin~aments are transform fault sys tcms' with a considerable venical component (dr P. Bellini, perso comm., 1999). The fauhs have been active since remote time, but their last intense activation was very recent (possi bly c\'en Holoccne in age) and they are stili \'cry activc today. Therefore, in conclusion the possibility of an extensive emerged plain stretched ali the \Vay down to Pescara or even farthcr south to the Gargano promontory, recently disrupled in its southern margin by tne mentioned fault systems, does not seem to be so remote.
Alternatively, the pacbyderms may have followed a horscshoe route from the north eaSlern Balkan areas southwards along the Adriatic sbores of Ilaly (Sala 1990). This is the classical model, \Vhich however does not cxplain the absolute abscnce 01' M primigeni
us and
C.
cm/iquita/is in centrai !taly as \Velias in O1osl of the somhern regions of thc peninsula. Thc palcobiogeographic and pale ogeographic sCllings seen so far lead to lhe
conc lusion that the Cardamone fauna is likely
the l'esuli of a O1ixture of typical elements of the J'vlammUllrus-Coelodon/a faunal complex
~
8\~
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r!gU'e 9 Schematic map
showr>&
O.. -12<:m to ,2000 m bsll50baths IO lhe Adnatìc: bilSlll of! Pesora and the Gargano prornoir'
'''Y.
mem
..,
5WNE-trendl1g trons[orm fatlt system~ ooe of the sugg...d ways of i:rmigrat"" of the MooYnu~1oct>nra 1à...1COIT'jllox elemento \OH." h Ap.JI.. (Indcated by the 'NJde dotted am:>w)DEINSEA 9, 2003
(M primigenius,
C.
an/iquila/is, E. fems and C crocu/a probably immigrated from tbe cast, along with red fox) with more tempera te, autochlhonous faunas (cha ractcrised , in particular, by B. primigenills, C. elaphus, L.europaell' and Q. clIl1iculus). IL apparently
reprcsents an t:cotone connecting the cold Balkan bioprovince with Ihe temperate !\'lt:diterranean and lhcrefore suggests that Apulia formed a paleobioprol'ince somehow climatically and paleogeograpically connec ted with the Balkaos. The combined absence of E. hydruntil1w' and D. dama may define the chronological position of this ccotonc, since it is widely agreed that the two species were absent al thc climax of the last glaeial, that is at the sccond W o..rmianl Weichselian Pleniglacial, which is dated some 22 -18 ky before presen l.
ACKNOWLEDGEMENTS The authors are deeply indebted to prof. Nicola Greco and to mr Giancarlo De L.lImé for grahting access, respectively, to the matc riai kept at thc Istiluto Tecnico O.G. Costa and to the Museo Civico di Paleontologia e Paletnologia di Maglie.
RUSTION/ e! al;venebrate fauna [rom Cardamone REFERENCES
BanolomC'i, G., 1980 -Micromarnmiferi dci Plio
Ph:istoccne - I Vcnebrali FossiJi Italiani, catalogo
della InOSl f1!, Vernna, pp. 249-258
Bonfiglio, L. & Burgio, E., 1992 - Significato poleonm bienlalt: c:: crollologico delle mammalofaune pldstoce
nir.:he della Sicilltl in relazione all'evoluzione paleo
geografica - Il QUaiemario 5 (2): 223-234
Botti,
u.,
1890 -La grona ossifcra di Cardamone inTerreS d'Olramo (lecce) - Bollettino della Società
Oeologica llalion. 9 (3): 3-30
C:t.vul!ini, P., 1994 - Variazioni nella biologia ddla volpI: Vulp~vulpa (Linn:n:us) -Doctoml Ihcsis IJniv~rsily of Siena
Cho .., M., Ch.ng, Y.P., Hu, C.K., Li\!, H.l., l'l,u, Y.H.,
Chou, P.H., Lee, Y.C. & IIsicn, I U I., 1959
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Ferretti, i\·1.P., 1998 - Gli elefunti dci Plio-pieistocenc
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