Microsoft Word - 00_indice_LM04.docx Il Quaternario IT ISSN 039-3356 Italian Journal of Quaternary Sciences 24 (1), 2011 - 93-101 A PATHOLOGIC STRAIGHT-TUSKED ELEPHANT FEMALE FROM NEUMARK NORD (GERMANY) Federica Marano1 & Maria Rita Palombo1,2 1Università degli Studi di Roma Sapienza, Roma 2Università degli Studi di Roma Sapienza, IGAG - CNR, Roma Corresponding author: F. Marano ABSTRACT: Marano F. & Palombo M.R., A pathologic straight-tusked elephant female from Neumark Nord. The elephant bones collected during more than ten years (1985-1996) in the interglacial (MIS 7 or MIS 5) lacustrine deposits exposed in the open-cast lignite mine of Neumark Nord (Germany), represent one of the most outstanding samples of straight-tusked elephants in the Pleistocene of Europe. In 1994, a skeleton of an adult female (E8 HK 2007:55) was found in nearly anatomical connection. The postcranial bones were almost all preserved, the skull was partially destroyed, and the tusks and mandible were missing. Wearing stage of molars and fusion of bones, as well as the pelvis morphology indicate a 39-40 years old female for which a height at the shoulders of about 2,9 m and a weight of about 5000 kg have been calculated. The most peculiar traits of this female are the serious scoliosis that affected its vertebral column and the invasive arthritis that deformed its femur and pelvis girdle on the right side. RIASSUNTO: Marano F. & Palombo M.R., Una femmina di “elefante antico” con malformazioni ossee del giacimento di Neumark Nord (Germania). I resti di Palaeoloxodon raccolti per più di dieci anni (1985-1996) nei depositi lacustri (MIS 7 o MIS 5) nella miniera di lignite di Neumark Nord (Germania), rappresentano una tra le più ricche popolazioni di “elefante antico” nel Pleistocene Europeo. Nel 1994, fu rinvenuto uno scheletro pressoché completo e in parziale connessione anatomica di una femmina adulta (E8 HK 2007:55). Il cranio è parzialmente danneggiato, mancano le difese e la mandibola, mentre le ossa dello scheletro postcraniale sono in discreto stato di conservazione. Il grado di usura dei molari superiori, lo stato di ossificazione delle ossa degli arti e la morfologia del bacino indicano che lo scheletro ap- partiene ad una femmina di circa 39 - 40 anni di età, alta alla spalla circa 2,9 m con una massa corporea di circa 5000 kg. Questo esem- plare presenta interessanti patologie ossee, in particolare l’intera colonna vertebrale è stata affetta da scoliosi, mentre il femore destro e il bacino sono stati interessati da iperossificazione. Entrambe le patologie furono probabilmente conseguenti ad un trauma giovanile. Key Words: Palaeoloxodon, Age, Sex, Body Mass, Pathology, Pleistocene, Germany. Parole-chiave: Palaeoloxodon, Età, Sesso, Massa corporea, Patologie, Pleistocene, Germania. 1. INTRODUCTION The site of Neumark Nord is one of the richest in remains of palaeoloxodont elephants and undoubtedly the richest in Central Europe. Since the early 80’s, an extraordinary rich sample of fossil remains was retrieved from the lacustrine deposits exposed in the open-cast lignite mine of Neumark Nord 1 (Sachsen-Anhalt, Middle East Germany) (MANIA, 2010) (Fig. 1). At Neumark-Nord 1, a long stratigraphical succession documents the envi- ronmental evolution of an endorheic basin during a complete interglacial climatic cycle. The lacustrine de- posits have been supposed to be late Middle Pleisto- cene in age (MIS 7) on the basis of vegetational data, indicating subcontinental, mild climatic conditions (GRÜBE, 2003; SEIFERT-EULEN, 2010), as well as of the presence of Apodemus maastrichtiensis, a species present at Weimar-Ehringsdorf but not recorded from the Eemian deposits of the area (HEIRICH, 2001, 2010). This chronology has been questioned by some authors, who considered the interglacial to be the Eemian (MIS 5e) (KOLFSCHOTEN, 2000; KOENIGSWALD, 2007, BOETT- GER et al., 2009 and references in those papers). Re- cently ESR dating of few samples might support an Ee- mian age (SCHÜLER, 2010), whilst data obtained by amino-acid racemisation of the “intra-crystalline” fraction of Bithynia tentaculata opercula are consistent with a late Middle Pleistocene age (PENKMAN, 2010). The la- custrine sediments lie with no hiatus over a ground mo- raine ascribed to the “Drenthe-glaciation” by MANIA (1999), and are overlaid by thick loess layers, then the succession is closed by muddy interglacial sediments and glacial deposits, respectively, thought to be Eemian and Weichselian in age (MANIA, 1999, 2004) (Fig. 2). The rich palaeontological record comes from the first interglacial cycle, and counts about two hundreds of plant species, invertebrates, insects, and a diversified vertebrate fauna, including several, more or less com- plete, skeletons of large mammals (elephants, rhinoce- roses, fallow deer, red deer, aurochs as well as some carnivores such as wolf, lion and spotted hyaena). Ele- phant remains were retrieved from 6 fossil-bearing lay- ers within the lower, middle and upper »gyttja«, which represent successive episodes within the small lake fill- F. Marano & M.R. Palombo 94 ing (MANIA 1999, 2004) (Fig. 2). Taking into account the place the remains were collected from, their dimensions as well as the progression in tooth eruption and epiphy- sis fusion, the minimum number of elephant individuals found at Neumark Nord could be nearly 70, though the calculated number would be overestimated, because of the difficulty to assessing the skull fragments, and large- ly incomplete bones from “bone beds”, which were se- riously damaged by excavators (PALOMBO et al., 2010). According to the research performed to date, the mor- phology and biometry of skulls, tusks, molars and post- cranial bones indicate that the specimens fall within the variability of European straight-tusked elephants (see e.g. OSBORN, 1942; TREVISAN, 1949; MACCAGNO, 1962; AGUIRRE, 1968-1969; BEDEN, 1969; KROLL, 1991; TSOU- KALA & LISTER, 1998; DAVIES, 2002; PALOMBO et al., 2010). This paper is aimed at assessing the ontogeneti- cal age, sex and body size as well as to briefly de- scribe the pathological bones of an almost complete skeleton found in the October 1994 in the fossiliferous level 6.1, at a distance of about 150 m from the coas- tline of the lake basin (Fig. 2, 3). At the time of discov- ery, the skull was still in articulation with the vertebrae, the pelvis was in anatomical position, while the long bones were disarticulated and partially dislocated. The exceptionally well preserved stomach contents, with the remains of its last meal, were found under the chest (GRÜBE et al., 2010). The skull is partially de- stroyed, the maxilla is well preserved and the molari- form teeth are located in the alveoli, while tusks and jaw are missing. The postcranial bones are almost all preserved except for carpal and metacarpal bones, while the tarsals are represented by calcaneus, astra- galus, navicular, cuboid, right mesocuneiform and ec- tocuneiform bones, some sesamoids are also present (Fig.4). This specimen (E8: HK 2007: 55) is stored in the Landesmuseum für Vorgeschichte in Halle Saale (Germany). 2. METHODS The completeness of the remains has made it possible to determine both age and gender, as well as body mass and shoulder height of the studied E8 individual. The age was inferred according to the progression in tooth eruption and wear of plates of the upper molars, adapting to these teeth the method tested by LAWS (1966) on the lower molars of extant Afri- can elephants (Loxodonta africana) and also applied to fossil species (ROTH, 1984; HAYNES, 1991). The status of the epiphy- seal-diaphyseal junction of long bones has also been taken into account (ROTH, 1984; HAYNES, 1991). About the gender determination, sev- eral features have been regarded as of po- tential information about the sex of a pro- boscidean, e. g. body size, length and stoutness of long bones, morphology and size of skull and teeth (molars and tusks), atlas and epistropheum, pelvis and carpal bones (see PALOMBO & VILLA 2003, and references therein). The gender of E8 elephant was in- ferred based on the morphology and dimensions of the pelvis girdle, which have been demonstrated to be par- ticularly appropriate and have been largely applied to determine the gender in fossil proboscideans (KROLL 1991, LISTER 1996; GÖHLICH, 2000; PALOMBO & VILLA, 2003). Body mass was considered as the best proxy of body size, according to GINGERICH et al. (1982). Body mass of proboscideans has been estimated using dif- ferent allometric equations (e.g. ROTH 1990, PALOMBO & GIOVINAZZO, 2005; CHRISTIANSEN, 2004). Here, we cal- culated the body mass of E8 by means of the regression equations proposed by CHRISTIANSEN (2004), regarded as the most appropriate when either the actual shoulder height or the inferred circumference of manus are not available. Measurements were taken only on the left long bones because of the pathology affecting the right hind leg of E8 elephant. The stature was inferred combining the results of direct measurements of the forelimb (but acropodi- al bones) and the equation proposed by OSBORN (1942), this because of the poor preservation of the hu- merus, for which only the functional medial length is available. 3. DESCRIPTION AND RESULTS 3.1 Main morphological features Skull - According to the preserved fragments of skull, the nuchal fossa was deep and showed rough sur- face (Fig. 5b), the frontal bones flat and rather devel- oped antero-posteriorly. The fan, is moderately ex- panded; the tusk-sockets are rather small and gently bent towards the lateral side (Fig. 5e), the premaxillary fossa becomes rapidly shallow, broad and somewhat convex towards the distal end of the fan (Fig.5c). Maxil- lary bones have a thin and short pyramidal process, the last molariform teeth (see below) were not completely A pathologic straight-tusked elephant female … 95 erupted, and residual, small alveoli are present on both sides. Teeth - The not completely erupted molariform teeth have 8 plates in use, while a not evaluable number of plates, all fused together, occupy the entire free por- tion of alveoli, whose posterior portion is well ossified. The lateral side of the alveoli is locally broken, enabling us to see how the roots of molars are completely formed and fused to the plates (Fig. 5d). Accordingly, the teeth have been regarded as last molars (M3) (MARANO, 2009), consistently with the degree of epyphyseal- dyaphyseal fusion of long bones (see below). The morphology and dimension of the molars fall in the range of south European straight-tusked elephant populations. The molars are narrow and high crowned, the occlusal surface is ovate and slightly convex, plates are slightly bowed out on the lateral side, in the less worn plate, the enamel occlusal figures are formed by a large oval central loop and two small lateral loops. At an intermediate wear stage, the enamel figures show a moderate medial expansion. The enamel is thin, crin- kled, densely and regularly folded, and the folds extend on the entire length of the enamel figure. Stylohyoideum - In both right and left stylohyoi- deum the posterior ramus is broken, the superior ramus, the less developed, is moderately elongated, and the inferior ramus is slender, gently curved upwards and backwards, without any evident lateral deflection; in an- terior view it shows a gently concave lateral outline. Postcranial skeleton - The vertebral column pos- sesses 7 cervical, 19 thoracic (including ribs as well), 4 lumbar, 5 sacral vertebrae, while the maximum number of caudal vertebrae is unknown. The morphology of the limb bones and pelvis girdle is consistent with those known for the European straight-tusked elephants (e.g. MACCAGNO 1962, TREVISAN 1949, KROLL 1991, TSOUKA- LA E LISTER 1998): the humerus is characterized by a large and angled deltoid tuberosity and a more or less deep deltoid fossa; the ulna is massive with a robust olecranus and wide distal articular surface; the left fe- mur has a relatively short neck, angled on the diaphysis of about 40°, the diaphysis is moderately twisted as is commonly found in femora of adult individuals; the tibia possesses a weak cnemial crest, the distal epiphysis is smaller than, but nearly as massive as the proximal epiphysis. The general shape of the tarsal bones does not substantially differ from that of the actually quite va- riable European straight tusked elephants (TREVISAN, 1949; KROLL, 1991; FERRETTI, 2008). The pelvis shows a gently curved lateral profile of the ileum wing, less rounded than in Elephas maximus, but more rounded than in Loxodonta africana africana and Mammuthus primigenius (see PALOMBO & VILLA 2003 for a discus- sion). With the latter two species, the pelvis of E8, as well as that of straight tusked elephants, shares a ileum with a more transversally elongated wing. 3.2 Age, sex, body size Age - The peculiar progression in displacement of the molariform teeth, characterizing elephants, provides a particularly useful means for determining their age. Accordingly, the first step in determining the age of E8 elephant was assessing the age on the basis of the eruption progress and the degree of wear of its upper Fig. 2 - Neumark-Nord 1 basin, stratigraphical section. On the left the lake at the time of the deposition of lower (a), middle (b) and upper (c) »gyttja« (Mania, 2004); dots indicate the main find- ings of elephant remains, the star indicates the place where the skeleton of E8 was found (modified from PALOMBO et al., 2010). Sezione stratigrafica del bacino di Neumark-Nord. Sulla sinistra il lago al momento della deposizione del »gyttja« inferiore (a), medio (b) e superiore (c (Mania, 2004); i punti indicano i princi- pali ritrovamenti dei resti di elefanti; la stella indica il punto del ritrovamento di E8 (modificato da Palombo et al., 2010). molars. In E8 both molariform teeth (M3) are not com- pletely erupted, furthermore, the M2 were already ex- pelled, but residual, small alveoli of M2 are still pre- s s 96 Fig. 3 - Skeleton o Lo scheletro dell’e Fig. 4 - Skeletal served and studie Schema dello sch scheletrici trovati of elephant E 8: s esemplare E8: in plan of E8: in re ed. heletro dell’esemp e studiati. stomach content in nero il contenuto ed the skeletal el plare E8: in rosso n black (modified dello stomaco (m ements pre- gli elementi after Mania, 2010 modificato da Mani served. The vanced: the 8 plates, sho first five plat proposed by ing into acco and upper tusked elep Neumark No E8 specime corresponds The ag and wear o fusion of th vertebral bo indeed, are traces of on Gende consistent w wide and ro ameter of 5 and flattene of the pelvic is 3.1, while aperture ver values fall w TER, 1996; values of th males. 0). ia, 2010). e stage of wear occlusal surfac owing a comple tes (Fig. 5a). Ac y LAWS (1966) f ount the similar molariform tee phants, and in ord rich sample en falls in the LA s to an age rang ge inferred on of molars is co e axial and lim order of the sca almost complet going ossificatio er - The morph with that of a fe ounded pelvic a 540 mm, reduc ed ilium. The ra c aperture versu e that of the d rsus the minimu with in the varia PALOMBO & VIL he same ratios F. Maran r of M3 is not p ce (120 mm of ete fused enam ccording to the for the lowers m progression in eth often show particular obs e (PALOMBO et AWS’s XXII age ging from 37 to 4 the basis of th onsistent with th mb bones (ROT apula and the c tely fused, show on (Fig. 6). hology of the p male: it is chara aperture with a ced thickness o tio of the maxim us the minimum diagonal height um width of iliu ability range of LLA, 2003; KRO s being signific no & M.R. Palombo particularly ad- length) counts el loops in the pattern of age molars, but tak- the both lower n by straigth- served in the al., 2010), the e group, which 40 years. he progression he epiphyseal H, 1984). The caput femuris, wing no or faint pelvis girdle is acterized by a transverse di- of the ischium mum diameter m width of ilium t of the pelvic m is 3.4. Both f females (LIS- OLL, 1991), the antly lower in A t f m d s A pathologic straight Fig. 5 - The skull the penultimate h fused together an Il cranio di E8 in v molari sono già st altre lamine in for Fig. 6 - Scapula s distal cartilage. Scapola con ossif simale. t-tusked elephant fe of E8 elephant in have already bee nd occupies the en vista ventrale (a), tati espulsi dagli a rmazione. showing the still im ficazione quasi co male … ventral (a) poster en expelled from ntire portion of the posteriore (b) e la alveoli. Il particola mperfect ossificati ompleta dell’estrem rior (b), lateral (c) the alveoli. The p e alveoli. aterale (c). Sono v are degli alveoli, vi on of the mità pros- and frontal (e) vie postero-lateral vie visibili gli ultimi mo isti in norma poste Body s 8511,38 kg on the basis physis of th maximum le mass of abo using the m gives the be coefficient a falling close can and Asi The sh the forelimb possible hei it is about 2 OSBORN (19 10% higher about the sa 3.3 Patholo Some Nord show a frequent am such as the some cauda top, while in brae, arthrit ew showing the p ew of the alveoli olari non del tutto ero-laterale (d) mo size - The estim to 3162,27 kg. T s of the minimu e humerus, the ength of the tib out 5900 kg. T minimum circum est prediction th and low error ra e to the maximu an females (HA houlder height, i b long bones a ight of the miss 290 cm by usin 942). Accordin r than a fema ame age (LAWS ogy of the elephan anomalous ossi mong vertebrae a e atlas, the oss al vertebrae the n others, such is or arthrosis h artially erupted la (d) shows the pl fuoriusciti, mentr ostra come non si mated body mas The first value w um circumferen e second on th bia, yielding an The body mass mference of the hanks to the hi ate, is approxim um body mass AYNES, 1991). inferred by direc and tacking into sing more than ng the equation gly, E8 should le of Loxodont , 1996; HAYNES nt bones found fication/growth, and teeth. In so ification is inco neural channel as the lumbar had caused hy 97 ast molars, whilst lates completely re i penultimi iano presenti ss ranges from was calculated nce of the dia- e basis of the average body calculated by e ulna, which igh correlation ately 4900 kg, of extant Afri- ctly measuring o account the 250 cm, while n proposed by d be of about ta africana of , 1991). d at Neumark- which is most ome vertebrae, omplete and in is open at the r sacral verte- perossification w t t g s w f d i v g 98 and anomalous The skele which certainly bits of the eleph cimen are the s column and the and pelvis on th umn, the last th the lumbar and the vertebral co girdle, the righ shows a consp while the ilium mur, 7% shor Fig. 7 - Pelvic gir formation and hyp left: the correspo deformation. Cinto pelvico dell iperossificazione destro con il capu Fig. 8 - Detail of t Dettaglio dell’ace arthritis (Fig. 9) normally bent, versely, no evid epiphysis or th arthritis and art er, domesticate elephants, mos goad injuries c s fusion of verteb eton of E8 sho have affected hant. The most serious scoliosis e invasive arthrit he right side (F horacic vertebra sacral ones ar olumn is marke ht acetabulum picuous hyperos shaft is sclerou rter than the rdle of E8 elepha per-ossification of onding femur show ’esemplare E8 co dell’acetabolo des ut marcatamente d he right acetabulu tabolo destro dell ): the caput fem the neck lengt dent anomalies he zeugopodium throsis are, actu d Asian elephan st of pathologica caused by keep bral bodies. ows some mal the quality of li peculiar traits o s that affected it tis that deforme ig. 7). In the ve ae arch toward rch toward left. A edly twisted. In is anormously ssification along us (Fig. 7, 8). Th left, was def ant showing the e f the right acetabu wing the caput w on deformazione d stro. Sulla sinistra deformato. um of E8. l’esemplare E8. moris is elongate th strongly redu s affected eithe m and achropod ually, common a nts. In domestic al ossifications pers. Several c formations ife and ha- of this spe- ts vertebral ed its femur ertebral col- right, while As a result the pelvic large and g its edge, he right fe- formed by xtensive de- ulum. On the with a strong diffusa e a: il femore ed and ab- uced. Con- r the distal dium. Both among old- cated Asian are due to cases have also been re young anim difficulty but pear. Thus fects becom 4. DISCUSS Morph obtained by understandi variability of Althou laeoloxodon related to bu HASI, 2004; S 2010), in the the nomenc phants”. Dis onomic statu loxodon”, as as present elephants h the Asian sp fide species Fig. 9 - Fem showing the caused by the time. Femore dell’e marcata defo artrite l’esemp eported for zoo mals tolerate tra t, when they gro it would take a me clear. SION hological feature y biometrical an ng of the mor f European strai ugh several dat n as a bona fid ut separated fro SHOSHANI et al. e literature, ther clature for Eur sagreements e us (genus or su s well as on the in Eurasia. Th ave been cons pecies “Elephas s, Elephas antiq ur of E8 elephan extensive defo e severe arthritis esemplare E8 in n ormazione e ipero plare ha sofferto p F. Maran o elephants too. umatic injuries ow older, the p a long time befo es shown by E nalysis, contribu rphological and ight-tusked elep a concur, to de de elephantine g om Elephas (INU ., 2007; PALOMB re are different o rasian “Straigh ssentially cente ubgenus) ascrib e number of spe he European s idered as belon s namadicus” or quus/Palaeoloxo nt in anterior and ormation and hy the female had s norma anteriore e ossificazione. A c per lungo tempo. no & M.R. Palombo It seems that without much problems reap- ore arthritis ef- E8 and results ute to a better d dimensional phants. esignating Pa- genus, closely UZUKA & TAKA- BO & FERRETTI, opinions about ht-tusked ele- er on the tax- ed to “Palaeo- ecies regarded traight-tusked nging either to r, as one bona odon antiquus. d posterior view, yper-ossification, suffered for long e posteriore con causa della forte A pathologic straight-tusked elephant female … 99 Alternatively, both these species have been considered present in Europe. Two morphotypes have been recent- ly described by SAEGUSA & GILBERT (2008): the Italian “namadicus-like” morphotype (showing an extreme de- veloped parieto-frontal crest, nearly reaching the exter- nal choanae), and the “German-Stuttgart” morphotype (characterized by a weak development of the parieto- frontal crest). Although the parieto-frontal and nuchal crests of E8 skull were seriously damaged during the recovery, some preserved large fragments indicate that its fore- head could have been wide and gently convex. Accor- dingly, the morphology of the preserved parieto-frontal part of the skull of E8 seems to confirm the peculiar de- velopment of the parieto-frontal crest, shown by some adult males from Neumark Nord (PALOMBO et al., 2010), which differentiates the skulls of Central Europe (espe- cially from Germany) from those of Southern Europe. Although the development of the parieto-frontal crest of the skulls from Neumark Nord are variable, they are mainly characterized by a moderately extended fore- head and by a dorsally positioned parieto-frontal crest, as observed in skulls ascribed by SAEGUSA & GILBERT (2008) to the “Stuttgart” morphotype. Accordingly, most of the German palaeoloxodon populations do not present the morphological “namadicus” traits shown by the Italian continental adult elephants ascribed by SAE- GUSA & GILBERT (2008) to this morphotype e.g. skulls from La Polledrara Cecanibbio, (PALOMBO et al., 2003; PALOMBO & FERRETTI 2005) - as well as by the endemic Sicilian species Palaeloxodon mnaidriensis, showing a parieto-frontal crest strongly advanced and folded to cover the forehead (FERRETTI, 2008; PALOMBO et al., 2010). Whatever the significance of such a difference, the morphology inferred by the skull fragments of E8, seems to confirm that females possibly had a less de- veloped crest than males. The age of about 40 years, inferred on the basis of the degree of wear and number of plates of upper molars, is also supported by the degree of ossification of skeletal elements. This age confirms a prevalence of adults older than 45 years, in the elephant popula- tion of Neumark Nord while only the 3% of the popula- tion is younger than 6 years. The body mass of E8, 34% lower than average body mass estimated for Neumark Nord elephants, is consitent with the small size of this individual, confirming the great morphological and dimensional variability of this species and the significant dimensional gap be- tween male and female in fossil and extant elephant populations (PALOMBO et al., 2010). Moreover the con- cordance among dimensional data, pelvis morphology confirms the usefullness of some pelvis features (partic- ularly shape of pelvic aperture and proportion of the ilium shaft) as support to infer gender in proboscideans. 5. CONCLUSION The specimen E8, as are all the elephants of the population of Neumark Nord, is assigned to the species Palaeoloxodon “antiquus”, pending a systematic revi- sion on the European straigt-tusked elephants (PALOM- BO & FERRETTI, 2010). The analysis on the nearly com- plete skeleton, indicates that E8 was a small female of about 40 years, weighting about 5000 kg, with a height at the shoulder about 290 cm. Accordingly, E8 has al- most the same size of females, but is significantly smaller than males of the same age from the same population. Moreover, the presence of extensive bone abnormalities, indicats that the female was lame in its right leg and perhaps suffered a lot of pain during its life. The extensive sclerosis suggests that the arthritis de- veloped over a quite long time, and probably was caused by a traumatic injury when the female was young, albeit it is difficult to ascertain what event actual- ly caused these injuries. ACKNOWLEDGEMENTS We are indebt to Prof. H. Meller for inviting one of us (Maria Rita Palombo) to study the elephants from Neumark Nord. We thank everyone at the Landesmu- seum für Vorgeschichte of Halle for granting us access to the fossil material in their care and for their kindness and help. We are grateful to the anonymous reviewers for their comments on the manuscript. REFERENCES AGUIRRE, E., (1968/69). Revisión sistemática de los ‘‘Ele- phantidae’’ por su morfologıá y morfometrıá denta- ria. Estudios Geológocos, (Madrid), 24, 109-167; 25, 123-177, 317-367.BEDEN, M., (1969) - Etude et réconstruction des restes de Palaeoloxodon anti- quus du Chartelard (Charente). Bulletin Sciences de la Terre, Université de Poitier, 10, pp. 44-56. BOETTGER T., NOVENKOB E.Y., VELICHKO A.A., BORISO- VA O.K., KREMENETSKI K.V., KNETSCH S., JUNGE F.W. (2009) - Instability of climate and vegetation dynamics in Central and Eastern Europe during the final stage of the Last Interglacial (Eemian, Mi- kulino) and Early Glaciation. Quaternary Interna- tional, 207 (1-2), pp. 137-144. CHRISTIANSEN P. (2004) - Body size in proboscideans, with notes on elephant metabolis. Zoological Jour- nal of the Linnean Society, 140, pp. 523-549. DAVIES, P., (2002) - The Straight-tusked EIephant (Pa- laeoloxodon antiquus) in Pleistocene Europe. PhD unpublished thesis, University of London. FERRETTI, M.P. (2008) - The dwarf elephant Palaeo- loxodon mnaidriensis from Puntali Cave, Carini (Sicily; late Middle Pleistocene): Anatomy, syste- matic and phylogenetic relationships. Quaternary International, 182, pp. 90-108. GINGERICH, P.D., SMITH, B.H., ROSENBERG, K. (1982) - Allometric scaling in the dentition of primates and prediction of body weight from tooth size in fossils. American Journal of Physical Anthropology 58, pp. 81-100. GÖHLICH U.B. (2000) - On a pelvis of straight-tusked elephant Elephas antiquus (Proboscidea, Mamma- lia) from Binsfeld near Speyer (Rhineland-Palatine, Germany). Paläntologische Zeitschrift, 74, pp. 205-214. GRÜBE R. (2003) - Planzliche Nahrungsreste der fossilen F. Marano & M.R. Palombo 100 Elefanten und Nashorner aus dem Interglazial von Neumark - Nord (Geiseltal). Veroffentlichungen des Landesamtes für Archäologie Sachsen-An- halt, 57 (1) pp. 221-236. GRÜBE R., PALOMBO M.R., IACUMIN P. (2010) - What did Elephants from Neumark Nord eat? Kataloge zur Dauerausstellung im Landesmuseum für Vorge- schichte Halle: Elefantenreiche - Eine Fossilwelt in Europa. Landesmuseum für Vorgeschichte Halle (Saale), pp. 252-273. HAYNES G. (1991) - Mammoths, Mastodonts, and Ele- phants, Cambridge University Press, Cambridge, 395 pp. HEINRICH W.D. (2001) - Kleinsäugerreste aus intergla- zialen Ablagerungen von Neumark - Nord, Mittel- deutschland. Praehistoria Thuringica, 6/7 pp. 132- 138. HEINRICH W.D. (2010) - Klein aber oho: Zwergwaldmaus Apodemus maastrichtiensis. Kataloge zur Daue- rausstellung im Landesmuseum für Vorgeschichte Halle: Elefantenreiche- Eine Fossilwelt in Europa. Landesmuseum für Vorgeschichte Halle (Saale), pp. 79-81. INUZUKA N., TAKAHASHI, K. (2004) - Discrimination be- tween the genera Palaeoloxodon and Elephas and the independent taxonomical position of Palaeo- loxodon (Mammalia: Proboscidea). In: Baquedan, E., Rubio, S. (Eds), Miscelanea en homenaje a Emiliano Aguirre. Zona Arqueologica 4 (2), pp. 234-244. Museo Arqueologico Regional, Madrid. KOENIGSWALD W. VON (2007) - Mammalian Faunas From the Interglacial Periods in Central Europe and Their Stratigraphie CorrelationThe climate of past interglacials. In SIROCKO F. (Ed.) The Climate of Past Interglacials, Developments in Quaternary Sciences, 7, pp. 445-454 KOLFSCHOTEN T. VAN (2000) - The Eemian mammal fau- na of central Europe. Geologie en Mijnbouw, 79 (2/3), pp. 269-281. KROLL, W., (1991). Der Waldelefant von Crumstadt. Ein Beitrag zur Osteologie des Waldelefanten Elephas (Palaeoloxodon) antiquus Falc. and Cautl. (1847). Inaug. Diss., Tierärztl. Kak., Univ. München. LAWS R.M., (1966) - Age criteria for the African ele- phant, Loxodonta a. africana. East African wildlife Journal, 4, pp. 1-31. LISTER A.M. (1996) - Sexual dimorphism in the mam- moth pelvis: an aid to gender determination. In: J. SHOSHANI/P. TASSY (eds.), The Proboscidea (Ox- ford), pp. 254-259. MACCAGNO, A.M., (1962) - Gli elefanti fossili di Riano (Roma). Geologica Romana 1, pp. 33-132. MANIA D. (1999) - 125 000 Jahre Klima- und Umwel- tentwicklung im mittleren Elbe-Saale-Gebiet. Her- cynia 32, pp. 1-97. MANIA D. (2004) - Neumark Nord - Geologie eines Inter- glazials. Praehistoria Thuringica, 10, pp. 26-42. MANIA D. (2010) - Neumark Nord - Ein interglaziales Ökosystem des mittelpaläolithischen Menschen. Veröffentlichungen des Landesamtes für Denk- malplege und Elephas antiquus Falconer & Caut- ley, 1847 del Pleistocene medio superiore di Neu- mark Nord: implicazioni sistematiche e paleoeco- logiche. Archäologie, 6 (2), 527 pp. MARANO, F. (2009) - Elephas antiquus Falconer & Caut- ley, 1847 del Pleistocene medio superiore di Neuark Nord: implicazioni sistematiche e paleoe- cologiche. Università di Roma, Unpublished thesis. OSBORN H.F. (1942) - Proboscidea: a monograph of the discovery, evolution, migration and extinction of the mastodonts and elephants of the world. Ste- godontoidea, Elephantoidea, 2. New York. PALOMBO M.R., ALBAYRAK E., MARANO F. (2010) - The straight-tusked Elephants from Neumark Nord, a glance to a lost world. Kataloge zur Dauerausstel- lung im Landesmuseum für Vorgeschichte Halle: Elefantenreiche-Eine Fossilwelt in Europa. Lan- desmuseum für Vorgeschichte Halle (Saale), pp. 218-247. PALOMBO M.R., ANZIDEI A.P., ARNOLDUS-HUYZENDVELD A. (2003) - La Polledrara di Cecanibbio: one of the richest Elephas (Palaeoloxodon) antiquus sites of the late Middle Pleistocene in Italy. Deinsea 9, pp. 317-330. PALOMBO M.R, FERRETTI M.P. (2005) - Elephant fossil record from Italy: knowledge, problems, and pers- pectives. Quaternary International, 126-128, pp. 107-136. PALOMBO M.R., FERRETTI M.P. (2010) - What about the taxonomical status of European Straight-tusked elephants? Quaternaire, ors serie, 3 pp. 27-28 PALOMBO, M.R. & GIOVINAZZO, C. (2005) - Elephas fal- coneri from Spinagallo Cave (South-Eastern Sicily, Hyblean Plateau, Siracusa): brain to body weight comparison. Monografies de la Societat d’Història Natural de les Balears, 12, pp. 255-264; Palma de Mallorca. PALOMBO M.R., VILLA P. (2003) - Sexually dimorphic characters of Elephas (Palaeoloxodon) antiquus from Grotte Santo Stefano (Viterbo, Central Italy. Deinsea, 9, pp. 293-315. PENKMAN K. (2010) - Neumark Nord 1: preliminary re- sults of the amino acid analysis. Kataloge zur Dauerausstellung im Landesmuseum für Vorge- schichte Halle: Elefantenreiche-Eine Fossilwelt in Europa. Landesmuseum für Vorgeschichte Halle (Saale), pp. 75-78. ROTH V.L. (1984) - How elephants grow: heterochrony and the calibration of developmental stages in some living and fossil species. Journal of Verte- brate Paleontology, 4, pp. 126-145. ROTH, L. (1990) - Island dwarf elephants: a case study in body mass estimation and ecological inference. In: DAMUTH, J. & MACFADDEN, B. J. (Eds.): Body Size in Mammalian Paleobiology. Estimation and Biological Implications. 151-179; Cambridge (Cambridge University Press). SAEGUSA H., GILBERT W. H. (2008) - Elephantidae. In: W. H. GILBERT, B. ASFAW (eds.), Homo erectus in Africa, Pleistocene Evidence from the Middle Awash, Ethiopia. The Middle Awash Series 1, Chapter 9 (Berkeley and Los Angeles), pp. 195- 228. A pathologic straight-tusked elephant female … 101 SCHULER Y. (2010) - ESR-Datierung von Zahn- schmelzproben der archaologischen Fundhori- zonte von Neumark-Nord. Kataloge zur Daue- rausstellung im Landesmuseum für Vorgeschichte Halle: Elefantenreiche-Eine Fossilwelt in Europa. Landesmuseum für Vorgeschichte Halle (Saale), pp. 64-70. SEIFERT-EULEN M. (2010) - Die Vegetation der Warmzeit aus dem Becken NN1 von Neumark-Nord auf- grund der Pollenanalyse. Kataloge zur Daue- rausstellung im Landesmuseum für Vorgeschichte Halle: Elefantenreiche-Eine Fossilwelt in Europa. Landesmuseum für Vorgeschichte Halle (Saale), pp. 127-139. SHOSHANI, J., FERRETTI, M.P., LISTER, A.M., SAEGUSA, H., AGENBROAD, L.D., MOL. D., TAKAHASHI, K. (2007) - Relationship within the Elephantinae us- ing hyoid characters. Quaternary International 169-170, pp.174-185. TREVISAN, L., (1949) - Lo scheletro di Elephas antiquus italiquus di Fonte Campanile (Viterbo). Paleonto- graphia Italica 44 (1948), pp. 2-78. TSOUKALA, E. LISTER, A. (1998) - Remains of straight- tusked elephant, Elephas (Palaeoloxodon) anti- quus Falc. & Caut., 1847 ESR-dated to oxygen isotope Stage 6 from Grevena (W. Macedonia, Greece). Bollettino della Società Paleontologica Italiana 37 (1), pp. 117-139. Ms. ricevuto il 25/05/2010 Testo definitivo ricevuto il 8/10/2010 Ms. received: May 25, 2010 Final text received: October 8, 2010