10
Pal~iontologische Zeitschrift 74 (1/2) 205-214 5 Abb., 3 Tab. Stuttgart, Mai 2000 On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld near Speyer (Rhineland-Palatinate, Germany) URSULA B. GOHLICH, Mtinchen With 5 figures and 3 tables Kurzfassung: Fin nahezu vollst/indiges Becken eines Ele- phantiden aus den jungpleistoz~inen Ablagerungen der Ober- rhein-Ebene, gefunden in der Gemarkung ,Im Binsfeld' n6rd- lich von Speyer, wird vorgestellt. Die Abhandlung enthalt eine osteologische Beschreibung und vor allem metrische Verglei- che mit zahlreichen Objekten der pleistoz~inen Arten Elephas antiquus (FALCONER • CAUTLEY), Mammuthus meridionalis (NESTI), M. trogontherii (POHLIG) und M. primigenius (BLUMENBACH). Unter Beriicksichtigung von Geschlechts- und Altersbestimmung sowie bio- und lithostratigraphischen Aspekten ergab die Diskussion zur Systematik eine wahr- scheinliche Zugeh6rigkeit zu Elephas (Palaeoloxodon) anti- quus (FALCONER & CAUTLEY). Anhand der Beckenausbildung kann rtickgeschlossen werden, dab es sich um ein adultes und riesiges Weibchen gehandelt hat. Trotz geringer Besch~idigung ist es eines der vollst~indigsten und besterhaltenen Becken des Waldelefanten. Abstract: This paper documents the discovery and study of an almost complete elephantid pelvis, found in Late Pleistocene deposits in the Upper Rhine valley in the administrative dis- trict ,Im Binsfeld' north of Speyer. The paper contains an os- teological description and metrical comparison with numerous specimens of the Pleistocene species Elephas antiquus (FAL- CONER & CAUTLEY), Mammuthus meridionalis (NESTI), M. tro- gontherii (POHLIG) and M. primigenius (BLUMENBACH). The systematic discussion, taking account of sex, age and bio- and lithostratigraphy, shows that the specimen probably belongs to Elephas (Palaeoloxodon) antiquus (FALCONER & CAUTLEY). It is concluded that the pelvis belonged to an adult female of huge size. In spite of an unimportant injury, it is one of the most complete and best preserved pelves of the straight-tusked el- ephant. Introduction The pelvis studied here was discovered in November 1989 by the diving enthusiast MARTIN DANZ in the ,G~insdreck-Weiher' - a former gravel pit - of the recrea- tional resort of Binsfeld near Binshof south of Otterstadt and north of Speyer (Fig. 1). An elephantid skull, which was discovered at the same time, could not be recovered despite a systematic search. On 6.1.1990 the pelvis was Fig. 1. Map showing the site where the pelvis of Elephas antiquus (FALCONER & CAUTLEY) was found. salvaged from clay deposits (testimony of the divers) at a depth of 14.5 m below lake level by the discoverers Mr. DANZ and Mrs. and Mr. GOLTL. The specimen is housed in the Landessammlung fur Naturkun- de Rheinland-Pfalz (Inv. Nr. PW 1998 5064). Elephants constituted one of the most characteristic ele- ments of the mammalian fauna during the Pleistocene. In Europe they were represented by four species (excluding the Mediterranean dwarf elephants): the southern ele- phant (Mammuthus [Archidiskodon] meridionalis [NESTI]), the steppe elephant (Mammuthus trogontherii [POHLIG]), the woolly mammoth (Mammuthus primi- genius [BLUMENBACH]) and the straight-tusked or forest elephant (Elephas [Palaeoloxodon] antiquus [FALCONER & CAUTLEY]). These species belong to two contempora- neous evolutionary lineages. The mammoth lineage first appeared in Eurasia in the Late Pliocene, about 2.5 Ma (LISTER 1996a: 203) with M. meridionalis, a representa- tive of warm-temperate climates. During the early Mid- dle Pleistocene (c. 0.7-0.5 Ma, LISTER 1996a: 209) it was replaced by the more cold-adapted steppe elephant M. Address of the author: Dr. URSULA B. GOHLICH, Institut ftir Pal~iontologie und Historische Geologie, Richard-Wagner-Str. 10, D- 80333 Miinchen, Germany. e-mail: [email protected] 0031-0220/00/0074-0205 $ 2.50 9 2000 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart

On a pelvis of the straight-tusked elephant Elephas …...On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld 207 of RiB-Wtirm-interglacial

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Page 1: On a pelvis of the straight-tusked elephant Elephas …...On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld 207 of RiB-Wtirm-interglacial

Pal~iontologische Zeitschrift 74 (1/2) 205-214 5 Abb., 3 Tab. Stuttgart, Mai 2000

On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld near Speyer (Rhineland-Palatinate, Germany)

URSULA B. GOHLICH, M t i n c h e n

With 5 figures and 3 tables

Kurzfassung: Fin nahezu vollst/indiges Becken eines Ele- phantiden aus den jungpleistoz~inen Ablagerungen der Ober- rhein-Ebene, gefunden in der Gemarkung ,Im Binsfeld' n6rd- lich von Speyer, wird vorgestellt. Die Abhandlung enthalt eine osteologische Beschreibung und vor allem metrische Verglei- che mit zahlreichen Objekten der pleistoz~inen Arten Elephas antiquus (FALCONER • CAUTLEY), Mammuthus meridionalis (NESTI), M. trogontherii (POHLIG) und M. primigenius (BLUMENBACH). Unter Beriicksichtigung von Geschlechts- und Altersbestimmung sowie bio- und lithostratigraphischen Aspekten ergab die Diskussion zur Systematik eine wahr- scheinliche Zugeh6rigkeit zu Elephas (Palaeoloxodon) anti- quus (FALCONER & CAUTLEY). Anhand der Beckenausbildung kann rtickgeschlossen werden, dab es sich um ein adultes und riesiges Weibchen gehandelt hat. Trotz geringer Besch~idigung ist es eines der vollst~indigsten und besterhaltenen Becken des Waldelefanten.

Abstract: This paper documents the discovery and study of an almost complete elephantid pelvis, found in Late Pleistocene deposits in the Upper Rhine valley in the administrative dis- trict ,Im Binsfeld' north of Speyer. The paper contains an os- teological description and metrical comparison with numerous specimens of the Pleistocene species Elephas antiquus (FAL- CONER & CAUTLEY), Mammuthus meridionalis (NESTI), M. tro- gontherii (POHLIG) and M. primigenius (BLUMENBACH). The systematic discussion, taking account of sex, age and bio- and lithostratigraphy, shows that the specimen probably belongs to Elephas (Palaeoloxodon) antiquus (FALCONER & CAUTLEY). It is concluded that the pelvis belonged to an adult female of huge size. In spite of an unimportant injury, it is one of the most complete and best preserved pelves of the straight-tusked el- ephant.

Introduction The pelvis studied here was discovered in November 1989 by the diving enthusiast MARTIN DANZ in the ,G~insdreck-Weiher' - a former gravel pit - of the recrea- tional resort of Binsfeld near Binshof south of Otterstadt and north o f Speyer (Fig. 1). An elephantid skull, which was discovered at the same time, could not be recovered despite a systematic search. On 6.1.1990 the pelvis was

Fig. 1. Map showing the site where the pelvis of Elephas antiquus (FALCONER & CAUTLEY) was found.

salvaged f rom clay deposits (testimony of the divers) at a depth o f 14.5 m below lake level by the discoverers Mr. DANZ and Mrs. and Mr. GOLTL.

The specimen is housed in the Landessammlung fur Naturkun- de Rheinland-Pfalz (Inv. Nr. PW 1998 5064).

Elephants constituted one of the most characteristic ele- ments o f the mammalian fauna during the Pleistocene. In Europe they were represented by four species (excluding the Mediterranean dwarf elephants): the southern ele- phant (Mammuthus [Archidiskodon] meridionalis [NESTI]), the steppe elephant (Mammuthus trogontherii [POHLIG]), the wool ly mammoth (Mammuthus primi- genius [BLUMENBACH]) and the straight-tusked or forest elephant (Elephas [Palaeoloxodon] antiquus [FALCONER & CAUTLEY]). These species belong to two contempora- neous evolutionary lineages. The mammoth lineage first appeared in Eurasia in the Late Pliocene, about 2.5 Ma (LISTER 1996a: 203) with M. meridionalis, a representa- tive of warm-temperate climates. During the early Mid- dle Pleistocene (c. 0.7-0.5 Ma, LISTER 1996a: 209) it was replaced by the more cold-adapted steppe elephant M.

Address of the author: Dr. URSULA B. GOHLICH, Institut ftir Pal~iontologie und Historische Geologie, Richard-Wagner-Str. 10, D- 80333 Miinchen, Germany. e-mail: [email protected]

0031-0220/00/0074-0205 $ 2.50 �9 2000 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart

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206 URSULA B. GI3HLICH

trogontherii - transitional to M. primigenius, the late Mid- dle (0.4-0.3 Ma, LISTER 1996a: 210) to Late Pleistocene glacial species. Contemporaneous with the late M. meri- dionalis and the following Mammuthus chronospecies is the only representative of the second lineage, the straight- tusked or forest elephant E. antiquus. It inhabited Europe during warm-temperate times from the Early to the Late Pleistocene (DUBROVO 1977: 1085; YON KOENIGSWALD 1988: 230), having immigrated through central Europe during the early Middle Pleistocene Cromerian Complex (VON KOENIGSWALD 1994: 191). The different ecological adaptions of the forest elephant and the steppe elephants (M. trogontherii, M. primigenius) lead in general to al- ternating occurrence of these elephant groups in Middle to Late Pleistocene deposits (although in some early Mid- dle Pleistocene deposits [e.g. middle stage of the Mos- bach Sands] M. trogontherii and E. antiquus apparently occur together [ADAM 1961: 5f] --probably in different habitats). In the late Middle to Late Pleistocene the alter- nation of the lineages becomes notably marked during the climatic changes of those times. In the glacials the forest elephant withdrew to the south of Europe while the area north of the Alps was dominated by mammoth. The most northern occurrences ofE. antiquus are known from Eng- land, Denmark and Poland at about 54-55~ During the interglacial-glacial transitions mammoth probably coex- isted with E. antiquus (YON KOENIGSWALD 1988: 217, 234).

Contrary to the mammoth-lineage, which is distin- guished by progressive adaptation (and therefore by mor- phological skeletal and dental changes) to colder climates during the Pleistocene, the straight-tusked elephant re- mains 'conservative' and barely shows any morphologi- cal changes in the course of the following 500.000 years. Therefore its remains are not suitable for interpreting stratigraphic succession in the Pleistocene deposits of Europe, but they can be used as an ecological indicator.

The European straight-tusked elephant is termed here Elephas antiquus (FALCONER & CAUTLEY 1847) although its affiliation to Elephas or to Palaeoloxodon is as yet undecided (see e.g. DUBROvo 1977; MAGLIO 1973; YON KOENIGSWALD 1988: 2290. In the opinion of some au- thors E. antiquus and the South and East Asian E. namadicus (FALCONER & CAUTLEY 1845) are synony- mous (e.g. MAGLIO 1973).

Abbreviations

AMNH American Museum of Natural History, New York, USA

BMNH The Natural History Museum, London, United Kingdom

GPIUM Geologisch-Pal~iontologisches Institut der Universit~it Mtinster, Germany

LfA Landesamt fiir Arch~iologie Sachsen-Anhalt, Halle

LfD Landesamt ftir Denkmalpflege Rheinland- Pfalz, Ref. Erdgeschichtliche Denkmalpflege, Mainz, Germany

LfN Landesammlung fiir Naturkunde Rheinland- Pfalz, Mainz, Germany

LMVH Landesmuseum fiir Vorgeschichte in Halle, Germany

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain

MZP Muzeum Ziemi Polska Akademia Nauk, Warzawa, Poland

NKM Novosibirsk Museum of Regional Ethnology, Novosibirsk, Russia

NMMS Naturkunde- und Mammut-Museum Siegsdorf, Germany

PIN Palaeontological Institute of the Russian Academy of Sciences, Moscow, Russia

SMNS Staatliches Museum flit Naturkunde Stuttgart, Germany

SMS Spengler-Museum Sangerhausen, Germany UA Department of Historical Geology and

Paleontology of the University of Athens, Greece

ZM Zoological Museum of the Russian Academy of Sciences, St. Petersburg, Russia

CH = Switzerland, E = Spain, F -- France, D = Germany, GB = Great Britain, GR = Greece, I = Italy, NL = The Netherlands, PL = Poland, RUS = Russia, USA = United States of America

Litho- and biostratigraphic context In the southern part of the northern Oberrhein-Ebene a cyclic aggradation of three thick gravel- and coarse sand layers, deposited during several glacials from the Elster (Minded to the end of the Weichsel (Wtirrn) glacial are interlayered with two clay-silt beds, which are supposed to have been deposited during the Holsteinian and the Eemian (= Ril3/Wtirm) interglacials respectively (ScHwElSS 1988: 22).

The outcrop in the region south of Otterstadt is a Late Pleistocene sediment (SCHWE~SS 1988:22). It is separated by the Eemian interglacial bed, the so called 'Oberer Ton', from underlying Early to Middle Pleistocene sedi- ments. The 'Oberer Ton' often underlies the horizons used for commercial gravel exploitation.

According to SCHARPF (1977: 41) the 'Oberer Ton' is mostly found at a depth of 20-30 m. In the southern part of north Oberrhein-Ebene LOSCHER (1988: 85) could verify that the ,Oberer Ton' west of the Rhine is only 9- 10 m below ground-water level while to the east of the Rhine it is 16-20 m below the ground-water level.

The hydrogeological working group 'Arbeitsgruppe 1980' concluded that the sediments directly above the 'Oberer Ton' are of glacial origin whereas, according to palaeontological data, the basal 3-5 m of the overlying layer is supposed to be interglacial (LOSCHER 1988: 85).

According to records and specimens in the Staatliches Museum ftir Naturkunde in Stuttgart there is proof of M. primigenius and E. antiquus from Binsfeld (pers. comm. Dr. R. ZIEGLER 1998). These taxa are represented at the sites Binsfeld SW and Binsfeld SE, whereas at Binsfeld N only M. primigenius is known. At Binsfeld SW the 'Oberer Ton' is found at a depth of 15 m under ground level. Accordingly the clayey bed, where the specimen described was discovered at a depth of 14.5 m under the ground-water level, presumably corresponds to the so- called ,Oberer Ton'. These sediments are supposed to be

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On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld 207

of RiB-Wtirm-interglacial age. This would fit well with the statement of the divers (pers. comm. Mr. DANZ to Dr. WUTTKE 1998) that they also found fossil tree trunks with diameters of about 2 m, which - according to LOSCHER (1988: 81, fig. D2) - supports an interglacial age for the deposit.

The pelvis from Binsfeld - osteological description

The pelvic girdle (Fig.2) is almost completely preserved. Only the tuber coxae of the right ilium wing and the cau- dal ends of both ischia (tubera ischiadica) are lacking. For measurements see Fig.3 and Tab. 1.

The 3 bones of the innominate are totally fused. The pelvic symphysis (symphysis pelvina) is ossified, but between the fused innominates (ossa coxae) there is a symphyseal fissure.

The laterally extending ilium wing (ala ossis ilii) is slightly concave ventrally. Ventrally in the medial half of the wing there is a longish concavity oriented cranio- medially to caudolaterally. The crista iliaca - craniolate- rally-convex - is thickened, rough and partly dorsally upturned. The lateral tuber coxae is also thickened, with a dorsal swelling. The craniomedial sacral tuber (tuber sacrale) is upturned dorsally. It is thickened cranially and thinner and sharper caudally. On the medial side of the upturned sacral tuber a tuberositas iliaca is manifest. Starting at the caudal end of the sacral tuber, the inner margin of the pelvic aperture is sharp-edged with a low crista ending at the corpus ossis ilii. A weak and short linea glutea can be observed dorsally in the medial and caudal half especially of the right wing.

The corpus ossis ilii is flattened dorsoventrally. Crani- ally to the hip socket (acetabulum), the lateral surface of the corpus is rugose.

The ventrolaterally orientated, hemispherical acetabu- lure is nearly circular. Mediocaudally it is notched into a narrow (c. 1.5 cm) incisura acetabuli which opens to a longish, relatively slim (c. 3.5 cm) and short (c. 10 cm) acetabular fossa, not reaching the centre of the depres- sion. The fossa is subdivided by three low ridges arising in the middle of the fossa and extending cranially, dorsally and ventrally.

The pubic symphysis is fused and ventrally thickened to form a strong ventral pubic tubercle (tuberculum pubicum ventrale). The cranial crest (pecten ossis pubis) is sharp-edged. Also the eminentiae iliopubicae on both sides are well-marked. The transverse cranial branch of the pubis (ramus cranialis ossis pubis) has an oval cross- section and is dorsoventrally compressed. Ventrally, on both sides of the pubic tubercle and extending a few cm medial of the acetabulum, there is a rounded to trans- verse-oval (c. 5x4 cm) protuberance.

The obturate foramen (foramen obturatum) is of long- ish-oval shape. Its caudal end is separated by a constric- tion consisting of two opposing projections, a pointed

Tab. 1. Measurements (mm) of the pelvis of Elephas antiquus (FALCONER & CAUTLEY) from Binsfeld, ( ) estimated measure of incomplete bone, [ ] real measure of fragmentary bone.

site Binsfeld (D) museum LfN Col.Nr. PW 1998 5064 ~ender female measurements

sin. dext. 1 2 3 4 5 6 7 8 9

10 11 12 13 14 15 16

(1580) 600 810 510

[1060] [450]

(600) 600 590

[970] 1050 207 207 545 540 290 273 248 248 23 0 240 185 185

380

spur laterally and, opposite to it, a little projection devel- oped along the caudal half of the medial inner margin of the foramen. Ventrally on the corpus ossis pubis a sulcus arises from the caudal end of the obturate foramen and flattens caudally.

Caudally the ischia contact each other at an angle of 90-100 ~ The caudal end (tuber ischiadica and caudal parts of tabula ossis ischii) of each side is lacking. The medial branches of the ischia (ramus ossis ischii) are ex- tremely flattened dorsoventrally. Between them there is a symphyseal fissure of c. 10 cm length and 1.0-1.3 cm width - about level with the distal half of the obturate foramen. The caudal part of the corpus ossis ischii is of an oval shape - dorsomedially-ventrolaterally flattened. Dorsally on its cranial part there is a well-marked, sharp- edged spine (spina ischiadica) reaching from the corpus of the ilium to the corpus of the ischium and ending about half way along the length of the obturate foramen. Dorsolaterally on the corpus ossis ischii, between the ischiadical spine and the caudal half of the acetabulum, there passes a sulcus of c. 7 cm length, aligned parallel to the acetabular fossa. Within the sulcus some longish crests can be observed.

After the study and drawing of the specimen the miss- ing part of the crista iliaca of the right ilium wing was restored and completed.

Discussion

Species distinction and determination of Pleistocene el- ephants usually relies on dental and cranial characters; diagnostic features on the morphology of postcranial

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208 URSULA B. GOHLICH

Fig. 2a, b. Elephas antiquus (FALCONER & CAUTLEY), pelvis from Binsfeld. - a: cranial view, b: caudodorsal view.

material is much more difficult (see e.g. DUBROVO & JAKUBOWSKI 1988). Due to the rarity of pelvic remains little work has been carried out on species identification, so that species comparison has not been possible hith- erto, nor studies of inter- and intraspecific variation in pelvic shape.

For metrical comparisons with pelves of M. meridionatis, M. trogontherii and M. primigenius meas- urements in Tab. 3 were taken from the study of LISTER (1996b). Additional pelvic measurements of E. antiquus (sexed by cited authors in Tabs. 2 and 3) come from my own studies and several publications (Tabs. 2, 3).

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On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld 209

Fig. 2c, d. Elephas antiquus (FALCONER • CAUTLEY), pelvis from Binsfeld. - c: cranioventral view, d: laterocranioventral view.

For metrical comparisons on remains of proboscideans, knowledge of sex and approximate age is neccessary. This is particularly important in elephants as they grow throughout nearly their whole lives. The older the ani- mal, the bigger it and its skeletal remains are. Addition- ally there is sexual dimorphism; males are larger than females. These aspects have to be considered when com- paring skeletal measurements of specimens or taxa.

Gender determination

After skeletal size and robusticity and morphology of the skull and tusks (see e.g. AVERIANOV 1996; KROLL 1991: 43ff), the pelvic morphology of elephantids provides in-

formation for the determination of the sex, as DERANIYAGALA (1955) and KROLL (1991) have shown for the Asian elephant (Elephas maximus), HAYNES (1990) and KROLL (1991) for the African elephant (Loxodonta africana) and LISTER & AGENBROAD (1994) and LISTER (1996b) for the mammoths. Gender determination is an important contribution to understanding aspects like the taphonomy of sites, interpretation of size, or social struc- ture.

Elephants are uniparous mammals, normally bearing only a single young, which is relatively large at the time of birth. The mother's pelvis - hormonally influenced during the pregnancy - changes in that bone is resorbed along the pelvic aperture to widen the birth canal (KROLL

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210 URSULA B. GOHLICH

Tab. 2. Comparison of the measurements (mm) of pelves of E. antiquus (FALCONER ~z CAUTLEY). ( ) estimated measure of incomplete bone, [ ] real measure of fragmentary bone.

si te Crums tad t (D) G r 6 b e m G r 6 b e m Brfihl (D) Brfihl (D) Brfihl (D) Bue lna (E) II (D) I (D) Kiesficker 72 Rheingewann Schlangen Asturias

winkel museum HLMD LfA LfA SMNS SMNS SMNS MNCN? Col.Nr. RS 3014 RS 3015 HK 95: 6517.5.7.72.27 6616.2.12.88.3 6616.17.11.80.19 ?

4213 +20 gender female (juvenile) female male male ? ? male? measurements * KROLL 1991, Tab.65, 66 and ~ SMNS SMNS MAZO 1998: from 274

sin. dext. dext. dext. dext. sin. sin. dext. sin. dext. ? l (1260) (1610) 2 *465 "515 3 *660 *890 4 *360 530 5 *790 *780 (1260) - - 6 *280 (640) - (420) - 7 (439) *560/ *660 *690/ 520 - -

440 535 8 470 (625) (640) - - 9 *755 *1100 "910 - -

10 150 153 "195 *280 260 *265 - 247 251 305 11 395 390 ~ *655 - 620 613 12 205 "215 *250 *340 332 271 270 13 190 " 1 9 0 *260 *320 284 282 260 254 14 " 1 7 2 " 1 7 0 *255 "215 227 222 199 197 15 " 1 7 0 "167 *205 "210 - 206 210 160-170 1 6 (310) (340) (420) - -

site U p n o r M e g a l o p o - M e g a l o p o - Mega lopo- Mega lopo- V i t e r b o ( I ) Warsaw Chatelard (GB) lis ( G R ) lis ( G R ) lis (GR) lis (GR) (PL) (F)

museum BMNH UA1960/143 UA 1960/87 UA 1960/89 UA 1960/90 Pisa MZP VIII/ (destroyed) Vm 248-310

gender ? male ? ? ? ? ? ? measure- ANDREWS & MELENTIS 1963, Tab. 16, 17 TREVlSAN JAKUBOVSrd BEDEN 1969 ments from COOPER 1928 1947,from a et al. 1968

drawing sin.+dext, dext. sin. dext. sin. dext. dext. ?

1833 ? - - 1 2 3 945 4 5 6 (570) 7 710 8 9 1100 ? 1022

10 11 611 12 304 13 290 14 15 250 210 16

[460]

588 28O 228

208

[505] [783] (1080) 1000 ? (280) 244

480 486 251 258 210

190 180 270

200 210

1991: 45, 47). Wi th succe s s ive p r e g n a n c i e s the inner

marg in o f the p e l v i c aper ture b e c o m e s m o r e accen tua ted ,

the corpus ossis ilii b e c o m e s th inner and the e m i n e n t i a e

i l i o p u b i c a e b e c o m e m o r e e n h a n c e d . KROLL'S s tudies

(1991 : 45f) c o n f i r m these gende r - spec i f i c m o r p h o l o g i c a l

and also me t r i ca l p e l v i c cha rac te r s in both r ecen t e l -

ephants and E. antiquus. A c c o r d i n g to LISTER & AGENBROAD (1994) and LISTER

(1996b), the ra t io o f e i the r the wid th (here m e a s u r e m e n t

2) or the d i a g o n a l h e i g h t (here m e a s u r e m e n t 7) o f the

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On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld 211

Tab. 3. Comparison of some measurements (cm) and indices of pelves of E. antiquus FALCONER & CAUTLEY, M. meridionalis (NESTI), M. trogontherii (POHLIG) and M. primigenius (BLUMENBACH) (measures and indices of M. meridionalis, M. trogontherii and M. primigenius from LISTER 1996b).

site Museum gen- measurements indices of measurements Col. Nr. der 1 7 2 8 10 10/1 2/1 2/7 2/8 2/10 7/10

E. antiquus Binsfeld(D) LtN f (158) 60 60 59 20.7 -0.131 -0 .380 1.0 1.017 2.90 2.90 Crumstadt (D) HLMD RS3014-5 f (126) 56 46.5 47 15.2 --0.121 --0.369 0.83 0.989 3.06 3.68 Gr6bem II (D) LfA f 19.5 - Gr6bern I (D) LfA HK 95:4213 m 66 28.0 2.36 Kies/icker 72 (D) SMNS m (161) 53 51.5 (63) 26.5 -0.165 -0.320 0.97 0.82 1.94 2.0 Buelna (E) MMEE m? 30.5 - Upnor (GB) ? ? 183 Megalopolis UA 1960/143 ? 71 (GR) Viterbo (I) Pisa (destr.) ? (28) Warsaw (PL) MZP Vllt/Vm ? 24.4 M. meridionalis Valdarno (I) 1GF, 14838 m 160 40 44 27.0 0.169 0.275 1.10 1.63 1.48 Valdarno(1) IGF, 10791 m 188 47 54 28.4 0.151 0.287 1.15 1.90 1.65 Valdarno(1) 1GF, 1057 m? 178 47 49 25.1 0.141 0.275 1.04 - 1.95 1.87 Valdarno (1) 1GF, 1050 f 130 45 47 17.0 0.131 0.362 1.04 - 2,76 2.65 Valdarno(I) 1GF, 13730 f 138 45 44 15.6 0.113 0.319 0.98 2.82 2.88 M. trogontherii Edersleben (D) SMS f 154 63 57 63 19.2 0.125 0.370 0.90 0.904 2.97 3.28 M. primigenius Praz Rodet (CH) Lausanne m 42 20.0 2.10 Steinheim (D) SMNS m 48 5 1 25.0 1.06 - 2.04 1.92 Siegsdorf(D) NMMS m 160 44 50 26.0 0.163 0.313 1.14 1.92 1.69 Ahlen (D) GP1UM m 137 40 50 62 0.365 1.25 0.806 Condover (GB) Shropshire m 143 43 46 -49 20.2 0.141 0.322 1.07 0.939 2.28 2.13 Gewande(NL) Hoofddorp m? (119) 42 42 - 46 19.0 -0.160 -0.353 1.00 -0.913 2.21 2.21 Beresovka(RUS) ZM, N5315 m 134 33 41 52 18.3 0.137 0.306 1.24 0.788 2.24 1.8 Taimyr(RUS) ZM, N2710 m 125 41 43 47 19.4 0.155 0.344 1.05 0.915 2.22 2.11 LiakhovIs.(RUS) MNHN m (112) 33 43 -47 20.8 - 0 . 1 8 6 - 0 . 3 8 4 1.30 -0.915 2.07 1.59 Gydan (RUS) P1N m 130 46 46 19.3 0.148 0.354 1.000 2.38 LenaR.(RUS) ZM, 71911 m 137 44 48 55 18.2 0.133 0.350 1.09 0.872 2.64 2.42 Aa(F) Boulogne f (134) 54 - 40 19.4 -0.145 0.403 -1.350 2.78 Oyosh(RUS) NKM f 117 39 44 -45.2 -15.4 0.132 0.377 1.13 0.973 2.86 2.53 Rochester (USA) AMNH f 46 50 19.0 1.09 2.63 2.42 Kerkdriel (NL) Empel f?. 128 43 45 42 16.0 0.125 0.352 1.05 1.07 2.81 2.69

pelvic aperture, to the m i n i m u m width of the corpus ossis ilii (here measurement 10), allows a distinction between the sexes to be made. Both the morphology of the pelvis, especially around the pelvic aperture - with crests along the inner margin, and the accentuated eminent iae il iopubicae - and the metrical proportions of the pelvis (Fig. 4), indicate that the Binsfeld specimen belonged to a female.

Size and an at tempt at age determination

The metrical study shows that our specimen is relatively large, especially considering that it is a female. In com- parison with the female pelves of the compared species and specimens it seems to be one of the biggest and even surpasses several males (Fig. 5).

But as ment ioned above, the individual age should also be considered because of the continuous growth observed in elephants. Age determinat ion can be undertaken accu- rately with the help of the dentit ion, but this, unfortu- nately, is not preserved in this case.

An age determinat ion by means of the pelvis is hardly practicable. Studies on recent African elephants (HAYNES

1991:351, tab. A15) showed that the three bones of the innominate fuse at the age of 8 to 12 years. The fusion of the sacrum to the innomina te seem to vary from 18-26 years in females and to take place later than the age of 50 years in males ofLoxodonta africana. The age of fusion of elephantid innominates in the pelvic symphysis is not studied as yet. Personal observat ions of LISTER suggest that in male mammoth the two halves may fuse between 30 and 40 years (pers. comm. Dr. A. LISTER 1998). Fe-

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212 URSULA B. GOHLICH

male specimens are more rare, but there is a fused pelvis of a female M. primigenius f rom Oyesh, Sibir ia whose age is est imated by AVERIANOV (1996: 263) to be 35-40 years.

We have to take into account that all these observations are made in different taxa and often are based only on a few specimens. The combina t ion o f the states of fusion at the pelvis from Binsfe ld ( fusion of three bones of the innominate, fusion o f pe lv ic symphys i s , no fusion to the sacrum) does not fit with these observat ions. Accordingly an age determinat ion seems not to be poss ible by means of present data. But the fus ion o f the three bones of the innominate and espec ia l ly the fus ion of the left and right halves strongly suggest , that the an imal was adult.

Fig. 3. Elephas antiquus (FALCONER & CAUTLEY), pelvis mea- surements (according to YON DEN DRIESCH 1976, KROLL 1991, LISTER 1996b). (1) maximum horizontal width of pelvic girdle; (2) maximum horizontal width of pelvic aperture; (3) maximum horizontal width between the outer margins of the acetabula; (4) width between eminentiae iliopubicae; (5) maximum length of pelvic girdle; (6) length of the symphysis; (7) diagonal height of pelvic aperture, taken from the pubic symphysis to lowest point of the sacral attachement; (8) width of ilium wing from tuber coxae to nearest point of pelvic aperture; (9) direct maximum length of ilium, taken from tubercoxae to tuber sacrale; (10) minimum width of ilium shaft (corpus ossis ilii), taken parallel to the plane of the shaft; (11) minimum perimeter of ilium shaft (corpus ossis ilii); (12) minimum perimeter of the pubis shaft (corpus ossis pubis); (13) minimum perimeter of the ischium shaft (corpus ossis ischii); (14) maximum inner length of foramen obturatum; (15) maximum length of acetabulum; (16) horizontal distance between outer margin of acetabulum and tuber coxae.

Results

Morphological and metr ica l s tudies alone would not have al lowed a systematic de te rmina t ion closer than the speci- men is an elephantid. The size of the specimen (Fig. 5) indicates that it is too big to be M. primigenius. There is only one pelvis of M. primigenius (Siegsdorf) which reaches the d imensions o f the one s tudied here, and that is a male; all others of whatever sex are dist inctly smaller. The size of the spec imen in this s tudy is very s imilar to that of the only de t e rmined female specimen of M. trogontherii. Male spec imens o f M. meridionalis and E. antiquus compared in this pape r are bigger, whereas fe- males are smaller. But it has to be remembered that the only female E. antiquus noted here (Crumstadt) , is ap- parent ly from a younger ind iv idua l with an unfused pu- bic symphysis .

The litho- and b ios t ra t ig raph ic si tuation at Binsfeld suggests a Late P le is tocene age and the remarkably good preservat ion of the spec imen supports a more or less au- tochthonous deposi t ion. M. meridionalis and M. trogon- therii can be exc luded in all probabi l i ty , being Early and Middle Pleis tocene species. This is conf i rmed by the oc- currence of dental remains of only M. primigenius and E. antiquus at the site o f Bins fe ld (pers. comm. Dr. R. ZmGLER 1998).

Fig. 4. Comparison of some gender dependent measurement ratios (see Tab. 3) of elephantids (symbols see Fig. 5).

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On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld 213

Fig. 5. Comparison of the size of elephantid pelves (by means of measurement 1: maximum horizontal width of the pelvis).

By the means of the pelvic morphology and dimensions the specimen can best be determined as a female of E.

antiquus. Although the pelvic sample of E. antiquus is very

small, the specimens available appear to follow the same metrical pattern as LISTER (1996b) pointed out for Mammuthus, suggesting that the gender distinction may be valid for straight-tusked elephants, too.

For confirming this and for considering the intraspe- cific variation of pelvic morphology and size of E. anti- quus in comparison with the representatives of the Mammuthus-lineage more material is needed.

Acknowledgements I wish to thank Dr. M. WUTTKE and the Landesamt ftir Denk- malpflege Rheinland-Pfalz, Referat Erdgeschichtliche Denk- malpflege (Mainz), for placing the specimen at my disposal and for their financial support, Dr. A. LmTER (University Col- lege London) for some helpful information, Mr. R DAVIES (University College London) and Dr. A. GENTRY (London) for critically reading the manuscript and improving the English, Prof. Dr. W. YON KOENIGSWALD (Univ.-Institut ftir Pal~iontolo- gie Bonn) for regional indications and Dr. H.-J. DOHLE (LfA) for further measures on the pelves of Gr6bern. Special thanks are due to Dr. R. ZIEGLER (SMNS) for decisive litho- and bio- stratigraphic information and for supplying further measures of some pelvic specimens from Briihl.

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Eingang des Manuskriptes am 12. Januar 1999; Annahme durch die Schriftleitung am 5. Januar 2000.