On a pelvis of the straight-tusked elephant Elephas antiquus

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 Elephantiden aus den jungpleistoz~inen Ablagerungen der Oberrhein-Ebene, gefunden in der Gemarkung ,Im Binsfeld' n6rdlich von Speyer, wird vorgestellt. Die Abhandlung enthalt eine
osteologische Beschreibung und vor allem metrische Vergleiche 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 wahrscheinliche Zugeh6rigkeit zu Elephas (Palaeoloxodon) antiquus (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 district ,Im Binsfeld' north of Speyer. The paper contains an osteological description and metrical comparison with numerous
specimens of the Pleistocene species Elephas antiquus (FALCONER& CAUTLEY), Mammuthus meridionalis (NESTI), M. trogontherii (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 elephant.
Introduction
The pelvis studied here was discovered in N o v e m b e r
1989 by the diving enthusiast MARTIN DANZ in the
,G~insdreck-Weiher' - a former gravel pit - of the recreational 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 from 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 Naturkunde Rheinland-Pfalz (Inv. Nr. PW 1998 5064).
Elephants constituted one of the most characteristic elements o f the mammalian fauna during the Pleistocene. In
Europe they were represented by four species (excluding
the Mediterranean dwarf elephants): the southern elemeridionalis
phant (Mammuthus [Archidiskodon]
[NESTI]), the steppe elephant (Mammuthus trogontherii
[POHLIG]), the woolly m a m m o t h (Mammuthus primigenius [BLUMENBACH]) and the straight-tusked or forest
elephant (Elephas [Palaeoloxodon] antiquus [FALCONER
& CAUTLEY]). These species belong to two contemporaneous evolutionary lineages. The m a m m o t h lineage first
appeared in Eurasia in the Late Pliocene, about 2.5 Ma
(LISTER 1996a: 203) with M. meridionalis, a representative of warm-temperate climates. During the early Middle 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. URSULAB. GOHLICH,Institut ftir Pal~iontologie und Historische Geologie, Richard-Wagner-Str. 10, D80333 Miinchen, Germany. e-mail: u.goehlich@lrz.uni-muenchen.de
0031-0220/00/0074-0205
$ 2.50
9 2000 E. Schweizerbart'scheVerlagsbuchhandlung,D-70176 Stuttgart
206
URSULAB. GI3HLICH
trogontherii - transitional to M. primigenius, the late Middle (0.4-0.3 Ma, LISTER 1996a: 210) to Late Pleistocene
glacial species. Contemporaneous with the late M. meridionalis and the following Mammuthus chronospecies is
the only representative of the second lineage, the straighttusked 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 KOENIGSWALD1994: 191). The different ecological
adaptions of the forest elephant and the steppe elephants
(M. trogontherii, M. primigenius) lead in general to alternating occurrence of these elephant groups in Middle
to Late Pleistocene deposits (although in some early Middle Pleistocene deposits [e.g. middle stage of the Mosbach Sands] M. trogontherii and E. antiquus apparently
occur together [ADAM 1961: 5f] --probably in different
habitats). In the late Middle to Late Pleistocene the alternation 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 England, Denmark and Poland at about 54-55~ During the
interglacial-glacial transitions mammoth probably coexisted with E. antiquus (YON KOENIGSWALD1988: 217,
234).
Contrary to the mammoth-lineage, which is distinguished by progressive adaptation (and therefore by morphological skeletal and dental changes) to colder climates
during the Pleistocene, the straight-tusked elephant remains 'conservative' and barely shows any morphological 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 authors E. antiquus and the South and East Asian E.
namadicus (FALCONER & CAUTLEY 1845) are synonymous (e.g. MAGLIO 1973).
Abbreviations
AMNH
BMNH
GPIUM
LfA
LfD
LfN
American Museum of Natural History, New
York, USA
The Natural History Museum, London,
United Kingdom
Geologisch-Pal~iontologisches Institut der
Universit~it Mtinster, Germany
Landesamt fiir Arch~iologie Sachsen-Anhalt,
Halle
Landesamt ftir Denkmalpflege RheinlandPfalz, Ref. Erdgeschichtliche Denkmalpflege,
Mainz, Germany
Landesammlung fiir Naturkunde RheinlandPfalz, 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 sediments. 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 910 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 socalled ,Oberer Ton'. These sediments are supposed to be
On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld
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 caudal 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 craniomedially to caudolaterally. The crista iliaca - craniolaterally-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. Cranially to the hip socket (acetabulum), the lateral surface of
the corpus is rugose.
The ventrolaterally orientated, hemispherical acetabulure 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 depression. 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 crosssection 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 transverse-oval (c. 5x4 cm) protuberance.
The obturate foramen (foramen obturatum) is of longish-oval shape. Its caudal end is separated by a constriction consisting of two opposing projections, a pointed
207
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
museum
Col.Nr.
~ender
measurements
Binsfeld (D)
LfN
PW 1998 5064
female
sin.
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
dext.
(1580)
600
810
510
[1060]
[450]
1
(600)
[970]
207
545
290
248
23 0
185
600
590
1050
207
540
273
248
240
185
380
spur laterally and, opposite to it, a little projection developed 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 extremely 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, sharpedged 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 missing part of the crista iliaca of the right ilium wing was
restored and completed.
Discussion
Species distinction and determination of Pleistocene elephants usually relies on dental and cranial characters;
diagnostic features on the morphology of postcranial
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 hitherto, nor studies of inter- and intraspecific variation in
pelvic shape.
For
metrical
comparisons
with
pelves
of
M.
meridionatis, M. trogontherii and M. primigenius measurements 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).
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 animal, the bigger it and its skeletal remains are. Additionally there is sexual dimorphism; males are larger than
females. These aspects have to be considered when comparing skeletal measurements of specimens or taxa.
Gender determination
After skeletal size and robusticity and morphology of the
skull and tusks (see e.g. AVERIANOV1996; 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 structure.
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
210
URSULAB. 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.
site
museum
Col.Nr.
gender
measurements
from
C r u m s t a d t (D)
Gr6bem
II (D)
HLMD
RS 3014
RS 3015
LfA
Brfihl (D)
Kiesficker 72
Brfihl (D)
Rheingewann
LfA
SMNS
HK 95:
6517.5.7.72.27
4213
female (juvenile)
female
male
male
* KROLL 1991, Tab.65, 66 and ~
SMNS
6616.2.12.88.3
sin.
l
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
site
museum
gender
measurements from
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
dext.
(1260)
*465
*660
*360
*790
*780
*280
(439)
*560/
440
470
*755
150
153
395
390
205
"215
190
"190
"172
"170
"170
"167
(310)
(340)
Upnor
(GB)
BMNH
Megalopolis ( G R )
UA1960/143
?
ANDREWS
&
COOPER1928
male
sin.+dext,
1833 ?
dext.
dext.
Gr6bem
I (D)
dext.
*1100
*280
~
"195
*250
*260
*255
*205
Megalopolis ( G R )
UA 1960/87
?
SMNS
dext.
sin.
(1610)
"515
*890
530
(1260)
(640)
*690/
520
535
(625) (640)
"910
260
*265
*655
*340
*320
"215
"210
(420)
*660
Megalopolis ( G R )
UA 1960/89
?
?
MELENTIS 1963, Tab. 16, 17
Brfihl (D)
B u e l n a (E)
Schlangen
Asturias
winkel
SMNS
MNCN?
6616.17.11.80.19
?
+20
?
male?
SMNS
MAZO 1998:
274
sin.
dext.
?
sin.
dext.
-
284
227
Megalopolis ( G R )
UA 1960/90
?
sin.
-
dext.
sin.
-
1022
[460]
[505]
[783]
611
304
290
588
28O
228
480
251
210
486
258
210
208
190
180
332
282
222
-
(420)
247
620
271
260
199
206
Viterbo(I)
Pisa
(destroyed)
?
TREVlSAN
1947,from a
drawing
dext.
251
613
270
254
197
210
-
305
160-170
Warsaw
Chatelard
(PL)
(F)
MZP VIII/
Vm 248-310
?
?
JAKUBOVSrd BEDEN1969
et al. 1968
dext.
?
945
(570)
710
1100 ?
250
1991: 45, 47). W i t h s u c c e s s i v e p r e g n a n c i e s the i n n e r
m a r g i n o f the p e l v i c a p e r t u r e b e c o m e s m o r e a c c e n t u a t e d ,
the c o r p u s ossis ilii b e c o m e s t h i n n e r 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 studies
(1991 : 4 5 f ) c o n f i r m these g e n d e r - s p e c i f i c m o r p h o l o g i c a l
(1080)
(280)
1000 ?
244
200
270
210
and also m e t r i c a l p e l v i c c h a r a c t e r s in b o t h r e c e n t elephants and E. antiquus.
A c c o r d i n g to LISTER & AGENBROAD (1994) and LISTER
(1996b), the ratio o f e i t h e r the w i d t h (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 ( h e r e m e a s u r e m e n t 7) o f the
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
Col. Nr.
gender
LtN
HLMD RS3014-5
LfA
LfA HK 95:4213
SMNS
MMEE
?
UA 1960/143
f (158)
f (126)
f
m
m (161)
m?
?
183
?
1
measurements
7
2
8
10
10/1
indices o f measurements
2/1
2/7
2/8
2/10 7/10
E. antiquus
Binsfeld(D)
Crumstadt (D)
Gr6bem II (D)
Gr6bern I (D)
Kies/icker 72 (D)
Buelna (E)
Upnor (GB)
Megalopolis
60
56
60
46.5
59
47
66
53
51.5
(63)
20.7
15.2
19.5
28.0
26.5
30.5
-0.131 - 0 . 3 8 0 1.0
--0.121 --0.369 0.83
-
1.017
0.989
2.90
3.06
2.90
3.68
0.82
1.94
2.36
2.0
1.63
1.90
1.95
2,76
2.82
1.48
1.65
1.87
2.65
2.88
-0.165
-0.320
-
0.97
27.0
28.4
25.1
17.0
15.6
0.169
0.151
0.141
0.131
0.113
0.275
0.287
0.275
0.362
0.319
1.10
1.15
1.04
1.04
0.98
19.2
0.125
0.370
0.90
0.904
2.97
3.28
1.06
1.14
1.25
1.07
1.00
1.24
1.05
1.30
-
2.10
2.04
1.92
1.92
1.69
71
(GR)
Viterbo (I)
Warsaw (PL)
Pisa (destr.)
MZP Vllt/Vm
?
?
(28)
24.4
M. meridionalis
Valdarno (I)
Valdarno(1)
Valdarno(1)
Valdarno (1)
Valdarno(I)
1GF,
IGF,
1GF,
1GF,
1GF,
14838
10791
1057
1050
13730
m
m
m?
f
f
160
188
178
130
138
40
47
47
45
45
44
54
49
47
44
f
154
63
57
m
m
48
m
160 44
m
137 40
m
143 43
m? (119) 42
m
134 33
m
125 41
m (112) 33
m
130
m
137 44
f (134)
f
117 39
f
46
f?.
128 43
42
51
50
50
46
42
41
43
43
46
48
54
44
50
45
-
M. trogontherii
Edersleben (D)
SMS
63
M. primigenius
Praz Rodet (CH)
Steinheim (D)
Siegsdorf(D)
Ahlen (D)
Condover (GB)
Gewande(NL)
Beresovka(RUS)
Taimyr(RUS)
LiakhovIs.(RUS)
Gydan (RUS)
LenaR.(RUS)
Aa(F)
Oyosh(RUS)
Rochester (USA)
Kerkdriel (NL)
Lausanne
SMNS
NMMS
GP1UM
Shropshire
Hoofddorp
ZM, N5315
ZM, N2710
MNHN
P1N
ZM, 71911
Boulogne
NKM
AMNH
Empel
pelvic aperture, to the m i n i m u m width of the corpus ossis
ilii (here m e a s u r e m e n t 10), allows a distinction between
the sexes to be made. Both the morphology of the pelvis,
especially a r o u n d the pelvic aperture - with crests along
the inner m a r g i n , and the accentuated e m i n e n t i a e
iliopubicae - and the metrical proportions of the pelvis
(Fig. 4), indicate that the Binsfeld specimen belonged to
a female.
Size and an a t t e m p t at age determination
The metrical study shows that our specimen is relatively
large, especially considering that it is a female. In comparison 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).
20.0
25.0
26.0
62
-49
-46
52
47
-47
46
55
-40
-45.2
42
0.163
0.313
0.365
20.2 0.141
0.322
19.0 -0.160 -0.353
18.3 0.137
0.306
19.4 0.155
0.344
20.8 - 0 . 1 8 6 - 0 . 3 8 4
19.3 0.148
0.354
18.2 0.133
0.350
19.4 -0.145 0.403
-15.4 0.132
0.377
19.0
16.0 0.125
0.352
1.09
1.13
1.09
1.05
0.806
0.939
-0.913
0.788
0.915
-0.915
1.000
0.872
-1.350
0.973
1.07
2.28
2.21
2.24
2.22
2.07
2.38
2.64
2.78
2.86
2.63
2.81
2.13
2.21
1.8
2.11
1.59
2.42
2.53
2.42
2.69
But as m e n t i o n e d above, the individual age should also
be considered because of the continuous growth observed
in elephants. Age d e t e r m i n a t i o n can be undertaken accurately with the help of the dentition, but this, unfortunately, is not preserved in this case.
A n age determination by m e a n s of the pelvis is hardly
practicable. Studies on recent African elephants (HAYNES
1 9 9 1 : 3 5 1 , tab. A15) showed that the three bones of the
i n n o m i n a t e fuse at the age of 8 to 12 years. The fusion of
the sacrum to the i n n o m i n a t e 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 i n n o m i n a t e s in the pelvic symphysis is not
studied as yet. Personal observations of LISTER suggest
that in male m a m m o t h the two halves may fuse between
30 and 40 years (pers. c o m m . Dr. A. LISTER 1998). Fe-
212
URSULAB. GOHLICH
male specimens are m o r e rare, but there is a fused pelvis
o f a female M. primigenius f r o m Oyesh, Sibiria whose
age is estimated b y AVERIANOV (1996: 263) to be 35-40
years.
We have to take into account that all these observations
are m a d e in different t a x a and often are based only on a
few specimens. The c o m b i n a t i o n o f the states of fusion
at the pelvis from B i n s f e l d (fusion o f three bones o f the
innominate, fusion o f p e l v i c s y m p h y s i s , no fusion to the
sacrum) does not fit with these observations. Accordingly
an age determination s e e m s not to be possible by means
of present data. But the fusion o f the three bones o f the
innominate and e s p e c i a l l y the fusion o f the left and right
halves strongly suggest, that the a n i m a l was adult.
Results
Fig. 3. Elephas antiquus (FALCONER& CAUTLEY),pelvis measurements (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.
Morphological and metrical studies alone would not have
allowed a systematic d e t e r m i n a t i o n closer than the specimen is an elephantid. The size o f the specimen (Fig. 5)
indicates that it is too big to be M. primigenius. There is
only one pelvis o f M. primigenius (Siegsdorf) which
reaches the d i m e n s i o n s o f the one studied here, and that
is a male; all others o f w h a t e v e r sex are distinctly smaller.
The size of the s p e c i m e n in this study is very similar to
that of the only d e t e r m i n e d f e m a l e specimen of M.
trogontherii. M a l e s p e c i m e n s o f M. meridionalis and E.
antiquus c o m p a r e d in this p a p e r are bigger, whereas females are smaller. But it has to be r e m e m b e r e d that the
only female E. antiquus noted here (Crumstadt), is apparently from a y o u n g e r i n d i v i d u a l with an unfused pubic symphysis.
The litho- and b i o s t r a t i g r a p h i c situation at Binsfeld
suggests a Late P l e i s t o c e n e age and the remarkably good
preservation o f the s p e c i m e n supports a more or less autochthonous deposition. M. meridionalis and M. trogontherii can be e x c l u d e d in all probability, being Early and
M i d d l e Pleistocene species. This is c o n f i r m e d by the occurrence o f dental r e m a i n s o f o n l y M. primigenius and E.
antiquus at the site o f B i n s f e l d (pers. c o m m . Dr. R.
ZmGLER 1998).
Fig. 4. Comparison of some gender dependent measurement ratios (see Tab. 3) of elephantids (symbols see Fig. 5).
On a pelvis of the straight-tusked elephant Elephas antiquus (Proboscidea, Mammalia) from Binsfeld
213
DERANIYAGALA,P. E. R 1955. Some extinct elephants, their
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 c o n f i r m i n g this and for considering the intraspecific variation of pelvic m o r p h o l o g y and size of E. antiquus in c o m p a r i s o n with the representatives of the
Mammuthus-lineage more material is needed.
Acknowledgements
I wish to thank Dr. M. WUTTKEand the Landesamt ftir Denkmalpflege Rheinland-Pfalz, Referat Erdgeschichtliche Denkmalpflege (Mainz), for placing the specimen at my disposal
and for their financial support, Dr. A. LmTER(University College 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.-Institutftir Pal~iontologie 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 biostratigraphic information and for supplying further measures
of some pelvic specimens from Briihl.
References
ADAM, K.D. 1961. Die Bedeutung der pleistoz~inen S~iugetierfaunen Mitteleuropas ftir die Geschichte des Eiszeitalters.- Stuttgarter Beitr~ige zur Naturkunde 78: 1-34,
Stuttgart.
ANDREWS, C. W. & COOPER, C. E 1928. On a specimen of
Elephas antiquus from Upnor. - 25 pp., London (British
Museum of Natural History).
AVERIANOV,A. O. 1996. Sexual dimorphism in the mammoth
skull, teeth, and long bones. - [In:] SHOSHANI,J. & TASSY,
P. [eds.] The Proboscidea. Evolution and Palaeoecology of
Elephants and their Relatives: 260-267, New York (Oxford
University Press).
BEDEN, M. 1969. Etude et r6construction des restes de Palaeoloxodon (Elephas) antiquus du Chartelard (Charente).Bulletin des Sciences de la Terre de l'Universit6 de Poitiers
10: 44-56, Poitiers.
relatives and the two living species.- 161 pp., Colombo
(Ceylon National Museum).
DRIESCH,A. VON DEN 1976. Das Vermessen von Tierknochen
aus vor- und frtihgeschichtlichen Siedlungen.- 114 pp.,
Miinchen (Institut far Pal/ioanatomie, Domestikationsforschung und Geschichte der Tiermedizin der Universit/it
Mtinchen).
DUBROVO, I. A. 1977. Origin and migration of palaeoloxodont
elephants. - International Geology Review 19 (9): 10851088, Washington.
DUBROVO, I. A. & JAKUBOVSKI,G. 1988. The carpus morphology of the forest elephant (Palaeoloxodon) and its
significance for taxonomy. - Prace Muzeum Ziemi 40: 6583, Warszawa.
HAYNES, G. 1990. The mountains that fell down: life and death
of heartland mammoths. - [In:] AGENBROAD,L. D.; MEAD,
J. I. & NELSON,L. W. [eds.] Megafauna and man: discovery
of America's heartland. - Hot Springs Scientific Papers 1:
22-31, Hot Springs.
1991. Mammoths, mastodonts, and elephants: biology,
behavior, and the fossil record. - 413 pp., New York (Cambridge University Press).
JAKUBOWSKI,G.; KRYSIAK,K. & ROSKOSZ,T. I968. The forest
elephant - Palaeoloxodon antiquus (FALCONER& CAUTLEY,
1847) fi-om Warsaw. - Prace Muzeum Ziemi 12: 187-215,
Warszawa.
KOEN1GSWALD,W. VON [ed.] 1988. Zur Pal~ioklimatologie des
letzten Interglazials im Nordteil der Oberrheinebene.[In:] FRENZEL,B. [series ed.] Pal/ioklimaforschung 4:327
pp., Stuttgart (Gustav Fischer Verlag).
- - 1994. Mensch und Tier im Rheinland w/~hrend der vielfNtigen Klimaschwankungen des Eiszeitalters.- [In:]
KOENIGSWALD,W. VON & MEYER, W. [eds.] Erdgeschichte
im Rheinland: 189-208, Mtinchen (Friedrich Pfeil).
KROLL, W. 1991. Der Waldelefant von Crumstadt. Ein Beitrag
zur Osteologie des Waldelefanten, Elephas (Palaeoloxodon) antiquus FALCONER& CAUTLEY(1847).- Doctoral
thesis, Tier~irztliche Fakult~it der Ludwig-MaximiliansUniversitiit Miinchen: 104 pp. [unpublished].
LISTER,A. 1996a. Evolution and taxonomy of Eurasian mamm o t h s . - [In:] SHOSHANI,J. & TASSY, P. [eds.] The Proboscidea. Evolution and palaeoecology of elephants and
their relatives: 203-213, New York (Oxford University
Press).
1996b. Sexual dimorphism in the mammoth pelvis: an aid
to gender determination. - [In:] SHOSHANI,J. & TASSY, R
[eds.] The Proboscidea. Evolution and palaeoecology of
elephants and their relatives: 254-259, New York (Oxford
University Press).
LISTER, A. & AGENBROAD,L. D. 1994. Gender determination
of the Hot Springs mammoths. - [In:] AGENBROAD,L. D.
& MEAD, J. I. [eds.] The Hot Springs mammoth site: 208214, Hot Springs / South Dakota (The Mammoth site of Hot
Springs).
LOSCHER, M. 1988. Stratigraphische Interpretation der jungpleistoz~inen Sedimente in der Oberrheinebene zwischen
Bruchsal und Worms. - [In:] KOENIGSWALD,W. VON [ed.]:
Zur Pal/ioklimatologie des letzten Interglazials im Nordteil
der Oberrheinebene: 79-104, Stuttgart (Gustav Fischer
Verlag).
MAGLIO,V. J. 1973. Origin and evolution of the Elephantidae. Transactions of the American Philosophical Society, New
Series, (3) 63: 1-149, Philadelphia.
MAZO, A.V. 1998. E1 elefante f6sil de Buelna (Asturias). Nota
preliminar. - Estudios Geol6gicos 54: 273-274, Madrid.
MELENTIS, J. K. 1963. Studien tiber fossile Vertebraten Griechenlands, 3. Die Osteologie der pleistoz~inen Proboscidier
des Beckens von Megalopolis im Peloponnes (Griechen-
-
-
-
214
URSULAB. GOHLICH
land). - Annales G6ologiques des Pays Hell6niques 14: 1107, Athen.
SCHARPF, H.-J. 1977. Erl~iuterungen zur geologischen Karte
von Hessen 1:25.000, Blatt 6316 W o r m s . - 282 S., Wiesbaden (Hessisches Landesamt fur Bodenforschung).
SeHWElSS, D. 1988. Jungpleistozfine Sedimentation in der n6rdlichen Oberrheinebene. - [In:] KOENIGSWALD,W. VON [ed.]
Zur Palfioklimatologie des letzten Interglazials im Nordteil der Oberrheinebene: 19-78, Stuttgart (Gustav Fischer
Verlag).
TREVISAN, L. 1948. Lo scheletro di Elephas antiquus italicus
di Fonte Campanile (Viterbo).- Palaeontographia Italica
44: 1-77, Pisa.
Eingang des Manuskriptes am 12. Januar 1999;
Annahme durch die Schriftleitung am 5. Januar 2000.