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A stylohyoideum of Palaeoloxodon antiquus from Gesher Benot Ya'aqov, Israel: morphology and functional inferences

Authors:
Gesher Benot Ya'aqov (GBY), located south
of the Hula Valley, Dead Sea Rift, a segment of
the Great African Rift System, is known for its
early to middle Pleistocene (780,000 years ago)
sedimentary sequence comprised of several
Acheulian archaeological horizons, rich in lithic
assemblages as well as in fauna and flora
(Goren-Inbar et al. 2000). Material referable to
extinct straight-tusked elephant, Palaeoloxodon
antiquus (family Elephantidae, order Pro-
boscidea) includes a cranium, tusk fragments,
limb bones, and one left stylohyoideum (Goren-
Inbar et al. 1994). Some authors prefer to use
Elephas (Palaeoloxodon) antiquus instead of
Palaeoloxodon antiquus.
The stylohyoid (Hebrew University catalogue
no. GBY #215) was found in sediments proba-
bly originating from Layer II-6 (Trench II, Area
B), the same layer where the cranium material
of P. antiquus was found. One of a set of little
studied bones, the stylohyoid is found deep in
the throat (the hyoid apparatus comprised of
two stylohyoidea, two thyrohyoidea, and one
basihyoideum) of elephants, living and extinct
(Shoshani 1986). They are rarely collected
from extant or extinct proboscideans. Unfamil-
iarity with these bones has resulted in describ-
ing a stylohyoid as an antler of an extinct deer
(Green 1956). The stylohyoid is a Y-shaped
bone, one “arm” of the Y is the superior ramus,
the other “arm” is posterior ramus, and the third
“arm” is the inferior ramus (terminology after
Inuzuka et al. 1975). In this specimen, the in-
ferior ramus is broken, otherwise it is in good
condition; the combined superior-posterior
length is 135 mm. This specimen does not have
the typical “angulus” (a depression on the su-
perior-posterior rami as observed in side view)
that has been observed on several stylohyoid
bones of P. naumanni (Inuzuka et al. 1975).
This difference could be a species difference
between P. naumanni and P. antiquus. GBY
#215 has a tubercle on the superior ramus that
may be homologous to tubercles observed on
some P. naumanni stylohyoidea. It is noted that
only one stylohyoid of P. antiquus was exam-
665
A stylohyoideum of Palaeoloxodon antiquus from
Gesher Benot Ya’aqov, Israel: morphology
and functional inferences
J. Shoshani1, N. Goren-Inbar2, R. Rabinovich3
1Department of Biology, University of Asmara, Asmara, Eritrea (Horn of Africa)
hezy@eol.com.er
2Institute of Archaeology, Hebrew University, Mt Scopus, Jerusalem, Israel
goren@cc.huji.ac.il
3Department of Evolution, Systematics and Ecology, Hebrew University, Jerusalem, Israel
- rivka@vms.huji.ac.il
SUMMARY: Among the mammal remains discovered at Gesher Benot Ya'aqov (south of the Hula Valley,
Dead Sea Rift, Israel) were cranium, tusk fragments, and limb bones that were assigned to an extinct straight-
tusked elephant, Palaeoloxodon antiquus. One stylohyoideum was also collected. Based on inferences from
data on living elephants, it is hypothesized that P. antiquus had a tongue about 80 cm long used to grasp leaves
and grasses. Further, P. antiquus lived in small herds, about 5-15 individuals, and that herd members could
have communicated with infrasonic calls with other herds, a few kilometers away. All in all, the hyoid appa-
ratus has been a pivotal structure for adaptation in the course of proboscidean evolution.
ined compared to several stylohyoids of P. nau-
manni.
Based on the work of Inuzuka (1977a, b) and
our morphological observations (in the context
of archeological settings of GBY, as well as
other observations of skeletons of P. nauman-
ni), it is suggested that the genus Palaeolox-
odon be considered a bona fide taxon and not a
subgenus of Elephas. Detailed long-term study
(about 25 years) of processes and grooves for
muscle attachment and twisting of bones on the
hyoid apparatus (mostly stylohyoidea) enables
the authors to infer functionality for these
bones. Elephantid taxa, for example, possess a
posterior ramus which is absent in some early
proboscideans, e.g., Mammut americanum
(Tassy & Shoshani 1988; Saegusa & Shoshani
1992). This ramus serves for attachment of the
digastricus muscle that helps to open the jaw
(Eales 1926; Garrod 1875; Gasc 1967). Identi-
fication of GBY 215 was made by comparing it
to 194 stylohyoid hyoid elements, representing
151 individuals of living and extinct pro-
boscideans, namely: Mammoths (e.g., Mam-
muthus primigenius), Asian elephants (Elephas
maximus), extinct elephantids (e.g., E. recki,
Palaeoloxodon naumanni), African elephants
(Loxodonta cyclotis and L. africana), stegodons
(e.g., Stegodon aurorae), gomphotheres (e.g.,
Gomphotherium productum, Amebelodon flori-
danus), mammutids (Mammut americanum),
and a deinothere (Deinotherium giganteum).
Based on gular musculature of living elephants
and morphology of hyoid apparatus (e.g., Eales
1926), it is hypothesized that the functional
anatomy of hyoid of P. antiquus was similar to
that of extant elephants. It is suggested that P.
antiquus had a tongue about 80 cm long that
could be projected a short distance from the
mouth to grasp leaves and grasses. The tongue
along with a flexible trunk, enabled straight-
tusked elephants to graze or to browse on Pleis-
tocene foliage about 8 meters above ground
[trunk flexibility is deduced from size and posi-
tion of external naris and from size and num-
bers of infraorbital canals (Shoshani 1986);
height above ground is estimated from data on
living elephants when standing on hind legs
(Shoshani et al. 1987). It is suggested that P.
antiquus lived in small herds, about 5-15 indi-
viduals, and that herd members could have
communicated with infrasonic calls with other
herds, perhaps a few kilometers away (commu-
nication hypothesis is based on hyoid and
cochlear anatomy; Meng et al. 1997). In addi-
tion, it is proposed that the hyoid apparatus sup-
ported a pharyngeal pouch used as a resonating
chamber (similar to the condition in howler
monkey; Vaughan et al. 2000). At other periods,
this pouch was used to store water for drinking
or dousing in time of stress (Shoshani 1998). It
appears that the hyoid apparatus has been a piv-
otal structure for adaptation to newly available
ecological niches of in their long geological
history.
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A stylohyoideum of Palaeoloxodon antiquus from Gesher Benot Ya’aqov, Israel...
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