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

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-
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)
2Institute of Archaeology, Hebrew University, Mt Scopus, Jerusalem, Israel
3Department of Evolution, Systematics and Ecology, Hebrew University, Jerusalem, Israel
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-
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
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Goren-Inbar, N., Lister, A., Werker, E. &
Chech, M.A. 1994. A butchered elephant
skull and associated artifacts from the
Acheulian site of Gesher Benot Ya'aqov, Is-
rael. Paleorient 20/1: 99-112.
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Melamed, Y., Kislev, M.E., Tchernov, E., &
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oloxodon naumanni from saruyama, Shi-
mosa-machi, Chiba Prefecture, central
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Kamei, T. 1975. On the stylohyoid bone of
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and hearing in proboscideans. Journal of
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godontidae) from the Pleistocene of Java
and its phylogenetic significance. Journal of
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A stylohyoideum of Palaeoloxodon antiquus from Gesher Benot Ya’aqov, Israel...
... We registered modifications such as striations, scratches, cut marks, tooth marks, gnaw marks, burnt elements and pathology in their anatomical location along the element. Our identification of surface damage is based on published criteria (carnivore tooth marks: Binford, 1981;Selvaggio, 1988, 1991;Blumenschine, 1995;cut-, hack-, and percussion marks: Binford, 1981;Shipman, 1981;Shipman and Rose, 1983), as well as comparisons with assemblages from known taphonomic origins in the HUJ collections; assemblages of rodents (Rabinovich, 1990), porcupines (Rabinovich and Horwitz, 1994) and carnivores (Rabinovich, 1990) in captivity, and collected material from recent Table 2 Body size groups (kg) of mediumelarge mammals from GBY (after Rabinovich, 1998). BSGA (Elephant, > 1000 kg); BSGB (Hippopotamus, rhinoceros approx. ...
... In Area C there are also only a few elephant bones, but they include more body elements in addition to teeth and tusk fragments: Layer V-6 contained a cervical vertebra, an astragalus and a carpal; Layer V-5 contained a posterior ramus of a hyoid bone (Shoshani et al., 2004(Shoshani et al., , 2007, and a carpal; and JB yielded a pisiform and an intermedium carpal. A complete M 2 (r) from JB indicates that the elephant was at least 26 years old when it died (Maglio, 1973). ...
... Both, however, are a product of the agent of accumulation. Body part representation and surface modification of the studied fauna suggest that the reptiles and most of the amphibians at GBY represent Biocoenose d local life assemblages, and Thanatocoenose d a death assemblage comprised of bones of animals that died locally (Shipman, 1981). Taphocoenose d an assemblage of bones brought together by a taphonomic process/agent is represented by some of the anurans and by the mediumelarge mammals. ...
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This study investigates faunal remains from the site of Gesher Benot Ya'aqov, analyzing how hominins may have utilized vertebrates and exploring paleoenvironments during the Early and Middle Pleistocene. Based on analyses of a range of vertebrates, results show that the species distribution of terrestrial vertebrates (microvertebrates and mammals) at the site of Gesher Benot Ya'aqov shows relative stasis above the MatuyamaeBrunhes Boundary (MBB) (0.78 Ma). However, consistency in faunal remains at the site reflecting stasis does not seem to reflect accurately the paleoenvironment. Marked changes are demonstrable in the lake-margin sedimentary background, archaeological remains, and agents of accumulation and damage, as well as in the density of mediumelarge mammals. This study emphasizes the significance of studying assemblages as a whole rather than the species representation on its own.
... The area is tectonically active, which results in the creation and rejuvenation of complex landscape features that have been shown to be particularly favourable for hominin occupation. It is also relatively rich in Palaeolithic remains and has been extensively studied, so that there is a database of observations on which synthetic research can draw (Bar-Yosef, 1995;Tchernov, 1999;Turner, 1999;Goren-Inbar et al., 2000;O'Regan et al., 2005;King and Bailey, 2006;Shea, 2008;Belmaker, 2010;Fleagle et al., 2010;Issar, 2010;Bar-Yosef and Belmaker, 2011;Winder et al., 2012). All of this makes the region an interesting regional 'laboratory' for the examination of the various factors that may have promoted or constrained hominin occupation and dispersal. ...
... But GBY also offers other advantages to hominins (Stekelis et al., 1937(Stekelis et al., , 1938Alperson-Afil et al., 2009;van Zeist and Bottema, 2009). Unlike many potential migration-hunting sites, the migratory animals are not the only resource (Goren-Inbar et al., 1992, 2000Shoshani et al., 2001;Rosenfeld et al., 2004;Rabinovich et al., 2011). Because of its lake-side/riverine location, non-migratory animals (pigs, fallow deer, even hippopotamus) were available locally, so that hominin groups arriving to exploit the spring or autumn herd migration could not only support themselves while they waited, but support themselves if the migration, for whatever reason, failed to materialise (Rabinovich and Biton, 2011). ...
We explore the relationship between the edaphic potential of soils and the mineral properties of the underlying geology as a means of mapping the differential productivity of different areas of the Pleistocene landscape for large herbivores. These factors strongly control the health of grazing animals irrespective of the particular types of vegetation growing on them, but they have generally been neglected in palaeoanthropological studies in favour of a more general emphasis on water and vegetation, which provide an incomplete picture. Taking the CarmeleGalileeeGolan region as an example, we show how an understanding of edaphic potential provides insight into how animals might have exploited the environment. In order to simplify the analysis, we concentrate on the Lower Palaeolithic period and the very large animals that dominate the archaeofaunal assemblages of this period. Topography and the ability of soils to retain water also contribute to the differential productivity and accessibility of different regions and to patterns of seasonal movements of the animals, which are essential to ensure a supply of healthy fodder throughout the year, especially for large animals such as elephants, which require substantial regions of good grazing and browsing. Other animals migrating in groups have similar needs. The complex topography of the Southern Levant with frequent sudden and severe changes in gradient, and a wide variety of landforms including rocky outcrops, cliffs, gorges, and ridges, places major limits on these patterns of seasonal movements. We develop methods of mapping these variables, based on the geology and our substantial field experience, in order to create a framework of landscape variation that can be compared with the locations and contents of archaeological sites to suggest ways in which early hominins used the variable features of the landscape to target animal prey, and extend the analysis to the consideration of smaller mammals that were exploited more intensively after the disappearance of the elephants. We consider some of the ways in which this regional-scale approach can be further tested and refined, and advocate the development of such studies as an essential contribution to understanding the wider pattern of hominin dispersal.
... The geological sequence of Revadim as published in 1999 Wieder and Gvirtzman, 1999), comprises an Table 1 Pleistocene proboscideans from the Levant (see location of sites in Fig. 1 Lister, 2004 Previously defined by Hooijer (1961) and Guérin et al., 1993Palaeoloxodon antiquus Gesher Benot Ya'aqov, Jordan Valley, Israel Goren-Inbar et al., 1994Shoshani et al., 2001;Biton, 2011 Revadim, Central Coastal Plain, Israel Marder et al., 1999;Rabinovich et al., 2005; Table 2). In general, the paleosurface (i.e. the original topography of the site) is characterized by undulating topography, a combination of rills and gullies. ...
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Lower Paleolithic faunal and lithic assemblages serve as a major source of information on the behavior and capabilities of Early- and Middle Pleistocene hominins. The multi-layered Late Acheulian site of Revadim Quarry provides a rare opportunity to study hominin–elephant interactions during the Late Lower Paleolithic period in the Levant. A large proportion of this open-air site was excavated (ca. 250 m2) and yielded a wealth of lithic and faunal remains. In this paper the proboscideans from Revadim are presented for the first time within the broader geomorphological, stratigraphic and archaeological context in order to allow a better understanding of elephants within the Acheulian in the southern Levant. The unprecedented quantity of elephant remains at the site is accompanied by large and rich lithic assemblages. Of special interest are several elephant bones with cut marks, and the earliest appearance in the southern Levant of bones that seem to have been shaped to resemble tools. The site bears testimony to complex exploitation of proboscideans.
... The stylohyoidea of Eurasian straight-tusked elephants are little known, and reported from a very few sites. The stylohyoidea of straight-tusked elephants, thus far described, are those of Palaeoloxodon naumanni from Japan (Inuzuka et al., 1975;Inuzuka, 1977), of a primitive Palaeoloxodon, closer to Palaeoloxodon recki than to Palaeoloxodon antiquus, from the early Middle Pleistocene site of Gesher Benot Ya'aqov (Israel) (Goren-Inbar et al., 1994;Shoshani et al., 2001), and of Palaeoloxodon antiquus from the late Middle Pleistocene site of Neumark Nord (Saale, Germany) . Conversely, no stylohyoidea of Italian Palaeoloxodon antiquus have hitherto been described. ...
The site of La Polledrara di Cecanibbio (Latium, Italy) is related to deposits of the PG6 Sequence (Middle Pleistocene, Aurelia Formation, MIS 10 and 9). The sediments are mainly volcaniclastic in composition, and constitute the filling of incised valleys, mainly characterized by fluvial deposits at the base, passing upward to fluvio-lacustrine and palustrine deposits containing abundant fossil mammal remains and artifacts. The arrangement of the specimens and taphonomic observations suggest that most of the transport of the bones occurred during flooding events, followed by progressive swampy phases, resulting in the formation of areas with stagnant and muddy waters where some elephants became trapped, as indicated by remains in partial anatomical articulation. Recent excavations carried out at the site permit a better definition of the palaeoenvironmental reconstruction, already partially outlined in previous publications. In particular, an area showing a close correlation between the skeleton of an elephant and human activity, allows documentation and better understanding of some aspects of human eelephant interaction, probably mainly represented by scavenging activity.
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Results are based on ten hyoid characters of seven proboscidean taxa; four belong to the subfamily Elephantinae. Using PAUP we generated six equally parsimonious trees. The consensus tree yields two polytomies: in the basal polytomy the relationships among Mammut, gomphotheres, Stegodon, and Loxodonta are not resolved; in the other polytomy Palaeoloxodon, Mammuthus, and Elephas are grouped in a cluster. The joining of Palaeoloxodon and Mammuthus has not been suggested previously, and there are ample non-hyoid data of grouping Loxodonta, Elephas, and Mammuthus. For this reason and since a major focus of this study has been to test relationships among Loxodonta, Elephas, and Mammuthus, we relied on studies of other workers and rearranged the consensus cladogram to unite Loxodonta with other members of Elephantinae (Palaeoloxodon, Elephas, and Mammuthus). The hypothesis that Mammuthus is more closely related to Elephas than to Loxodonta is more parsimonious (by two evolutionary steps) than when Loxodonta joins Mammuthus. This finding is provisional and should be retested with additional data, especially with taxa with small sample size, and with specimens of extinct Elephantinae taxa. Results also corroborate other findings that Palaeoloxodon is a bona fide elephant genus; we classify it in the new subtribe Palaeoloxodontina Zhang and Zong [1983. Genus Palaeoloxodon of China. Vertebrata PalAsiatica 21(4), 301–312].
Elephas (Palaeoloxodon) antiquus is a well-known elephant species of the Middle and Late Pleistocene of Europe, but few skeletons so far have been described in detail. Here we present a detailed account of a partial skeleton in good condition from the alluvial sands of the Amyntaio coal mines, Macedonia, Greece. It represents a large male aged in its forties. Based on extant and extinct elephant specimens, the Amyntaio's elephant estimated height at the shoulder is 3.5 m and its weight close to 9 tonnes. A CT scan was performed on the deformed fifth metacarpal which was diagnosed with osteomyelitis, probably rendering the animal lame. No signs of further biologically induced ante- or post-mortem modifications were detected. From that skeleton the first known basihyoid bone of E. antiquus is recovered; comparisons with homologous bones of other elephantid taxa show it has a very distinct morphology and can be used in phylogenetic studies of the Elephantidae family.
This volume, the first in a series devoted to the paleoanthropological resources of the Middle Awash Valley of Ethiopia, studies Homo erectus, a close relative of Homo sapiens. Written by a team of highly regarded scholars, this book provides the first detailed descriptions, photographs, and analysis of the fossil vertebrates-from elephants and hyenas to humans-from the Daka Member of the Bouri Formation of the Afar, a place renowned for an abundant and lengthy record of human ancestors. These fossils contribute to our understanding human evolution, and the associated fauna provide new information about the distribution and variability of Pleistocene mammals in eastern Africa. The contributors are all active researchers who worked on the paleontology and geology of these unique deposits. Here they have combined their disparate efforts into a single volume, making the original research results accessible to both the specialist and the general reader. The volume synthesizes environmental backdrop and anatomical detail to open an unparalleled window on the African Pleistocene and its inhabitants.
This chapter is dedicated to a detailed paleontological description of the medium- to large-sized mammal fauna discovered at Gesher Benot Ya‘aqov (GBY), taking into account previous paleontological studies undertaken in this sector of the Dead Sea Rift.
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Le crâne d'un elephant a defense droite, Palaeoloxodon antiquus, a ete decouvert en 1989 a Gesher Benot Ya'aqov, site du Pleistocene moyen, en Israel. Ce crâne etroitement associe a des artefacts en pierre et en bois fait partie integrante de la composition du sol d'occupation acheuleen. Son emplacement et les dommages qu'il a subis suggerent des fractures intentionnelles pour l'extraction de la cervelle. Un nucleus en basalte, un bloc de pierre et un morceau de chene se trouvaient sous le crâne et ont pu etre utilises pour le retourner. On decrit ici un scenario possible de chasse et de demembrement, en s' inspirant de paralleles archeologiques et modernes. Ces trouvailles corroborent d'autres elements mis en evidence sur le site qui indiquent un comportement sophistique des habitants de cette region, il y a 500.000 ans.
A new approach to proboscidean evolution depicts taxa in three major radiations. This approach highlights general proboscidean evolutionary trends and origins more than the specific relationships among them. Data from more than 55 million years of evolution help to interpret how the integration of primitive and derived characters was essential to proboscidean success. Only two, or perhaps three, species remain of approximately 164 that lived in the past. Extinct forms were extremely cosmopolitan, occupying a variety of habitats, from deserts to mountain tops, on all continents except Australia and Antarctica. Challenges for future investigators include a better understanding of structure and function of infrasonic call production and perception, brain features, and reproductive biology in extinct proboscideans based on inferences from living forms.
Typescript. Thesis (Ph. D.)--Wayne State University, 1986. Includes bibliographical references (leaves 572-635).
The Acheulean site of Gesher Benot Ya'aqov in the Dead Sea Rift of Israel documents hominin movements and technological development on a corridor between Africa and Eurasia. New age data place the site at 780,000 years ago (oxygen isotope stage 19), considerably older than previous estimates. The archaeological data from the site portray strong affinities with African stone tool traditions. The findings also reflect adroit technical skills and in-depth planning abilities, more advanced and complex than those of earlier archaeological occurrences in the Levant.