Content uploaded by Jan van der Made
Author content
All content in this area was uploaded by Jan van der Made on Dec 20, 2020
Content may be subject to copyright.
137
Jan van der Made1*, Mohamed Sahnouni2, 3, 4, Kamel Boulaghraief4
1 CSIC, Museo Nacional de Ciencias Naturales, c. José Gutierrez Abascal 2, 28006 Ma-
drid, Spain; mcnjv538@mncn.csic.es
2 Centro Nacional de Investigación sobre la Evolución Humana, Paseo Sierra Atapuerca
3, 09002, Burgos, Spain; mohamed.sahnouni@cenieh.es
3 Stone Age Institute & Anthropology Department, Indiana University Bloomington, IN,
USA
4 Centre National de Recherches Préhistoriques, Anthropologiques et Historiques (CNR-
PAH), 3 rue Franklin Roosevelt, 16000 Algiers, Algeria ; boulaghraifkamel@yahoo.fr
*Corresponding author
ABSTRACT
Archaeological excavations at the site of El Kherba, an Early Pleisto-
cene site in Algeria, have yielded Hippopotamus fossils, including a skull.
El Kherba is a site known for producing Oldowan stone artifacts and cut-
marked bones, dated to about 1.8 Ma. e elevated occipital and orbits of
the skull, the short temporal fossa and the backward sloping nasal profile
suggests that it belongs to Hippopotamus gorgops. e same species of Hi-
ppopotamus is present in the nearby locality of Ain Hanech, which is of the
same age as El Kherba. e oldest known Hippopotamidae are from the
Early Miocene of sub-Saharan Africa, where they reached their major di-
versity with five or six coeval species being common during the Plio-Pleis-
tocene. From there they dispersed northwards to North Africa and Euro-
pe, and eastwards to the Indian subcontinent and SE Asia. e most likely
Hippopotamus gorgops
from El Kherba (Algeria)
and the context
of its biogeography
Jan van der Made et al.
138
origin of the European Hippopotamus is North Africa and it seems that in
both areas the same three or very similar species appeared one after the
other. e first Hippopotamus from North Africa is not well known, but it
may have originated from the East African H. kaisensis and may have given
rise to H. antiquus. is species, which lived from 2 till shortly after 1.8 Ma,
is well documented in Europe. Hippopotamus gorgops dispersed around 1.8
Ma to North Africa and around 1.2 Ma to Europe, where it is known as H.
tiberinus. During the late Middle or Late Pleistocene Hippopotamus am-
phibius dispersed to North Africa and Europe. e dispersal of H. gorgops
across the Sahara towards North Africa around 1.8 Ma is well documented.
e dispersal of the species which gave rise to H. antiquus may have occu-
rred as early as 2.5 Ma, when lakes were extensive in East Africa. e same
climatic circumstances, that caused more extensive lakes and rivers in East
Africa, may have created similar environments in the Sahara, allowing hi-
ppos and other fauna, including humans, to spread northwards.
Keywords: Hippopotamus, dispersal, Sahara, Maghreb, environment
RESUMEN
Las excavaciones arqueológicas en El Kherba, un yacimiento del
Pleistoceno inferior de Argelia, han proporcionado fósiles de Hippopota-
mus, incluyendo un cráneo. El Kherba es un yacimiento conocido por ha-
ber revelado la presencia de artefactos líticos olduvayenses y huesos con
huellas de corte datados hace unos 1.8 Ma. El elevado occipital y órbitas,
la fosa temporal corta y los nasales inclinados hacia atrás sugieren que el
cráneo pertenece a Hippopotamus gorgops. Esta misma especie de Hippo-
potamus está presente en el cercano yacimiento de Ain Hanech, contem-
poráneo a El Kherba. Los Hippopotamidae más antiguos conocidos son
del Mioceno del África sub-sahariana, donde alcanzaron su mayor grado
de diversidad con cinco o seis especies coetáneas durante el Plio-Pleisto-
ceno. Desde aquí estas especies se dispersaron hacia el norte, alcanzando
el norte de África y Europa, y hacia el este, hasta el subcontinente indio y
el sureste de Asia. El origen más probable de los hipopótamos europeos
es el norte de África, y parece que las mismas especies, o especies muy
próximas, vivieron en ambas zonas. No se conoce bien la primera especie
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
139
de Hippopotamus del norte de África, pero pudo haberse originado del
H. kaisensis del este de África, dando lugar a la especie Hipopotamus
antiquus, que vivió en Europa entre unos 2 y 1,8 Ma. Hace unos 1,8 Ma,
H. gorgops se dispersó hacia el norte de África y hace alrededor de 1,2
Ma a Europa, donde se conoce como Hippopotamus tiberinus. Durante
el Pleistoceno Medio tardío o Superior, Hippopotamus amphibius se dis-
persó hacia el norte de África y Europa. La dispersión de los hipopótamos
a través del Sahara hacia el Norte de África ocurrió probablemente hace
alrededor de 1,8 y 2,5 Ma. En este tiempo existían extensos lagos en el
este de África y las mismas circunstancias climáticas puedieron haber
dado lugar a ambientes parecidos con lagos extensos y ríos en el norte
de África. Estos ambientes pudieron haber permitido la dispersión hacia
el norte de los hipopótamos y de otra fauna, incluyendo a los humanos.
1. INTRODUCTION
e Hippopotamidae have a remarkable biogeographical history. Hi-
ppos were more diverse in sub-Saharan Africa than in North Africa and
dispersed several times into Europe and Southern Asia. Most probably the-
se dispersals into Europe took place through North Africa.
Most researchers consider the initial dispersal of hippos into Europe
as an important event of stratigraphic importance (e.g. Rook & Martínez
Navarro, 2010), and consider this event to be related to climatic change
and even to the first human dispersal to Europe (e.g. Martínez Navarro,
2010). e taxonomy (Caloi et al., 1980; Faure, 1985; Mazza, 1991; 1995;
Guérin, 1996; Kahlke, 1997; 2001; 2006), the number of dispersals, and the
timing of the first of these towards Europe (e.g. Faure, 1985; Mazza, 1991;
1995; Van der Made, 2005a; 2011; Martínez-Navarro, 2010, Rook & Martí-
nez-Navarro, 2010; Bellucci et al., 2012; 2014) have been contentious issues
in paleontology. A recent proposal is that there were three Pleistocene dis-
persals: 1) Hippopotamus antiquus (= H. major) at about 2 Ma; 2) Hippo-
potamus tiberinus shortly before 1.2 Ma; and 3) Hippopotamus amphibius
in the late Middle or early Late Pleistocene (Van der Made et al., in press).
Most likely, the hippo species which reached Europe or their imme-
diate ancestors passed through or at least reached the Maghreb. Pleistoce-
ne fossils of Hippos have been described or mentioned from North Africa
Jan van der Made et al.
140
(Pomel, 1890; 1896; Arambourg, 1970; 1979; omas, 1977; Geraads, 1980;
2002; Geraads et al., 1992; Chaid-Saoudi et al., 2006; Van der Made & Sah-
nouni, 2013).
Here we report on some morphological features of a hippo skull re-
covered from the El Kherba archaeological site (Algeria), and discuss their
significance to systematics and the dispersal of hippo species from sub-Sa-
haran Africa towards the north.
2. THE EL KHERBA ARCHAEOLOGICAL SITE
e El-Kherba site is situated on the edge of the north-eastern Alge-
rian High Plateau, north of the city of El-Eulma (Province of Sétif). Disco-
vered in 1992, it is a part of the Ain Hanech Plio-Pleistocene site complex,
representing a lateral extension of the classic site of Ain Hanech. Both El
Kherba and Ain Hanech are contemporary and belong to the Ain Hanech
Formation. e Ain Hanech Formation is 30 m thick and formed of several
cyclothemic units of fluvial origin (from O to T) (Sahnouni & de Heinzelin
1998). e stratigraphic profiles of the Ain Hanech and El-Kherba sites are
correlated with the base of Unit T based on stratigraphic and altimetric
evidence.
Paleomagnetic studies undertaken in the Ain Hanech Formation, in-
cluding El Kherba, indicate a succession of magnetic polarities (R-N-R),
beginning with reversed polarity in Units O, P, and Q, followed by normal
polarity in Units R and S, and a reversed polarity in Unit T and the cal-
crete deposits sealing the sequence. e normal polarity at the bottom of
the sequence, identified as the Olduvai subchron, is 6 m thick with the El
Kherba and Ain Hanech archaeological sites situated near the top. ere-
fore, an age of 1.7 Ma is estimated for both localities (Parés et al., 2014),
which is also corroborated by biostratigraphic data from East Africa (Sa-
hnouni & Van der Made 2009). us, currently the El Kherba and Ain
Hanech sites document the oldest archaeological occurrences known in
North Africa.
e excavations at El Kherba (Figure 1) yielded a rich fauna associa-
ted with Oldowan stone tools. e fauna is savanna-like and comprises of
Proboscidea, Equidae, small and large Bovidae, Girafidae, Suidae, Carnivo-
ra, Crocodilia, and Lagomorpha. e stone artifacts include unifacial and
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
141
bifacial choppers, polyhedrons, subspheroids, spheroids, whole flakes, re-
touched pieces, and various fragments. e lithic industry and associated
fauna are contained in three distinct levels (A, B, and C). However, the bulk
of the archaeological materials, mainly accumulated in Level B including
the Hippopotamus skull described in this study, were recovered within a
floodplain setting consisting of silt and clay with gravel and calcic grains,
deposited on top of a 20 cm thick conglomerate characterized by gravels of
various sizes and shapes.
3. BIOGEOGRAPHIC HISTORY OF THE HIPPOPOTAMIDAE
e origin of Hippopotamidae has been under discussion since the
nineteenth century (see reviews by Pickford, 2008; Orliac et al., 2010). In
recent times, the Palaeochoeridae (Pickford, 1983; 2008; 2015) and An-
thracotheriidae (Gentry & Hooker, 1988; Van der Made, 1999; Boisserie
et al., 2005a; 2005b; Orliac et al., 2010; Lihoreau et al., 2015) have been
proposed as their ancestors. If hippos originated from the Palaeochoe-
ridae, they did so during the Early Miocene, either already in Eurasia or
Figure 1: A view of excavation at El Kherba in 2006. (Photo Jordi Mestre)
Jan van der Made et al.
142
shortly after their dispersal into Africa. If they originated from the An-
thracotheriidae, they may have done so during the Late Eocene or Early
Oligocene (Lihoreau et al., 2015). For a long time fossil teeth from the
Early Miocene site of Rusinga (Kenya) were considered to represent the
oldest evidence for hippo (Coryndon, 1978; Van der Made, 1999). Later,
Pickford (2007) named those teeth Kulutherium and placed this genus in
the Anthracotheriidae, but Orliac et al (2010) reaffirmed the identity of
the teeth as hippo. e slightly younger Morotochoerus ugandensis was
named on the basis of fossils from Moroto (Uganda) and four other lo-
calities and was placed in the Tayassuidae (Pickford, 1998). Later, it was
recognized to be a gryphon taxon based on fossils of different taxa, which
were identified as Albanohyus, Kenyasus, Lopholistriodon and prossibly
Nguruwe (Van der Made, 2003; 2010). e holotype from Moroto was
subsequently placed in the Hippopotamidae by Orliac et al. (2010) and in
the Anthracotheriidae by Pickford (2011). However, there seems to be a
consensus that by the Middle Miocene, Hippopotamidae occurred only
in sub-Saharan Africa and that the Hippopotaminae and more specifica-
lly the genus Hippopotamus originated there.
Hippopotaminae spread out of Africa during the latest Miocene.
Hexaprotodon reached the Middle East, the Indian subcontinent and as far
away as Java in SE Asia (Falconer & Cautley, 1836; Colbert, 1938; Hooi-
jer, 1950; Dubois, 1908; Van der Maarel, 1932; Faure, 1986; Gentry, 1999;
Boisserie, 2005). At about the same time the hippo species Hexaprotodon?
pantanelli reached Europe, where it is known from about nine latest Mio-
cene and one early Pliocene locality. is species has simple molars like
Hexaprotodon and only four lower incisors like Hippopotamus and its affi-
nities have been disputed (see review by Van der Made, 1999 and the cited
references). While Hexaprotodon has a long temporal range in southern
Asia, the incursion in Europe was short-lived, from about 6 Ma till shortly
after 5 Ma.
During the Plio-Pleistocene, Hippopotamidae diversified in sub-Saha-
ran Africa, where at certain times existed six or more contemporaneous
species (Figure 2) (Weston & Boisserie, 2010; Van der Made, 2014a, figure
19; 2014b, figure 21). By contrast, the diversity seems to have been much
lower in North Africa. Several species were named on the basis of fossi-
ls from North African localities while still more fossils were described or
mentioned from others.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
143
Gaudry (1876) named a small species based on some teeth from Pont
de Duvivier across the river Seybouse near Annaba (formerly Bône) in Al-
geria (the geographic positions of the localities are indicated in Figure 3
and the approximate ages in Figures 4 & 5). e specimens have a primitive
morphology (Pomel, 1890), are believed to be of Pliocene age and currently
placed in a different genus as Hexaprotodon? hipponensis (Weston & Bois-
serie, 2010).
e oldest North African localities where Hippopotamus has been
reported are the following three localities. Arambourg (1970) mentioned
Hippopotamus sp. from Ain el Bey and Hippopotamus amphibius from
Ain Boucherit (pages 8, 131) and Lac Ichkeul (page 131), and described
Hippopotamus (Hexaprotodon) hipponensis from Ain Boucherit (page 7),
but indicated its presence a question mark in the summary table (page 131)
Figure 2: The chronological distribution of the African Plio-Pleistocene Hippopotamidae (adap-
ted from Van der Made, 2014 and ultimately based on Weston & Boisserie, 2010). Archaeopo-
tamus harvardi reconstruction by Mauricio Antón; recent Choeropsis liberiensis (IVAU); “Trilo-
bophorus” afarensis from Hadar (ARCCH AL109/319); Hexaprotodon protamphibius from Omo
(MNHN 1933-9-777); recent Hippotamus amphibius (CNRPAH); Hippopotamus gorgops holo-
type from Olduvai (MNB).
Jan van der Made et al.
144
(Figure 4). One of us (J. van der Made) reviewed the fossils from these loca-
lities in the MNHN, but only nine teeth and two bones from Ain Boucherit
were identified as hippos and none from Ichkeul. Our work at Ain Bou-
cherit yielded few hippo specimens, which are too poor for an unequivocal
species designation.
Figure 3: The geographic location of the hippopotamus localities mentioned in the text.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
145
Hippos were also reported from a group of slightly younger sites in
North Africa (Figure 4). Arambourg (1979) reported Hippopotamus amphi-
bius from Mansoura and Ain Hanech. Geraads (1980) considered the Early
Pleistocene hippos to belong to the “group of H. amphibius”, that many of
them might be H. gorgops, which he suggested to be a junior synonym of
H. sirensis. Chaid-Saoudi et al (2006) assigned the fossils from Mansoura to
Hippopotamus cf. sirensis.
A group of still younger localities in North Africa includes Tighennif
(formerly Ternifine, formerly Palikao). Pomel (1890) named the species Hi-
ppopotamus sirensis on the basis of fossils from this locality. Later he des-
cribed the species in more detail (Pomel, 1896). Geraads (1980) reported
poor material of Hippopotamus from omas Quarry 1 level L and ho-
minid level, similar to the living species. Later, he assigned the material to
Hippopotamus cf. sirensis (Geraads, 2002). Richer materials from Aïn Maa-
rouf, but still lacking the relevant skull parts were assigned to H. c.f. sirensis
(Geraads et al., 1992). e fossils assigned to H. sirensis (or H. cf. sirensis)
cover a very long temporal range.
Hippopotamus amphibius, the living species, is reported from the
youngest North African localities (Figure 4), including Tihodaine (o-
mas, 1977) and Gisement des Phacochères (Hadjouis, 1990). e fossi-
ls from the cave of Pointe Pescade and Beni Saf (Algeria), described by
Pomel (1896) as Hippopotamus icosiensis, probably belong to the living
species of hippo.
e fossil evidence seems to point to two or three Pleistocene Nor-
th African species of Hippopotamus, which appeared one after the other
(Figure 4). North African hippo diversity seems to have been low, as was
also the case in Europe. European hippos are likely to have come from or
through North Africa, either crossing the Gibraltar or the Sicily Straits, or
following the route around the eastern end of the Mediterranean.
Concerning the systematics and temporal distribution of European Hi-
ppopotamus, there are two main opinions, and recently a third has been
published (Figure 5). Traditionally, two species or subspecies were recog-
nized: the early Middle Pleistocene Hippopotamus antiquus (= H. major or
H. amphibius antiquus), which is large, and H. amphibius from the early
Middle to Late Pleistocene, which is smaller, but still larger than the li-
ving hippos (e.g. Hooijer, 1942; 1950; Kurtén, 1968; Blandamura & Azza-
roli, 1977; Caloi et al., 1980). Faure (1984) named the latter H. incognitus.
Jan van der Made et al.
146
e two species were supposed to have been sympatric virtually during the
entire Middle Pleistocene (Figure 6) (Faure, 1985; Faure & Guérin, 1992;
Guérin, 1996). Kahlke (1997; 2001; 2006) recognized the same taxa, but as
subspecies of H. amphibius. Generally, it was believed that H. antiquus (=
H. major) was closely related to, or evolved from, Hippopotamus gorgops
(Coryndon, 1977; 1978; Blandamura & Azzaroli, 1977; Faure, 1985).
Figure 4: The approximate chronological position of the North African localities with hippos
(solid squares indicate presence, open squares indicate uncertainty: cf., aff., sp., ?), compared
to the East African record (thick lines). Hippopotamus sp. from Ain Boucherit unit P/Q upper
molar (MNHN 195A-13-94); H. gorgops skulls from El Kherba (KH06-G30-149), Ain Hanech
(photograph from Arambourg, 1979) and Olduvai (Photographs from Mazza, 1991); H. amphi-
bius (= H. icosiensis) from Pointe Pescade (Pomel, 1896).
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
147
e second main opinion is from Mazza (1991; 1995; figure 6), who ar-
gued that there were two dispersals from Africa. First, H. gorgops dispersed
to Europe, giving rise to H. antiquus. Subsequently, Hippopotamus tiberinus
evolved from H. antiquus, but also coexisted with that species. Later, H. am-
phibius dispersed from Africa and coexisted with the latest populations of H.
tiberinus. In this model, H. incognitus is just a large H. amphibius.
ere is confusion on the species names Hippopotamus major and H.
antiquus. For a discussion see Hooijer (1942), Blandamura and Azzaroli
(1977), Caloi et al. (1980) and Faure (1985). e informal names “Grand
hippopotame fossile” and “Ippopotamo maggiore” used by Cuvier in 1804
and Nesti in 1820, respectively, are not valid in a nomenclatorial sense, whi-
le Desmarest named H. antiquus in 1822 and Cuvier named H. major in
1824. e name given by Desmarest has thus priority and should be used.
Both names are based on fossils from the Upper Valdarno, stored in the
Muséum National d’Histoire Naturelle (MNHN) in Paris.
Not only the systematics and affinities of the European hippos were
controversial, but also the date of their first arrival. Central in the debate is
the age of the hippo fossils from Valdarno (Italy). Blandamura and Azzaroli
(1977) believed that the fossils originate from the Tasso Faunal Unit and
that their age is about 1.5 Ma, while Mazza (1991) believed their age to be
1.5-1.2 Ma. However, no other hippo occurrences were known from Eu-
rope that are this old, and Faure (1985) considered that these fossils come
from higher up in the local sequence and have an age of close to 1 Ma,
corresponding to the early Middle Pleistocene. At present the beginning of
the Middle Pleistocene is taken at the beginning of the Brunhes at 0.78 Ma.
Currently, the age of the Tasso Faunal Unit is estimated to be 1.8 till about
1.6 Ma (Gliozzi et al., 1997). Since Faure (1985), numerous Early Pleistoce-
ne Hippopotamus fossils have been found in Europe, notably from Coste
San Giacomo (Bellucci et al., 2012; 2014; also referred to as Costa San Gia-
como by Gliozzi et al., 1997), dated to older than 2 Ma.
Considering these new data and the fact that, H. antiquus from Upper
Valdarno has the morphology of the occipital, temporal fossa and orbits
more like in H. amphibius than in H. tiberinus and H. gorgops, a third model
of the systematics and temporal distribution of the European hippos was
proposed (Figure 6) (Van der Made, in press). According to the new propo-
sal, H. antiquus dispersed shortly before 2 Ma from Africa to Italy and went
extinct shortly after 1.8 Ma, while at or shortly before 1.2 Ma H. tiberinus
Jan van der Made et al.
148
dispersed from Africa to Europe and during the late Middle Pleistocene H.
amphibious did so.
With the currently available information, it seems that during the
Pleistocene, both North Africa and Europe were colonized three times by
different hippo species. In the case of Africa these are: Hippopotamus sp.,
H. gorgops/H. sirensis, and H. amphibius. In the case of Europe these are: H.
antiquus, H. tiberinus and H. amphibius.
Figure 5: The chronological distribution of the European Pleistocene species of Hippopotamus
(particularly the oldest localities) and the approximate age of the hippo localities (solid squa-
res indicate presence, open squares indicate uncertainty: cf., aff., sp.?). Ranges and localities
largely after Van der Made et al. (2015); ages of Tor di Quinto, Maglianella, Sant’Oreste very
approximate. Dorsal and lateral views of Hippopotamus skulls: H. antiquus from Upper Valdarno
(MNHN) and Figline (U Valdarno; IGF 1043, photograph from Mazza, 1991); H. tiberinus from
Untermassfeld (IQW 1991/23909 Mei 23438, photographs from Kahlke, 2001) and from Maglia-
nella (CC C601, photograph from Mazza, 1991); H. amphibius from Tor di Quinto (USR).
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
149
4. MATERIAL
Excavations at the site of El Kherba have yielded Hippopotamus fossils, in-
cluding a skull, which is stored at the Musée Public National de Sétif (MPNS),
Sétif in Algeria. In addition to the hippo skulls described and figured in the
literature, the following specimens were used for comparison in this study:
Choeropsis liberiensis: A recent skull from the comparative collections of
the Instituut voor Aardwetenschappen (IVAU) of the University of Utrecht,
Netherlands.
“Trilobophorus” afarensis: Skull AL109/319 at the National Museum of
Ethiopia, Addis Ababa, Ethiopia.
Hippopotamus protamphibius: Skulls 1933-9-777 and 1933-33-768 from
the Omo (Ethiopia) in the Muséum National d’Histoire Naturelle (MNHN),
Paris, France.
Figure 6: Three models of the origin, systematics, and temporal distribution of the European
Hippopotamidae.
Jan van der Made et al.
150
Hippopotamus antiquus: Two skulls from the Upper Valdarno (Italy) on
display in the MNHN, and another one on display at the Museo di Storia
Naturale in Florence, Italy (formerly Istituto di Geologia, IGF).
Hippopotamus gorgops: e holotype skull from Olduvai housed at the Mu-
seum für Naturkunde (MNB) in Berlin, Germany.
Hippopotamus amphibius: A fossil skull from Tor di Quinto in the Univer-
sitá La Sapienza in Rome, Italy (USR) and recent skulls in the Institut Ca-
talà de Paleoecologia Humana i Evolució Social, (IPHES) Tarragona, Spain,
and the Centre National de Recherches Préhistoriques Anthropologiques
et Historiques (CNRPAH), Algiers, Algeria.
Hippopotamus sp.: Bones and teeth from Ain Boucherit in the MNHN, Pa-
ris, France.
5. DESCRIPTION AND COMPARISON
e skull from El Kherba is fairly complete, but even while embedded in
the sediment it was in a bad state (Figure 7/1). e postero-dorsal part of the
skull is preserved and shows several of the features that are of interest here.
e orbits are much elevated with respect to the rest of the frontal bone. is
is well seen in frontal view (Figure 8). If seen from the side, the orbit is well
above the dorsal surface of the skull just before and behind the orbit. is
feature is as in Hippopotamus gorgops and H. tiberinus, while in H. amphibius
and H. antiquus the orbit tends to be lower, though occasionally there are
individuals with a somewhat more elevated orbit (Figures 5 & 8). In more
primitive or older African hippos the orbits tend to be less elevated (Figure 2).
When seen in side view and if the occlusal surface is taken as horizon-
tal, the line along the dorsal surface of the nasals slopes backwards or is
horizontal in H. gorgops, H. tiberinus and in the skull from El Kherba, while
in H. amphibius and many older or primitive hippos it slopes anteriorly
(Figures 5 & 8). is suggests that in H. gorgops and H. tiberinus the nostrils
are more elevated than in H. amphibius. Still in side view, the dorsal profile
of the parietals and occipital rises steeply backwards in H. gorgops, H. tibe-
rinus and in the El Kherba skull (Figures 5 & 8). is profile also tends to be
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
151
concave up in the El Kherba skull. In H. amphibius and many older or more
primitive species, this profile is straight or convex up and does not rise as
much as in the El Kherba skull.
In H. amphibius, the dorsal profiles of the nasals, parietals and occiput
form nearly a straight line which passes through the middle of the orbit.
In H. gorgops, H. tiberinus and the skull from El Kherba, the line along the
dorsal surface of the nasals passes below the orbit and makes a marked
angle with the dorsal profile of the parietals and occiput (Figures 5 & 8).
Figure 7: Hippopotamus gorgops from El Kherba. 1) KH06-G30-149 - skull still in the excavation
(photograph by Z. Harichane). 2) KH08-E34-231 - left lower third molar: a) buccal, b) occlusal,
and c) lingual views.
Jan van der Made et al.
152
e dorsal surface of the nasals and brain case are flexed. In most hippos,
including H. amphibius, the temporal fossa is long, while in H. gorgops, H.
tiberinus and the skull from El Kherba it is short (Figures 2, 4, 5 & 8). is
seems to be related to the forward movement of the occiput.
6. DISCUSSION
6.1. Taxonomic assignment
With the extremely elevated orbits, the skull from El Kherba differs
from the more primitive or older species of hippos, including from the li-
ving Choeropsis and the fossil Hippopotamus protamphibius. e orbits are
also more elevated than in Hippopotamus amphibius. Other hippos with
similarly elevated orbits include Hippopotamus gorgops, Hippopotamus ka-
rumensis and Hexaprotodon palaeindicus (Boisserie, 2005, figure 8). e
latter is a Hexaprotodont Indian species, while H. karumensis is reported to
have molars with lower crowns than H. gorgops. e hippo from El Kherba
has high crowned molars (Figure 7b). It seems justified to assign the fossils
from El Kherba to Hippopotamus gorgops.
Geraads (1980) suggested that H. gorgops could be a junior synonym
of H. sirensis. is suggestion is interesting, but H. sirensis from the type
locality at Tighenif, described and figured by Pomel (1890; 1896), does not
include the relevant skull parts. e excavations by Arambourg have in-
creased the collections from that locality greatly, but we had no access to
the Algerian fossils stored in the MNHN, thus we cannot discuss the pro-
blem here.
e locality of Ain Hanech is located a very short distance from El
Kherba, in the same stratigraphic unit and approximately at the same
topographic height, the bedding being more or less horizontal. ere-
fore the age is the same. Arambourg (1979) mentioned or described and
figured hippo fossils from Ain Hanech, including two skulls, which he
assigned to Hippopotamus amphibius. If correct, this would be one of the
oldest records of that species. However, the orbits are very elevated, and
the dorsal profile of the nasals slopes backwards (Figure 8). is feature
would rather suggest the presence of H. gorgops, not only in El Kherba,
but also in Ain Hanech.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
153
6.2. Functional morphology
Hippos have an amphibious life style, which is reflected in various
morphological adaptations. Two of these are relevant here.
Firstly, extant Hippopotamuses have eyes which are elevated above
the skull. is is a common adaptation of animals that live at or close to
Figure 8: Morphological features that distinguish Hippopotamus amphibius and Hippopotamus
gorgops (shown on the type skull from Olduvai in the MNB) and, as well as the state in skull
KH06-G30-149 from El Kherba. The skull of H. amphibius is a recent skull in the IPHES and for
H. gorgops the holotype skull from Olduvai in the MNB and NHM M14957 also from Olduvai
(photograph from Mazza, 1991) are shown.
Jan van der Made et al.
154
the water surface, like frogs and crocodiles, but also in fishes that live
near the bottom, like flounder and sole. e elevated eyes allow the Hi-
ppopotamus to see above the water, while being largely submerged (Figu-
re 9-1) and sole to see when hidden in the sand at the bottom. In the skull
from El Kherba, this adaptation is seen in the orbits which are elevated
above the dorsal surface of the skull. In hippos and anthracotheres, eleva-
ted orbits are considered to be an adaptation to an amphibious life style.
e feature of elevated orbits has been used to link Anthracotheriidae to
Hippopotamidae (Colbert, 1935), but today this is understood to be due
to convergence (Boisserie, 2005). Within Hippopotamidae, this feature
evolved in parallel in the south Asian Hexaprotodon and in Hippopota-
mus, but if current phylogenetic models are correct, H. karumensis and
H. gorgops also acquired extremely elevated orbits in parallel. e more
elevated orbits in H. gorgops are generally interpreted as a more extreme
aquatic adaptation than in the living species of hippo.
Secondly, living hippos have nostrils and ears that can be closed while
diving (Nowak, 1991). e nostrils are on top of the nose and, like the eyes,
may be above water while most of the body and head is submerged (Figure
9-1). e position of the nostrils cannot be judged from the shape of the
nasals. However, the anteriorly rising nasals in H. gorgops, suggest that the
nostrils were more elevated than in the living species of Hippopotamus. It
seems plausible that the greater degree of elevation of the anterior part of
the nose in H. gorgops is another aquatic adaptation that is more developed
than in H. amphibius.
Above, it has been observed that the temporal fossa of H. gorgops is
shorter than in H. amphibius and, as a result, there is less distance between
the very elevated orbits and occiput. It should be possible to establish whe-
ther this resulted from a simple forward displacement of the occiput or
from a forward rotation of the brain case. Whatever the cause, the result is
that, the temporal fossa is shorter and the posterior most place of attach-
ment of the temporal muscles moved forwards. Hippopotami are known
to open their jaws up to 150º in antagonistic display (Nowak, 1991). An
extreme opening of the jaws could be facilitated by the change in skull ar-
chitecture in H. gorgops. ese morphological differences are clearly visible
in the fossils. However, most probably they would not have resulted in a
very different aspect in a living individual (see tentative reconstruction in
Figure 9-3).
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
155
6.3. The Hippopotamus record from sub-Saharan and North Africa,
and from Europe compared
Unlike sub-Saharan Africa, where there was a high diversity of hippos
(Figure 2), North Africa (Figure 4) and Europe (Figure 5) may have had
just one species of Hippopotamus at a time. As observed above, European
hippos are likely to have come from or through North Africa, and being
amphibious either crossed the Straits of Gibraltar or Sicily, or came around
the eastern end of the Mediterranean. us, it would not be surprising if
both areas shared the same species.
Hippopotamus antiquus may have arrived in Europe shortly before 2
Ma (Figure 5). It was a large hippo with relatively low orbits and occiput
and a short temporal fossa. e European hippo differed from H. gorgops
and H. karumensis in having less elevated orbits and occiput, but the or-
bits are clearly more elevated than in H. protamphibius, and it is much
larger than H. aethiopicus. Of the sub-Saharan species (Figure 2), the one
which is most similar to H. antiquus is H. amphibius and probably, becau-
se of its phylogenetic position, not the well known H. kaisensis. Hippo-
potamus amphibius seems to have originated from H. kaisensis after the
arrival of H. antiquus in Italy. e Hippopotamus from Ain Boucherit is
close in age to the earliest European H. antiquus, but its skull shape is not
known. It is possible that H. kaisensis: 1) dispersed towards North Africa;
2) is the hippo documented in Ichkeul, Ain el Bey and Ain Boucherit; and
3) dispersed to Italy before 2 Ma, where it went extinct shortly after 1.8
Ma (Figure 10).
Figure 9: Hippopotamus amphibius in Bioparco, the zoological garden of Rome (1) and in the
zoo of Zürich (2). Reconstruction of the appearance of Hippopotamus gorgops by: elevating the
eyes, occiput and ears; shortening the distance between occiput and ears and the eyes; and a
clockwise rotation of the dorsal surface of the nasals.
Jan van der Made et al.
156
e European Hippopotamus tiberinus appeared some time before 1.2
Ma (see discussion on the ages of the localities by Van der Made et al., In
Press) and lived there well into the Middle Pleistocene (Figure 5). It is a
very large hippo with elevated orbits and occiput, a short temporal fossa
and nasals that anteriorly slope upwards. ese features are shared with the
sub-Saharan H. gorgops, which appeared at or shortly after 2 Ma ago and
survived till well into the Middle Pleistocene (Figure 4). e North African
hippo fossils with ages between roughly 2 and 0.5 Ma include the type ma-
terial of H. sirensis and other specimens which have tentatively been assig-
ned to H. sirensis, but also the skull from El Kherba, which is here assigned
to H. gorgops. ese two species might be synonymous (Geraads, 1980). In
any case, on the basis of the skull from El Kherba, it is possible that H. gor-
gops dispersed shortly after 2 Ma to North Africa and much later, around
1.2 Ma, to Europe (Figure 10).
e latest record of H. tiberinus is controversial. Mazza (1991, plate 2,
figure 2) assigned a skull fragment from the Rhine gravels at Eich to H. tibe-
rinus. But this skull has a convex up lateral profile of the parietal and occipi-
tal, as in H. amphibius. e fossils from these deposits are considered to be
Eemian in age (early Late Pleistocene or about stage 5) (Von Koenigswald &
Heinrich, 1999). Mazza and Bertini (2013, figure 1) indicated the age of the
Rhine gravels at Eich as Late Pleistocene, but did not even discuss the hippo
fossils from there. Instead, these authors (figure 2) marked the last occurren-
ce of “H. gr. H. antiquus (= H. tiberinus)” as being in Isotope Stage 7 based on
materials from the locality of Castel di Guido. Previously, the age of Castel di
Guido was considered to be stage 9 by Mazza (1995), who also argued that “a
possible attribution to H. tiberinus should not be ruled out”. Despite this sug-
gestion, these fossils are too poor for a secure attribution. Mazza (1995) also
considered H. tiberinus to be present in Mosbach-2 and Jockgrim (500-600
ka, Germany). Pandolfi and Petronio (2015) considered the latest H. tiberinus
to be from Maglianella (Italy) and another locality correlated to Stage 15. Sta-
ge 15 or 500 ka may well be the age of the latest occurrence of the species.
e first record of H. amphibius in Europe is also controversial. Mazza
and Bertini (2013, figure 2) indicated the first appearance of H. amphibius
to be in Stage 5, and in particular in Barrington (UK). is is the type loca-
lity of H. incognitus and is considered to be Late Pleistocene in age (Ipswi-
chian or about Stage 5; Faure, 1985; Stuart, 1982). However, Pandolfi and
Petronio (2015) considered the first record of H. amphibius to be in Cava
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
157
Nera Molinario and Fontana Ranuccio (both in Italy), correlated to Stage
11 or 13. Caloi et al, (1980) suggested a skull from Tor di Quinto (Italy)
might be from Cave Nera Molinario. In this quarry, there are various units
of different ages, the youngest being 432-422 ka (Di Stefano & Petronio,
1993). Mazza (1995) considered the age of this skull unknown, and Mazza
and Bertini (2013), have not even discussed it. e skull has a dorsal profile
as in H. amphibius (Figure 5) and may well be the oldest European record
of this species, though its age is still problematic. e replacement of H.
tiberinus by H. amphibius may have happened sometime between 500 and
420 ka. In North Africa, the timing of the replacement is more imprecisely
known. H. amphibius arose much earlier in sub-Saharan Africa and seems
to have dispersed during the late Middle Pleistocene to North Africa and
subsequently to Europe (Figure 10).
It is peculiar that H. gorgops, or at least hippos with a similar mor-
phology, have been replaced more or less simultaneously in Europe, Nor-
th Africa and sub-Saharan Africa by H. amphibius, which remained as the
dominant or only hippo (Figure 10). It seems difficult to explain this with
climatic or environmental change. Possibly H. amphibius evolved a feature,
which gave it an edge over H. gorgops, possibly it is the robusticity of its
limb bones. H. amphibius (or H. incognitus) tends to have very short limb
bones (Faure, 1985). It is not clear when this feature evolved in Africa. As
yet, differences in the robusticity of the bones have not been studied rela-
tive to this question, and there are only a few localities that have yielded
relevant skull materials.
e living H. amphibius is smaller than the fossil H. amphibius from
Europe, but it is not known when the size decrease occurred in Africa. e
European hippos might be recognized as a different subspecies as H. a.
incognitus. However, it is possible that the North African H. icosiensis is
identical to H. a. incognitus, and thus a senior synonym.
e last European Hippopotamus may have been from level G at Gro-
tta Romanelli (Italy) and level C2 at Canale delle Acque Alte (Italy), corres-
ponding to Isotope Stage 4 or even Stage 3 (Pandolfi & Petronio, 2015).
Early Holocene (11-8 ka) hippo fossils are known from the southern half of
what is today the Sahara, and even from the Nile delta (Drake et al., 2011).
Rock art suggests even a presence in these areas after 4000 BP (Le Quellec,
1999). At present, the geographical distribution of H. amphibius is restric-
ted to sub-Saharan Africa.
Jan van der Made et al.
158
6.4. Hippos and human dispersal
at the dispersal of Hippopotamus into Europe is related to human
dispersal has been suggested by various authors. An elaborate description
is provided by Martínez Navarro (2010). He proposed that, humans, H. an-
tiquus and other species including eropithecus, dispersed into Europe
at the Plio-Pleistocene transition (at about 1.8 Ma) which he argued was
favoured by the northward expansion of African mixed habitats savannas
and gallery forest. He also proposed that hippos could have been easily
Figure 10: The chronological distribution of the Pleistocene species of Hippopotamus and the
approximate age of the hippo localities (solid squares indicate presence, open squares indicate
uncertainty: cf., aff., sp.?) in sub-Sharan Africa, North Africa, the Levantine Corridor and Europe.
Arrows indicate possible dispersal events.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
159
hunted from the shores by throwing stones on them. However, there are
some flaws in this model. First of all, hippos are not defenceless animals
that can be easily killed. Hippos are among the most dangerous animals in
Africa, killing hundreds of humans each year (Treves & Naughton-Treves,
1999; Le Bel et al., 2011; Chomba et al., 2012). Actually, the estimated num-
ber of humans killed annually by hippo vary from 3001, up to nearly 30002.
A hippo is reported to have killed 13 persons in a single attack3. us, it
does not seem likely that hippos, with a possible body weight of some four
tonnes, were easy to kill by throwing stones from the bank of a river. Ano-
ther flaw is that all of these species did not disperse to Europe at the same
time. At the time Martínez Navarro (2010) wrote this, some vertebras from
Pirro Nord (Italy) were identified as eropithecus (Rook et al., 2004; Rook
& Martínez-Navarro, 2013), but now these fossils are recognized to belong
to a porcupine (Alba et al., 2014). As a result the fossils from Cueva Victoria
(Spain) remain the only known European eropithecus (Gibert et al., 1995;
Ferràndez-Cañadell et al., 2014). is locality is believed to be a little older
than 0.78 Ma (Gibert & Scott, 2014). Human dispersal into Western Europe
is much older than 0.78 Ma, though far less than 1.8 Ma. Given the current
evidence, the dispersal of H. tiberinus into Europe is probably a little older
than that of Homo, but the appearance of H. antiquus in Europe is certainly
much earlier. e dispersals of all of these species and humans did not oc-
cur in a single, but in many events widely separated in time. Finally, there is
no evidence for the expansion of African habitats to Europe.
e dispersal of H. tiberinus to Europe shortly before 1.2 Ma seems
to be close in age to the arrival of the first humans there, but no likely
causal link has been identified. H. tiberinus may have arrived a little ear-
lier and its dispersal may have been favoured by different environmental
conditions. Instead, the dispersal of hippos across the Sahara towards
North Africa could prove to be related to periods of increased humidify,
i.e. when lakes and rivers were much larger. ough human and hippo
environmental requirements and ecology are fundamentally different, in-
creased humidity may have also favoured the dispersal of humans across
the Sahara.
1 http://www.animaldanger.com/most-dangerous-animals.php
2 https://www.youtube.com/watch?v=cvoEj1_yOrU
3 http://www.abc.net.au/news/2014-11-20/hippopotamus-attack-kills-13-in-boat-in-ni-
ger/5904646
Jan van der Made et al.
160
e distribution of the two living species of hippo, and especially the
common hippo, seems to depend on the vicinity of a permanent water body.
While the pygmy hippo Choeropsis liberiensis lives in forested environments
and eats a variety of food, including fruit, the common hippo H. amphibius is
a grazer that lives in open habitats close to a lake or river. It spends the day in
or near the water and submerges in water to regulate its body temperature,
and during the night it forages at a distance of up to some 3 km from the water
(Nowak, 1991). is implies that it needs a permanent body of water with a
minimum depth of about 1.5 metre for survival. Judging from skull morpho-
logy, all of the species of the genera Hippopotamus and Hexaprotodon seem to
be more adapted to an amphibious life style than Choeropsis, and probably all
had similar environmental requirements as the common living hippo. In order
to reach North Africa and Europe, hippos needed rivers or lakes to cross the
Sahara and to disperse along the north coast.
ere are indications that large rivers were flowing across the Sahara
during the Eemian and the Early Holocene (Drake et al., 2011; Coulthard et
al., 2013; Skonieczny et al., 2015). ere may have been rearrangements of
watersheds by river capture, allowing aquatic and amphibious fauna to enter
extensive new river basins. In addition, there are Holocene fossils and rock art
depicting hippos in several parts of the Sahara, dating from before and after
4 ka (Le Quellec, 1999). Large rivers and lakes and northward expansion of
hippos may have occurred during some of the earlier interglacials, but may-
be not during all of them. e extension of lakes has been studied in a long
sedimentary sequence in the Rift Valley (Trauth et al., 2005), where extensive
lakes occurred close to 2.5 Ma, and between about 1.9 and 1.7 Ma. e occu-
rrence of extensive lakes in the Rift is believed to be related to global clima-
tic events, such as the increasing importance of the 40 ka Milankovich cycle
around 2.5 Ma and the development of the Walker Circulation from about
1.9 to 1.7 Ma. If global climate affected the extension of lakes in East Africa, it
may have also been the case in North Africa. Extensive lakes and associated
rivers are the preferred habitats of hippos and may have allowed H. kaisensis
(around 2.5 Ma?) and later H. gorgops (around 1.9 Ma?) to cross the Sahara.
Based on the above we could infer that hippos may have lived in the area of
the present day Sahara during parts of at least some of the climatic cycles and
may have spread in this way to the North. e subsequent dispersal of hippos
from North Africa to Europe depended probably on quite different condi-
tions, though we do not know exactly which conditions these were.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
161
Human north ward expansion, as well as other faunal exchange across
the Sahara, may have been permitted by the more humid climate and ex-
tensive lakes and rivers, which allowed hippos to disperse northward. Ba-
sed on current evidence it seems unlikely for early humans to have cros-
sed the Straits of Gibraltar or Sicily (Villa, 2001; Van der Made, 2005b).
In addition, the human fossils or indications of human presence in Sou-
th-West and Southern Asia are older than in Europe (e.g. Dennell et al.,
1988; Bar-Yosef & Goren-Inbar, 1993; Gabunia & Vekua, 1995; Larick et
al., 2001; Dennell & Roebroeks, 2005) (Gibert et al., 1999; Carbonell et
al., 2008; Toro-Moyano et al., 2013). erefore, it seems more likely that
humans dispersed out of Africa through the Middle East (e.g. Sirakov et
al., 2010; Van der Made, 2011; 2013; Bar-Yosef & Bellmaker, 2011). Primi-
tive features of the humans in Dmanisi and Flores (Martinón-Torres et al.,
2007; Lordkipanidze et al., 2007; Argue et al., 2009; Jungers et al., 2009)
suggest the possibility of a dispersal from sub-Saharan Africa into Eurasia
anterior to that of Homo erectus, which means one dispersal before and
one around 1.8 Ma. It is of interest to note here that, human remains from
Rabat and Tighennif, for which the name Homo mauritanicus is availa-
ble, seem to be more primitive than H. erectus (Martinón-Torres et al.,
2007). e human crossing of the Sahara might correspond to two north
ward dispersal events. e first one before 1.8 and possibly around 2.5 Ma,
when the hippo species present in Ain Boucherit reached the Maghreb
and gave rise to H. antiquus of Europe. At the same time a primitive spe-
cies of Homo dispersed to the Maghreb, giving rise to H. mauritanicus,
and to the south of Asia, giving rise to the species present in Dmanisi and
Flores. e second event around 1.8 Ma, when H. gorgops reached North
Africa and H. erectus dispersed into southern Asia.
7. CONCLUSIONS
e following conclusions are drawn:
1) H. gorgops is present in El Kherba. It dispersed around 1.9 Ma
into North Africa when lakes and rivers were more extensive.
is may have coincided with the dispersal of H. erectus out of
Africa.
Jan van der Made et al.
162
2) H. gorgops gave rise to, or is identical with, the European H. tibe-
rinus around 1.2-1.4 Ma.
3) Before 2 Ma, possibly around 2.5 Ma, a species of hippo, perhaps H.
kaisensis, dispersed to North Africa.
4) e species H. antiquus may have originated from H. kaisensis or a
similar form in North Africa when it dispersed to Italy shortly befo-
re 2 Ma.
5) H. amphibius dispersed during the Middle Pleistocene to North
Africa and subsequently to Europe.
6) e dispersals of Hippopotamus to North Africa may have been
possible due to the presence of extensive lakes and rivers in the area
which presently is the Sahara. e same environmental conditions
may have allowed human North-ward dispersal.
e following questions remain open:
• It is possible that Hippopotamus sirensis, H. gorgops, H. georgicus
and H. tiberinus are synonyms.
• It is also possible that H. antiquus and H. kaisensis are synonyms.
• To document the evolution (and timing) of shorter limb bones in
African H. amphibius.
• It is possible that H. amphibius incognitus and H. icosiensis may be
synonyms for the early stage (large) of H. amphibius.
• To document the decrease in size (and timing) leading to the living
H. amphibius amphibius in Africa.
ACKNOWLEDGEMENTS
e following persons and institutions provided access to the spe-
cimens studied: C. Argot, G. Billet, P. Brewer, E. Cioppi, M. Esteban, T.
Getachaw, O. Hampe, N. Ibañez, M.R. Palombo, P.Y. Sondaar, CNRPAH,
Musée National d’Archéologie de Sétif, Musée National Public de Cirta
de Constantine. V. González Gascón helped us with cartography. e fo-
llowing persons allowed us to use photographs of fossils from their arti-
cles in our figures: R.D. Kahlke, P. Mazza. Financial support which made
this study possible was received for research projects: HAR2013-41351-P
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
163
(MINECO, Madrid) and CGL2012-38434-C03-02 (MICINN, Madrid);
Synthesys grants DE-TAF-668 and GB-TAF-4119; A1/035657/11 (AE-
CID, Madrid); CNRPAH (Algeria); Marie Curie PCIG09-GA2011-293581
(Brussels), National Science Foundation (grants # BCS-0517984), L.S.B.
Leakey Foundation, Wenner-Gren Foundation, National Geographic So-
ciety, Stone Age Institute (USA).
REFERENCES
Alba D. M., Colombero S., Delfino M., Martínez-Navarro Pavia M., Rook L., 2014. A thorny
question: e taxonomic identity of the Pirro Nord cervical vertebrae revisited. Journal
of Human Evolution, 76, 92-106.
Arambourg C., 1970. Les Vertébrés du Pléistocène de l’Afrique du Nord. Archives du Mu-
séum National d’Histoire Naturelle, 10.
Arambourg C., 1979. Vertébrés Villafranchiens d’Afrique du Nord (Artiodactyles, Carnivo-
res, Primates, Reptiles, Oiseaux). Fondation Singer-Polignac, Paris.
Argue D., Morwood M. J., Sutikna T., Jatmiko, Saptomo E.W., 2009. Homo floresiensis: A
cladistic analysis. Journal of Human Evolution, 57, 623-639.
Bar-Yosef O., Goren-Inbar N., 1993. e Lithic Assemblage of Ubeidiya. A Lower Palaeoli-
thic Site in the Jordan Valley. Institute of Archaeology, Hebrew University of Jerusalem,
Jerusalem.
Bar-Yosef O., Belmaker M., 2011. Biogeography and climatic change as a context to human
dispersal out of Africa and within Eurasia. Quaternary Science Reviews, 30, 1318-1337.
Bellucci L., Mazzini I., Scardia G., Bruni L., Parenti F., Segre A. G., Segre Naldini E., Sardella
R., 2012. e site of Coste San Giacomo (Early Pleistocene, central Italy): Palaeoenviron-
mental analysis and biochronological overview. Quaternary International, 267, 30-39.
Bellucci L., Bona F., Corrado P., Magri D., Mazzini I., Parenti F., Scardia G., Sardella R., 2014.
Evidence of late Gelasian dispersal of African fauna at Coste San Giacomo (Anagni Ba-
sin, central Italy): Early Pleistocene environments and the background of early human
occupation in Europe. Quaternary Science Reviews, 96, 72-85.
Blandamura F., Azzaroli A. 1977. L”Ippopotamo maggiore” di Filipo Nesti. Atti della Acca-
demia Nazionale dei Lincei. Memorie, Classe die Scienze fisiche, matematiche e natura-
li, serie 8, vol. 14, sezione 2, no. 5, pp. 169-188.
Boisserie J. R., 2005. e phylogeny and taxonomy of Hippopotamidae (Mammalia: Artio-
dactyla): A review based on morphology and cladistic analysis. Zoological Journal of the
Linnean Society, 143, 1-26.
Boisserie J. R., Lihoreau F., Brunet M., 2005a. Origins of Hippopotamidae (Mammalia, Ce-
tartiodactyla): Towards resolution. Zoologica Scripta, 34 (2), 119-143.
Boisserie J. R., Lihoreau F., Brunet M., 2005b. e position of Hippopotamidae within Ce-
tartiodactyla. Proceedings of the National Academy of Science, 102 (5), 1537-1541.
Caloi L., Palombo M. R., Petronio C., 1980. Resti cranici di Hippopotamus antiquus (= H.
major) e Hippopotamus amphibius conservati nel Museo di Paleontologia dell’università
di Roma. Geologica Romana, 19, 91-119.
Jan van der Made et al.
164
Carbonell E., Bermúdez de Castro J. M., Parés J.M., Pérez-González A., Ollé A., Mosquera M.,
Cuenca-Bescós G., García N., Granger D. E., Huguet R., Van der Made J., Martinón-To-
rres M., Rodríguez X. P., Rosas A., Sala R., Stock G. M., Vallverdú J., Vergès J. M., Allué E.,
Burjachs F., Cáceres I., Canals A., Benito A., Díez C., Lozano M., Mateos A., Navazo M.,
Rodríguez J., Rosell J., Arsuaga J.L., 2008. e first hominin of Europe. Nature, 452, 465-469.
Chaid-Saoudi Y., Geraads D., Raynal J. P., 2006. e fauna and associated artefacts from the
Lower Pleistocene site of Mansourah (Constantine, Algeria). Comptes Rendus Palevol,
5, 963-971.
Chomba C., Senzota R., Chabwela H., Mwitwa J., Nyirenda V., 2012. Patterns of human
wildlife conflicts in Zambia, causes, consequences and management responses. Journal
of Ecology and the Natural Environment, 4 (12), 303-313.
Colbert E. H., 1935. Distributional and phylogenetic studies on Indian fossil mammals. IV
e Phylogeny of the Indian Suidae and the origin of the Hippopotamidae. American
Museum Novitates, 799, 1-24.
Colbert E. H., 1938. Fossil mammals from Burma in the American Museum of Natural His-
tory. Bulletin of the American Museum of Natural History, 74, 255-436.
Coryndon S. H., 1977. e taxonomy and nomenclature of the Hippopotamidae (Mam-
malia, Artiodactyla) and a description of two new fossil species. Proceedings of the Ko-
ninklijke Akademie van Wetenschappen, Series B, 80 (2), 61-88.
Coryndon S. H., 1978. Hippopotamidae. In: Maglio V. J. & Cooke H.B.S. (Ed.), Evolution of
African Mammals. Harvard University Press, Cambridge & London, pp. 483-495.
Coulthard T. J., Ramirez J. A., Barton N., Rogerson M., Brücher T., 2013. Were rivers flowing
across the Sahara during the Last Interglacial? Implications for human migration throu-
gh Africa. Plos One, 8 (9), 1-12.
Dennell R.W., Rendell H. R., Hailwood E., 1988. Early tool-making in Asia: Two-million-
year-old artefacts in Pakistan. Antiquity, 62, 98-106.
Dennell R. W., Roebroeks W., 2005. An Asian perspective on early human dispersal from
Africa. Nature, 438, 1099-1104.
Di Stefano G., Petronio C., 1993. A new Cervus elaphus subspecies of Middle Pleistocene
age. Neues Jahrbuch für Geologie und Paläontologie, Abhandlngen, 190 (1), 1-18.
Drake N. A., Blench R. M., Armitage S. J., Bristow C. S., White K. H., 2011. Ancient water-
courses and biogeography of the Sahara explain the peopling of the desert. Proceedings
of the National Academy of Science, 108 (2), 458-462.
Dubois E., 1908. Das geologische Alter der Kendeng oder Trinil Fauna. Tijdschrift van het
Nederlandsch Aardrijkskundig Genootschap, series 2, 25, 1235-1270.
Falconer H., Cautley P. T., 1836. Note on the fossil hippopotamus of the Siwalik Hills. Asiatic
Research Calcutta, 19 (3), 3953.
Faure M., 1984. Hippopotamus incognitus nov. sp., un hippopotame (Mammalia, Artiodac-
tyla) du Pléistocène d’Europe occidentale. Geobios, 17 (4), 427-434.
Faure M., 1985. Les hippopotames Quaternaires non-insulaires d’Europe occidentale. Nou-
velles Archives du Muséum d’Histoire naturelle de Lyon, 23, 13-79.
Faure M., 1986. Les hippopotamidés du Pléistocène ancien d’Oubeidiyeh (Israël). Mémoires
et Travaux du Centre de Recherches français de Jérusalem, 5, 107–142.
Faure M., Guérin C. 1992. La grande faune d’Europe occidentale au Pléistocène moyen et
supérieur et ses potentialités d’information en préhistoire. Mémoires de la Société Géo-
logique de France, 160, 77-84.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
165
Ferràndez-Cañadell C., Ribot F., Gibert Ll. 2014. New fossil teeth of eropithecus oswaldi
(Cercopithecoidea) from the Early Pleistocene at Cueva Victoria (SE Spain). Journal of
Human Evolution, 74, 55-66.
Gabunia L., Vekua A., 1995. A Plio-Pleistocene hominid from Dmanisi, east Georgia, Cau-
casus. Nature, 373, 509-512.
Gaudry A., 1876. Sur un Hippopotame fossile découvert à Bone (Algérie). Bulletin de la
Société Géologique de France, 3 (4), 147–154.
Gentry A. W., 1999. A fossil hippopotamus from the Emirate of Abu Dhabi, United Arab
Emirates. In: Whybrow P. J. Hill A. (Ed.), Fossil vertebrates of Arabia. Yale University
Press, New Haven, pp. 271–289.
Gentry A.W., Hooker J. J., 1988. e phylogeny of the Artiodactyla. In: Benton M. J. (Ed.),
e phylogeny and classification of the tetrapods 2: Mammals. Clarendon Press, Oxford,
pp. 235-272.
Geraads D., 1980. La faune des sites à ‘Homo erectus’ des carrières omas (Casablanca,
Maroc). Quaternaria, 22, 65-94.
Geraads D., 2002. Plio-Pleistocene mammalian biostratigraphy of Atlantic Morocco. Qua-
ternaire, 13 (1), 43-53.
Geraads D., Amant F., Hublin J. J., 1992. Le gisement pléistocène moyen de l’Aïn Maarouf
près d’El Hajeb, Maroc: Présence d’un hominidé. Comptes Rendus de l’Académie des
Sciences, Paris, 314, Série II, 319-323.
Gibert J., Ribot F., Gibert Ll., Leakey M., Arribas A., Martínez B., 1995. Presence of the
cercopithecid genus eropithecus in Cueva Victoria (Murcia, Spain). Journal of Human
Evolution, 28, 487-493.
Gibert J., Gibert Ll., Albadalejo S., Ribot F., Sánchez F., Gibert P., 1999. Molar tooth fragment
BL5-O: e oldest human remain found in the Plio-Pleistocene of Orce (Granada pro-
vince, Spain). Human Evolution, 14, 3-19.
Gibert Ll., Scott G.E., 2014. Edad del yacimiento de Cueva Victoria y su relación con otros
yacimientos de la Península Ibérica. Mastia, 11-13, 85-100.
Gliozzi E., Abbazzi L., Argenti P., Azzaroli A., Caloi L., Capasso Barbato L., Di Stefano G.,
Esu D., Ficcarelli G., Girotti O., Kotsakis T., Masini F., Mazza P., Mezzabotta C., Palombo
M.R., Petronio C., Rook L., Sala B., Sardella R., Zanalda E., Torre D., 1997. Biochronolo-
gy of selected mammals, molluscs and ostracods from the Middle Pliocene to the Late
Pleistocene in Italy: e state of the art. Rivista Italiana di Paleontologia e Stratigrafia,
103 (3), 369-388.
Guérin C., 1996. Famille des Hippopotamidae. In: Guérin C. & Patou-Mathis M. (Ed.), Les
Grands Mammifères Plio-Pléistocènes d’Europe. Masson, Paris, pp. 44-47.
Hadjouis D., 1990. Megaceroides algericus (Lydekker, 1890), du gisement des Phacochères
(Alger, Algérie). Étude critique de la position systématique de Megaceroides. Quater-
naire, 1 (3-4), 247-258.
Hooijer D. A., 1942. On the nomenclature of some fossil hippopotami. Archives Néerlanda-
sies de Zoologie, 6 (2-3), 279-282.
Hooijer D. A., 1950. e fossil Hippopotamidae of Asia with notes on the recent species.
Zoologische Verhandelingen, 8, 1-124.
Jungers W. L., Harcourt-Smith W. E. H., Wunderlich R.E., Tocheri M. W., Larson S. G., Su-
tikna T., Rhokus Awe Due, Morwood M. J., 2009. e foot of Homo floresiensis. Nature,
459, 81-84.
Jan van der Made et al.
166
Kahlke R. D., 1997. Die Hippopotamus-Reste aus dem Unterpleistozän von Untermassfeld.
In: Kahlke R. D. (Ed.), Das Pleistozän von Untermassfeld bei Meinigen (üringen), Teil
1. Römisch-Germanisches Zentralmuseum Forschungsinstitut für Vor- und Frühges-
chichte. Monographiën, 40 (1), pp. 277-374.
Kahlke R. D., 2001. Schädelsreste von Hippopotamus aus dem Unterpleistozän von Un-
termassfeld. In: Kahlke R. D. (Ed.), Das Pleistozän von Untermassfeld bei Meiningen
(üringen). Teil 2. Römisch-Germanisches Zentralmuseum Forschungsinstitut für
Vor- und Frühgeschichte. Monographiën, 40 (2), pp. 483-500.
Kahlke R. D., 2006. Untermassfeld. A Late EarlyPleistocene (Epivillafranchian) fossil site
near Meiningen (uringia, Germany) and its position in the development of the Euro-
pean mammal fauna. British Archaeological Reports, International Series, 1578.
Koenigswald W. von, Heinrich W. D., 1999. Mittelpleistozäne Säugetierfaunen aus Mitteleu-
ropa - der Versuch einer biostratigraphischen Zuordnung. Kaupia, 9, 53-112.
Kurtén B., 1968. Pleistocene mammals of Europe. Weidenfeld & Nicolson, London.
Larick R., Ciochon R. L., Zaim Y., Sudijono, Suminto, Rizal Y., Aziz F., Reagan M., Heizler
M., 2001. Early Pleistocene 40Ar/39Ar ages for Bapang Formation hominins, Central
Java, Indonesia. Proceedings of the National Academy of Sciences, 98 (9), 4866-4871.
Le Bel S., Murwira A., Mukamuri B., Czudek R., Taylor R., La Grange M., 2011. Human
wildlife conflicts in Southern Africa: Riding the whirl wind in Mozambique and in Zim-
babwe. In: López-Pujol J. (Ed.), e Importance of biological interactions in the study of
biodiversity. In Tech, Rijeka, pp. 283-322.
Le Quellec J. L., 1999. Répartition de la grande faune sauvage dans le nord de l’Afrique du-
rant l’Holocène. L’Anthropologie, 103/l, 161-176.
Lihoreau F., Boisserie J.-R., Manthi F. K., Ducrocq S., 2015. Hippos stem from the longest
sequence of terrestrial cetartiodactyl evolution in Africa. Nature Communications, 6
(6264), 1-8.
Lordkipanidze D., Jashashvili T., Vekua A., Ponce de León M. S., Zollikofer C. P. E., Rightmi-
re G. P., Pontzer H., Ferring R., Oms O., Tappen M., Bukhsianidze M., Agusti J., Kahlke
R., Kiladze G., Martinez-Navarro B., Mouskhelishvili A., Nioradze M., Rook L., 2007.
Postcranial evidence from early Homo from Dmanisi, Georgia. Nature, 449, 305-310.
Maarel F. H. van der, 1932. Contribution to the knowledge of the fossil mammalian fauna of
Java. Wetenschappelijke Mededelingen, Dienst van den Mijnbouw in Nederlansch-In-
dië, 15.
Made J. van der, 1999. Superfamily Hippopotamoidea. In: Rössner G. & Heissig K. (Ed.), e
Miocene land mammals of Europe. Verlag Dr. Friedrich Pfeil, München, pp. 203-208.
Made J. van der, 2003. Suoidea (pigs) from the hominoid locality of Çandir in Turkey. Cou-
rier Forschungs-Institut Senckenberg, 240, 149-179.
Made J. van der, 2005a. La fauna del Pleistoceno europeo. In: Carbonell E. (Ed.), Homínidos:
Las primeras ocupaciones de los continentes. Ariel, Barcelona, pp. 394-432.
Made J. van der, 2005b. Considerations on dispersals between Africa and Europe across the Strait
of Gibraltar. In: Rodríguez Vidal J., Finlayson C., Giles Pacheco F. (Ed.), Cuaternario Medi-
terraneo y poblamiento de hominidos. AEQUA & Gibraltar Museum, Gibraltar, pp. 91-92.
Made J. van der, 2010. e pigs and “Old World peccaries” (Suidae and Palaeochoeridae,
Suoidea, Artiodactyla) from the Miocene of Sandelzhausen (southern Germany): Phylo-
geny and an updated classification of the Hyotheriinae and Palaeochoeridae. Paläonto-
logische Zeitschrift, 84, 43-121.
Hippopotamus gorgops from El Kherba (Algeria) and the context of its biogeography
167
Made J. van der, 2011. Biogeography and climatic change as a context to human dispersal
out of Africa and within Eurasia. Quaternary Science Reviews, 30, 1353-1367.
Made J. van der, 2013. Faunal exchanges through the Levantine Corridor and human disper-
sal: e paradox of the late dispersal of the Acheulean industry. In: Sahnouni M. (Ed.),
Proceedings of the International Symposium Africa, Cradle of Humanity: Recent Disco-
veries. Travaux du Centre National de Recherches Préhistoriques, Anthropologiques et
Historiques, Nouvelle série, 18, pp. 255-296.
Made, J. van der, 2014a. La evolución de los macromamíferos africanos del Plio-Pleis-
toceno. The Plio-Pleistocene large mammals of Africa - Why they evolved like they
did. In: Domínguez Rodrígo M., Baquedano E. (Ed.), The Craddle of Humankind, La
Cuna de la Humanidad. Museo Arquelógico Regional, Alcalá de Henares & Museo
de la Evolución Humana, Burgos, pp. 308-323 (English), 137-177 (Spanish), 360-362.
Made J. van der, 2014b. e large mammals of the Plio-Pleistocene of Africa: Afrotheria, Pe-
rissodactyla and Artiodactyla I. / Los grandes mamíferos del Plio-Pleistoceno africano:
Afrotheriua, Perissodactyla y Artiodactyla I. In: M. Domínguez Rodrígo, E. Baqueda-
no (Eds), e Craddle of Humankind / La Cuna de la Humanidad. Museo Arquelógico
Regional, Alcalá de Henares & Museo de la Evolución Humana, Burgos, pp. 324-336
(English), 179-215 (Spanish), 362-364.
Made J. van der, Sahnouni M., 2013. Updated Plio-Pleistocene faunal lists for Ain Boucherit,
Ain Hanech, and El Kherba sites, Algeria. In: Sahnouni M. (Ed.), Proceedings of the
International Symposium Africa, Cradle of Humanity: Recent discoveries. Travaux du
Centre National de Recherches Préhistoriques, Anthropologiques et Historiques, Nou-
velle série, 18, pp. 223-242.
Made J. van der, Rosell J., Blasco R., In Press. Faunas from Atapuerca at the Early-Middle
Pleistocene limit: e ungulates from level TD8 in the context of climatic change. Qua-
ternary International, http://dx.doi.org/10.1016/j.quaint.2015.09.009
Martínez-Navarro B., 2010. Early Pleistocene faunas of Eurasia and hominin dispersals. In:
Fleagle, J. G., Shea, J. J., Grine, F. E., Baden, A. L., Leakey, R. E. (Ed.), Out of Africa I: e
first hominin colonization of Eurasia. Springer, Dordrecht, pp. 207-224.
Martinón-Torres M., Bermúdez de Castro J. M., Gómez-Robles A., Arsuaga J. L., Carbonell
E., Lordkipanidze D., Manzi G., Margvelashvili A., 2007. Dental evidence on the homi-
nin dispersals during the Pleistocene. Proceedings of the National Academy of Sciences,
104 (33), 13279-13282.
Mazza P., 1991. Interrelations between Pleistocene hippopotami of Europe and Africa. Bo-
lletino della Società Paleontologica Italiana, 30 (2), 153-186.
Mazza P. 1995. New evidence on the Pleistocene hippopotamuses of Western Europe. Geo-
logica Romana, 31, 61-241.
Mazza P. P. A., A. Bertini, 2013. Were Pleistocene hippopotamuses exposed to climate-dri-
ven body size changes? Boreas, 42, 194-209.
Nowak R. M., 1991. Walker’s Mammals of the World. Fifth Edition. Volume 2. e Johns
Hopkins University Press, Baltimore & London, pp. 643-1629.
Orliac M., Boisserie J. R., MacLatchyd L., Lihoreau F., 2010. Early Miocene hippopo-
tamids (Cetartiodactyla) constrain the phylogenetic and spatiotemporal settings
of hippopotamid origin. Proceedings of the National Academy of Sciences, 107(26),
11871-11876.
Pandolfi L., Petronio C., 2015. A brief review of the occurrences of Pleistocene Hippopota-
mus (Mammalia, Hippopotamidae) in Italy. Geologia Croatica, 68 (3), 313–319.
Jan van der Made et al.
168
Parés J. M., Sahnouni M., Made, J. van der, Pérez-González A., Harichane Z., Derradji A.,
Medig M., 2014. Early human settlements in Northern Africa: Paleomagnetic evidence
from the Ain Hanech Formation (northeastern Algeria). Quaternary Science Reviews,
99 (1), 203-209.
Pickford M., 1983. On the origins of the Hippopotamidae together with a description of two
new species, a new genus and a new subfamily from the Miocene of Kenya. Geobios, 16,
193-217.
Pickford M., 1998. A new genus of Tayassuidae (Mammalia) from the Middle Miocene of
Uganda and Kenya. Annales de Paléontologie, 84 (3-4), 275–285.
Pickford M., 2007. A new suiform (Artiodactyla, Mammalia) from the Early Miocene of East
Africa. Comptes Rendus Palévol, 6, 221-229.
Pickford M., 2008. e myth of the hippo-like anthracothere: e eternal problem of homo-
logy and convergence. Revista Española de Paleontología, 23 (1), 31-90.
Pickford M., 2011. Morotochoerus from Uganda (17.5 Ma) and Kenyapotamus from Kenya
(13-11 Ma): Implications for hippopotamid origins. Estudios Geológicos, 67 (2), 523-540.
Pickford M., 2015. Large ungulates from the basal Oligocene of Oman: 3 - Anthracotherii-
dae. Spanish Journal of Palaeontology, 30 (2), 257-264.
Pomel A., 1890. Sur les Hippopotames fossiles de l’Algérie. Comptes Rendus de l’Académie
des Sciences, Paris, 110, 1112-1116.
Pomel A., 1896. Les Hippopotames. Carte géologique de l’Algérie. Paléontologie Monogra-
phies. Imprimerie P. Fontana, Alger.
Rook L., Martínez-Navarro B., Howell F. C., 2004. Occurrence of eropithecus sp. in the
Late Villafranchian of southern Italy and implication for Early Pleistocene “out of Africa”
dispersals. Journal of Human Evolution, 47, 267-277.
Rook L., Martínez-Navarro B., 2010. Villafranchian: e long story of a Plio-Pleistocene
European large mammal biochronologic unit. Quaternary International, 219, 134–144.
Rook L., Martínez-Navarro B., 2013. e large sized cercopithecoid from Pirro Nord and
the importance of eropithecus in the Early Pleistocene of Europe: Faunal marker for
hominins dispersal outside Africa. Palaeontographica, Abteilung A, 298, 107-112.
Sahnouni M., de Heinzelin J., 1998. e site of Ain Hanech revisited: new investigations at
this Lower Pleistocene site in Northern Algeria. Journal of Archaeological Science, 25,
1083-1101.
Sahnouni M., Made J. van der, 2009. e Oldowan in North Africa within a biochronologi-
cal framework. In: Schick K. & Toth N. (Ed.), e cutting edge: New approaches to the
archaeology of human origins. Stone Age Institute Press, Gosport, pp. 179-210.
Sirakov N., Guadelli J. L., Ivanova S., Sirakova S., Boudadi-Malinge M., Dimitrova I., Fe-
randez Ph., Ferrier C., Guadelli A., Iordanova D., Iordanova N., Kovatcheva M., Kru-
mov I., Leblanc J.C., Miteva V., Porpov V., Spassov R., Taneva S., Tsanova T., 2010.
An ancient continuous human presence in the Balkans and the beginnings of human
settlement in western Eurasia: A Lower Pleistocene example of the Lower Palaeolithic
levels in Kozarknika cave (North-western Bulgaria). Quaternary International, 223-
224, 94-106.
Skonieczny C., Paillou P., Bory A., Bayon G., Biscara L., Crosta X., Eynaud F., Malaizé B.,
Revel M., Aleman N., Barusseau J. P., Vernet R., Lopez S., Grousset F., 2015. African hu-
mid periods triggered the reactivation of a large river system in Western Sahara. Nature
Communications, 6 (8751), 1-6.
Proceedings of the II Meeting
of African Prehistory
Burgos 15-16 April, 2015
Actas de las II Jornadas de Prehistoria Africana
Edited by:
Mohamed Sahnouni
Sileshi Semaw
Joseba Rios Garaizar
Photo Credit - Cover
Rock art painting from «Ti-n-Aressou » rock shelter in Tassili-n-Ajjer (Algeria).
Photo by Malika Hachid
KNM-ER 1813 skull.
Photo from cast by CENIEH
Trihedron from the Acheulean hominin site of Tighennif (ex. Ternine, Algeria).
Photo by Jordi Mestre
ISBN: 978-84-946649-1-5
Depósito legal: AS 1258 - 2017
Edita: Consorcio para la construcción, equipamiento y explotación del
Centro Nacional de Investigación sobre la Evolución Humana, Burgos (Spain)
© Consorcio CENIEH, 2017
Imprime: Grácas Eujoa
Hippopotamus gorgops
from El Kherba (Algeria)
and the context
of its biogeography
Chapter 05
Figure 1
Excavation at El Kherba (photograph J. Mestre).
Ma
0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
4.5
5.5
5.0
H. gorgops
H. gorgops
Hippopotamus amphibius
Hippopotamus kaisensis
Hippopotamus aethiopicus
Hippopotamus karumensis
Hippopotamus protamphibius
H. amphibius
Hexaprotodon? sahabiensis
Hexaprotodon bruneti
Hexaprotodon? hipponensis
H. protamphibius
Hexaprotodon? imaguncula
Hippopotamus? dulu
Hippopotamus? coryndonae
o
bophorus”
e
nsis
Choeropsis liberiensis
Hippopotamus? afarensis
eropsis
“Tril
o
afar
e
A. harvardi
Pleistocene
Pliocene
Early Early Middle
Late Holoc.
L
Mioc.
Saotherium mingoz
Archaeopotamus harvardi
Cho
Figure 2
The chronological distribution of the African Plio-Pleistocene Hippopotamidae (adapted from Van der Made, 2014 and
ultimately based on Weston & Boisserie, 2010). Archaeopotamus harvardi reconstruction by Mauricio Antón; recent
Choeropsis liberiensis (IVAU); “Trilobophorus” afarensis from Hadar (ARCCH AL109/319); Hexaprotodon
protamphibius from Omo (MNHN 1933-9-777); recent Hippotamus amphibius (CNRPAH); Hippopotamus gorgops
holotype from Olduvai (MNB).
Ma
0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
4.5
5.0
5.5
19
18
36
25
15
14
17
16
6
5
24
23
20
19
18
21
26
32
3334
35
11
10
2
9
8
13 12
127
31
29
28 27
26
30
3
8
47
26
14 Akh lk l ki
1Bô
Hexaprotodon? H. georgicus / tiberinus H. amphibius / icosiensis
19 Untermassfeld
18 Colle Curti
26 Ata
p
uerca
6 Upper Valdarno
5 Coste S Giacomo
21 Mauer
15 Libakos / Aliakhmon Q-Profil / Kapetianos
32 Tor di Quinto
20 Mosbach
16 Maglianella
17 Sant’Oreste
2 Lac Ichkeul
3 Ain el Bey
4 Ain Boucherit
26 Tihodaine
28 Pointe Pescade
27 G. des Phacochères
30 Beni Saf
31 Latamne
14
Akh
a
lk
a
l
a
ki
1
Bô
ne
H. antiquus / kaisensis
H.
g
or
g
ops / sirensis 22 Jockgrim
29 Ain Maarouf
H. amphibius / incognitus
25 St. Privat
24 Sainzelles
p
27 Venta Micena / Barranco León / Fuente Nueva 3
32
Tor
di
Quinto
7 Ain Hanech / El Kherba
10 Thomas Quary / Ain Maarouf
11 Ubeidiyah
12 G. Benot Yaakov
13 Evron
gg
8 Manssoura
9 Tighenif
36 Barrington
35 Eich
33 Castel di Guido
34 Fontana Ranuccio
Figure 3
The geographic position of the hippopotamus localities mentioned in the text.
0
i
s
Temporal range
(lines) as in sub-
Saharan Africa
North African
localities
G des Phacochères
0
0.5
H. gorgops / sirens
i
Thomas Q., H. level
Tihodaine
Pointe Pescade
G
.
des
Phacochères
Beni Saf Hippopotamus amphibius (=H. icosiensis) Pointe Pescade
1.0
ibius
Tighenif
Thomas Q., level L
Ain Maarouf Hippopotamus gorgops Tighenif
Hippopotamus gorgops Olduvai
1.5
H. kaisensis
H. amph
Mansoura
Ain Hanech
El Kherba
Hippopotamus gorgops Aïn Hanech
2.0
25
Ain el Bey
Ain Boucherit unit R
Ain Boucherit unit P/Q
El Kherba puits
El
Kherba
Hippopotamus gorgops El Kherba
2
.
5
Lac Ichkeul
Hippopotamus sp. Ain Boucherit P/Q
Figure 4
The approximate chronological position of the North African localities with hippos (solid quares indicates
presence, open squares indicate incertainty: cf., aff., sp., ?), compared to the East African record (thick
lines)
Hippopotamus
sp from Ain Boucherit unit P/Q upper molar (MNHN 195
-
13
-
94);
H gorgops
skulls
lines)
.
Hippopotamus
sp
.
from
Ain
Boucherit
unit
P/Q
upper
molar
(MNHN
195
13
94);
H
.
gorgops
skulls
from El Kherba (KH06-G30-149 ), Ain Hanech (photograph from Arambourg, 1979) and Olduvai
(Photographs from Mazza, 1991); H. amphibius (= H. icosiensis) from Pointe Pescade (from Pomel, 1896).
c
ognitus /
h
ibius
0
Europe
(oldest localities)
H. In
c
amp
h
r
gicus
Mauer
? Tor di Quinto
0.5
Mosbach
H. amphibius Tor di Quinto
Maglianella
Jockgrim
H. tiberinus / geo
r
Untermassfeld
Colle Curti
St. Privat
Sainzelles
Atapuerca TE7+14
Barranco León
Fuente Nueva 3
Venta Micena
Atapuerca TD8
Akhalkalaki
1.0
H. tiberinus Maglianella
Sant’Oreste
Venta
Micena
Upper Valdarno
Libakos
Aliakhmon Q-Profil
Kapetanios 1.5
H. tiberinus Untermassfeld
H. antiquus Figline,
U Valdarno
H.antiquus
Coste S Giacomo
2.0
2.5
H. antiquus U Valdarno
Figure 5
The chronolo
g
ical distribution of the Euro
p
ean Pleistocene s
p
ecies of
H
i
pp
o
p
otamus
(p
articularl
y
the
gpp
pp p
(p y
oldest localities) and the approximate age of the hippo localities (solid squares indicates presence, open
squares indicate uncertainty: cf., aff., sp., ?). Ranges and localities largely after Van der Made et al.
(2015); ages of Tor di Quinto, Maglianella, Sant’Oreste very approximate. Dorsal and lateral views of
Hippopotamus skulls: H. antiquus from Upper Valdarno (MNHN) and Figline (U Valdarno; IGF 1043,
photograph from Mazza, 1991); H. tiberinus from Untermassfeld (IQW 1991/23909 Mei 23438,
photographs from Kahlke, 2001) and from Maglianella (CC C601, photograph from Mazza, 1991); H.
amphibius from Tor di Quinto (USR).
Mazza 1991
EuropeAfric a
Van der Made et al. 2015
Afric a Europe
Faure 1985
EuropeAfric a
s
H. amphibius
H. tiberinus
a
ntiquus
H. gorgops
H. amphibius
H. amphibius
H. amphibius
H. tiberinus
Ma
0
.
antiquus
H. incognitus
H. amphibiu
s
H. gorgops
H.
a
H. gorgops
s
H
. antiquus
0.5
1.5
1.0
H
.
Hippopotamus
H. kaisensi
s
H
2.5
2.0
Figure 6
Three models of the origin, systematics, and temporal distribution of the European Hippopotamidae.
1
2a 2b 2c
5 cm
Figure 7
Hippopotamus gorgops from El Kherba. 1) KH06-G30-149 - skull still in the excavation (photograph by
Z. Harichane). 2) KH08-E34-231 - left lower third molar: a) buccal, b) occlusal, and c) lingual views.
Hippopotamus gorgops
Olduvai
Hippopotamus amphibius
Recent
Hippopotamus gorgops
El Kherba
Much elevated orbit
Much elevated orbit
Gently sloping occiput
Long parieto-occipital profile
Steep occiput
Long parieto-occipital profile
Steep occiput
Generally moderately
elevated orbit
Much elevated
orbit
Much
elevated
orbit
Generally little elevated orbit
Steep
occiput
Long parieto-occipital profile
Much elevated
orbit
Long temporal fossa Short temporal
fossa
Figure 8
Morphological features that distinguish Hippopotamus amphibius and Hippopotamus gorgops (shown on on
the type skull from Olduvai in the MNB) and, as well as the state in skull KH06-G30-149 from El Kherba. The
skull of
H amphibius
is a recent skull in the IPHES and for
H gorgops
the holotype skull from Olduvai in the
skull
of
H
.
amphibius
is
a
recent
skull
in
the
IPHES
and
for
H
.
gorgops
the
holotype
skull
from
Olduvai
in
the
MNB and NHM M14957 also from Olduvai (photograph from Mazza, 1991) are shown.
1
2
3
Figure 9
Hippopotamus amphibius in Bioparco, the zoological garden of Rome (1) and in the zoo of Zürich (2).
Reconstruction of the appearance of Hippopotamus gorgops by: elevating the eyes, occiput and ears;
shortening the distance between occiput and ears and the eyes; and a clockwise rotation of the dorsal
su
rf
ace
o
f
t
h
e
n
asa
l
s.
su ace o t e asa s.
s
/ incognitus
p
hibius /
e
nsis
Subsaharan
Africa
Levantine corridor EuropeNorth Africa
oldest localities
H. amphibiu
s
0
0.5
s
0
0.5
H. am
p
icosic
e
Tihodaine
Pointe Pescade
G. des Phacochères
Beni Saf
Latamne
H. amphibius
Thomas Q., H. level ?
i
s
Mosbach
Mauer
Maglianella
Tor di Quinto
Jockgrim
1.0
H
. tiberinus / georgicu
s
Untermassfeld
Colle Curti
St. Privat
Sainzelles
Atapuerca TE7+14
BLó
Fuente Nueva 3
Atapuerca TD8
Akhalkalaki
1.0
o
rgops / sirensis
H. gorgops / sirensis
Tighenif
G. Benot Yaakov
Evron
Ain Maarouf ?
Thomas Q., level L ?
H. gorgops / sirens
i
H. amphibius
Mauer
1.5
H
B
arranco
L
e
ó
n
Venta Micena
Valdarno Sup
Libakos
Aliakhmon Q-Profil
Kapetanios
1.5
H. g
o
Manssoura
Ain Hanech
El Kherba
Ubeidiyah
H. kaisensis
2.0
H
.antiquus
Valdarno
Sup
.
Coste S Giacomo
2.0
a
isensis?
Ain el Bey
Ain Boucherit R
Ain Boucherit P/Q
El Kherba puits
El
Kherba
Figure 10
The chronological distribution of the Pleistocene species of
Hippopotamus
and the approximate age of the hippo
2.5
H
2.5
H. k
a
Lac Ichkeul
Ain
el
Bey
The
chronological
distribution
of
the
Pleistocene
species
of
Hippopotamus
and
the
approximate
age
of
the
hippo
localities (solid quares indicates presence, open squares indicate incertainty: cf., aff., sp., ?) in Subsharan Africa, North
Africa, the Levantine Corridor and Europe. Arrows indicate possible dispersal events.
