Carnivores (Creodonta and Carnivora) from the basal middle Miocene of Gebel Zelten, Libya, with a note on a large amphicyonid from the middle Miocene of Ngorora, Kenya

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Bulletin of the Tethys Geological Society, Cairo, Volume 5, March 2010, P. 43-54
CARNIVORES (CREODONTA AND CARNIVORA) FROM THE BASAL MIDDLE MIOCENE OF GEBEL ZELTEN, LIBYA,
WITH A NOTE ON A LARGE AMPHICYONID FROM THE MIDDLE MIOCENE OF NGORORA, KENYA
J. Morales*, P. Brewer** and M. Pickford***
*Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, C/ José Gutiérrez Abascal,
2, 28006 Madrid, Spain <mcnm166@mncn.csic.es>
**Natural History Museum, Cromwell Road, London, SW7 5BD, England <p.brewer@nhm.ac.uk>
***Collège de France, and Département Histoire de la Terre, UMR 7202 (CR2P) du CNRS, case postale 38, 8,
rue Buffon, 75005 Paris, France <pickford@mnhn.fr>
ABSTRACT
Undescribed fossils from Gebel Zelten, Libya, belong to the poorly known amphicyonid Afrocyon burolleti, and to a
second indeterminate amphicyonid. The Family Amphicyonidae in Africa, although considerably less diverse than in
Eurasia and North America, nevertheless has a longer fossil record extending from the Early Miocene to the latest Miocene,
and spanning the entire continent. A large amphicyonid from Ngorora, Kenya, is described in order to contribute to the
knowledge of the family. A new species of Creodonta Anasinopa libyca from Gebel Zelten is somewhat smaller than the
type species Anasinopa leakeyi from East Africa, and difffers from it by its more hypocarnivorous dentition.
INTRODUCTION
We here describe and interpret unpublished carnivoran fossils from the basal Middle Miocene strata at Gebel Zelten,
Libya, in the collections of the Natural History Museum, London, collected by the late R.J.G. Savage, and fossils found in
1997 during the International Palaeontology Expedition led by the late R. Daams (Wessels et al., 2003). We include a
description and interpretation of an amphicyonid tooth from the late Middle Miocene deposits of the Ngorora Formation,
Kenya.
This material, although consisting of only a few specimens, is important as it throws light on the basal Middle
Miocene carnivoran fauna of Libya. Most of the previously published fossils were in a rather poor state of preservation,
especially the amphicyonid Afrocyon burolleti, the holotype and only known specimen of which is from Gebel Zelten
(Arambourg, 1961) and the machairodont felid Syrtosmilus syrtensis (Ginsburg, 1978) also based on a damaged mandible.
Creodonts from the strata were hitherto represented only by the huge species Megistotherium osteothlastes (Savage, 1973)
and an undescribed “proviverrinid” (Savage, 1971) or Anasinopa (Savage & Hamilton, 1973). Several post-cranial bones
attributed to Megistotherium, have, on closer inspection, turned out to belong to the medium sized anthracothere,
Sivameryx africanus (four metapodials, Savage, 1973 Fig. 17) and a perissodactyl (a supposed magnum attributed to
Megistotherium is in fact a rhinocerotid sesamoid, Savage, 1973, Fig. 16).
Thus even though the material described and interpreted in this paper comprises only a few specimens, the better
preservation of some of the fossils provides interesting information about the fauna, and helps to clarify its taxonomic
relationships. SYSTEMATICS
Carnivora Bowdich, 1821
Family Amphicyonidae Haeckel, 1866
Genus Afrocyon Arambourg, 1961
Species Afrocyon burolleti Arambourg, 1961
Holotype: MNHN 1961-5-7, left mandible with abraded cheek teeth (p/4-m/3) (Fig. 1).
New material: NHM M 82373, left mandible containing well preserved p/4-m/2, the alveoli for m/3, p/3 and p/2
(Fig. 2); NHM M 82374, left m/2 (18.2 x 13 mm) (Fig. 3).
Description
NHM M 82373, is a mandible in which the horizontal ramus is relatively shallow. The p/4 is eroded mesially, so it is
not possible to determine whether it possessed an anterior cuspid or not. The main cusp is strong and tall, the posterior
cusp is quite tall, and there is a strong cingulum which is linked to the base of the posterior cusp by a crest. The anterior
premolars are not preserved, the alveoli of the p/3 indicate that it was not reduced and was juxtaposed to the p/4, the p/2
was probably biradiculate, and was separated from the p/3 by a gap.
The m/1 has a relatively short trigonid relative to its height. The anterior wall of the paraconid is completely vertical,
the metaconid is relatively strong and is displaced posteriorly with respect to the protoconid. The talonid consists of a
large sectorial hypoconid, there is practically no entoconid, only an inclined surface sloping towards the interior of the
talonid. In the m/2, wear has eradicated the trigonid morphology, so it is not possible to determine whether it had a
paraconid. There is a strong basal expansion mesio-buccally. The metaconid is small and low compared to the protoconid.
The talonid is dominated by a strong hypoconid, and there is almost no entoconid, just a rounded peripheral crest.

J. Morales et al.
44
Fig. 1: MNHN 1961-5-7, holotype mandible of Afrocyon burolleti Arambourg, 1961, from Gebel Zelten Libya, A) lingual, B)
occlusal, and C) buccal views (scale : 5 cm).
NHM M 82374 (Fig. 3) is a well preserved, unworn, left m/2. It has a clear paraconid. As in the above m/2, the
protoconid is taller and stronger than the metaconid. The talonid is dominated by the hypoconid, the entoconid is small,
reduced to a lightly subdivided lingual crest.

Carnivores (Creodonta and Carnivora) from the Basal Middle Miocene of Gebel Zelten, Libya 45
Fig. 2: NHM M 82373, left mandible, Afrocyon burolleti, from Locality L 25,
Gebel Zelten, Libya, A) buccal, B) occlusal, and C) lingual views (scale : 10 mm)
Table 1: Measurements (in mm) of the teeth of Afrocyon burolleti and Amphicyon sp. indet. from Gebel Zelten, Libya
(e – broken or damaged tooth, measurement estimated).
Bucco-lingual breadth Mesio-distal length Specimen 11.2e 19.8e 1961-5-7 left p/4 15.3e 31.0e 1961-567 left m/1 13.9e 23.3e 1961-5-7 left m/2 9e 16e 1961-5-7 left m/3 9.1 15.7e NHM M 82373 left p/4 14.0 28.2 NHM M 82373 left m/1 13.2 19.3 NHM M 82373 left m/2 13.2 17.9 NHM M 82374 left m/2 5.4 8.2 BIZ.2A.15 right p/2 6.0 10.8 BIZ.2A.15 right p/3 8.7 17.3 BIZ.2A.15 right p/4
Genus Amphicyon Lartet, 1837 (in Blainville, 1837)
Species Amphicyon sp. indet.
Material: BIZ.2A.15, fragment of right mandible containing p/4-p/2, the alveoli of p/1 and the canine root (Fig. 4).
Locality: Gebel Zelten, Libya
Age: Basal Middle Miocene
Description
BIZ.2A.15, preserves the horizontal ramus which is deep and robust, with strong alveoli for the canine, which
retains part of the canine root, the p/1 is represented by an uniradiculate alveolus, the p/2 is reduced with a single low
cusp, the p/3 is larger than p/2, but it also consists of a single low cusp. The p/4 is strong, without an anterior cuspid, the
posterior cuspid is moderately strong, the central cuspid is moderately tall, quite a bit lower than in mandible NHM M
82373, and the posterior cingulum well marked. All the premolars are separated by diastemata.
Fig. 3: NHM M 82374, left m
/
2, Afrocyon burollet
i
,
from Gebel Zelten, Libya, A) occlusal, B) lingual
and C
)
buccal views
(
scale : 10 mm
)
.

J. Morales et al.
46
Fig. 4: BIZ.2A.15, fragment of right mandible containing p/4-p/2, from Gebel Zelten, Libya, Amphicyon sp. indet., A) lingual, B) occlusal,
and C) buccal views (scale bar : 10 mm)
Discussion
This mandible is similar in dimensions to the new specimen NHM M 82373. However, its horizontal ramus is quite a
bit deeper and more robust, approaching that of the holotype of A.burolleti (Arambourg, 1961). In contrast, the p/4 has
lower cusps than in the holotype and in NHM M 82373, such that, together with the reduction in size of the anterior
premolars and the development of diastemata leads us to exclude it from A. burolleti. However, these differences from
Afrocyon burolleti, approach it to several species of the genus Amphicyon, in particular Amphicyon major from Sansan
(Ginsburg, 1961, 1963) which is however somewhat larger than the African form. Ginsburg & Welcomme (2002) attributed
several fossils from Gebel Zelten, which were identified as Megistotherium osteothlastes by Savage (1973), to a species the
size of Amphicyon giganteus (Schinz, 1825). In the present state of knowledge it is not possible to attribute BIZ.2A.15 to
Amphicyon giganteus, not only because of its smaller dimensions, but also because of the reduction of the anterior
premolars and the development of diastemata between them, features which distance it from this species (Morales et al.,
1997). Thus it is not possible to exclude the possibility that at Gebel Zelten, there existed three different species of
Amphicyonidae (Afrocyon burolleti, Amphicyon giganteus or a species similar in dimensions to it, and Amphicyon sp.
smaller than Amphicyon major).
A problem is encountered when comparing these fossils from Gebel Zelten with the holotype of Afrocyon burolleti
Arambourg, 1961, due to the poor state of preservation of the teeth (p/4-m/3) in the holotype (Fig. 1). The mandible of the
type specimen is somewhat more robust and the ramus is deeper than the new specimen, NHM M 82373, and the dentition
is larger. But the two specimens have the same morphology of the p/4 with the anterior cuspid tall and well developed. The

Carnivores (Creodonta and Carnivora) from the Basal Middle Miocene of Gebel Zelten, Libya 47
m/1s of the two specimens have the vertical orientation of the anterior wall of the paraconid, and the talonid is dominated
by the strong hypoconid. The m/2 and m/3 in the two mandibles, are large and quite broad (in NHM M 82373, only the
alveolus of m/3 is preserved, but its dimensions show that it was a large tooth).
The mandible, BIZ.2A.15, possesses a dentition which is comparable in dimensions to the new specimen NHM M
82373. However, the horizontal ramus is quite a bit deeper and more robust than it, features by which it resembles the
holotype of Afrocyon burolleti. In contrast, the p/4 has lower cusps than the type specimen and NHM M 82373, but has the
same overall morphological pattern.
We consider that the new material belongs to Afrocyon burolleti, which permits refinement of the diagnosis. The
differences in size of the horizontal rami and the teeth can be considered to fall within the range of variation of large
species of the family Amphicyonidae, as is the case in Amphicyon major from Sansan (Ginsburg, 1961).
Apart from its smaller dimensions, Afrocyon burolleti differs from Amphicyon major and Amphicyon giganteus by
its clearly more hypercarnivorous dentition, which is more sectorial, the p/4 is reduced, and the talonids of m/1 and m/2 are
reduced although the hypoconid is more developed. Similarly, in the m/2 of Afrocyon burolleti (NHM M 82374) the
morphology is clearly more primitive than it is in Amphicyon major, as revealed by the retention of the paraconid, the taller
and larger dimensions of the protoconid with respect to the metaconid, and its retired position. Comparable differences
apply to Amphicyon giganteus, a species with larger premolars with barely any gaps between them.
This hypercarnivorous tendency of Afrocyon burolleti compared to Amphicyon major and Amphicyon giganteus,
approaches it to Ysengrinia ginsburgi from Arrisdrift, Namibia (Morales et al., 1998, 2003). However, Y. ginsburgi possesses
a more reduced p/4, with no diastema between it and p/3. But above all, the m/1 and m/2 are quite different, being much
more hypercarnivorous, with narrow talonids comprised almost solely of the hypoconid, as in species of the genus
Ysengrinia (Ginsburg, 1965; Hunt 2002). These differences are probably great enough to maintain the generic separation
between Afrocyon burolleti from Gebel Zelten and Ysengrinia ginsburgi from Arrisdrift (Fig. 4), but the two forms are closer
to each other than either is to the genus Amphicyon. Finally, Afrocyon differs from Hecubides euryodon Savage (1965), not
only by its greater dimensions, but mainly by its markedly more hypercarnivorous dentition.
Fig. 5: Ysengrinia ginsburgi from Arrisdrift, Namibia, deposits which are about the same age as those at Gebel Zelten. 1) PQAD 133, left
mandible, A) lingual, and B) occlusal views; 2) AD 604’92, right maxilla, occlusal view of P4/-M1/ (scale bar : 10 mm).
Amphicyonidae indet.
Material: OCO 3’09, left m/2 (23.2 mm x 16.5 mm) (Fig. 6).
Locality: Ngorora, Kenya.
Age: Late Middle Miocene (ca 12.5 Ma).

J. Morales et al.
48
Description
The trigonid of this m/2 is taller than the talonid (Fig. 6), and is dominated by a well developed protoconid fronting
onto a quite tall metaconid, these two cusps being linked by a peripheral cristid which encloses a small anterior valley. The
talonid has a small chip of enamel broken from its lingual margin, but it is clear that it is narrower than the trigonid. We
highlight the strong development of the hypoconid, that it is disposed parallel to the mesio-distal axis of the molar,
contacting the posterior cristid of the protoconid. The lingual part of the talonid is occupied by a lateral cristid which joins
the base of the hypoconid, delimiting a narrow talonid valley.
Fig. 6: OCO 3’09, Amphicyon sp., left m/2 from the Ngorora Formation, Kenya, stereo occlusal view (scale : 10 mm).
Discussion
This is a large molar, close in dimensions to that of Amphicyon major from the French locality of Sansan (Ginsburg,
1961). It differs from Afrocyon burolleti, not only by its greater dimensions, but also by its narrower talonid and stronger
hypoconid. In addition the Ngorora tooth differs from that of Gebel Zelten by the lower anterior valley of the trigonid.
Comparable differences are present in the aforementioned Amphicyon major, in particular concerning the morphology of
the talonid, which in the European species is not reduced, is almost quadrangular in outline and has a well developed
talonid valley. It could be close to the m/2 from Samburu, Kenya (Tsujikawa, 2005). However, the Ngorora molar differs from
that of Amphicyonidae indet. from Samburu by the less reduced talonid. Similar differences are met in the known species
of Magericyon, M. castellanus (Ginsburg et al., 1981) and M. anceps (Peigné et al., 2008). M. castellanus is more primitive
than M. anceps by the less reduced talonid of the m/2. However, additional material is required for confident identification
of the Ngorora tooth and the Samburu specimens. We note that from the Early Miocene to the Late Miocene, there was a
tendency to reduce the talonid of m/2, in an analogous manner to that observed in species of Magericyon (Vallesian of
Spain).
GENERAL DISCUSSION ON AMPHICYONIDAE
Knowledge about the African Amphicyonidae is overall deficient, partly because the quantity of material is low, and
partly because an overall revision of the material is required. Grosso modo it is possible to differentiate three stages in the
history of the group in the African continent (Fig. 7).
The first, which covers the Early Miocene and part of the Middle Miocene, has a relatively high diversity, with at
least 5 genera, although in general there were only two species that co-existed at any one locality, except at Gebel Zelten (a
succession that spans an appreciable period of time) and Kipsaraman, Kenya, in which there were two species of
Hecubides as well as Agnotherium kiptalami (Savage, 1965;Schmidt-Kittler, 1987, Morales & Pickford, 2005). Apart from
Agnotherium at Kipsaraman, three different lineages of Amphicyonidae can be recognised in this stage; 1. The first
comprises the species of Hecubides, which in general retain an incipiently hypercarnivorous dental morphology, modest
size, save for the poorly known Hecubides macrodon (Savage, 1965). 2.- A clearly hypercarnivorous lineage comprising
Ysengrinia ginsburgi from Arrisdrift, Namibia, and Afrocyon burolleti from Gebel Zelten, Libya. To this lineage could be
assigned the form described as Hecubides sp. by Morlo et al., (2007) from Wadi Moghara, Egypt, which in our opinion is
quite close to Ysengrinia ginsburgi. 3.- A lineage including Amphicyon giganteus from Arrisdrift, Namibia and Amphicyon
sp. from Gebel Zelten described in this paper, both of which are close to Amphicyon major.
A second stage extends from the late Middle Micene to the base of the Late Miocene, in which only the genus
Agnotherium (Kaup, 1833) is known, the other genera from the previous stage having gone extinct (Kurten, 1976, Ginsburg,
1977, Pickford & Senut, 1997, Morales & Pickford, 2005). A second genus Myacyon dojambir was defined in the Late
Miocene of Oued Mya 1, Algeria, by Sudre & Hartenberger (1992), on the basis of a mandible containing m/1-m/2. This form
needs to be revised in detail, because several features approach it to larger species of the genus Agnotherium, in particular
by the hypercarnivorous tendency of the molars, the reduction of the metaconid and the presence of a basal cingulum.
However, it is necessary to examine the original fossil, as the published figures are insufficiently clear.

Carnivores (Creodonta and Carnivora) from the Basal Middle Miocene of Gebel Zelten, Libya 49
Fig. 7: Biochronology of African Amphicyonidae. (1 – Ysengrinia ginsburgi, 2 – Hecubides euryodon, 3 – Hecubides macrodon,
4 – Hecubides minor, 5 – Afrocyon burolleti, 6 – Bonisicyon illacabo, 7 – Amphicyon giganteus, 8 – Amphicyonidae indet., 9 – Myacyon
dojambir, 10 – Agnotherium kiptalami or A. antiquus).
The last stage in the history of the group occurred during the Late Miocene, as indicated by the presence of an
indeterminate form in the Samburu Hills, Kenya (Tsujikawa, 2005), which could be related to Magericyon anceps from the
Late Miocene of Spain (Peigné et al., 2008) and several large isolated teeth from Lothagam, Kenya, identified as
Amphicyonidae species A by Werdelin (2003). From Lothagam, Goma (Ethiopia) and Lemudong’o (Kenya) there is one of
the last, if not the last, Amphicyonidae in the fossil record, comprising a modestly sized, but quite hypercarnivorous, form,
for which Werdelin & Simpson (2009) erected the genus and species Bonisicyon illacabo. Finally, we note that, although
Amphicyonidae have been reported to persist in Asia until 7 Ma (Barry & Flynn, 1990; Peigné et al., 2006), the age
determinations are not secure, the youngest well dated specimens being no younger than 9 Ma, as in North America (Hunt,
2005) and Europe (Peigné et al., 2008). As such, it is intriguing that the family appears to have survived in Africa for several
million years longer than it did in other continents.
Order Creodonta Cope, 1875
Family Hyaenodontidae Leidy, 1869
Subfamily Hyainailourinae Pilgrim, 1932
Genus Anasinopa Savage 1965
Species Anasinopa libyca nov.
Holotype: NHM M 82375, right mandible containing m/1-m/3 (Fig. 8.1 A, B & C).
Referred material: NHM M 82376 (Fig. 8.2 A, B & C), is a right mandible with the root of the canine and alveoli of p/1, roots
of p/2, complete p/3 and the anterior root of p/4; NHM M 82377 (Fig. 8.4 A, B & C), is a right mandibular symphysis without
teeth; NHM M 82378 (Fig. 8.3 A, B & C) right mandible with milk dentition (d/4 and d/3).

J. Morales et al.
50
Type locality: Gebel Zelten, Libya.
Age: Early Middle Miocene.
Diagnosis: Anasinopa 10-25% smaller than Anasinopa leakeyi the type species of the genus. It differs from the type species
by the better development of the talonid of m/3, which is morphologically close to the m/2, despite the fact that in
Anasinopa leakeyi the talonids of m/2 and m/3 are very different, that of m/3 being greatly reduced. In addition, the p/3
possesses a better developed posterior cusp than that of A. leakeyi.
Description
NHM M 82375 is a right mandible containing m/1-m/3 (m/1, L = 9.3 mm, W = 5.1 mm; m/2, L = 13 mm, W = 7.1 mm;
m/3 L = ca. 13.5, talonid W = 5.4 mm). The serial position of the teeth in this jaw was determined using x-ray imagery (Fig.
9), which show clearly that there was no alveolus behind the last erupted tooth, which therefore corresponds to the m/3.
The wear on the teeth, especially great on the m/1 confirms this attribution.
The m/1 is so deeply worn that no morphological details remain. The m/2 is a typical carnassial with the V-shaped
trigonid in which the metaconid is quite reduced, and the broadened paraconid is in an anterolingual position. The
protoconid is almost conical. The talonid is almost as broad as the trigonid, and is simple with a chisel-shaped hypoconid
and a raised disto-lingual border. The m/3 is missing its trigonid, but it was probably also a carnassial judging from the
talonid which is similar to that of the m/2, although it is slightly narrower and shorter.
NHM M 82376 (Fig. 8.2 A, B & C), is a right mandible with the root of the canine and alveoli of p/1,roots of p/2,
complete p/3 and the anterior root of p/4 (C, L = 8.4 mm, W = 5.5 mm; alveolus of p/2 L = 8.7 mm; p/3, L = 9.8 mm, W = 4.4
mm). The mandibular ramus is low with a strong, elongate symphysis. The p/1 and p/2 each have two roots, the p/2 was
almost as big as the p/3. The latter is a relatively robust tooth with a small but well defined posterior cuspid. The anterior
cristid swells basally. A cingulum completely surrounds the tooth, but it is low.
NHM M 82377 (Fig. 8.4 A, B & C), is a right mandibular symphysis without teeth. It corresponds to a larger individual
than the previous specimen, but probably with similar morphology.
NHM M 82378 (Fig. 8.3 A, B & C) is a left mandible containing deciduous teeth (d/4 and d/3). The teeth are poorly
preserved, showing advanced corrosion, but allow us to observe that the d/3 is an elongated and gracile premolariform
tooth, with the principal cusp in an anterior position, preceded by a well developed anterior cusplet. The d/4 is close in
morphology to permanent molars, differing from them by being longer and more gracile, and with the metaconid greatly
underdeveloped, which as a consequence makes the trigonid more sectorial than in the permanent molars.
DISCUSSION ON CREODONTA
The inclusion of the new species from Gebel Zelten in Anasinopa raises several problems which need to be
discussed. Certainly, the general morphology of the fossils described above, show close similarity to Anasinopa leakeyi
from the Early Miocene of Napak, Uganda (Savage, 1965). However, the most important difference between A. libyca and A.
leakeyi resides in the morphology of the talonid of the m/3, which in the Gebel Zelten species is well developed, with low
cusps which completely surround an ample talonid valley. This morphology is quite close to the m/2 in the same mandible,
in which it is broader and the hypoconid better defined and taller. In A. leakeyi the difference between the talonids of m/2
and m/3 is strong, being reduced in m/3. Certainly, the difference in morphology of the talonid of m/2 and m/3 is even more
accentuated in species of Dissopsalis (Pilgrim, 1910, Colbert, 1935, Savage, 1965) or Metasinopa (Andrews, 1906; Osborn,
1909; Savage, 1965), genera grouped in the subfamily Hyainailourinae along with Anasinopa by Peigné et al., (2007). That
the talonids of the m/2 and m/3 are not reduced and show similar morphology is widespread in Hyaenodontidae from the
Palaeogene of Eurasia, North America and Africa (Matthew, 1906; Peigné et al., 2007, Gheerbrant et al., 2006), but is
unknown among the Miocene African representatives of the group. Even though A. leakeyi indubitably possesses an m/3
talonid which is reduced compared to that of m/2, the reduction is less than that observed in species of Dissopsalis, and
could be more with respect to Buhakia moghraensis Morlo et al., (2007) judging by the morphology of the m/2.
Nevertheless, it is reasonable to classify the new species from Gebel Zelten in Anasinopa, and to interpret this genus as
comprising less hypercarnivorous species than the rest of the African Miocene creodonts, with the exception of the highly
specialised genus Teratodon (Savage, 1965).

Carnivores (Creodonta and Carnivora) from the Basal Middle Miocene of Gebel Zelten, Libya 51
Fig. 8: Anasinopa libyca nov. sp. from Gebel Zelten, Libya. 1) Holotype. NHM M 82375 right mandible with m1-m3 A) lingual, B) buccal, C)
occlusal views; 2) NHM M 82376 right mandible with the root of the canine, alveoli of p/1, roots of p/2, complete p/3 and the anterior root
of p4, A) lingual, B) buccal, C) occlusal views; 3) NHM M 82378 left mandible with deciduous dentition (d/4 and d/3). A) lingual, B) buccal,
C) occlusal views, 4) NHM M 82377 right mandibular symphysis lacking teeth. A) lingual, B) buccal, C) occlusal views (scale bars : 10
mm).

J. Morales et al.
52
Fig. 9: Anasinopa libyca nov. sp. Holotype. NHM M 82375 right mandible with m/1-m/3. Radiograph, buccal view.
CONCLUSIONS
The carnivoran fauna from the Early and basal Middle Miocene deposits of Gebel Zelten, Libya, is poorly preserved,
with just a few cranio-dental specimens of amphicyonids, felids, and creodonts described. Some of the post-cranial bones
previously attributed to Creodonta (Savage, 1973) belong instead to anthracotheres and rhinocerotids. Specimens from the
deposits described herein, reveal that there are two amphicyonids at Gebel Zelten, and that in addition to the gigantic
creodont, Megistotherium osteothlastes, there is a medium sized species of hyaenodontid, here attributed to a new species
of Anasinopa, A. libyca, which is somewhat smaller than the type species Anasinopa leakeyi from East Africa.
Table 2. Carnivoran fauna from Gebel Zelten, Libya
Species Family Megistotherium osteothlastes Savage, 1973 Anasinopa libyca sp. nov. Hyaenodontidae Afrocyon burolleti Arambourg, 1961 Amphicyon sp. indet. Amphicyonidae Syrtosmilus syrtensis Ginsburg, 1978 Felidae
The low diversity of the Gebel Zelten carnivores contrasts with that from Wadi Moghra, Egypt (Morlo et al., 2007)
which comprises three or four creodonts (depending on whether Hyainailourus fourtaui and Megistotherium osteothlastes
are synonyms or not), one amphicyonid (Cynelos sp.) and three small carnivores (Herpestides aegyptiacus, Ketketictis
solida, and Moghradictis nedjema). The different representations at the sites are probably due to taphonomic factors or
collecting bias (small carnivore post-cranial bones from Gebel Zelten are in the Paris collection, but in the absence of teeth,
cannot be securely identified).
ACKNOWLEDGEMENTS
We thank Andy Currant, NHM London, and Christine Argot, MNHN, Paris, for access to the collections in their care.
Funding was provided by the Collège de France, and Project CGL2008-05813-CO2-01, MICINN (Spanish Government) and
the Research Group CAM-UCM 910607.
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