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Anthracotheres from Wadi Moghra, early Miocene, Egypt
Author(s): Ellen R. Miller, Gregg F. Gunnell, Mohamed Abdel Gawad, Mohamed Hamdan, Ahmed N.
El-Barkooky, Mark T. Clementz, and Safiya M. Hassan
Source: Journal of Paleontology, 88(5):967-981. 2014.
Published By: The Paleontological Society
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ANTHRACOTHERES FROM WADI MOGHRA, EARLY MIOCENE, EGYPT
ELLEN R. MILLER,
GREGG F. GUNNELL,
MOHAMED ABDEL GAWAD,
AHMED N. EL-BARKOOKY,
MARK T. CLEMENTZ,
AND SAFIYA M. HASSAN
Department of Anthropology, Wake Forest University, Winston Salem, NC 27106, USA, ,firstname.lastname@example.org.;
Duke Lemur Center Division of
Fossil Primates, Durham, NC 27705, USA, ,email@example.com.;
Geology Department, Faculty of Sciences, Cairo University, Cairo, Egypt,
Department of Geology and Geophysics, University of Wyoming, Laramie, WY 82071, USA, ,firstname.lastname@example.org.
ABSTRACT—The early Miocene site of Wadi Moghra, Qattara Depression, Egypt, is important for interpreting anthracothere
(Mammalia, Artiodactyla) evolution, because the Moghra sediments preserve a higher diversity of anthracotheres than any
other pene-contemporaneous site. New specimens from Moghra are described and form the basis for the systematic
revision of Moghra anthracotheres provided here. Among the important discoveries recently made at Moghra is the first
complete skull of Sivameryx moneyi. Other new specimens described here include two new species of Afromeryx, and a
new genus and species, all of which are unique to Moghra. A review of biogeographic information supports the conclusion
that three of the Moghra anthracotheres (Brachyodus depereti, B. mogharensis, and Jaggermeryx naida, n. gen. n. sp.) are
members of late surviving lineages with a long history in Africa, while three other species (Afromeryx grex, n. sp.,A.
palustris, n. sp.,and Sivameryx moneyi) represent more recent immigrants from Eurasia.
THIS CONTRIBUTION provides new information from Wadi
Moghra, early Miocene, Egypt, toward a better under-
standing of anthracothere (Anthracotheriidae, Artiodactyla)
evolution and paleobiology. Wadi Moghra (¼Moghara) com-
prises a number of fossil localities occupying the northeastern
portion of the Qattara Depression in Egypt’s Western Desert
(Fig. 1). Occurrences of fossil vertebrates at Moghra have been
reported for over a century (Andrews, 1899; Blanckenhorn,
1901). Fourtau (1918) published the first treatise on the Moghra
vertebrate fauna, and then re-published the same monograph
two years later with additional appendices added (Fourtau,
1920). No work was then done in the area until the collecting
expeditions of E. L. Simons (Duke University) and colleagues,
conducted at sporadic intervals during the 1970s and 1980s
(Rasmussen et al., 1989; Simons, 1994), followed by the work
of E. R. Miller and a joint Egyptian-American team working at
Moghra from the mid-1990s until present (e.g., Abdel Gawad et
al., 2010, 2012; Hassan et al., 2012; Miller, 1999; Miller and
Simons, 1998; Miller et al., 2009; Morlo et al, 2007; Pickford et
al., 2010; Sanders and Miller, 2002).
Moghra is an important site for interpreting anthracothere
morphology, systematics, paleobiology, and biogeography,
because it yields a higher diversity of anthracotheres than any
other early Miocene site. Moghra also contributes to our
understanding of anthracothere evolution, because six species
in four genera can be identified from the locality, and all of
these taxa are either best known from or only known from
Andrews (1899) was the first to describe an anthracothere
from Moghra, identifying one mandible and two associated teeth
as representing Brachyodus africanus. Fourtau (1918, 1920),
working with a much larger collection, recognized three
anthracothere species in three genera: Masritherium depereti,
Brachyodus africanus,andHyoboops moneyi.Overtheyears,
additional material has been attributed to these three taxa,
including specimens reported by Forster-Cooper (1924), E. L.
Simons and colleagues in the 1970s and 1980s, and in recent
years by a joint Egyptian-American team.
Pickford (1991) included a fair portion of the known
anthracothere material from Moghra in his revision of the
Neogene Anthracotheriidae. At that time, Pickford (1991)
identified four species at Moghra: Brachyodus depereti,B.
mogharensis,Sivameryx moneyi,andAfromeryx africanus.In
recent reviews of Cenozoic Anthracotheriidae (Holroyd et al.,
2010; Lihoreau and Ducrocq, 2007) these same taxa were
recognized. This report includes new information about some of
these previously recognized anthracothere species at Moghra,
including identification of the first known skull of S. moneyi,
and specimens representing new taxa. In the following
document, all specimens with Duke University Lemur Center
(DPC) and Cairo University Wadi Moghra (CUWM) designa-
tions, as well as much of the anthracothere material housed at
the Cairo Geological Museum (CGM), is described and
discussed here for the first time.
ABBREVIATIONS AND NOMENCLATURE
Institution.—AM, Geological Survey of Namibia Museum,
Windhoek; BMNH, Natural History Museum, London (specimens
designated ‘‘ M’’ ); CGM, Cairo Geological Museum, Cairo;
CUWM, Cairo University Wadi Moghra, Cairo University, Cairo;
DPC, Duke University Lemur Center, Durham, NC; NMNH,
National Museum of Natural History, Washington, DC; SMNS,
Staaliches Museum fu¨r Naturkunde Stuttgart.
Dental.—An upper case letter denotes a tooth in the maxillary
row, a lower case letter denotes a tooth in the mandibular series,
and a ‘‘d’’ indicates a deciduous tooth, e.g., M1, upper molar, p3,
lower premolar, dp, lower deciduous premolar.
Synonymy.—Detailed synonymies for anthracothere taxa dis-
cussed in this paper can be found in Lihoreau and Ducrocq (2007)
and Pickford (1991).
Class MAMMALIA Linnaeus, 1758
Order ARTIODACTYLA Owen, 1848
Family ANTHRACOTHERIIDAE Leidy, 1869
Genus BRACHYODUS Dep´
BRACHYODUS DEPERETI (Fourtau, 1918)
Type.—CGM 30798, mandible with both dentaries including
Journal of Paleontology, 88(5), 2014, p. 967–981
Copyright Ó2014, The Paleontological Society
left i3, m2–3 and right i3, p4–m3 (Fig. 2A) (also figured in
Fourtau, 1920; 46, 50), M 29372 is a cast.
Diagnosis.—Revised: very large species of Brachyodus, m1–
m3 length about 143 mm, C–P1 diastema about 95 mm, i1–i2
absent, i3 tusk-like, at least in males, p1 occasionally absent.
Occurrence.—Early Miocene, Wadi Moghra, Egypt; Gebel
Zelten, Libya; Sperrgebiet, Namibia.
Material.—CGM 7210, right M; CGM 30762 right dentary,
roots of p3, p4–m3 (Andrews, 1899, pl. XXIII, see below); CGM
30764, left dentary, p3–m2; CGM, 30797, left dentary p2–m2;
CGM 30800, right dentary p2–m3; CGM 30801, left maxilla P1,
P4; CGM 32984, symphysis; CGM 32985, symphysis; CGM
73660, left dentary p1; CGM unnumbered, right dentary alveolus
p1, crown p2 (Andrews, 1899, pl. XXIII); CGM unnumbered,
right dentary m2–3; CGM unnumbered, left maxilla P3–M1;
CGM unnumbered, right dentary m1–2; CGM unnumbered, right
dentary m2–3 (Fourtau, 1920, figs. 32, 33); CGM unnumbered,
left dentary, worn m1–3. CUWM 10, right M3; CUWM 14, right
maxilla P2–3; CUWM 82, left m3, CUWM 69, right maxilla M2–
3; DPC 2518, left dentary m1, m2 erupting; DPC 4064, right M2;
DPC 4065, right maxillary fragment with dP3–M1; DPC 4068,
left M3; DPC 4107, left M3 fragment; DPC 4421, left dentary
dp4–m2, DPC 4426, right dentary p4–m2; DPC 4438, left M1;
DPC 4465, left dentary worn m3; DPC 4586, right dentary p4–
m3; DPC 4593; left and right dentaries worn right and left m1–3;
DPC 6238, left M2; DPC 6440, left dentary m2–3; DPC 6467, left
M2; DPC 6620, right dentary, m1 broken, m2–3 worn; DPC 7373,
right upper dP4; DPC 7385, left maxilla fragment; DPC 7501, left
dentary m2–3; DPC 7746, left dentary p2–m2; DPC 8930,
isolated i3; DPC 8965, right dentary broken m2, m3 erupting;
DPC 9028, left dentary m1–3; DPC 9046, isolated i3; DPC
12553, left dentary, p4–m3; DPC 12930, left dentary dp3–4; DPC
14540, molar fragment; DPC 14563, right M1; DPC 14580, left
P4, M3; DPC 17679, left maxilla P3–M3; DPC 17681, right
dentary m2–3; DPC 17683, right M3; DPC 17684, left maxilla
P2–4; DPC 17689, left M3; DPC 17697, left maxilla, P4–M3; M
11067, left P4–M3 (Tables 1, 2).
Remarks.—Brachyodus depereti has been described previously
(Dineur, 1982; Fourtau, 1920; Holroyd et al., 2010; Pickford,
1991). Premolars and molars are morphologically similar to the
type species, B. onoideus from Europe, but the teeth of B. depereti
are larger. Upper molars are five-cusped with pinched rather than
loop-like styles. The dental enamel is finely wrinkled and there is
a prominent lingual cingulum. No skull preserving the premaxilla
of B. depereti is known, but it is likely that the central upper
incisors were tusk-like and pointed anterioinferiorly, as both B.
onoideus and B. aequatorialis show a similar condition (Pickford,
1991). The i1–2 are absent in B. depereti and, at least in males, i3
is a robust, tusk-like tooth. Upper dP4 is molariform. Black
(1978) suggested that the canine is possibly absent in females but
no material is available from Moghra that would either support or
refute this suggestion (after Pickford, 1991).
Two specimens are identified here as B. depereti, CGM 30762,
a right dentary with p4–m3, and an unnumbered dentary fragment
with the alveolus for p1 and the crown of p2. These specimens
were originally described and figured by Andrews (1899, pl.
XXIII), and subsequently CGM 30762 was also figured by
Pickford (1991, pl. 3; Figs.1, 2). Both of these authors cited the
dentary with p4-m3 as CGM 2849, but the specimen now has the
number CGM 30762. How two different catalog numbers came to
be associated with this specimen is unknown, but we identify the
specimen using the number it has now.
In addition, both of these specimens have a complicated past.
Cairo Geological Museum 30762 was originally published by
Andrews 1899 as the holotype of Brachyodus africanus. Fourtau
(1918, 1920) and Forster-Cooper (1924) subsequently assigned a
small number of specimens from Moghra to this taxon. Pickford
(1991) felt that the holotype jaw of B. africanus did not belong in
Brachyodus, although the rest of the hypodigm did. To remedy
this, Pickford reassigned CGM 30762 to the genus Afromeryx,
designated it as the type of a new species, A. africanus, and
identified one additional sub-adult mandibular specimen (M
15021) as a member of the A. africanus hypodigm (see below).
Pickford then created a new species, Brachyodus mogharensis
(see below), to house the ‘‘genuine’’ Brachyodus material
formerly recognized as B. africanus. Based on size and overall
morphology, we believe that CGM 30762 can comfortably be
included as a specimen of Brachyodus depereti (see discussion
below under Afromeryx for more details) and support that
Brachyodus depereti is well represented at Moghra but is a
much rarer element among the Gebel Zelten (Libya) and
Sperrgebiet (Namibia) faunas. In Libya, the taxon is documented
by the presence of a very large talus in the size range for B.
depereti (Pickford, 2003), and in Namibia the presence of B.
depereti has been identified on the basis of a distal lateral
metacarpal (AM 1097), a femur lacking the head and neck (AM
02), and possibly a large talus (unnumbered) (Pickford, 2003). An
occurrence of B. depereti has also been reported from Siwa Oasis,
Egypt (M 11967) although this is likely an error. Hamilton (1973)
published a description of six early Miocene fossil specimens
housed in the Natural History Museum (London). A note in the
collection indicates that these fossils were purchased by the
Museum in 1920 from Lady Moon, who found them ‘‘near Siwa
Oasis’’. However, there appears to be no continental Miocene
deposits in or around Siwa. Instead, Siwa has long been known as
a crossroad for caravans traversing northern Africa, and the fact
that B. depereti is a common element in the Moghra fauna
suggests that Moghra may be the original source of the material
and that people may have carried material from Moghra to
somewhere ‘‘near Siwa’’ .
BRACHYODUS MOGHARENSIS Pickford, 1991
Type.—M 15020, right maxilla P1–M3 (Fig. 2B) (also figured
in Pickford, 1991 pl. 1, fig.1).
FIGURE 1––Star indicates location of Wadi Moghra.
968 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
FIGURE 2––1,Brachyodus depereti, type dentary (CGM 30798) in occlusal view; 2, maxilla of Brachyodus mogharensis in occlusal view, M 15020, holotype;
3, maxilla of Brachyodus depereti, M 11067, in occlusal view. Scale bars¼10 cm.
MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 969
Diagnosis.—Smaller than B. depereti, similar in size to
Brachyodus aequatorialis; differs from B. aequatorialis in having
smaller upper premolars, more distinct barrels on the buccal cusps
of upper molars, more strongly developed buccal cingula on the
upper premolars and molars; infraorbital foramen opens superior
to P4, length of the molar tooth row about 114 mm (after
Occurrence.—Early Miocene, Wadi Moghra, Egypt.
Remarks.—The type specimen of Brachyodus mogharensis is a
maxilla in which the upper molar metaconules lack an antero-
lingual crest, there are continuous cingula around P2–4, the
lingual portion of which is particularly sharp-edged and beaded,
P4 has a single lingual cusp and a strong barrel on the buccal
aspect of the tooth. Brachyodus mogharensis differs from B.
depereti by its smaller size (Table 1, Fig. 2B, 2C).
Brachyodus mogharensis is known from a single maxillary
specimen. When the taxon was described (Pickford, 1991), the
holotype maxilla was accompanied by a hypodigm comprised of
five unnumbered mandibular specimens in various stages of wear.
One of these specimens (CGM 30775, figured in Fourtau, 1920,
fig. 31), has been moved to a new taxon (see below), and the
remaining specimens could not be clearly distinguished from
Brachyodus depereti and so have been assigned to that species.
Field work at Moghra in recent years has not yielded any
additional specimens clearly attributable to B. mogharensis.
Genus SIVAMERYX Lydekker, 1878
SIVAMERYX MONEYI Fourtau, 1918
Type.—CGM 30780, left dentary with p3–m3, (Fig. 3A, 3B)
(also figured by Fourtau, 1920, fig. 1).
Diagnosis.—A small species of Sivameryx, length of M1–3 59
mm (S. africanus comparable measurement 80 mm), differs from
Asian Sivameryx species in having lower molar paracristid
extending to lingual margin andinpossessingalingual
postprotocrista on upper molars that extends distally to the
Occurrence.—Early Miocene, Wadi Moghra, Egypt.
Material.—CGM 30781, left dentary p3–m3 (m3 broken);
CGM 39779, left dentary m3; CGM 73692, left dentary m2; CGM
TABLE 1—Wadi Moghra anthracothere mandibular measurements in millimeters. Abbreviations: L¼length; W¼width; *¼type specimen;
¼from Andrews, 1899;
p1 p2 p3 p4 m1 m2 m3
B. depereti CGM 30798* 31.25 19.08 32.18 (26.08) 37.67 32.33 59.33 34.54
22 13 27 17 25 17 36 24 51 25
CGM 30797 28.4 18.63 32 23.79 39.53 28.41 44.72 31.85
CGM 30880 26.4 17.17 27.62 20.89 32.54 23.64 37.73 26.48 43.84 29.86 58.73 31.61
CGM 73660 13.62 8.62
CGM no#52.3 24.35
CUWM 82 51.96 27.49
DPC 2518 35.17 24.39
DPC 4421 33.53 22.24 37.92 25.85
DPC 4426 34.57 23.38 42.94 29.91
DPC 4438 35.93 23.9
DPC 4586 28.63 20.62 28.92 21.75 39.93 28.82 54.83 29.4
DPC 6440 42.9 29.05 51.71 29.94
DPC 6620 38.45 31.04
DPC 7501 49.08 25.65
DPC 7746 24.51 15.06 24.44 16.84 26.04 18.67 33.71 22.12 39.14 28.02
DPC 8965 35.83 25.9 53.84 29.23
DPC 9028 31.26 22.48 35.74 25.18 51.19 26.02
DPC 12553 29.52 18.09 31.14 23.33 35.01 27.67 54.93 25.73
DPC 17681 35.71 27.78 51.62 29.04
DPC 17697 35.87 25.28 41.09 28.26
S. moneyi CGM 20780* 14.6 7.9 14.21 8.8 14.37 10.77 19.1 12.12 28.44 13.1
CGM 30781 13.62 7.46 13.36 8.95 16.34 9.87 20.28 14.48
CGM 30779 28.52 12.84
CGM 73692 18.66 11.82
DPC 4067 19.26 11.12 29.75 12.35
DPC 4425 14.02 7.7 13.8 8.69
DPC 4452 20.95 11.57 29.62 13.76
DPC 5973 9.4 6.04 12.49 6.25
DPC 7281 22.45 13.85
DPC 8972 12.86 8.78 14.12 9.38 19.46 11.39 26.02 12.74
DPC 12546 18.56 12.13 27.69 13.05
DPC 12547 13.61 8.72
DPC 12605 13.69 8.56 21.54 12.07
DPC 12608 18.51 10.37 27.4 11.82
DPC 12609 13.68 9.34 19.65 12.29
DPC 12610 12.73 7.96 13.88 9.01 15.17 10.47 19.02 12.23 29.46 12.55
DPC 12940 11.05 5.49 13.79 8.17
DPC 14547 29.67 14.03
DPC 14559 8.96 5.58 11.71 5.4 12.3 6.68
DPC 17680 13.59 9.08 13.64 10.8 18.84 12.17
DPC 17682 14.97 9.27 20.09 12.8
DPC 17742 15.03 9.82 18.78 12.23 28.43 13.37
A. grex M 15021* 15.76 11.2 18.45 13.11 21.38 14.82 30.86 20.71 44.12
A. palustris CGM 83571* 24.01 15.48 27.64 17.77 31.8 (20.08)
DPC 2542 25.75 17.78 20.53 32.2 23.27
DPC 4244 20.42 14.71 25.45 16.68
DPC 4604 27.2 18.85 28.47 20.45 34.26 25.35 49.12 26.15
J. naida CGM 30775 21.72 12.92
CGM 2499 17.1 (11.58) 23.58 (14.48) 15.41
970 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
73679, edentulous left dentary; CGM 84434, edentulous right
dentary with symphysis; CUWM 30, left dentary, m2–3; CUWM
31, left maxilla M2–3; CUWM 64a, right maxilla P2-M3, CUWM
64b left M3; CUWM 70, right maxilla M1–2; CUWM 100a, left
dentary alveoli p1–m1, crowns m2–3; CUWM 100b, left dentary
m2–3; CUWM 172, skull; DPC 3421, right dentary m1–2
fragments; DPC 3932, right dentary eroded m2–3; DPC 4066,
right maxilla M2–3; DPC 4067, right dentary p3–m1 (abraded);
DPC 4425, right dentary m1–3: DPC 4434, right dentary with
broken m3; DPC 4452, left dentary with broken m1–3; DPC
4535, right maxilla P3, P4; DPC 5968, right m3 fragment; DPC
5973, left dentary p1–2; DPC 5979, left M2; DPC 6234, left M1;
DPC 6243, right maxilla P3–M2; DPC 6289, left maxilla M1–3,
M1–2 broken; DPC 7281, right M2; DPC 7659, left maxilla M3:
DPC 8931, right maxilla fragment; DPC 8972, right dentary p3–
m3; DPC 9039, left dentary p1–dp4 roots; DPC 12538, left
dentary alveolus for p1, broken crowns p2–4; DPC 12546, right
dentary m2–3; DPC 12547, right maxilla P4–M1 (abraded); DPC
12605, left dentary p4–m3, m1–2 broken; DPC 12608, left
dentary m1–3, m1 broken; DPC 12609, right dentary p4, m2;
DPC 12610, left dentary p3–m3; DPC 12632, left M3 fragment;
DPC 12940, right dentary p1–2, p4–m1, m3 abraded; DPC 14547,
right dentary m2–3; DPC 14559, right dentary p1–3; DPC 17680,
left dentary p4–m2; DPC 17682, left dentary m1–2; DPC 17685,
TABLE 2—Wadi Moghra anthracothere maxillary measurements in millimeters. Abbreviations: L¼length; W¼width; *¼type specimen.
B. depereti CGM 30801 21.95 18.65 26.97 21.07
DPC 4064 34.41 48.88
DPC 4065 34.13 35.22
DPC 4068 42.15 47.87
DPC 6238 38.61 45.74
DPC 14563 34.76 38.53
DPC 14580 23.44 28.31
DPC 17679 23.79 23.54 21.78 27.69 31.81 38.88 42.03 44.45 45.82 49.62
DPC 17683 24.99 24.66 26.37 28.95 26.61 31.75
B. mogharensis M 15020* 14.79 12.34 16.88 14.16 18.00 17.61 16.31 24.04 24.04 27.83 33.25 37.04 35.1 39.38
S. moneyi CUWM 64A,B 19.92 18.27 23.08 21.18 22.46 23.44
CUWM 70 17.5 17.52 22.11 21.55
DPC 4066 21.47 22.34 24.04 24.86
DPC 4535 12.14 12.61 12.07 10.71
DPC 5979 20.71 20.89
DPC 6243 13.56 11.09 12.12 14.37 15.5 16.35 23.12 20.81
DPC 6289 21.1 21.32
DPC 7659 25.35 22.89
DPC 17685 20.69 20.08 22.23 22.12
DPC 17714 10.4 13.71
DPC 21506 27.49 30.03
FIGURE 3––1,2,Sivameryx moneyi, type dentary (CGM 30780) in occlusal and lateral views, respectively. Scale¼10 cm.
MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 971
left maxilla M2–3; DPC 17687, left dentary fragments; DPC
17701 right m3 fragment; DPC 17710, dentary fragments; DPC
17711, right dentary roots of p4; DPC 17714, left maxilla P4–M3,
M1 and M3 broken; DPC 17742, right dentary p4–m3; DPC
21506, right maxilla M3 (Tables 1, 2).
Description.—Sivameryx has been described and discussed
previously (Black, 1978; Fourtau, 1918, 1920), but most recently
by Pickford (1991) who noted important distinguishing features
of the genus such as: quasi-penticuspidate upper molars featuring
loop-like parastyles and mesostyles, protoconule is almost fused
with the protocone, canines are markedly sexual dimorphic, p2 is
double-rooted, the dentition is selenodont, and the m3 talonid is
loop-like and strongly obliquely oriented. Pickford (1991, p.
1511) noted that S. moneyi was mostly represented by lower jaws
and the species appeared to be ‘‘in all its known parts, merely a
smaller version of S. africanus’’ from Gebel Zelten.
Recovery of CUWM 172, a nearly complete adult skull of S.
moneyi, greatly expands our understanding of the taxon. Because
the focus of this paper is on the diversity of anthracotheres found
at Wadi Moghra, we provide an initial description of this
Sivameryx skull here. Detailed study of the skull and its
implications for larger issues involving the functional morphol-
ogy, systematics, and paleobiogeography of anthracotheres in
general is beyond the scope of this work.
The skull is well preserved (Fig. 4A–4D) and cranial sutures
are distinct throughout allowing for clear identification of key
Nasal bones: the nasals are broken anteriorly but originate at
anterior margin of orbit and are relatively broad posteriorly but
tapering anteriorly as far as they are preserved.
Frontal bones: the frontal bones are relatively short and wide,
forming the roof of the skull and comprise about two-thirds of the
medial wall of the orbit. Orbital processes are broken but are
presumed to have been relatively robust judging by the remaining
portion of their bases. The frontals extend rostrally as bilateral
narrow splints separating the caudal part of the nasals from the
lacrimals. The frontal has considerable contact with the lacrimal,
palatine, and orbitosphenoid but only a very small area of contact
with the maxillary bone. Dorsally the frontal forms distinct ridges
that sweep caudally from the orbital processes to converge
towards the sagittal crest.
Orbit: the orbital mosaic is comprised of the lacrimal,
maxillary, palatine, frontal, and orbitosphenoid bones, but is
dominated by contributions from the frontal and palatine bones
with a lesser contribution from the maxilla. The lacrimal foramen
is small and located well within the orbit and the facial wing of
the lacrimal extends relatively far anteriorly. The optic foramen is
relatively large and enters the orbit through the orbitosphenoid.
There are two smaller foramina anterior to the canal, a very small
one in the frontal and a larger one formed on the maxillopalatine
Parietal bones: a pronounced sagittal crest is present superiorly
on the parietals, and the crest extends caudally onto the occiput
where it helps form the most caudal part of the skull. The parietals
are rounded laterally and demarcate a relatively small braincase
that tapers anteriorly towards the parieto-frontal contact as well as
caudally toward the parieto-occipital suture. Ventrally and
caudally, the parietals converge to help form part of the nuchal
crest, and large gutters for the temporalis muscle are found caudal
and lateral to the braincase, extending along the lateral margins of
the parietal and temporal bones.
Temporal bones: the temporal forms the basal and lateral
portion of the braincase. The temporal extends and contacts the
parietal dorsally, the orbital wing of the sphenoid rostrally, and
the basisphenoid ventrally. At the juncture of the basisphenoid
and temporal is the internal auditory meatus and slit-like formen
lacerum medium that is continuous with that for the internal
auditory meatus. The bony auditory bulla is large and bulbous,
and has a distinct and elongate external auditory tube that extends
caudally and dorsally, as in some suids, peccaries and
hippopotami. Just rostral to the bulla is a distinct foramen ovale
formed in the caudal-most aspect of the sphenoid. Caudal to the
bulla is a small posterior lacerate foramen. The temporal contacts
the lateral wing of the basioccipital in a relatively long
dorsoventrally extend suture. The glenoid surface is flat, and
almost as wide as it is long. There is a distinct postglenoid process
but both an anterior process and a postglenoid foramen are absent.
Dorsocaudally the temporal bone forms a robust portion of the
lateral aspect of the nuchal crest.
Occipital bone: the inferior portion of the occipital is occupied
by large, robust occipital condyles. Anterior and lateral to the
condyles, two bony wings sweep forward to form dorsoventrally
flattened surfaces. The rostral portion of these surfaces form a
wall caudal and lateral to the auditory structures. These structures
also extend inferiorly to form small anteroposteriorly compressed
mastoids, although no stylomastoid foramina are evident.
Hypoglossal foramina are present rostral to the condyles, and
small but distinct jugal foramina are located inferiorly to the
nuchal crest and lateral on the basioccipital.
Sphenoid bone: ventrally, the sphenoid has a distinct central
ridge that continues rostrally to meet a sharply defined and high
vomer. Caudally the basiosphenoid is bifurcate with bilaterally
developed low processes extending caudally to the internal
Maxillary bone: the maxilla is robust and forms a rostrum that
is dorsoventrally relatively deep. However, the maxilla is
transversely compressed just rostral to the zygoma, so that the
rostrum appears relatively deep but mediolaterally fairly narrow.
The rostrum is concave and narrowest superior to P4 and then
broadens modestly rostrally. The concave portion of the rostrum
is occupied by a small infraorbital foramen positioned superior to
the posterior root of P3. From the infraorbital foramen, the
maxilla extends a short distance caudally to a point superior to
M3, where the bone terminates in a pointed process that is
notched medially and ventrally, near where the maxilla meets the
palatine. The maxilla forms the floor of the orbit and encloses the
infraorbital foramen. The portion of the palate formed by the
maxilla is relatively long and narrow, but not as elongate as in
Afromeryx or Brachyodus. The incisive foramina are broken
rostrally so their extent is hard to estimate. The caudal margin of
these foramina does not extend past I2. There is a paired set of
palatal foramina present at the rostral margin of P3 and a second
very small pair of palatal foramina present at the rostral margin of
Zygomatic bones: the rostral roots of right and left zygoma are
present but most of the zygoma are absent.
Palatine bones: the palatine extends rostrally as a broad plate
with a narrow central ridge until it reaches the rostral margin of
M3, where it tapers to a point that extends to the rostral margin of
M2. The palatine extends caudally well past the tooth row to the
caudal rim of the orbit, and a distinct post-palatine process is
Dentition: the dentition is represented by right P4–M3, left P3
(broken), and P4–M3. All teeth except right P4 and right M3 are
heavily worn. The anterior premolars are represented only by
alveoli and it is difficult to be certain of homologies because
supernumerary teeth were present, not only a common occurrence
in Libycosaurus (Lihoreau et al., 2006; Pickford, 2006) but also in
anthracotheres in general (personal observ.). The right dentition
anterior to P4 has 9 alveoli. We interpret these to represent P?
972 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
FIGURE 4––1–4,Sivameryx moneyi, skull (CUWM 172) in right lateral, left lateral, superior, and inferior views, respectively. Scale¼10 cm.
MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 973
(double-rooted), P1 (double-rooted), P2 (double-rooted), and P3
(triple-rooted). The left side of the palate has six alveoli anterior
to P3 which we interpret to represent the homologs of the right
side. Recovery of CUWM 172 confirms a number of features
previously cited as characteristic of S. moneyi and also reveals
some new details. The new skull confirms that P1 and P2 are
double-rooted (Pickford, 1991). The occurrence of supernumerary
premolars in S. moneyi has not been documented previously but
extra premolars seem to be a fairly common occurrence among
anthracothere species in general (personal observ.).
Genus AFROMERYX Pickford, 1991
AFROMERYX GREX new species
Type.—M 15021, sub-adult left dentary, edentulous symphysis,
crowns of p2–m3, m3 in crypt (Table 1, Fig. 5A).
Diagnosis.—Differs from A. zelteni in being larger (m2 area
.60%larger), having a much longer molar series length (m1–3
~108 mm in A. grex versus ~58 mm in A. zelteni), p4 with a
well-developed talonid; differs from A. palustris (n. sp. below) in
having smaller premolars (~25–30%), lower premolars with
pustulate anterior and posterior crests, lowers premolars with
distinct and elevated cingulids, p2 lacks development of an
Etymology.—"Grex,’’ Greek for flock or herd, in recognition of
the herd behavior of many artiodactyls.
Occurrence.—Early Miocene, Wadi Moghra, Egypt.
Description.—Description of holotype specimen and occlusal
details of teeth are available in Pickford (1991).
Remarks.—Afromeryx grex resembles the better known A.
zelteni from Gebel Zelten, in having an unfused symphysis, dental
formula 220.127.116.11; no specializations of the anterior dentition, i1–3
are small spatulate teeth; no i3–c diastema, c–p1 diastema
present, p1 single rooted, and mental foramina below i1, p2, p4.
In all known comparable parts, A. grex seems to be a larger
version of A. zelteni. As Pickford (1991) noted, the presence of
pustulate crests on the premolars in species of Afromeryx is
reminiscent of the condition seen in Gonotelma shabazi from the
Bugti Beds, early Miocene, Pakistan, although what this
resemblance means regarding the extent of a possible relationship
between the two taxa has not been explored.
The taxonomic history of Afromeryx is complex. Pickford
(1991) named Afromeryx africanus on the basis of a right dentary
with roots of p3, p4–m3 (CGM 30762), and attributed one
additional sub-adult mandibular specimen (M 15021) to this
species. The type specimen of A. africanus had previously been
recognized as representing Brachyodus africanus (Andrews,
1899), but Pickford (1991) stated that, ‘‘The material does not
closely resemble Brachyodus, the molars and premolars being
bunodont, the jaw being short with an unspecialized symphyseal
region’’ (1991, p. 1503). In terms of size and degree of
bunodonty, we find that CGM 30762 cannot be distinguished
from members of Brachyodus depereti and so we have assigned
the specimen to that taxon. In addition, it is unknown how the
assessment was made that CGM 30762 represented a taxon with a
short jaw and an unspecialized symphyseal region. The dentary is
broken in front of p3 and so is missing the anterior portion of the
jaw. This is critical because many of the features that distinguish
anthracothere species are located in the anterior dentition. For
example, species of Afromeryx have an unspecialized symphyseal
region (i1–3 small, spatulate teeth, no i3–c diastema, p2 single-
rooted), whereas species of Brachyodus show a number of
extreme specializations (suppression of i1–2, i3 tusk-like, peg-
like canine, long diastema). The fact that the holotype of A.
africanus designated by Pickford (1991) is a dentary with only
roots of p3 and crowns p4–m3 means that most of the key
diagnostic criteria of Afromeryx are not visible on this specimen.
Therefore, we feel that CGM 30762 cannot serve as the holotype
of a species of Afromeryx. To rectify this situation we have
reassigned CGM 30762 to Brachyodus depereti where it fits
comfortably based on size and known morphology and we have
assigned M15021 as the holotype and currently only know
specimen of a new species of Afromeryx, A. grex.
AFROMERYX PALUSTRIS new species
Type.—CGM 83751, right dentary with symphysis, alveoli i1–
p1, crowns p2–4 (Table 1, Fig. 5B–5D).
Diagnosis.—Differs from A. zelteni and A. grex in being much
larger (m2 area .70%larger than A. zelteni, m2 area .25%
larger than A. grex); further differs from A. grex in having
premolars with weaker pustulate crest development, premolars
lacking distinct and elevated premolar cingulids, taller and more
sharply tapering p2–3, p2 with better developed anterior crest. All
premolar crowns have very finely wrinkled enamel.
Etymology.—"Palustris,’’ marshy or swampy, from the Latin
‘‘palus,’’ marsh, in recognition of the anthracothere habitat at
Occurrence.—Early Miocene, Wadi Moghra, Egypt.
Material.––DPC 2542, right dentary with symphysis, p3–m1;
DPC 4244 right dentary p2–3; DPC 4604, left dentary p4–m3;
DPC 12593, right dentary dp4–m1, m2 in crypt.
Description.—The dentary, as in other anthracotheres, has an
unfused symphysis. It angles inferioposteriorly and is relatively
short, not extending beyond p1. The dentary is deepest just
posterior to m3 and has a rounded angle. The dentary tapers
gently to the level of m1–p4 and then tapers more dramatically
Incisors and canine: alveoli for three lower incisors are present,
and judging from their size these would have been small teeth.
The incisors are followed by an alveolus for a relatively small
canine, which is separated from the single-rooted and relatively
small p1 (see Fig. 5D) by a modest diastema (ranging from 20–30
mm in length).
p2: relatively robust with a tall and tapering protoconid, a
convex labial surface, and concave lingual surface. The labial
aspect is covered with crenulated enamel that extends lingually
around the anterior and posterior surfaces. There is a well-
developed anterior crest extending from the tip of the protoconid
along the anterolingual margin—there are three well-developed
accessory cuspules arrayed down the lingual surface of the crest.
The posterior portion of the tooth is broken but it is clear that
there was no talonid development. A weak and low lingual
cingulid is present but it extends only about half the length of the
tooth, from the anterior crest posteriorly. Weak antero- and
posterolabial cingulids are present.
p3: is generally similar to p2 but differs in several important
respects. Both the anterior and posterior surfaces are more
extensive with the posterior surface having a strong ridge
extending down the crown to reach a relatively well developed
cingulid. In addition, the cingulid wraps around the base of the
tooth both labially and lingually, but it is not continuous with a
similarly developed but less extensive anterior cingulid. As on p2,
there is no talonid development but the posterior cingulid on p3 is
better developed and forms a weak shelf. A heavy anterior crest
extends to the base of the tooth—this crest is weakly cuspate,
showing development of bulbous but indistinct cuspules formed
along its length. There is no distinct paracristid extending to the
protoconid. A slightly weaker and less robust posterior crest
extends to the tooth margin. This posterior crest is not cuspate
and, while robust, is not as heavy as the anterior crest. A narrow,
deep, dorsoventrally elongate and mesiodistally constricted valley
extends between the anterior and posterior crests. A weak, low
974 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
FIGURE 5––1,Afromeryx grex, type specimen (M 15021) in lateral view; 2–4,Afromeryx palustris, type specimen (CGM 83751), in lateral, medial, and
occlusal views, respectively. Arrow indicates alveolus for single-rooted p1. Scale¼10 cm.
MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 975
and short lingual cingulid closes the base of this valley. The tooth
crown is covered by complex, rugose enamel.
p4: is a more elongate version of p3, differing principally by
having a distinct and broad talonid region that extends posteriorly
as a flat shelf. The talonid shelf is weakly divided by a vague crest
extending down the posterior surface and bifurcating with one
branch extending to the talonid and the other terminating at the
metaconid. The anterior and posterior lingual crests are slightly
less robust than on p3 and are more angled anteriorly and
posteriorly, respectively, producing a less deep, but equally
extensive, lingual valley (CGM 83751, DPC 2542). However, this
development appears variable as other specimens (e.g., DPC
4604) show more closely juxtaposed anterior and posterior crests
that turn toward each other dorsally, producing a very narrow and
restricted lingual valley. In DPC 2542 there are no cuspules
developed on the anterior and posterior crests, while in DPC 4604
and CGM 83751, bulbous cuspules are present. The lingual
cingulid is better developed on p4 than the other premolars, and
produces a sloping lingual shelf that closes the base of the lingual
valley. p4 has no crest connecting the paraconid and protoconid,
but there is a weak crest connecting the metaconid to the apex of
the protoconid. A much better developed anterior cingulid that
forms a low but distinct shelf is present on p4.
Molars: the molars of Afromeryx palustris are poorly
represented but, where known, differ little from a common
bunoselenodont anthracothere pattern. Lower first and second
molars have trigonids and talonids of equal size and disposition.
The cristid obliqua nearly reaches the lingual border producing a
deep hypoconid notch that is closed ventrally by a hypocristid.
Lingual cusps are aligned along the lingual margin, except for the
small hypoconulid that is positioned lingual of center. There are
weak pre- and postcristids but these are not labially continuous
with the hypocristid. No lingual cingulids are present. The third
lower molar appears to be a more robust version of m2, but with a
large hypoconulid extension that forms a weakly oblique, single
loop that is separated from the talonid by a deep labial valley.
Like the premolars, all molars are covered by finely wrinkled
Remarks.—Material assigned to A. palustris resembles other
species of Afromeryx in having an unspecialized symphyseal
region, a single-rooted p1, and in placement of the mental
foramina. At present there are few comparable parts across all
species, but the three species of Afromeryx (A. zelteni, A. grex, A.
palustris) differ greatly in size, premolar proportions, and
elaboration of the premolar series.
To investigate more fully the nature of the relationship between
the larger (A. palustris) and smaller (A. grex) specimens from
Moghra, we looked at m2 dimensions. Specifically we examined
the square root of m2 area in a range of extant male and female
artiodactyl species, to test hypotheses about whether the size
difference between the m2 of A. palustris and that of A. grex was
on a par with what might be expected for males and females
within a species, two species in a genus, or perhaps between
individuals in two different genera.
In general, artiodactyls are not particularly sexually dimorphic
in the size of their postcanine dentition. Instead, sexual
dimorphism in artiodactyls is primarily manifest in body size
differences, elaboration of horns or antlers, and sometimes also in
canine size and pelage characteristics (e.g., presence of a dewlap).
However, we chose to use m2 dimensions partially because m2
size was one of the few attributes available in both A. palustris
and A. grex, but also because the postcanine dentition in
artiodactyls is morphologically conservative with regard to sexual
dimorphism, so that a large degree of difference between males
and females in this feature would be a clear result.
Table 3 presents summary statistics involving m2 proportions
in 14 artiodactyl species compared with the same proportions in
A. grex and A. palustris. The raw data from which these
calculations are derived are available as Appendix 1. Table 3
shows the average m2 area square root for males divided by that
of females. This ratio is used to approximate the degree of m2
sexual dimorphism present among the extant sample. The values
for the extant sample are then compared with the m2 square root
ratio of A. palustris (DPC 4604) versus A. grex (M 15021).
Results show that the ratio of m2 size in A. palustris to A. grex is
outside what might be expected for males and females in a single
artiodactyl species. The ratio for A. palustris to A. grex is about
1.17 while members of the extant sample fall within a fairly
restricted range of 0.92 to 1.08. The ratio of A. palustris to A. grex
also exceeds what might be expected for two species in the same
genus, as the values for Kobus ellipsiprymnus versus K. kob were
0.92 to 1.04, and Hippotragus equinus versus H. niger ranged
from 0.99 to 1.03. In fact, based on the data presented here, the
ratio of A. palustris m2 area to that of A. grex is even greater than
might be expected for members of two different artiodactyl
genera. However, we recognize A. palustris and A. grex as two
species of Afromeryx on the basis of anterior tooth morphology,
and also because the ratio of m2 size in A. palustris versus A. grex
does not exceed that which might be expected between a large
male of one species and a small female of a second species within
TABLE 3—Male and female m2 proportions (in millimeters) for fourteen artiodactyl taxa compared with Afromeryx grex and A. palustris. Abbreviations: M and
F¼male and female sample sizes; M/F¼mean male m2 divided by mean female m2; SQRT¼square root of M/F; AP/AG¼m2 value for A. palustris (AP) divided
by the m2 value for A. grex (AG); SQRT¼square root of AP/AG.
Genus Species M/F
m2 area M/F M/F SQRT AP/AG
Bison bison 5/5 617.38 555.8 1.11 1.05
Alces alces 3/3 581.9 563.1 1.03 1.01
Cervus elaphus 5/5 529.37 464.42 1.14 1.07
Rangifer tarandus 5/5 216.21 199.51 1.08 1.04
Syncerus caffer 5/5 394.45 371.07 1.06 1.03
Oryx gazella 5/5 341.54 300.17 1.14 1.07
Ovis canadensis 5/5 180.8 164.89 1.1 1.05
Sus scrofa 5/5 288.93 263.2 1.1 1.05
Kobus ellipsiprymnus 4/4 240.97 281.82 0.86 0.92
Kobus kob 5/5 154.1 142.53 1.08 1.04
Hippotragus niger 5/5 314.37 298.18 1.05 1.03
Hippotragus equinus 4/4 352.59 356.71 0.99 0.99
Taurotragus oryx 3/3 603.64 519.30 1.16 1.08
Odocoileus virginianus 5/5 146.88 138.21 1.06 1.03
Afromeryx grex 1 639.11
Afromeryx palustris 1 868.49 1.36 1.17
976 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
the same genus. For example, Appendix 1 shows that the m2 ratio
of a large male Hippotragus equinus (NMNH 164790) compared
with that of a small female of H. niger (NMNH 545318) is about
1.2, and a large male Kobus ellipsiprymnus (NMNH 61731)
compared with a small female of K. kob (NMNH 164784) is about
1.54, ratios that are either on a par with or exceed the ratio of 1.17
observed for A. palustris to A. grex.
JAGGERMERYX new genus
Type species.—Jaggermeryx naida new species by monotypy.
Diagnosis.—As for species.
Etymology.—‘‘Jagger’’ for Sir Mick Jagger, in recognition of
his famous lips, and meryx’’ Gr. ‘‘ruminant,’’ a common suffix for
Occurrence.—Early Miocene, Wadi Moghra, Egypt.
JAGGERMERYX NAIDA new species
Type.—DPC 2499, left dentary, alveoli i3-c, and partial crowns
of p2–4 (Fig. 6A–6C).
Diagnosis.—Differs from all other anthracotheres in having
multiple (six to ten) mental foramina present on both sides of the
symphysis, inferior boarder of symphysis inflected inferiorly.
Further differs from Brachyodus in having dental formula 18.104.22.168,
and lacking specializations of the anterior dentition such as
suppression of i1–2, or development of a tusk-like i3. Differs
from Afromeryx in having a deeper mandibular symphysis and in
lacking p1. Further differs from Sivameryx in being of larger size,
and in having a bunodont rather than selenodont dentition.
Description.—The most characteristic feature of this taxon is
the presence of multiple mental foramina. These foramina vary in
size, but extend posteriorly from below i1/i2 to p4, most typically
with four in a rhomboid configuration under i1–3, two superiorly
and two inferiorly, and with the medial and inferior foramen
much larger than the other three. Three additional foramina are
usually present below c–p2, and at least one is positioned below
p4. The symphysis is unfused, rugose, heavily constructed and
broad across its base but tapers superioanteriorly. Interestingly,
although the symphysis remains unfused, one specimen (CUWM
115) shows an anomaly whereby plastic deformation during
FIGURE 6––1–3,Jaggermeryx naida, type specimen (DPC 2499) in lateral, medial, and occlusal views, respectively; 4, comparison of probable female CUWM
1578b (upper) and male CUWM 115 (lower) dentaries of Jaggermeryx naida;5,Bothriogenys fraasi (SMNS 44170) from the early Oligocene, Fayum, Egypt
(see text for discussion). Arrow indicates multiple mental foramina. Scale¼10 cm.
MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 977
fossilization pressed the right and left mandibular halves together
in such a way that, at first glance the specimen appears to have
four left incisors and two right ones. The ventral boarder of the
symphysis in J. naida curves slightly ventrally so that the
symphysis is angled ventrally at about 45–488. The symphysis
(see Fig. 6D) can be twice as deep in males (~65–80 mm) as in
females (~42 mm).
Incisor alveoli are of approximately equal size, although in
some cases the i3 alveolus is slightly larger than for i1–2, and is
about the same size as the canine alveolus. Small diastemata
separate alveoli of i1–3 (~3 mm), and i3–c (9 mm). The species is
sexually dimorphic with females having a shallower symphysis
and a shorter c–p2 diastema (ranging from ~16–21 mm, e.g.,
CUWM 1578b), and males having a deep symphysis and a c–p2
diastema up to twice the length of that seen in females (~45 mm,
e.g., CUWM 115).
As represented on the type specimen, some but not all
individuals attributed to this taxon have an unusual morphology
associated with the anterior dentition, whereby the lower incisors
and canine seem to be ‘‘perched’’ on the lateral edge of the
dentary rather than situated perpendicular to the cheek tooth row
(Fig. 6C). Also, where known, lower i1–2 alveoli are slightly
procumbent anteriorly, and i3 is procumbent anterolaterally. One
explanation for the ‘‘splay’’ of the lower anterior teeth is that, in
addition to the normal cropping function of artiodactyl incisors,
the anterior dentition may have had a scoop-like or sieve-like
Etymology.—"Naida’’ derived from the Greek ‘‘Naias,’’ water-
nymph, in recognition of the species’ semi-aquatic habitat.
Material.—CGM 30775, edentulous left dentary symphysis,
alveoli for i2–3,c, p2–4; CGM 30885, right symphysis fragment;
CGM 83752, left dentary alveoli i3–c, roots of p2–3, partial roots
of p4; CUWM 115 edentulous dentary complete symphysis,
alveoli left i1–p2, alveoli right i1–p2; CUWM 157a, b right
dentary p3–4, m3; DPC 17702, right symphysis fragment; DPC
17738, right dentary alveoli i1–3,c, p2–4 (Table 1).
Remarks.—The type specimen of J. naida preserves only part
of the premolar series and is otherwise edentulous as is most of
the hypodigm, which is less than desirable. However, all of the
specimens assigned to J. naida show a number of distinctive
morphologies that individually—and certainly collectively—are
unknown among other anthracothere taxa.
Interestingly, one of the J. naida specimens has been known for
a long time because it was figured in Fourtau (1920, Fig. 31) as an
unnumbered dentary of Brachyodus africanus. Pickford (1991)
subsequently included this specimen as part of the hypodigm of B.
mogharensis, but we assign this specimen here (now carrying the
number CGM 30775) to J. naida, owing to the presence of
multiple mental foramina arranged in the characteristic pattern, as
well as the unusual canted morphology of the symphyseal region.
When only one unnumbered specimen collected almost a century
ago was known, it was possible to consider the unusual
morphology as representing idiosyncratic variation. However,
sustained collecting efforts at Moghra over the last decade have
recovered additional specimens with the same unusual morphol-
ogy, making clear that these specimens represent a new
The occurrence of a large number of mental foramina present
bilaterally in this species is unique among the Moghra
anthracotheres. Because the mental foramen transmits the mental
nerve, which provides sensory innervation to the chin and lower
lip, we interpret the morphology of J. naida as indicating that the
animal must have had either a very sensitive lower lip or an
acutely sensitive tactile snout, perhaps combined with perceptive
vibrissae. Also, the spacing of the lower incisor alveoli, and the
fact that the symphyseal region shows slight splay anteriorly,
suggests that the anterior lower dentition may have functioned as
a kind of scoop-like or sieve-like structure. This morphological
information, together with sedimentological data showing that the
Moghra deposits formed under wet conditions, and preliminary
results from stable isotope analyses indicating that many of the
Moghra anthracotheres, including J. naida, were semiaquatic,
leads to the conclusion that J. naida may have foraged along the
river banks, using its sensitive tactile snout, and employing the
lower anterior teeth in a cropping and scoop-like fashion.
ANTHRACOTHERIIDAE species indeterminate
The following specimens from Wadi Moghra are placed in an
indeterminate group because either the fragmentary nature of the
material, extreme wear, serious abrasion, or a combination of all
three prevents assigning the specimen to a known taxon. DPC
2504, left dentary fragment; DPC 3629, left maxilla M2–3; DPC
4245, edentulous partial palate left P2–3, right P2–4; DPC 4421,
left dentary dp4–m2, p4, p4 in crypt; DPC 4434 right dentary m3
fragment; DPC 4437, left dentary p2, dp3–4; DPC 4585, left
dentary fragment with tooth fragments; DPC 6426, right m3; DPC
8932, right edentulous dentary fragment; DPC 8933, right MX;
DPC 8952, right edentulous dentary fragment; DPC 12593, left
dentary symphysis, c, dp4–m1, m2 in crypt; DPC 12930, left
dentary dp3–4; DPC 14576, left dentary dp3–4, m1 in crypt; DPC
14577, right m3 fragment; DPC 17705, left edentulous dentary
fragment; DPC 17734, right dentary with m3.
Six anthracothere species in four genera are known from
Wadi Moghra. The two species of Brachyodus;B. depereti,and
B. mogharensis likely represent late surviving relatives of the
older African anthracothere lineage Bothriogenys known from
late Eocene–early Oligocene deposits of the Fayum, Egypt (see
also Pickford, 1991). Brachyodus depereti is a very large form
with a highly specialized anterior dentition, and from what is
known about B. mogharensis, the species appears to be a smaller
version of B. depereti, which suggests the two species share a
Jaggermeryx naida is also a very specialized anthracothere.
Although the unusual features of the dentary have not been
documented in any other Moghra species, SMNS 44170, a left
dentary with i3, p2–3 from the Fayum, identified as Bothriog-
enys fraasi, has multiple mental foramina in a pattern similar to
that observed for J. naida. This suggests that J. naida may also
be rooted among earlier African anthracotheres from the Fayum
Sivameryx moneyi,Afromeryx grex,andA. palustris are likely
early Miocene immigrants from Eurasia. As Pickford (1991)
noted, the pustulate crests on the premolars of Afromeryx are
reminiscent of those observed in Gonotelma from Pakistan, and
Sivameryx has clear relatives in India. Both of these taxa likely
arrivedinAfricaaspartofthewell-documented early Miocene
faunal influx of Eurasian animals permitted by the collision of
Afro-Arabian and Turkish plates.
Diagnostic features of African anthracotheres have been
detailed most recently in Holroyd et al., (2010) (see also,
Pickford, 1991 and Black, 1978). However, it is useful to
emphasize here that within Anthracotheriidae, species differ
from each other primarily in specializations of the anterior
dentition, size, and degree of brachyodonty versus selenodonty.
Anthracothere taxa certainly differ in molar morphology, but
overall, molar morphology can be conservative even when size
differences and specializations of the anterior dentition are
considerable. In addition, although the situation is improving,
little concrete information is available about the extent of sexual
978 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
dimorphism among anthracothere species, although the litera-
ture contains some cautionary tales about underestimating the
effects of dimorphism (e.g., Pickford, 2006).
In general, the Moghra fauna, including the suite of
anthracothere species discussed here, appears to have been
adapted to very wet conditions. Sedimentological work indicates
that the fossiliferous deposits at Moghra were formed in a
marginal marine coastal realm, at the distal end of a large
fluviatile system (Abdel Gawad, 2010; Abdel Gawad et al.,
2012; Hassan et al., 2012; Miller et al., 2006). Also taphonomi-
cally, anthracothere remains are commonly associated with
abundant coprolites (mainly crocodile), as well as freshwater
(catfish, gavial) and marine elements (shark teeth), indicating
that at least some Moghra sediments must represent slow-
moving fluviatile, estuarine or lagoonal deposits. Preliminary
results from stable isotope analyses also support the interpre-
tation of the Moghra environment based on other information
(Clementz et al., 2010), with initial results indicating that
Brachyodus,Sivameryx Afromeryx,andJaggermeryx all occu-
pied closed and semiaquatic environments.
We thank the people and Government of Egypt for permission
to work at Wadi Moghra, as well as the Director and staff of the
Cairo Geological Museum for access to specimens in their care.
We thank J. M. Plavcan for statistical advice regarding the extant
artiodactyl sample, J. Hooker for help in the collections, R.
Ciochon and J. Nichols for help with photographs, B. Miljour for
help with preparation of Figure 1, W. J. Sanders for his skill in
fossil preparation of the skull, and the editor and two anonymous
reviewers for productive comments. NSF grants OISE-0403472
and EAR-0808283 supported this research. This Duke Lemur
Center publication #1259.
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MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 979
APPENDIX—Artiodactyl sample used for comparison with Afromeryx grex and A. palustris. All specimens measured (mm) at NMNH. Abbreviations: L¼length;
Genus Species Specimen Sex
Bison bison 125357 M 34.86 19.92 694.411
Bison bison 123686 M 34.73 15.9 552.207
Bison bison 115282 M 32.71 21.25 695.088
Bison bison 122699 M 31.61 17.31 547.169
Bison bison 63363 M 31.18 19.18 598.032
Bison bison 120578 F 31.87 16.29 519.162
Bison bison 120580 F 30.62 15.64 478.897
Bison bison 102039 F 33.14 20.08 665.451
Bison bison 122606 F 29.86 17.62 526.133
Bison bison 126622 F 31.6 18.65 589.34
Alces alces 201331 M 29.93 20.55 615.062
Alces alces 218000 M 28.06 19.95 559.797
Alces alces 218792 M 26.99 21.15 570.839
Alces alces 15556 F 29.36 19.96 586.026
Alces alces 243970 F 29.87 18.87 563.647
Alces alces 218794 F 26.74 20.18 539.613
Cervus elaphus 1881 M 29.43 18.2 535.626
Cervus elaphus 2016 M 30.41 16.72 508.455
Cervus elaphus 2917 M 29.23 17.26 504.51
Cervus elaphus 2011 M 29.4 19.62 576.828
Cervus elaphus 2617744 M 31.62 16.49 521.414
Cervus elaphus 2896 F 27.5 16.68 458.7
Cervus elaphus 3155 F 27.39 16.71 457.687
Cervus elaphus 26021 F 28.61 16.75 479.218
Cervus elaphus 199490 F 27.63 17.1 472.473
Cervus elaphus 265025 F 27.94 16.25 454.025
Rangifer tarandus 205754 M 21.02 10.51 220.92
Rangifer tarandus 235361 M 17.98 11.41 205.152
Rangifer tarandus 206348 M 21.78 10.01 218.018
Rangifer tarandus 223750 M 20.44 10.45 213.598
Rangifer tarandus 199466 M 19.06 11.72 223.383
Rangifer tarandus 268011 F 18.39 10.76 197.876
Rangifer tarandus 174506 F 18.13 10.08 182.75
Rangifer tarandus 205755 F 19.66 10.38 204.071
Rangifer tarandus 199543 F 18.61 10.64 198.01
Rangifer tarandus 217428 F 20.04 10.72 214.829
Syncerus caffer 220129 M 25.85 14.21 367.329
Syncerus caffer 220131 M 26.11 14.54 379.639
Syncerus caffer 220134 M 22.25 14.08 313.28
Syncerus caffer 220295 M 28.06 14.59 409.395
Syncerus caffer 161943 M 28.72 17.5 502.6
Syncerus caffer 218478 F 23.64 12.54 296.446
Syncerus caffer 219060 F 25.72 13.06 335.903
Syncerus caffer 220130 F 25.89 13.79 357.023
Syncerus caffer 220137 F 23.76 15.68 372.557
Syncerus caffer 163311 F 26.89 18.35 493.432
Oryx gazella 368528 M 27.57 14.82 408.587
Oryx gazella 199091 M 24.17 12.87 311.068
Oryx gazella 267605 M 22.79 12.87 293.307
Oryx gazella 296154 M 26.83 14.08 377.766
Oryx gazella 163216 M 23.53 13.47 316.949
Oryx gazella 163210 F 22.48 13.22 297.186
Oryx gazella 184804 F 23.32 12.65 294.998
Oryx gazella 182130 F 23.83 11.51 274.283
Oryx gazella 163220 F 23.41 13.15 307.842
Oryx gazella 163213 F 24.08 13.56 326.525
Ovis canadensis 174512 M 17.25 9.54 164.565
Ovis canadensis 205154 M 18.53 8.76 162.323
Ovis canadensis 205155 M 19.81 11 219.297
Ovis canadensis 210208 M 18.34 9.38 172.029
Ovis canadensis 209416 M 18.16 10.23 185.777
Ovis canadensis 205157 F 19.02 11.2 213.024
Ovis canadensis 205156 F 18.59 7.58 140.912
Ovis canadensis 81803 F 17.71 8.73 154.608
Ovis canadensis 174513 F 17.72 8.7 154.164
Ovis canadensis 240319 F 17.89 9.04 161.726
Sus scrofa 143541 M 20.16 14.87 299.779
Sus scrofa 144365 M 18.35 13.27 243.505
Sus scrofa 144302 M 18.63 15.13 281.872
Sus scrofa 538818 M 18.11 14.17 256.619
Sus scrofa 283111 M 22.14 16.39 362.875
Sus scrofa 114179 F 17.59 13.28 233.595
Sus scrofa 141166 F 20.89 15.71 328.182
Sus scrofa 141028 F 19.05 13.81 263.081
Sus scrofa 114177 F 18.04 12.97 233.979
Sus scrofa 114282 F 19.95 12.89 257.156
Kobus ellipsiprymnus 20851 M 19.31 9.95 192.135
Kobus ellipsiprymnus 61731 M 24.49 12.15 297.554
980 JOURNAL OF PALEONTOLOGY, V. 88, NO. 5, 2014
Genus Species Specimen Sex
Kobus ellipsiprymnus 164689 M 21.71 11.36 246.626
Kobus ellipsiprymnus 14997 M 21.55 10.56 227.568
Kobus ellipsiprymnus 173877 F 22.7 11.73 266.271
Kobus ellipsiprymnus 20853 F 20.35 13.47 274.115
Kobus ellipsiprymnus 164793 F 21.42 11.94 255.755
Kobus ellipsiprymnus 237000 F 21.2 15.62 331.144
Kobus kob 199248 M 14.78 10.06 148.687
Kobus kob 199230 M 15.58 9.34 145.517
Kobus kob 199208 M 17 9.39 159.724
Kobus kob 239166 M 14.15 9.76 138.104
Kobus kob 164774 M 16.42 10.87 178.485
Kobus kob 164784 F 14.71 8.54 125.623
Kobus kob 164787 F 14.5 10.91 158.195
Kobus kob 164778 F 14.89 9.56 142.348
Kobus kob 164781 F 14.93 9.36 139.745
Kobus kob 164499 F 14.75 9.95 146.763
Hippotragus niger 61736 M 24.13 13.42 323.825
Hippotragus niger 21637 M 25.73 14.52 373.6
Hippotragus niger 252518 M 24.23 12.91 312.809
Hippotragus niger 198367 M 21.15 13.02 275.373
Hippotragus niger 218780 M 21.41 13.37 286.252
Hippotragus niger 384208 F 22.47 14.55 326.939
Hippotragus niger 196922 F 22.61 13.78 311.566
Hippotragus niger 580474 F 23.26 13.38 311.219
Hippotragus niger 545318 F 22.73 11.52 261.395
Hippotragus niger 396597 F 22.4 12.49 279.776
Hippotragus equinus 164790 M 23.96 15.68 375.693
Hippotragus equinus 181885 M 24.65 13.88 342.142
Hippotragus equinus 197982 M 23.27 14.48 336.95
Hippotragus equinus 199100 M 25.04 14.2 355.568
Hippotragus equinus 252707 F 24.76 15.59 386.008
Hippotragus equinus 164806 F 24.44 14.46 353.402
Hippotragus equinus 164805 F 22.85 15.57 355.775
Hippotragus equinus 164740 F 23.39 14.18 331.67
Taurotragus oryx 122741 M 35.99 18.45 664.016
Taurotragus oryx 163225 M 30.04 20.06 602.602
Taurotragus oryx 163308 M 31.59 17.23 544.296
Taurotragus oryx 162015 F 30.58 17.84 545.547
Taurotragus oryx 163224 F 27.53 17.98 494.989
Taurotragus oryx 260287 F 29.82 17.35 517.377
Odocoileus virginianus 285519 M 14.58 9.57 139.531
Odocoileus virginianus 217267 M 14.05 10.18 143.029
Odocoileus virginianus 285517 M 13.85 9.24 127.974
Odocoileus virginianus 271872 M 15.42 10.1 155.742
Odocoileus virginianus 254657 M 15.37 10.94 168.148
Odocoileus virginianus 274606 F 14.56 9.67 140.795
Odocoileus virginianus 285516 F 14.97 9.44 141.317
Odocoileus virginianus 265634 F 15.61 9.78 152.666
Odocoileus virginianus 265593 F 13.82 9.16 126.591
Odocoileus virginianus 265592 F 13.75 9.43 129.663
Afromeryx grex 15021 30.86 20.71 639.111
Afromeryx palustris 83751 34.26 25.35 868.491
MILLER ET AL.—ANTHRACOTHERES FROM MOGHRA 981