Palaeontologia Electronica

palaeo-electronica.org

Deciduous dentition and dental eruption sequence of

Bothriogenys fraasi (Anthracotheriidae, Artiodactyla)
from the Fayum Depression, Egypt
Hesham M. Sallam, Afifi H. Sileem, Ellen R. Miller, and Gregg F. Gunnell

ABSTRACT

Paleogene anthracotheres are poorly documented from Afro-Arabian localities.

This is due, in large part, to the fragmentary nature of the specimens that have been
described. However, sediments in the Jebel Qatrani Formation, Fayum Depression,
Egypt, preserve the richest anthracothere assemblage in all of Afro-Arabia. Unlike
other samples, the Fayum collection includes many complete dentitions, skulls, and
partial skeletons. Based on these extensive collections, this study provides the first
description of the complete deciduous dentition and dental eruption sequence for the
early Oligocene anthracothere Bothriogenys fraasi. A detailed discussion concerning
the pattern and timing of dental growth in B. fraasi is provided, and the ontogenetic
sequence documented for B. fraasi is compared with those available for suoids and
hippos, the two extant groups currently considered as possible sister taxa to anthraco-
theres. Results show that anthracotheres and suoids share a more similar dental emer-
gence pattern, and one that may be close to the primitive condition for artiodactyls,
while hippos have a very different dental eruption sequence as a consequence of their
highly divergent life history pattern. As a growing body of life history research indicates
that taxa in close phylogenetic proximity may be expected to share features of their
dental developmental pattern, this finding suggests a useful test of competing hypothe-
ses of a relationship between Anthracotheriidae and either Hippopotamidae or
Suiformes can potentially be developed based on eruption patterns.

Hesham M. Sallam, Mansoura University Vertebrate Paleontology Center, Department of Geology,

Mansoura University, Mansoura, 35516, Egypt, and Department of Evolutionary Anthropology, Duke
University, Durham, NC 27708, USA sallam@mans.edu.eg
Afifi H. Sileem, Vertebrate Paleontology Section, Cairo Geological Museum, Cairo, Egypt,
afifi.sileem@yahoo.com
Ellen R. Miller, Department of Anthropology, Wake Forest University, Winston-Salem, NC 27106, USA,
millerer@wfu.edu
Gregg F. Gunnell, Division of Fossil Primates, Duke Lemur Center, Durham, NC 27705, USA,
gregg.gunnell@duke.edu

Keywords: Fayum; Egypt, Oligocene; juvenile; dentition; Artiodactyla

Submission: 13 February 2016 Acceptance: 16 August 2016

Sallam, Hesham M., Sileem, Afifi H., Miller, Ellen R., and Gunnell, Gregg F. 2016. Deciduous dentition and dental eruption sequence
of Bothriogenys fraasi (Anthracotheriidae, Artiodactyla) from the Fayum Depression, Egypt. Palaeontologia Electronica 19.3.38A: 1-17
palaeo-electronica.org/content/2016/1595-anthracotheriidae-from-egypt

Copyright: Palaeontological Association October 2016

SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

INTRODUCTION Anthracotheriidae is an extinct family of artio-

dactyls known from Eocene – Miocene deposits
The terrestrial mammal-bearing localities of
across Laurasia and into Africa (but not elsewhere
the Jebel Qatrani Formation, Fayum Depression,
in Gondwana). Behaviorally, they are thought to
Egypt, including the well-known quarries L-41, A,
have occupied a range of browsing niches over
B, E, V, I, and M, are among the most productive
both their long temporal and broad geographic
Paleogene fossil localities known anywhere in
ranges, although whatever their specific feeding
Africa (Figure 1). The Jebel Qatrani deposits range
ecologies were they are widely acknowledged to
in age from late Eocene through early Oligocene
have been semi-aquatic based on skeletal propor-
(~34 to ~30 Ma) (Kappelman et al., 1992; Seiffert,
tions and inferred habitats where fossils are found
2006; Seiffert et al., 2008), and fossils recovered
(e.g., Black, 1978; Pickford, 1991; Holroyd et al.,
from the Jebel Qatrani have profoundly influenced
2010; Miller et al., 2014). As a group, identification
our understanding of primate (e.g., Seiffert et al.,
of the closest living anthracothere relative within
2010; Seiffert, 2012), and mammalian evolution
Artiodactyla is contested, with some researchers
(e.g., Simons and Rasmussen, 1990; Simons,
favoring a suiform affiliation (e.g., Pickford, 2007),
2008). However, relatively few of these mammalian
and others supporting a closer relationship with
studies have focused on anthracotheres (Andrews,
extant Hippopotamidae (e.g., Black, 1978; Bois-
1906; Schmidt, 1913; Black, 1978; Holroyd et al.,
serie and Lihoreau, 2006; O’Leary et al., 2012).
1996; Ducrocq, 1997; Sileem et al., 2015; Sileem
Much of the recent work on anthracotheres
et al., 2016), despite the fact that anthracotheres
has been concerned with establishing their alpha
are among the most abundant fossils recovered
taxonomy, as reflected in observable differences
from Fayum deposits.

FIGURE 1. 1, location map of the Jebel Qatrani area, Fayum Depression. 2, stratigraphic positions and age esti-
mates for major mammal-bearing fossil localities, following Seiffert (2006), EOB is abbreviation for Eocene Oligocene
Boundary. 3, map of Jebel Qatrani area, showing different rock units, common landmarks and the approximate posi-
tion of anthracothere-bearing fossil localities.

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 PALAEO-ELECTRONICA.ORG

among adult dental morphologies (e.g., Lihoreau material, there is no evidence for P1/1 replacement
and Ducrocq, 2007, but for exceptions see Pick- suggesting that Bothriogenys fraasi retained a dP1/1
ford, 2006; 2008). However, many of these investi- into adulthood. Morphology is not especially helpful
gations have been conducted without adequate to determine whether or not the retained tooth is
material. The mandibular and maxillary material
deciduous or permanent because, unlike dP3/3 and
available in many museum collections preserves
jaws with only molars and premolars, while many dP4/4, dP2/2 is neither molariform or of an exagger-
of the most striking differences among anthracoth- ated form but more like permanent P2/2 suggesting
ere taxa are manifest in their anterior dentition that the first deciduous premolar would be simple
(e.g., presence/absence of diastemata, reduced v. as well.
caniniform canines, relative size, and orientation of It is not uncommon for mammals (some pigs,
incisors), even when the morphology of the cheek hippopotamus, horses, hyraxes) to retain decidu-
teeth remains more conservative. ous first premolars that are not replaced (Zeigler,
In contrast, the collection of anthracothere 1971; Luckett, 1993; Smith, 2000). Only Tapirus
material from the Jebel Qatrani Formation is com- and maybe some archaeocete whales show evi-
prised of a large number of anthracothere speci- dence of replacement of the first premolar among
mens (N= ~2270, including isolated teeth and mammals and in the case of the latter the evidence
postcranial elements), that are currently recog- is not compelling (Luckett, 1993; Uhen, 2000).
nized as representing six species in three genera. Therefore, the first upper and lower premolars of
A fairly large proportion of these preserve at least Bothriogenys fraasi are here considered to be
part of the anterior dentition and 12% (N=33/270) retained deciduous teeth.
of the specimens attributed to B. fraasi are juve- All specimens are cataloged in the Duke University
niles. Lemur Center, Division of Fossil Primates and desig-
Here we provide the first documentation of the nated with the prefix DPC. The studied materials were
deciduous dentition and dental eruption sequence scanned using a Nikon XT H 225 ST micro-CT scanner
of the anthracothere, Bothriogenys fraasi, a taxon housed at Duke University’s Shared Materials Instru-
known from the early Oligocene upper sequence in mentation Facility, and three-dimensional reconstruc-
the Jebel Qatrani Formation. This work represents tions were rendered in Avizo v.8. Dental measurements
the first detailed account of an ontogenetic were taken from digital surface models (in Stanford “ply”
sequence for any anthracothere taxon and is made format). All three dimensional (3D) digital materials are
possible by the decades of fieldwork in the Fayum available for viewing and direct download at www.mor-
that recovered such a large sample of remarkably phosource.org, (morphosource.org/index.php/Detail/Pro-
well-preserved juvenile specimens. Results from jectDetail/Show/project_id/224). See also Table 1 for
this investigation into the pattern and timing of den- digital object identifiers, DOIs that directly link to 3D dig-
tal growth in B. fraasi will serve as a reference for ital media associated with each specimen. More details
comparison with other anthracothere species, and yet are provided in Supplemental Material. Reuse of
provides empirical information from growth and these data should cite MorphoSource, the DOI and this
development towards resolving the phylogenetic paper.
relationships within anthracotheres, and between
anthracotheres and other artiodactyl groups. DESCRIPTION
Upper Deciduous Teeth
MATERIAL AND METHODS
A complete upper and lower deciduous pre-
Tooth position is referenced as dI, dC, dP, P, molar dentition for Bothriogenys fraasi is available
and M (for deciduous incisors, canine, premolars, as a composite from specimens recovered from
and permanent premolars and molars, respec- Fayum Quarries I, M, and O. DPC 5167 (Figure
tively), with upper and lower teeth designated by 3.1-4, Table 1) is a right maxillary fragment with
superscript and subscript numbers (respectively), light post-mortem distortion that has led to minor
such that, for example, dP4 is the fourth upper surface cracks and displacements. However, the
deciduous premolar. We follow the nomenclature specimen clearly preserves the alveolus for dP1,
of Bärmann and Rössner (2011) for dental features
and crowns of dP2-M1, with M1 half exposed. The
(cusps and crests) of upper and lower deciduous
upper deciduous second premolar in DPC 5167 is
premolars (Figure 2).
triangular, has two roots, and is separated from the
In the following we refer to the first premolar of
mesial aspect of dP3 by a very short diastema. The
Bothriogenys as dP1/1. Based on the available

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SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

FIGURE 2. Dental terminology used to describe features of the deciduous premolars of Bothriogenys fraasi, following
Bärmann and Rössner (2011).

crown of the tooth is convex labially and concave the posterior cusp, creating a small inverted V-
lingually, is longer than it is wide, and has an oval shape on the labial border of the tooth. The disto-
occlusal outline, owing to a slightly wider posterior lingual crest of the anterior cusp is straight and
region. The crown bears two main cusps, the ante- short and runs distolingually. The distal crest of the
rior of which is larger and occupies the mesial part posterior cusp has the same length as the mesial
of the tooth, and the posterior cusp is smaller and crest, and runs distally to end as a small projection
occupies the distolabial portion of the crown. The in the distolabial corner of the crown. There is a low
anterior cusp has a pyramid shape with three and weakly developed but broad cingulum extend-
crests, the mesial of which is curved and joins a ing around the lingual margin of the posterior cusp
very short and low anterocingulum. The distolabial that connects distally with the distal crest of the
crest of the anterior cusp is longer than the mesial posterior cusp, and joins the distolingual crest of
crest, and slopes down to meet the mesial crest of

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TABLE 1. Length and width of teeth (in millimeters) of juvenile specimens of Bothriogenys fraasi from the upper most
terrestrial mammal-bearing localities of the Jebel Qatrani Formation, Fayum Depression, Egypt. DOIs representing
microCT data and associated mesh files and photographs on MorphoSource. The DOIs can be also found in Table S1
and should be cited along with this paper and MorphoSource if reused.
Bothriogenys fraasi
Upper dP1 dP2 dP3 dP4 M1
DPC 3224 Right -- -- -- -- -- -- 15.13 15.55 -- --
doi.org/10.17602/M2/M12690
DPC 5167 Right -- -- 11.22 5.18 15.21 10.40 15.00 13.66 18.90 ?
doi.org/10.17602/M2/M12646
DPC 20439 Right -- -- -- -- 16.27 10.65 15.29 14.40 -- --
doi.org/10.17602/M2/M12647

Lower dP1 dP2 dP3 dP4 M1

DPC 2705 Right -- -- 8.86 4.35 16.25 6.89 21.93 9.17 20.82 11.77
doi.org/10.17602/M2/M12688
DPC 3606 Right -- -- -- -- 17.17 6.29 21.49 8.59 -- --
doi.org/10.17602/M2/M12653
DPC 3947 Left -- -- -- -- 17.03 6.23 21.40 8.69 -- --
doi.org/10.17602/M2/M12639
DPC 6147 Left -- -- -- -- -- -- 20.45 8.23 -- --
doi.org/10.17602/M2/M12649
DPC 7706 Right -- -- 8.26 4.08 17.76 6.47 21.56 8.72 20.18 11.81
doi.org/10.17602/M2/M17034
DPC 7730 Left -- -- 9.43 4.41 17.77 6.94 22.54 9.16 21.48 12.15
doi.org/10.17602/M2/M12655
DPC 8638 Right -- -- 7.91 4.05 15.75 6.26 20.31 7.99 19.08 10.29
doi.org/10.17602/M2/M12684
DPC 10616 Left -- -- -- -- -- -- 20.32 7.66 19.41 10.20
doi.org/10.17602/M2/M12650
DPC 10633 Right -- -- -- -- 15.72 5.77 20.27 7.93 -- --
doi.org/10.17602/M2/M12654
DPC 11280 Left -- -- -- -- -- -- 21.24 8.44 19.15 10.64
doi.org/10.17602/M2/M12645
DPC 11407 doi.org/10.17602/M2/ Left -- -- -- -- -- -- 20.38 8.18 18.68 10.63
M12651
DPC 11412 Right -- -- -- -- 15.11 6.51 21.50 8.41 -- --
doi.org/10.17602/M2/M12648
DPC 11416 Left 7.01 2.96 9.20 4.67 16.87 6.54 22.63 8.83 -- --
doi.org/10.17602/M2/M12641
DPC 13570 Left -- -- 9.27 4.30 16.05 6.33 21.17 8.46 -- --
doi.org/10.17602/M2/M17039
DPC 13562 Right -- -- -- -- 15.22 6.07 21.26 8.58 19.06 10.71
doi.org/10.17602/M2/M12638

the anterior cusp mesially. The dP2 is the least what smaller. The anterior cone occupies the most
worn of the upper deciduous dentition. mesial portion of the crown and bears three crests.
The dP3 is well-preserved in DPC 5167 and The lingual crest originates at the apex and runs
DPC 20439 (Figure 3.1-5). The dP3 is longer than lingually, curving distally to end at the base of the
wide, and has a roughly triangular outline with a protocone and so enclosing a lingual anterior val-
narrower anterior portion and a broader posterior ley. There are two labial crests running from the
one. The occlusal surface has five major cusps anterior cone, which together delimit a small fovea
(anterior cone, protocone, paracone, metacone, on the labial border of the anterior cone. The proto-
and metaconule), all of which are more or less cone is the highest cusp in the crown and is situ-
equal in size, and all of which are about the same ated distal to the anterior cone. The protocone has
height, except for the paracone, which is some- a conical shape with a mesial crest that joins the

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SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

FIGURE 3. 1-4, right partial maxilla of DPC 5167 with dP2-M1, in (1) lateral, (2) medial, (3) anterior, and (4) occlusal
views; 5, right partial maxilla of DPC 20439 (doi.org/10.17602/M2/M12647) with dP3- dP4, in occlusal view; 6-8,
right dP4 of DPC 3224 (doi.org/10.17602/M2/M12690), in (6) occlusal, (7) lingual, (8) ventral views; 9, DPC
11416 (doi.org/10.17602/M2/M12641) with dP1- dP4, in occlusal view.

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distal crest of the anterior cone at a level lower base of the paraconule, and there are two postpro-
than the apex, creating a V-shaped crest that helps tocristae that run distolabially. The metaconule
close the anterior valley labially. The postparacrista bears three cristae; the premetacristule runs mesi-
slopes down distolabially to merge with the meso- olabially toward the main valley and connects with
style, and the premetacrista slopes down mesola- the most lingual postprotocrista, interrupting the
bially to connect with the mesostyle. Together course of the main valley. The postmetacristule is
these crests form an inverted V- shape that pro- well developed, runs distolabially, and fuses with
trudes as a mesostyle and weakly closes the pos- the distal cingulum. The lingual metacristule
terior valley labially. The mesostyle is crestiform, becomes a continuation of the mesial portion of the
weakly developed, and has a worn surface on both lingual cingulum. The distal cingulum is moderately
specimens. The metacone is pyramid shaped and developed, extends lingually, and courses around
bears three crests, the distal of which runs distally the lingual base of the metaconule, forming the dis-
and interrupts the posterior cingulum. The tal portion of the lingual cingulum. The mesial cin-
premetacrista is the mostly heavily worn crest on gulum is well developed but low, and runs lingually
the crown. The metaconule is transverse to the from the parastyle. In DPC 5167, the cingulum
metacone and connected to it via a short crest. The courses around the lingual base of the protocone
postmetaconulecrista runs distolabially and ends at and merges with the lingual metacristule, while in
the midpoint of the posterior cingulum. The DPC 3224 and 20439, the cingulum terminates at
premetaconulecrista runs mesially and terminates the mesial base of the protocone. The main valley
at the base of the protocone, closing the posterior is closed lingually either via a low connection
valley lingually. The posterior valley of dP3 is between the base of the protocone and the
broader and longer than the anterior valley. The metaconule, or by a continuous lingual cingulum.
posterior cingulum is relatively well developed, in Lower Deciduous Teeth
particular the labial aspect and starts from the base
of the mesostyle, courses around the posterior DPC 11416 (Figures 3.9, 4) preserves the
margin of the tooth, and terminates at the base of complete lower deciduous premolars series (dP1-4)
the protocone. and several other juvenile mandibular specimens
The dP4 occlusal surface is nearly identical to of Bothriogenys fraasi preserve parts of the decidu-
that of the first upper molar, differing only in being ous series (Table 1).
relatively smaller and having less robust cusps and The dP1 is visible in DPC 11416 but the tooth
crests. DPC 3224 (Figure 3.6-8) is the most well- is not completely erupted. The dP1 is a small, peg-
preserved dP4 in the available material. The dP4 like tooth, with a labiolingually compressed crown
has a semi-quadrate outline and four main cusps and an oval base. The dP1 crown has one main
(paracone, protocone, metacone, and cusp, from which run mesial and distal cristids, the
metaconule), all of which are about equal in size. distal one being longer than the mesial one. The
The mesostyle is well developed, placed slightly tooth is single-rooted, convex laterally, concave
distal of center on the labial wall of the tooth, and medially, and curves distally.
extends farther labially than the parastyle. The lin- The dP2 is a double-rooted tooth with one
gual side of the mesostyle is invaded by the post- main cusp that has its apex in the middle of the
paracrista and premetacrista, extending the crown. The tooth has a triangular shape in lateral
protofossa labially in a V-shape between the view. The labial surface of the crown is convex,
paracone and metacone. There is no cingulum that and the lingual surface is concave. The crown
runs distally from the mesostyle to meet the post- bears two main cristids. The distal cristid in some
metacrista as is seen in M1. The parastyle is dis- cases branches into two cristids, leading to a rela-
tinct and crestiform and forms the mesiolabial tively narrow groove. On the distal border, there is
corner of the crown. The paraconule is distinct, tri- a weakly developed cingulid that extends mesially
angular in shape, and situated at the midpoint onto the lingual border of the crown.
between the paracone and the protocone, from The dP3 is double-rooted molariform with a
which it is separated by deep, narrow notches. The crown length greater than width and has a gener-
preparacristule runs down mesiolabially toward the ally rhomboidal outline. The occlusal surface is
base of the paracone, whereas the postparacris- occupied by four major cusps (anteroconid, proto-
tule runs distally and fades before reaching the conid, entoconid, and hypoconid). The anteroconid
main valley of the crown. The preprotocrista is is concave lingually and convex labially. In some
short and runs mesially to connect with the lingual cases, it bears three cristids, the two mesial ones

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SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

FIGURE 4. Left mandibular fragment of DPC 11416 (doi.org/10.17602/M2/M12641) with dP1- dP4, showing stage I
of the eruption sequence of Bothriogenys fraasi, in (1) lateral, (2) internal, (3) medial, and (4) occlusal views.

of which delimit a small depression similar to the what convex, with low and well-developed lingual
corresponding fovea on the dP3. The mesial cristid cingulid. The hypoconid is relatively large, occu-
of the anteroconid slopes down toward the mesio- pies the distolabial corner of the crown, and has
lingual corner of the crown and forms the very relatively well-developed prehypocristid and post-
mesial tip of the tooth. The distal cristid of the hypocristid, which run mesiolingually and distolin-
anteroconid is relatively weakly developed when gually, respectively, from its tip. The entoconid is
compared with the mesial cristid. The labial cin- the smallest cusp and it is positioned transversely
gulid is low and well developed, and runs from the relative to the hypoconid, from which it is separated
labial base of the protoconid coursing around the by a deep and narrow valley. The preentocristid
anteroconid (in some specimens), and meets with meets the prehypocristid mesially, both of which
the distal cristid of the anteroconid. The protoconid join the postprotocristid at the posterior valley. The
is the largest and tallest cusp and is situated distal postentocristid runs distolabially from the entoco-
to the anteroconid, from which it is separated by a nid to connect with the posthypocristid distally, and
narrow and deep notch. The labial side of the pro- both continue as a single cristid that connects to
toconid is concave, and the lingual side is some- the distal cingulid, forming the distal tip of the

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crown. In some specimens, the entoconid lacks a the distal V-shape pattern of the tooth via a short
postentocristid. The distal cingulid is relatively crestid.
broad and limited to the posterior border of the
tooth. TOOTH ERUPTION SEQUENCE
The dP4 is larger than dP3, it is longer than it is AND COMPARISONS
wide, and the tooth is rhomboidal in occlusal out-
A nearly complete lower tooth eruption
line, with a wide talonid and a narrow trigonid. The
sequence can be documented for Bothriogenys
tooth bears six major cusps (anterolingual conid,
fraasi (Figures 4, 5, 6, 7, 8, Table 2). This eruption
anterolabial conid, metaconid, entoconid, protoco-
sequence can be compared with that of the extant
nid, and hypoconid). On the occlusal surface, the
Hippopotamus (Laws, 1968), and the European
lingual cusps align mesodistally, and are slightly
wild boar Sus scrofa (Matschke, 1967). Information
higher than the labial cusps. The mesial sides of
about the dental eruption sequence for both taxa is
the latter cusps show some wear in specimens of
available, and both taxa represent higher level
more advanced aged. On the lingual border of the
groups that have been frequently cited as being
tooth, there are two deep V-shaped notches sepa-
sister taxa to or derived from anthracotheres (Geis-
rating the lingual cusps, while on the labial border,
ler and Uhen, 2003; Pickford, 2008; Boisserie et
two wide and deep sinuses separate the labial
al., 2010; Lihoreau et al., 2015).
cusps. The entostylid is distinct and the smallest
In the developmentally earliest stage (Figure
cusp of the crown. The anterolingual conid is
4.1-4), Bothriogenys fraasi exhibits an exposed
placed mesial to the anterolabial conid, from which
dP1 crown tip, a nearly completely erupted dP2,
it is separated via a narrow and deep notch. The
fully erupted dP3-4, and an M1 visible in the crypt
cusp has three cristids, the most mesial of which
but not yet having begun to erupt. There is no evi-
runs mesiolingually, forming the mesial tip of the
dence of a C1-dP1 diastema at this point, which
crown. The labial cristid of the anterolingual conid
runs mesiolabially to join the mesial cristid of the must represent a very early post-partum stage.
anterolabial conid, forming a V-shape at the mesio- This stage roughly corresponds to Laws’ (1968)
labial corner of the crown. The distal cristid is Hippopotamus groups I and II, which he interprets
directed toward the base of the metaconid. The to represent ages between birth and six months,
anterolabial conid has two cristids, the distal cristid respectively. Laws (1968) noted that in hippos only
of which slopes down distolingually to attach to the the second through fourth deciduous premolars
junction of the preprotocristid, the premetacristid, were replaced but that there was no evidence that
and the distal cristid of mesiolingual conid. The the first deciduous premolar was replaced, only
metaconid has three cristids (mesial, distal, labial), lost through ontogeny.
is situated transverse to the protoconid, and is sep- There are some differences between Both-
arated from the protoconid by a deep, narrow val- riogenys and Hippopotamus during these early
ley. The postprotocristid meets with the labial stages of development. The first deciduous premo-
cristid of the metaconid at a high level on the lar in Hippopotamus erupts before dP2, while it has
crown. The postmetacristid is distally oriented and clearly just begun to erupt when dP2 is nearly fully
slopes down from the tip of the cusp to reach the erupted in Bothriogenys. As in Hippopotamus, dP1
distal basin of the crown. The entoconid has three is not replaced in Bothriogenys (or in any other
main cristids; the preentocristid slopes down from Fayum anthracothere for which there is data) but
the tip of the tooth to end at the base of the post- unlike Hippopotamus, Bothriogenys does not lose
metacristid, leaving the distal basin open lingually; dP1 later in ontogeny, typically retaining it through-
the postentocristid is directed distally and curves to out life.
connect with the posthypocristid, forming a V- When compared with Sus, Bothriogenys stage
shape and closing the basin between hypoconid I roughly corresponds to the documented
and entoconid distally; the labial cristid of the ento- sequence of eruption for the extant taxon but there
conid runs mesiolabially and ends at the base of are differences. Bothriogenys stage I documents
the hypoconid. The prehypocristid is well devel- the beginning of eruption of dP1, full eruption of the
oped and slopes down from the hypoconid toward other three deciduous premolars, and M1 has yet to
the junction between the postprotocristid and the appear. In Sus, following the full eruption of dP2-4
postmetacristid. The distal cingulid is well devel- (which occurs by 100 days), M1 is in place at ~172
oped but short, and bears a distinct small ento- days followed by P1 (or dP1) at ~204 days. Thus,
stylid, which is connected mesially with the tip of the first permanent molar erupts before P1 (or dP1)

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SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

FIGURE 5. Right mandibular fragment of DPC 7706 (doi.org/10.17602/M2/M17034) with dP2- M1, showing
stage II of the eruption sequence of Bothriogenys fraasi, in (1) lateral, (2) medial, (3-4) internal, and (5) occlusal
views.

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 PALAEO-ELECTRONICA.ORG

FIGURE 6. Right mandibular fragment of DPC 2705 (doi.org/10.17602/M2/M12688) with dP2- M1, showing stage
III of the eruption sequence of Bothriogenys fraasi, in (1) lateral, (2) medial, (3-4) internal, and (5) occlusal views.

in Sus while the opposite is true in Bothriogenys. indicate its presence. As in Hippopotamus, dP4
Bothriogenys stage I also shows no wear on the shows relatively heavy wear at this stage but unlike
deciduous premolars. in Hippopotamus dP3 remains unworn in Bothriog-
Stage II in Bothriogenys (Figure 5) documents enys. In Sus, the equivalent of Bothriogenys stage
the beginning of the development of the C1-dP1 II is reached by an age of about 33 weeks (roughly
diastema and roughly corresponds to Laws’ (1968) six months of age).
Hippopotamus group III, which occurs at approxi- Stage III in Bothriogenys (Figure 6) shows the
mately one year of age. In Bothriogenys, dP1-4 are continuation of mandibular growth as the C1-dP1
fully erupted as is M1, while M2 is just appearing in diastema approaches adult length. The early
the crypt but has not yet begun to erupt, and there beginning of crown formation for permanent P2-4
is no opening on the surface of the mandible to can be seen in scanned images and their apices

11
SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

FIGURE 7. Right mandibular fragment of DPC 13562 (doi.org/10.17602/M2/M12638) with P2, dP3-4, M1-M2,
showing stage IV of the eruption sequence of Bothriogenys fraasi, in (1) lateral, (2, 4) internal and X-ray, and (3)
medial views.

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 PALAEO-ELECTRONICA.ORG

FIGURE 8. Right mandibular fragment of DPC 6207 (doi.org/10.17602/M2/M17032) with P2- M3, showing stage V
of the eruption sequence of Bothriogenys fraasi, in (1) lateral, (2) occlusal, (3) medial, (4) internal views; 5-6, right par-
tial maxilla of DPC 10677 (doi.org/10.17602/M2/M12698), in (5) occlusal and (6) medial views.

point horizontally. The M2 crypt is open on the man- erupted permanent P2-4 yet but does have a fully
dibular surface so that M2 is visible. This stage cor- erupted M2, which does not occur in Hippopotamus
responds roughly to Laws’ (1968) Hippopotamus until group VIII. By Hippopotamus group VIII, only
groups IV and V (ages three to four years, respec- dP4 remains from the deciduous dentition, dP1 is
tively). Like in Hippopotamus group IV, dP3, and lost (may happen as early as group VI), and M3 is
dP4 both show relatively heavy wear, and M1 has still not visible on the surface of the mandible.
also begun to wear but M2 is not visible on the Bothriogenys stage IV documents an eruption
mandibular surface in Hippopotamus group IV. By pattern similar to that seen in Sus. At this stage,
Hippopotamus group V, the M2 is visible in its crypt Bothriogenys has P2 erupting but retains dP3-4 and
but this group differs from Bothriogenys stage III in has a fully erupted M1-2. In Sus, at the point where
having both permanent P2 and P3 visible on the M2 is fully erupted (at ~385 days) dP2-4 are still
surface whereas they have barely begun to form in retained, and there are no signs of permanent pre-
Bothriogenys. This stage corresponds to roughly molars (except for P1) or M3. In Sus the permanent
one year of age in Sus (Matschke, 1967). premolars are fully erupted by ~490 days.
In Bothriogenys stage IV (Figure 7) perma- Stage V in Bothriogenys (Figure 8) documents
nent P2 is now exposed on the surface, P3 and P4 the appearance of the full adult dentition and the
have not yet begun to erupt but their crowns are completion of mandibular growth wherein the C1-
fully formed beneath dP3-4, M1-2 are fully erupted, dP1 diastema reaches its maximum length. DPC
and M3 is visible in its crypt. There is moderate to 10677, right partial maxilla with P2-M1-2, shows
heavy wear present on dP3-4 and M1, but M2 that P3 and P4 are exposed simultaneously, which
remains unworn or only slightly worn at this stage. represents the early phase of stage V. At this stage
This stage corresponds to Hippopotamus groups only M1 exhibits moderate wear, while all other
VII and VIII (ages 8 to 11 years, respectively) as cheek teeth essentially have no evidence of wear.
documented by Laws (1968). The main differences This stage is reached by group XII in Hippopota-
with Hippopotamus are that Bothriogenys has not mus, which occurs at 22 years. At this point in Hip-

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SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

TABLE 2. Juvenile dental eruption sequence of Bothriogenys fraasi from the upper most terrestrial mammal-bearing
localities of the Jebel Qatrani Formation, Fayum Depression, Egypt.
Eruption
stage Eruption sequence of Bothriogenys fraasi juveniles Specimen no. Quarry
Stage I dP1 exposed; dP2, dP3 and dP4 erupted; M1 opened DPC 3606 M
DPC 3947 M
DPC 11416 I
DPC 13570 I
DPC 16633 I

Stage II dP1 exposed; dP2, dP3, dP4, M1 erupted; M2 opened DPC 7706 M
DPC 8638 M
DPC 11280 M
DPC 11407 M

Stage III P3 and P4 start to form; dP3, dP4, M1 present; M2 exposed DPC 2705 M
DPC 7730 I
DPC 10616 O

Stage IV P2 exposed; P3 and P4 formed; dP3, dP4, M1, M2 present M3 opened DPC 13562 I

Stage V Adult form i with P3 and P4 half exposed DPC 10677 I

Adult form ii with M3 half exposed DPC 6207 I
Adult form iii with M3 fully exposed DPC 20950 M

popotamus, M1 and M2 are exhibiting heavy wear before M1 in Bothriogenys, and Sus doing the
and cusp tips are beginning to wear on P2-4 and M3. opposite.
In Sus, a fully adult dentition is acquired by ~755 It is unclear why Hippopotamus exhibits an
days with the eruption of M3. accelerated eruption of its permanent premolars
relative to that seen in Bothriogenys and Sus, how-
DISCUSSION AND CONCLUSIONS ever, this is a somewhat common phenomenon in
longer lived species Smith (2000), referred to the
From the previous description, it is clear that pattern of permanent teeth erupting earlier in more
the Bothriogenys tooth eruption sequence (Figure slow growing, longer-lived species as Schultz’s
9) differs from both those of Hippopotamus and Rule. She found evidence to support Schultz’s
wild boars (Sus). The main difference with Hippo- Rule across a broad array of mammalian taxa
potamus is that the anthracothere retains all decid- including many species of ungulates, hippopota-
uous premolars through the period when the third mus among them. Based on Smith’s 2000 study, it
molar becomes visible in its crypt on the mandibu- is reasonable to postulate that artiodactyl groups
lar surface. Only the crown of permanent P2 that show this shift in eruption patterns (such as
becomes visible at the end of this sequence (Fig- that seen in Hippopotamus) may represent a spe-
ure 9, Stage IV). In Hippopotamus, no deciduous cialization within the order. Slower loss of the
premolars remain by the time M3 has appeared in deciduous cheek tooth dentition (relative to perma-
its crypt on the mandibular surface, and all three nent tooth eruption) therefore may be primitive for
permanent premolars (and dP1) are fully erupted. artiodactyls, given that both Bothriogenys and Sus
With regard to the overall sequence of premolar exhibit this pattern.
versus molar eruption, Bothriogenys is more simi- Part of the explanation for these differences in
lar to Sus than it is to Hippopotamus, except for the eruption pattern probably has to do with differences
sequence of erupting the first deciduous premolar in longevity as suggested by Schultz’s Rule (Smith,
2000). Based on differences in body mass (Eisen-

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 PALAEO-ELECTRONICA.ORG

FIGURE 9. Comparison of the different stages (I-V) of the eruption sequence of Bothriogenys fraasi.

berg, 1981), members of Hippopotamus almost by the middle of the second year of life and had a
certainly have longer life spans than did members life span of about 12 years (Myers et al., 2016),
of Bothriogenys, and because of this it may benefit quite different from that seen in Hippopotamus
members of Hippopotamus to have permanent pre- (reproductive age at three and half years and life
molars in place to function in conjunction with the span of ~55 years).
molars in a manner different from that of anthraco- Examining Laws (1968) Hippopotamus
theres, especially later in their lives. Given the sim- groups XII through XX, it is clear that premolars,
ilarity in body size between Bothriogenys fraasi based on increasing wear, begin to participate in
and Sus scrofa, it is a reasonable assumption that food processing by age 22 and become important
Bothriogenys probably reached reproductive age alternative agents for processing as molars

15
SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

become heavily worn. Given that anthracotheres ing access to the fossil collection at the Division of
did not live as long as hippos, the permanent pre- Fossil Primates, Duke Lemur Center and J. Tho-
molars were probably less important as alternative stenson and C. Crawford (Duke University) for pro-
food processing agents and may have been more viding access to micro-CT scanning facilities. This
important in food acquisition and initial preparation. research was funded by U.S. National Science
Further evidence that Hippopotamus and Foundation grants BCS-0416164 to E.R. Seiffert
anthracotheres use their permanent premolars dif- (ERS) and E.L. Simons, BCS-0819186 (ERS) and
ferently can be seen in the wear patterns of premo- BCS-1231288 to ERS, J.G. Fleagle, G.F. Gunnell,
lars versus molars in the two groups. In and D.M. Boyer. Funding was also provided by
Hippopotamus of advanced age (past the age of grants from The Leakey Foundation to E.R.S. This
35) all cheek teeth show signs of heavy apical is Duke Lemur Center publication #1327 and NSF
wear such that all teeth become flattened and den- BCS 1552848 to D.M. Boyer. This paper is a contri-
tine becomes exposed on the surface. In anthraco- bution to project BR/121/A3/PALEURAFRICA of the
theres (at least in all documented Fayum forms), Belgian Science Policy Office.
similar wear can be found on the molars but the
permanent premolars never exhibit apical wear REFERENCES
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SUPPLEMENTAL MATERIAL

Scanning information for Bothriogenys fraasi specimens used in this study. Cite DOI and spec-
imen number when utilizing scans from this table. All files are available directly from:
www.MorphoSource.org.

Specimen DOI File size (MB) File type

doi.org/10.17602/M2/M13080 2.30 MB Digital camera image
DPC-2705 doi.org/10.17602/M2/M16982 410.32 MB Mesh file
doi.org/10.17602/M2/M12688 502.22 MB Zipped tiff stack
doi.org/10.17602/M2/M13082 2.56 MB Digital camera image
DPC-3224 doi.org/10.17602/M2/M16983 47.15 MB Mesh file
doi.org/10.17602/M2/M12690 77.86 MB Zipped tiff stack
doi.org/10.17602/M2/M13075 2.68 MB Digital camera image
DPC-3606 doi.org/10.17602/M2/M16986 761.77 MB Mesh file
doi.org/10.17602/M2/M12653 779.97 MB Zipped tiff stack
doi.org/10.17602/M2/M13060 2.71 MB Digital camera image
DPC-3947 doi.org/10.17602/M2/M17001 153.51 MB Mesh file
doi.org/10.17602/M2/M12639 259.23 MB Zipped tiff stack
doi.org/10.17602/M2/M13063 2.68 MB Digital camera image
DPC-4894 doi.org/10.17602/M2/M17002 256.65 MB Mesh file
doi.org/10.17602/M2/M12644 206.21 MB Zipped tiff stack
doi.org/10.17602/M2/M13066 2.64 MB Digital camera image
DPC-5167 doi.org/10.17602/M2/M17003 163.21 MB Mesh file
doi.org/10.17602/M2/M12646 275.07 MB Zipped tiff stack
doi.org/10.17602/M2/M13070 2.56 MB Digital camera image
DPC-6147 doi.org/10.17602/M2/M17004 43.24 MB Mesh file
doi.org/10.17602/M2/M12649 363.82 MB Zipped tiff stack
doiorg/10.17602/M2/M17033 2.74 MB Digital camera image
DPC-6207 doi.org/10.17602/M2/M17031 2.740 GB Mesh file
doi.org/10.17602/M2/M17032 769.88 MB Zipped tiff stack
doi.org/10.17602/M2/M17037 2.72 MB Digital camera image
DPC-7706 doi.org/10.17602/M2/M17035 217.59 MB Mesh file
doi.org/10.17602/M2/M17034 513.22 MB Zipped tiff stack
doi.org/10.17602/M2/M13078 2.30 MB Digital camera image
DPC-7730 doi.org/10.17602/M2/M17029 298.16 MB Mesh file
doi.org/10.17602/M2/M12655 435.92 MB Zipped tiff stack
doi.org/10.17602/M2/M13079 2.75 MB Digital camera image
DPC-8638 doi.org/10.17602/M2/M17030 270.24 MB Mesh file
doi.org/10.17602/M2/M12684 202.75 MB Zipped tiff stack
doi.org/10.17602/M2/M13071 2.70 MB Digital camera image
DPC-10616 doi.org/10.17602/M2/M16969 456.16 MB Mesh file
doi.org/10.17602/M2/M12650 614.44 MB Zipped tiff stack
doi.org/10.17602/M2/M13076 2.64 MB Digital camera image
DPC-10633 doi.org/10.17602/M2/M16963 573.96 MB Mesh file

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SALLAM ET AL.: ANTHRACOTHERIIDAE FROM EGYPT

Specimen DOI File size (MB) File type

doi.org/10.17602/M2/M12654 732.29 MB Zipped tiff stack
doi.org/10.17602/M2/M13084 2.64 MB Digital camera image
DPC-10677 doi.org/10.17602/M2/M16966 500.91 MB Mesh file
doi.org/10.17602/M2/M12698 436.46 MB Zipped tiff stack
doi.org/10.17602/M2/M13065 2.65 MB Digital camera image
DPC-11280 doi.org/10.17602/M2/M16970 215.02 MB Mesh file
doi.org/10.17602/M2/M12645 235.51 MB Zipped tiff stack
doi.org/10.17602/M2/M13074 2.71 MB Digital camera image
DPC-11407 doi.org/10.17602/M2/M16971 559.86 MB Mesh file
doi.org/10.17602/M2/M12651 782.23 MB Zipped tiff stack
doi.org/10.17602/M2/M13068 2.74 MB Digital camera image
DPC-11412 doi.org/10.17602/M2/M16972 422.1 MB Mesh file
doi.org/10.17602/M2/M12648 511.58 MB Zipped tiff stack
doi.org/10.17602/M2/M13062 2.61 MB Digital camera image
DPC-11416 doi.org/10.17602/M2/M16974 225.28 MB Mesh file
doi.org/10.17602/M2/M12641 282.98 MB Zipped tiff stack
doi.org/10.17602/M2/M13059 2.77 MB Digital camera image
DPC-13562 doi.org/10.17602/M2/M16975 351.17 MB Mesh file
doi.org/10.17602/M2/M12638 443.05 MB Zipped tiff stack
doi.org/10.17602/M2/M17042 2.98 MB Digital camera image
DPC-13570 doi.org/10.17602/M2/M17039 234.05 MB Mesh file
doi.org/10.17602/M2/M17038 325.26 MB Zipped tiff stack
doi.org/10.17602/M2/M13067 2.63 MB Digital camera image
DPC-20439 doi.org/10.17602/M2/M16979 492.04 MB Mesh file
doi.org/10.17602/M2/M12647 723.26 MB Zipped tiff stack

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