8.

1
Elephantoidea from Lothagam
Pascal Tassy

Ten elephantoid taxa are described from the Late MioceneEarly Pliocene strata at Lothagam. Six are
identified or tentatively identified at the species level. From the Lower Nawata come Anancus kenyensis,
Stegotetrabelodon orbus, Primelephas gomphotheroides, and an elephantid (previously described as Stegotetrabelodon orbus or Primelephas gomphotheroides) that is also present in the Apak Member. Associated
in the Upper Nawata are Anancus kenyensis, a trilophodont gomphothere, Stegotetrabelodon orbus, and
an unidentified elephantid (species A). A new early Elephas species, also from the Upper Nawata and
represented by a juvenile mandible, demonstrates that Elephantinae differentiation occurred during the
Late Miocene. From the Apak Member of the Nachukui Formation have been recovered Anancus kenyensis, Stegotetrabelodon orbus, Elephas cf. E. ekorensis, Loxodonta ?aff. L. exoptata, and unidentified elephantids, Elephas species A and B.

Figure 8.1 Restoration of Stegotetrabelodon orbus by Mauricio Anton. Shoulder height estimated at about 3 meters.

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Lothagam proboscideans provide a key to understanding the early differentiation of elephantids and were
originally studied by Vincent J. Maglio in the 1970s
(Maglio 1970, 1973; Maglio and Ricca 1977). Most of
Maglios material was recovered from the Lothagam region during the late 1960s by a team led by Brian Patterson of Harvard University. Maglio described two
primitive elephantids from Lothagam, Stegotetrabelodon
orbus and Primelephas gomphotheroides, in 1970. In the
same paper, Maglio described a primitive member of
the loxodont elephantine lineage, Loxodonta adaurora,
from the Early Pliocene locality of Kanapoi and from
Lothagam. Maglio thus established that elephantines
originated in the Late Miocene and differentiated during the Early Pliocene.
The Lothagam sequence was subdivided by Behrensmeyer (1976) into three units: Lothagam 1 and Lothagam 2, both of Miocene age, and Lothagam 3 of
Pliocene age. Member 1B of Lothagam 1 yielded four
proboscideansDeinotherium, Anancus, and the elephantids Primelephas and Stegotetrabelodonwhile
Member 1C lacked Stegotetrabelodon orbus (Maglio
1973). The elephant Loxodonta adaurora was described in the Pliocene Lothagam 3 faunal unit (Maglio 1973).
The stegotetrabelodont was subsequently recognized
from Mpesida in the Late Miocene of the Baringo Basin,
Kenya (Tassy 1986:106: Stegotetrabelodon sp., vraisemblablement St. orbus), and in the lower Adu-Asa
Formation of the Awash valley, Ethiopia (Kalb and
Mebrate 1993:40). The species Primelephas gomphotheroides, first conceived as the ancestor of the three
elephantine lineages (Loxodonta, Elephas, and Mammuthus), has also been described from the Lukeino Formation of the Baringo Basin (Tassy 1986), from the
Adu-Asa Formation of the Awash Valley (Kalb and Mebrate 1993:50), from the lower Oluka Formation of
Uganda as cf. P. gomphotheroides (Tassy 1994), and
from the Manonga-Wembere Formation of Manonga
Valley, Tanzania (Sanders 1997).
Fieldwork at Lothagam by National Museums of
Kenya Expeditions from 1989 to 1993 provided a new
stratigraphic framework (Leakey et al. 1996; McDougall
and Feibel 1999) and, relevant to elephantoid systematics, several new fossils that allowed revision of the
status of the previously described species and new hypotheses about the timing of elephantine differentiation. New fossils and new age documentation for many
of the older specimens modify the previously estimated
range of the taxa. Primelephas gomphotheroides is now
restricted to the Lower Nawata (ca. 76.7 Ma). Stegotetrabelodon orbus is recognized from both the Lower
and Upper Nawata and extends into the Apak Member
of the Nachukui Formation. The gomphothere Anancus
kenyensis, although rare, ranges from the Lower Nawata
to the upper part of the Apak Member (ca. 74.2 Ma).

Interestingly, fossils from the Upper Nawata and Apak

show a greater diversity than previously thought. At
least four elephantids are associated in this interval of
time: S. orbus (Lower Nawata to Apak), and three elephantines: a new species of the Elephas clade (Upper
Nawata), a taxon described here as Elephantidae, gen.
and sp. indet (Lower Nawata and Apak) and another
as Elephantidae, gen. and sp. incertae sedis A (Upper
Nawata to Apak). From higher in the Apak Member are
described Elephas cf. E. ekorensis, Loxodonta sp. indet.
(?aff. L. exoptata), and Elephantidae, gen. and sp. incertae sedis B (middle Apak). Last but not least, a gomphothere that is clearly different from Anancus is present at the top of the Upper Nawata (5 Ma).

Abbreviations
KNM National Museums of Kenya, Nairobi
HI height index ( 100H/W)
L length
W width (in parentheses, number of the plate where
the width was measured)

Systematic Description
Proboscidea Illiger, 1811
Elephantoidea Gray, 1821
Gomphotheriidae Cabrera, 1929
This paraphyletic family consists of bunodont mastodonsthat is, stem elephantids, the definition of
which is open to discussion (i.e., Shoshani 1996; Tassy
1996b). Two gomphotheres are recognized at Lothagam: the previously described Anancus kenyensis and a
probable trilophodont gomphothere not recorded previously from the Late Miocene of East Africa.

Anancus Aymard in Dhorlac, 1855

Anancus kenyensis (MacInnes, 1942)
(Figure 8.2 [14]; table 8.1)

Lothagam Material

Lower Nawata: 346, Rt. P4.

Upper Nawata: 340, a Rt. maxilla with Rt. M1 or 2;
23781, Lt. dP4.

Apak Member: 341, associated portions of Rt. M3, Lt.
M3, and Lt. M3; 23790, anterior portion Lt. M1 or 2;
28567, Lt. dP4.

Horizon indet: 361, portion Rt. M1; 383, portion Rt.
M1.

Elephantoidea from Lothagam

333

Figure 8.2 Gomphotheriid dentitions: 1 KNM-LT 340, Anancus kenyensis, right maxilla fragment with right M1 or 2 (Apak
Member), occlusal view; 2 KNM-LT 341, Anancus kenyensis, associated portions of right M3 (Apak Member), occlusal view;
3 KNM-LT 23790, Anancus kenyensis, anterior portion left M1 or 2 (upper Apak Member), occlusal view; 4 KNM-LT
28567, Anancus kenyensis, left dP4 (middle Apak Member); 5 KNM-LT 26324, Gomphotheriidae, gen. and sp. indet., right
M1 (Apak Member), occlusal view. Scales 5 cm.

LT 28567, a left dP4 from the middle Apak Member is

broken, and the anterior part is missing (figure 8.2 [4]).
This tooth displays derived characters. Supplementary
accessory cusps are present. The postcingulum is inflated and consists of an outlined fifth loph and two
posterior cuspules connected to the last loph. This arrangement (outlined fifth loph with primitive postcin-

gulum) can be consistent with either a tetralophodont

or pentalophodont grade: a dP4 with a complicated posterior area can be associated with more primitive intermediate molars (M12).
LT 23781, a left dP4 (Upper Nawata), has four lophids with no trace of a fifth. The crown is heavily
eroded and does not display distinctive traits.

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Pascal Tassy

An isolated premolar from the old collection (LT

346) was identified by Tassy (1986:95) as a right P?4 of
Anancus kenyensis. This tooth comes from the Lower
Nawata. Its bunodonty is indicated by the round shape
of the wear facets. It has four main cusps, and the pretrite cusps are enlarged. Pretrite and posttrite half lophs
are separated. The postcingulum is inflated. It is very
reminiscent of another premolar, from the Lukeino
Formation, also allocated to Anancus kenyensis (Tassy
1986:94, plate 13, figure 5). The dislocation of the lophs
gives an anancoid pattern to these teeth. However, no
premolar of Anancus has yet been found to be associated with anancoid molars.
The anterior portion of upper left first or second
molar (23790; Upper Apak) shows a complete trefoil on
the second loph, a feature characteristic of a complex
crown pattern (figure 8.2 [3]). Complex crowns previously recovered from undetermined horizons at Lothagam include 361 (portion of right M1, rather worn)
and 383 (portion of right M1 with two posterior pretrite
conules on the first loph, big posterior posttrite conule
on the second loph). However, a partial right maxilla
from the Upper Nawata (LT 340) displays a very simple
right first or second molar with no posterior pretrite
conule on the second loph and no accessory cusps (figure 8.2 [1]); cement is present, although not plentiful.
This tooth is remarkably low crowned and narrow.
LT 341 (Apak Member) constitutes three fragments
of upper and lower third molars from the same individual. The right M3 (341A) comprises fragments of the
four posterior lophids (figure 8.2 [2]). The left M3
(341C) consists of the posterior area (last lophid and
postcingulum), and the left M3 (341B) constitutes the
posterior half of the crown (three lophs and the postcingulum). The associated portions of the M3 show
many posttrite accessory cusps: three posttrite conules
on the most anterior preserved loph (probably the third
loph), one big anterior pretrite conule on the same
loph, and two on the posterior loph. The postcingulum
is complicated and forms two transverse lines. The successive lophs show no anancoid pattern, a condition
reminiscent of the Anancus specimens described from
Lukeino (KNM-LU 532, Tassy 1986: plate 13, figure 2).
The M3s have supplementary posttrite cusps. The enamel is thick (6.8 mm on the right lower molar). Cement is present but not plentiful on 341AB. These
teeth appear distinctively smaller and narrower than
those described from Lukeino, but their weathering precludes precise measurements.

Discussion
In the late 1970s, Smart (1976) cited a primitive form
of the genus Anancus at Lothagam; this was interpreted

as Anancus kenyensis by Coppens et al. (1978). Tassy

recognized two morphs among molars of Anancus kenyensis: the primitive kenyensis morph based on the
holotype of A. kenyensis from Kanam (MacInnes 1942)
and the derived petrocchi morph based on the sample
from Sahabi (Libya) described by Petrocchi (1954). In
East Africa the kenyensis morph was recognized at
Kanam, Lukeino, Mpesida, and Lothagam, whereas the
petrocchi morph was described from Sahabi (Libya),
Kanapoi, Aterir, and the Chemeron Formation of Baringo Basin. However, both morphs are known to be
associated at Lothagam, Lukeino, Chemeron, and
Kanam, and this association suggests that evolutionary
implications of these morphs may not be meaningful
without supporting biostratigraphic evidence.
Three specimens were recovered during the
19891993 expeditions: 23790, portion of M1 or 2; 23781,
left dP4; and 28567, left dP4. These clarify neither the
status of the species nor the respective evolutionary level
of Late Miocene and Early Pliocene molars. However,
they do provide an opportunity to define the distinctive
traits of the two morphs. Simple molars (kenyensis
morph) display plesiomorphic characters such as lack
of supplementary cusps and tetralophodonty of intermediate molars; in complex molars (petrocchii morph)
accessory cusps are associated with pentalophodonty.
The partial upper molar LT 23790 (upper Apak) has
supplementary cusps that are not present in the tetralophodont M1 or 2 LT 340 from the Upper Nawata. The
dP4 (LT 23781; Upper Nawata) is too worn and weathered to show any character except the lack of a fifth
lophid. The dP4 (LT 28567; middle Apak) is complex
with accessory cusps and an incipient fifth loph. This
combination would fit with the petrochii morph, but on
this tooth the cingular cusps are connected to the fifth
loph. This last trait is more primitive than the latest
known molars of Anancus where the postcingulum is
fully separated from the last loph (M2 from Sagantole
Formation, Beeryada beds, Ethiopia, described by Kalb
and Mebrate 1993:39).
Furthermore, it seems erroneous to expect clearcut association of the pentalophodont grade with
supplementary accessory cusps. In the important population of Anancus from the Early Pliocene of Dorkovo
(Bulgaria), rare pentalophodont M2s are found among
much more common tetralophodont ones (MetzMuller 1997). The holotype of Anancus kenyensis from
Kanam shows the complex morph (supplementary accessory cusps) with no fifth loph. An intermediate form
from Nkondo in Uganda has a pentalophodont M2
without postcingulum.
Thus, without a large collection of teeth from one
site, it is difficult to precisely determine the evolutionary
level of Anancus molars. What can be safely concluded
is that the molars from the Apak Member show distinct

Elephantoidea from Lothagam

complex traits. But these molars are not among the

most complicated of Anancus (Kanapoi, Chemeron,
and Aterir in Kenya; the Beearyada beds in Ethiopia).
At Kanapoi (4.2 Ma; Leakey et al. 1996) two M2s
(KNM-KP 384 and KNM-KP 410) are nearly identical
and combine all derived traits: accessory conules, fifth
lophid, strong anancoidy. Moreover no tooth at Lothagam matches those found at Sagantole, Ethiopia (Kalb
and Mebrate 1993:37), or Baards Quarry at Langebaanweg (Hendey 1978:10), all of which show an extremely
serrated enamel and very thin loph(id)s and are perhaps
the ultimate evolutionary level of Anancus in eastern
and southern Africa at around 4 Ma. Based on all the
available morphological evidence, all teeth from Lothagam, including those from the Apak Member, would
be older than 4 Ma.

Gomphotheriidae gen. and sp. indet.

(Figures 8.2 [5], 8.3; table 8.2)

Lothagam Material

Upper Nawata: 26324, Rt. M1.
This lower molar is heavily worn (figures 8.2 [5] and
8.3). The crown seen in occlusal view has the primitive
shape of Miocene gomphothere molars. It is trilophodont. The third lophid is enlarged, and the wear patterns of the lophids are typical of bunodont teeth. Pretrite and posttrite halves of the lophids are well
separated, and the posttrite half-lophids are located
posteriorly to the pretrite half-lophids. Because the
tooth is heavily worn, the central conules are contiguous with the lophids and all three lophids are confluent.
Dentine indentations in the enamel of the posterior side
of the second posttrite half-lophid indicate the presence
of two small conules. Although the third lophid of the
tooth is damaged, it is seen to be separated labially from
the postcingulum. The posterior side of the tooth is

Figure 8.3 Gomphotheriidae gen. and sp. indet., right M1,

KNM-LT 26324 (Upper Nawata), occlusal view. Scale
5 cm.

335

preserved at the level of the cervix. Consequently, even

if it could be deduced that the postcingulum was inflated, there is no room for a fourth lophid.

Discussion
The geographic and stratigraphic provenance of this
tooth is unequivocal. According to M. G. Leakeys files
(personal communication 1997) it was excavated in
1992 from the lower white bed of the Purple Marker,
a horizon that denotes the top of the Upper Nawata
(Leakey et al. 1996:557). This tooth belongs to a trilophodont gomphothere because no fourth lophid can be
reconstructed from what remains of the posterior area
of its crown. When LT 26324 is compared with a tetralophodont M1 or dP4 of Anancus kenyensis such as
LT 23781, it clearly cannot belong to Anancus kenyensis
because no trilophodont dP4 is known in Anancus.
Morphological variation seen in the dP4 of Anancus affects the size of the postcingulum but not the presence
or absence of a fourth lophid.
This M1 has a generalized morphology that could
belong to any trilophodont Miocene species. For example, if it came from Middle Miocene strata in East
Africa it could well belong to the trilophodont amebelodontid Protanancus macinnesi Arambourg 1945. However, Alengerr, a Middle Miocene locality, is probably
the youngest locality to yield P. macinnesi (Tassy
1986:57). Thus P. macinnesi is not found in the latest
Miocene. The only other Late Miocene trilophodont
species from East Africa is Choerolophodon ngorora
(Maglio 1974), whose latest occurrence at Ngorora and
Nakali dates back to at least 10 Ma (Tassy 1986:69),
thereby predating the upper member of the Nachukui
Formation by about 4 million years. Moreover, although very worn, it is certain that the M1 from the
Upper Nawata cannot belong to a choerolophodont because it has neither wrinkled enamel nor a choerolophodont wear pattern.
Other than Anancus, the only gomphothere known
from the latest Miocene of Africa (ca. 75 Ma) is Amebelodon cyrenaicus from Sahabi, Libya (Gaziry 1987). I
consider this a junior synonym of Mastodon grandincisivus Schlesinger 1917 from Maragah in Iran, which is
also present in the contemporaneous locality of Jebel
Barakah in Abu Dhabi (see discussion by Tassy 1999).
LT 26324 could well belong to this species and would
be its first record from East Africa. The M2 of this species is known to have reached the tetralophodont grade,
but an association of trilophodont M1 (and dP4) with
tetralophodont M2 is not unexpected in amebelodontid
species such as that from Libya. This condition is also
displayed, for example, by Platybelodon grangeri from
the Middle Miocene of Mongolia and China. Thus even

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Pascal Tassy

in well-prospected areas, unexpected discoveries such

as this worn small molar of a hitherto unrepresented
species may still occur.

Elephantidae Gray, 1821

Stegotetrabelodon Petrocchi, 1941
Stegotetrabelodon orbus Maglio, 1970
(Figures 8.1, 8.4, 8.5 [1]; tables 8.38.7)

Holotype

KNM-LT 354, mandible with Lt. M2 and M3, Lt. and

Rt. lower tusks, and associated Rt. M2 and partial Lt.
M3.
Type locality

Lothagam, Apak Member of the Nachukui Formation.

Lothagam Material

Lower Nawata: 343, ?Rt. P4; 344, Lt. and Rt. M3; 347,
Lt. M3; 349, Rt. M3; 360, Lt. M3; 23791, Lt. M3; 26318,
cranium with Rt. upper tusk, Rt. and Lt. M3; 26334,
anterior portion of Lt. M1 or 2 and isolated plate of
indeterminate molar.

Upper Nawata: 352, Lt. M3.

Upper Nawata or Apak Member: 355, two portions
of Rt. and Lt. M3; 359, Rt. M2, Lt. M3, Rt. and Lt.
M3s.

Apak Member: 354, holotype (see above); 363, partial
Rt. M3.

Horizon indet.: 365, Rt. dP3; 366, Rt. M3; 367, Rt. M3.
The description and measurements of the original collection from Lothagam were published by Maglio
(1970, 1973). Of the material previously described
(Maglio 1970; 1973:18; Maglio and Ricca 1977:12), one
specimen (LT 316, M3) was not seen, one described as
M2 (LT 366) is here considered M3, and one specimen
(LT 350, two partial mandibles with right and left lower
M2, erroneously listed by Maglio as LT 342) is not
identified as S. orbus (see section Elephantidae, gen.
and sp. indet.). Accordingly, the range of variation of
M2 and M3 is reevaluated (tables 8.38.6). In this section
I emphasize two new specimens (LT 26318 and LT
26334) recovered between 1989 and 1993 from the
lower part of the Nawata Formation.
Partial cranium LT 26318 is from an adult. The alveoli of M2s are partly resorbed, and the three anterior
plates of both right and left M3s are partly worn (figure
8.4 [3]). The dorsal part of the skull is missing, and the
basicranium is weathered and compressed under the

palate so that the condyles are situated close to the choanae. Although the precise height of the maxilla cannot
be measured, the area of the maxilla between the processus zygomaticus and the alveolar arch is large. This
trait (plus the size of the M3s) is consistent with an
identification of this cranium as male (Tassy 1996a).
The preserved portion of the right upper tusk, partly in
the premaxilla, is straight, but the portion is too short
(400 mm) to give a precise idea of the orientation of
the entire tusk. The M3s are large, with seven widely
spaced thick plates, each made of five main cusps (figure
8.4 [45]). The laminar frequency is low (2.662.86).
The cement is plentiful but does not embed the rear
half of the teeth. The first three plates are worn. The
first and second plates display a digitation that indicates
the presence of a posterior pretrite conule in the first
two interlophs. Conules are either lacking on the other
plates or are confined to the base of the posterior side
of the plates and hidden by a coating of cement. The
cusps forming the plates are nearly equal in size. The
plates are moderately high (HI 73.7 on the fifth
plate). In labial view, the anterior plates are anteriorly
inclined, whereas the last two plates (sixth and seventh)
are posteriorly inclined.
LT 26334A comprises the anterior three plates of a
left M1 or 2 (figure 8.4 [67]). A big, round conule is
present at the base of the posterior sides of the first and
second plates. A smaller conule is present on the third
plate. The shape of the plates is asymmetrical: the second is concave-concave, and the third is convexconvex.
LT 23791 is a left M3 with seven plates and a distinct
postcingulum (figure 8.4 [1]). The first five plates are
worn. A posttrite conule is present on the posterior side
of fourth and fifth plates, but no pretrite is present. This
unusual feature is probably attributable to individual
variation. The tooth is small, its size less than 80 percent
of the holotypea difference that is consistent with sexual dimorphism seen in proboscideans, although it can
also be explained by size variation between populations.

Discussion
There is no major difference between specimens from
the Lower Nawata and from higher in the succession.
Only three specimensall from the old collection
come from the later horizons: the holotype (M3 and
associated M3s) from the Apak Member plus LT 352
(M3) from the Upper Nawata and LT 355 (posterior
portions of associated right and left M3) that is either
from the Upper Nawata or the Apak Member. In particular, the teeth of the holotype match well with the
M3s of cranium LT 26318 from the Lower Nawata. Size,
proportions, height, lamellar frequency, and plate mor-

Elephantoidea from Lothagam

337

Figure 8.4 Stegotetrabelodon orbus dentitions: 1 KNM-LT 23791, left M3 (Lower Nawata), occlusal view; 2 KNM-LT 366,
right M3 (horizon indet.), occlusal view; 3 KNM-LT 26318, palate with right and left M3 (Lower Nawata), occlusal view;
4 KNM-LT 26318, left M3 (Lower Nawata), occlusal view (same individual as 3); 5 KNM-LT 26318, left M3 (Lower
Nawata), labial view (same individual as 3 and 4); 6 KNM-LT 26334A, associated anterior portion of left M1 or 2 (Lower
Nawata), occlusal view; 7 KNM-LT 26334A, associated anterior portion of left M1 or 2 (Lower Nawata), lingual view (same
individual as 6). Scales 5 cm, except for figure 3.

phology are nearly identical in the molars of these two

individuals that are perhaps separated by two million
years. The variation in size and morphology displayed
in the Lower Nawata remains unchanged in specimens
from higher in the sequence. The M3s have either seven

plates (LT 26318 and LT 360 from Lower Nawata, and

LT 367, horizon unknown) or six (LT 347 and LT 359
from the Lower and Upper Nawata, respectively, and
LT 366 of unknown horizon). LT 347 and LT 366 are
atypical. LT 347 is large (L 210.0, W 105.9 [3])

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Pascal Tassy

and is of typical M3 size, but its posterior end is abbreviated so that the shape of the crown is closer to that of
a M2 in occlusal view. However, the holotype M2 has
five plates and a postcingulum connected to the fifth
plate, whereas in LT 347 the postcingulum is well separated from the sixth plate and forms the posterior end
of the crown. LT 366 (figure 8.4 [2]) is small (L
173.1, W 86.0 taken at the third plate). Maglio
(1973:18) interpreted LT 366 as an M2. This tooth is
indeed short (only slightly longer than the M2 of the
holotype (LT 354, L 157.8, W 92.4 taken at the
fourth plate, FL 3.25). But other characters match
better with those of M3s. The posterior end of this tooth
is narrow, and the posterior root is subdivided. The
wear pattern is asymmetrical: more pronounced lingually on the first two plates (normal condition) and

more pronounced labially on the posterior plates. When

present in elephants, such asymmetry of wear is usually
characteristic of the M3. A second example of unusual
asymmetry in wear is provided by the cranium LT
26318: the labial half of the second plate of the M3s is
more worn than the lingual half. If LT 366 is an M3, it
is indeed a small tooth, probably that of a female,
whereas large M3s such as LT 367 belong to males (figure 8.5 [1]).
Specimen LT 363 (Apak Member) was allocated to
P. gomphotheroides by Maglio (1973:21) as partial right
M3 and mandibular symphysis. It is here allocated to
S. orbus. The partial M3 has plates made of big, round
cusps closer to those of S. orbus. The mandibular symphysis does not belong to the same individual but is
that of a suoid.

Figure 8.5 Elephantidae dentitions: 1 KNM-LT 367, Stegotetrabelodon orbus, right M3 (horizon indet.), occlusal view; 2
KNM-LT 375, Primelephas gomphotheroides, right M1 and first plate of M2 (horizon indet.), occlusal view; 3 KNM-LT 358,
Primelephas gomphotheroides, associated right M1 and M2 (Lower Nawata), occlusal view; 4 KNM-LT 342, Elephantidae gen.
and sp. indet., left M2 (?Lower Nawata), occlusal view; 5 KNM-LT 350, Elephantidae gen. and sp. indet., left mandible
fragment with M2 (Upper Nawata/?Apak), occlusal view. Scale 5 cm.

Elephantoidea from Lothagam

Sexual dimorphism is also seen on M3s. Although it

has seven plates, LT 23791 (figure 8.4 [1]) from Lower
Nawata is smaller than the other known M3s from Lothagam such as LT 359 (with seven plates and a reduced
postcingulum; L 259.0, W 108.0[3]) or the holotype (with seven plates and a huge postcingulum
forming an incipient eighth plate L 266.0, W
87.9[3]).
So-called intermediate molars (dP4, M1, M2) of Stegotetrabelodon orbus are rare. The holotype with associated upper and lower second molars is the only example from Lothagam. The partial germ LT 26334A,
allocated to M1 or M2, shows an asymmetry in the shape
of the plates: the second is concave-concave, and the
third is convex-convex. This asymmetry is found in
lower molars of S. orbus (e.g., LT 354 and LT 359).
Two lower M3s (LT 349 from Lower Nawata and LT
352 from Upper Nawata), allocated to S. orbus by Maglio, contradict some previous hypotheses on evolutionary characters and trends. According to Maglio
(1973:18), a diagnostic feature of S. orbus is the presence
of posterior columns ( conules) only behind the first
two plates. However, LT 349 and LT 352 display conules
on plates posterior to the second plate.
Kalb and Mebrate (1993) and Kalb et al. (1996) emphasized the importance of the shape of the plates in
erecting evolutionary hypotheses for elephantid molars
where convex-convex plates is the primitive condition
and concave-concave is the derived condition. In this
feature, most of the molars of S. orbus are primitive,
including LT 352 from the Apak Member. Some variation is seen: a concave-concave shape is present on the
second plate of the holotype or on a second and third
plate associated with the straight fourth plate in LT 359.
However, LT 349 from the Lower Nawata is more derived. Although fragmentary, the first four plates of this
tooth are concave-concave. Curiously, the primitive
condition observed on LT 352 is associated with a derived trait, which makes this tooth unique in the collection of S. orbus. Numerous apical digitations are seen
on the third and fourth plates (seven digitations on the
fourth plate) and are associated with an anteroposterior compression of the upper part of the plates.
Variation exists, and associations of character states
are sometimes self-contradictory. As a consequence, no
clear-cut evolutionary separation can be observed between the molars found in Lower Nawata and those
documented as Apak or Upper Nawata or ?Apak.
Morphological differences are few and merely due to
individual variation. Size differences are conceivably
due to sexual dimorphism. In this respect the holotype
mandible would be that of a male. The relative shortening of the symphysis, compared to that of Stegotetrabelodon syrticus from Libya (also a male), would be
characteristic of the species S. orbus. The hypothesis for

339

sexual dimorphism in one species explaining the size

differences of the symphyseal rostrum and tusks between the mandibles from Sahabi and Lothagam can be
ruled out. The species S. orbus is valid and is clearly
distinguishable by derived traits from the roughly contemporaneous (Late MioceneEarly Pliocene) species S.
syrticus from Sahabi, Libya (Petrocchi 1954; Maglio
1973), and from Abu Dhabi (Tassy 1999).

Primelephas Maglio, 1970

Primelephas gomphotheroides Maglio, 1970
(Figures 8.5 [23], 8.6 [5]; tables 8.68.9)

Holotype

KNM-LT 351, associated Rt. and Lt. M3, and fragment

of palate.
Type locality and horizon

Lothagam, Nawata Formation, Lower Member.

Lothagam Material

Lower Nawata: 351, holotype (see above); 358, Rt.
and Lt. M2, Rt. M2, portion of Lt. M2, Rt. and Lt. M1,
portions of mandibular ramus and symphysis, portion of lower tusk; 23782, Lt. P4.

Horizon indet.: 375, Rt. M1 and first plate of M2.
The only new specimen allocated to Primelephas gomphotheroides is an isolated premolar, probably left P4
(table 8.8 and figure 8.6 [5]). This tooth is broken anteriorly and consists of the rear part of the first loph,
the second loph, and a strong postcingulum. The first
loph is made of two separate halves from the two round
main cusps. The second loph and postcingulum are linear. Although larger, this tooth is reminiscent of the left
P?4 (KNM-LU 730) from Lukeino (Tassy 1986:111) and
of the P4s of a cranium from the Late Miocene of
Uganda, allocated to P. gomphotheroides (Tassy 1994:
plate 3, figure 3).

Discussion
No new important specimens of Primelephas gomphotheroides were recovered from Lothagam by the National Museum expeditions, but deciduous teeth possibly belonging to this species are described in the
section labeled Stegotetrabelodon or Primelephas. Maglio
(1973:21) assumed that about fifteen additional specimens belong to P. gomphotheroides, from Lothagam,

340

Pascal Tassy

Figure 8.6 Elephantidae dentitions: 1 KNM-LT 26332, Stegotetrabelodon or Primelephas, associated left dP2dP3 (Lower Nawata), occlusal view; 2 KNM-LT 22866, Stegotetrabelodon or Primelephas, left dP3 (Lower Nawata), occlusal view; 3
KNM-LT 26329, Stegotetrabelodon or Primelephas, ?right P3 (Lower Nawata), occlusal view; 4 KNM-LT 26339, Stegotetrabelodon or Primelephas, left P3 (Lower Nawata), occlusal view; 5 KNM-LT 23782, Primelephas gomphotheroides, ?left P4 (Lower
Nawata), occlusal view; 6 KNM-LT 405, Elephantidae gen. and sp. incertae sedis A, partial right dP3 (Upper Nawata), occlusal view. Scale 5 cm.

Chemeron Formation, and Kanam East (Kenya) and

from the Kaiso Formation (Uganda). However, from
Lothagam I retain only four individuals: the holotype
and LT 358 (nine associated specimens that belong to
the same individual) from the Lower Nawata; LT 375
(M1 of unknown horizon not included in the hypodigm
by Maglio); and LT 23782the newly found isolated
broken P4 also from the Lower Nawata. Another specimen listed by Maglio, LT 363 (partial right M3 with
mandibular symphysis) is here allocated to S. orbus (the
symphysis being that of a suoid). Including LT 375 (the
M1) in the hypodigm implies some variation in molar
morphology.

LT 358 is crucial for understanding Primelephas gomphotheroides and keeping the species separate from S.
orbus (figure 8.5 [3]). This individual comprises associated upper and lower intermediate molars and mandibular symphysis and tusk. The relative states of wear
of the M1s and upper and lower second molars confirm
that these teeth belong to only one individual and, consequently, that the broken mandible and tusk also belong to the same individual. Erupting lower and upper
second molars show a slight wear on the anterior cingulum and first plate (but no dentine is seen). The symphysis and tusk were described in detail by Maglio and
Ricca (1977:25, plate 2 [8]) under the field number 290

Elephantoidea from Lothagam

67-K. It was assumed from this specimen that P. gomphotheroides had lower tusks, which is a primitive feature among the Elephantinae. But there is no anatomical evidence to support the association of the portion
of tusk and the mandibular fragment. The reconstruction proposed by Maglio can be refuted; nonetheless,
this does not mean that the tusk LT 358 does not belong
to that individual. Although a trough is preserved in the
symphyseal area, extending posterior to the symphyseal
border, Maglio and Riccas (1977) description is not
accurate. Because of its position in the mandible, this
trough, into which the incisor was put by Maglio and
Ricca, cannot be the alveolus but is instead the mandibular canal. In fact, that portion of the symphysis
where the alveolus for the incisor should be is not preserved on this specimen. Consequently, it is not possible
to prove or disprove the association of the tusk and the
mandibular fragment. There is also no connection between the symphysis and the portion of ramus where
the mandibular canal is clearly seen. The preserved part
of the tusk, 95 mm long, must be close to the origin
because of the large size of the pulp canal (the diameter
of which is 37.6 mm). These proportions are too small
for an upper tusk associated with erupting M2s and
match best with these of a lower tusk of a stegotetrabelodont. However, the molars of LT 358 do not belong
to S. orbus. Because there is no reason to assume that
the different specimens labeled LT 358 were not really
associated, Maglios conclusion that P. gomphotheroides
had lower tusks is consistent although, again, it should
be emphasized that there is no anatomical evidence to
support Maglios reconstruction. Now, as twenty-five
years ago, a complete mandible is needed to confirm
retention, orientation, and size of the lower tusks in
primitive elephantines.
Direct association of second and third molars is not
known in P. gomphotheroides, but the allocation of LT
358 to this species is based on several coherent dental
characters of the intermediate molars (shape of crown,
number and shape of plates) that confirm Maglio and
Riccas (1977) description. The association of M1 and
M2 (figure 8.5 [3]) is important to identify the evolutionary level of P. gomphotheroides. The M1s have five
plates and a small postcingulum connected to the fifth
plate. Both anterior and posterior columns are present.
Plates 35 are convex-convex; plate 2 is straightconcave, and the posterior side of plate 1 is concave.
The left M2 consists of two broken plates. The right M2
is very long; it is broken posteriorly and had at least six
plates. It becomes wider posteriorly, a primitive feature
more accentuated than on M1s where maximal width is
taken on the fourth plate. The plates are convex-convex,
and anterior and posterior columns are present on the
six preserved plates. The M2s (right M2 figured by Maglio 1973:plate 3, figure 3; left M2 figured by Tassy

341

1986:plate 14, figure 2) have an incipient sixth plate that

is narrow but well separated from the fifth plate (on the
right M2, an additional cingular cuspule is present on
the posterior end), so that the associated M2 could well
have had seven plates. The plates are convex-convex.
This character is more primitive here than in the holotype left M3 (LT 351), where plates 24 are concaveconcave, plates 5 and 6 are convex-convex (primitive),
and plates 7 and 8 are concave-concave. This variation
is perhaps significant for the evolutionary level of P.
gomphotheroides with incipient elephantine derived
traits in the shape of the plates.
On LT 358, plates are more numerous, more spaced,
and more subdivided apically than in S. orbus. Although
the differences are not very profound, they bring LT
358 closer to the holotype M3s of P. gomphotheroides
(especially the M2s) than to the collection of molars of
S. orbus.
LT 375 consists of a right M1 and anterior portion
(anterior cingulum and first plate) of right M2 (figure
8.5 [2]). The anterior end of M1 is broken, and four
plates and a narrow postcingulum are preserved. The
first anterior preserved plate has a concave posterior
side, the second preserved plate is convex-straight, and
the posterior two plates are convex-convex. A straight/
concave shape was noticed on the first two plates of the
M1 of LT 358. The anterior end of the M2 of LT 375 is
less derived than that of the M2 of LT 358: the anterior
cingulum is more bulbous, and the first plate is less
rectilinear. In contrast, the M1 of LT 375 has more
widely spaced plates than in other molars of P. gomphotheroides. If the partial M2 could be allocated to S.
orbus, the M1 with thinner plates and more reduced
columns cannot. The allocation of LT 375 to P. gomphotheroides is perhaps a better bet.

Stegotetrabelodon orbus or
Primelephas gomphotheroides
(Figure 8.6 [14]; table 8.10)

Lothagam Material

Lower Nawata: 434, Rt. dP4; 22866, Lt. dP3; 26325,
partial dP3; 26326, Lt. dP?2; 26329, ?Rt. P3; 26332,
associated Lt. dP2dP3; 26339, Lt. P3.
LT 26326, a dP2, consists of four main cusps, each subdivided into at least two cusps arranged in a linear fashion. Pretrite and posttrite half-lophs are separated by a
deep median sulcus. The postcingulum is inflated and
tends to form two half-lophs separated by a median
sulcus. This tooth resembles KNM-LU 699 from the
Lukeino Formation allocated to cf. Primelephas by
Tassy (1986: plate 9, figure 5).

342

Pascal Tassy

LT 26332 is an anterior left dP2 associated with the

anterior portion of a dP3 (figure 8.6 [1]). The small dP2
is complete. The primitive pattern of the crown is that
of an elephantoid. The anterior cusp is inflated. The two
posterior cusps are smaller; the lingual cusps, slightly
worn, are transversely enlarged, whereas the smaller
round labial cusp is connected to a postcingulum cuspule and the labial cusp. This tooth resembles that of
Miocene gomphotheres rather than that of elephantids,
such as the dP2 from the Lukeino Formation described
by Tassy (1986:111, figure 43) as cf. Primelephas. The
associated dP3 is broken posteriorly. The first lophid
lacks the pretrite (lingual) cusp. The second and third
lophids are anteroposteriorly compressed (or platelike), with numerous apical digitations. Conules are
present in the two preserved interlophids. In the second
interlophid, the posterior central conule of second lophid is weak and the anterior central conule of the third
lophid is larger. The lophids are linear, much more so
than those of the dP3 of S. orbus (LT 365; Maglio and
Ricca 1977: plate 1, figure 8; Tassy 1986: plate 14, figure
5), thus making this tooth clearly elephantine. The association of a small gomphothere-like dP2 with an elephantine dP3 can only be explained by an important
variation of the morphology of dP2.
LT 22866 and LT 26325, two dP3s, share the platelike loph pattern of elephantids. The complete left dP3
(LT 22866; figure 8.6 [2]) is trilophodont with a linear,
low postcingulum. A contact between the posttrite halfloph 1 and the pretrite half-loph 2 can be seen; this is
a derived character for gomphotheres that persists in
early elephantids (Tassy 1986) but is lost in later elephants. A contact also exists between a tiny posttrite
cuspule on the posterior side of the second loph and
pretrite cuspules on the anterior side of the third loph,
but no anancoid dislocation is seen. The incomplete
dP3, LT 26325, consists of the posterior lingual region.
Its inflated postcingulum is more complicated and less
linear than that of LT 22866.
LT 434, a right dP4 from the earlier collection (Lothagam 1B), consists of four entirely worn lophs. It is
short. The wear pattern of the lophs reflects the plate
shape of elephantids.
LT 26339 and LT 26329, left and right small lower
third premolars, are nearly trilophodont and very reminiscent of the P3s of tetralophodont gomphotheres.
LT 26339 (figure 8.6 [4]) has a strong third lophid that
is better developed than that in LT 26329 (figure 8.6
[3]). Both have a distinct thin postcingulum connected
to the posttrite half of the third lophid. The first lophid
is made of two cusps, and the second is made of four.
The third lophid is low but distinct from both the second loph and the postcingulum; it is more linear on
LT 26339 and more strongly connected to the pretrite
half of second lophid on LT 26329. These two

gomphothere-like premolars with no anancoid trait can

only be interpreted as primitive elephantid premolars.
They confirm the hypothesis of persistence of premolars
in both Stegotetrabelodon orbus and Primelephas gomphotheroides, first based on P4s from Lothagam and Lukeino (Maglio 1973; Tassy 1986).

Discussion
As the species Stegotrabelodon orbus is more common
than Primelephas gomphotheroides, the specimens described in this section belong perhaps to the former,
but there are no anatomical characters that unequivocally support this view. Because the two known P3s are
slightly different and because these differences lie in the
respective states of elephantid features, the more derived LT 26339 could be allocated to P. gomphotheroides
and LT 26329 to S. orbus. The same approach for dP3
would determine LT 22866 as P. gomphotheroides and
LT 26325 as S. orbus. This is mere speculation, however.
Associated material, with deciduous teeth, premolars
and molars, is the only clue to a safe systematic identification.

Elephas Linneaus, 1758

Elephas nawataensis sp. nov.
(Figures 8.7, 8.8 [14]; tables 8.11, 8.12)

Holotype

KNM-LT 23783, mandible with Rt. and Lt. dP4s and

M1s, and various mandibular fragments including the
tip of the symphysis.
Type locality and horizon

Lothagam, upper member of Nawata Formation.

Lothagam Material

Upper Nawata: 23783, holotype (see above); 26327,
portion of left M1 or 2.

Diagnosis
Elephantid with a brevirostrine mandible without lower
tusks. In lateral view, ventral border of the symphysis
convex and not bent downward. dP4s have x6x plates
with slight posterior widening. M1s narrow with x5x or
x6 plates. No protruding enamel loops on dP4 and no
columns on M1, except on first plate. Low laminar frequency of M1 (3.54.4). Enamel of M1 thick (3.9 mm).

Elephantoidea from Lothagam

Crown of M1 low (HI 74.3). Plates 14 of M1 with both

convex-convex and concave-concave orientations
(plates 14 concave-concave, plates 56 [or 5x] convexconvex).
The mandible LT 23783 (figure 8.7) belongs to a young
individual (dP4 in wear, M1 erupting, unworn). It is
fairly well preserved and not distorted, although the
symphysis is broken in the middle without contact between the horizontal rami and the tip of the symphyseal

343

rostrum. Both ascending branches are lacking. Even if

the total length of the rostrum cannot be directly deduced, the mandible is of the brevirostrine type. From
its preserved parts, the symphyseal rostrum was probably short. In occlusal view the horizontal rami are very
convergent so that there is space only for a narrow rostrum well individualized from the rami. The preserved
parts of the symphysis are low. All these traits are those
of a brevirostrine mandible. The symphyseal rostrum
lacks any trace of incisor. Its tip is smoothly spatulate.

Figure 8.7 Elephas nawataensis sp. nov., holotype. KNM-LT 23783: 1 mandible with right and left dP4 and M1 (Upper Nawata), occlusal view; 2 lateral view of left ramus without M1. Scales in cm.

344

Pascal Tassy

The symphyseal trough is not deep. In lateral view the

ventral angle of the rostrum is weak. This angulation is
anteriorly placed, not at the posterior border of the
symphysis, and this is a derived trait. On the contrary,
in lateral view (figure 8.7 [2]), the ventral border of the
ramus is convex at the level of the posterior symphyseal
border; this trait is associated with a brevirostrine mandible that has a true elephantine beak. The mandibular
canal is well preserved on this mandible. The small anterior mandibular foramen is anterior to the symphyseal posterior border. The large posterior mandibular
foramen is situated at the level of the first plate of dP4
that is, at the anterior end of the dental arch.
The dP4s are one plate longer than M1s. They have
six plates and a postcingulum. The number of plates of
M1 can be expressed in alternate ways: M1s either have
five plates and a strong, well-separated postcingulum or
six plates and no postcingulum. In either case, M1s have
a less derived laminar formula than dP4 has.
In occlusal view, the crown of dP4 is narrow, with an
attenuated posterior widening at the level of the fifth
plate; this is a derived condition. The dP4s are fully
worn, the occlusal plane is concave, and the first three
plates are excavated. The enamel band is much wrinkled
(a trait of deciduous teeth). The wear facets indicate the
presence of weak columns ( conules), mostly integrated in the wrinkling made by the digitations of the
plates. Consequently, the enamel band is rather straight
without a loop. These columns appear on the posterior
face of the first plate (both right and left sides), on the
posterior face of the second plate (left side), and on the
anterior face of the fifth plate (right side). In occlusal
view, the plates vary from straight-concave (plates 1 and
2) to convex-convex (plates 36).
The erupting M1s (figure 8.8 [13]) are narrow without posterior widening; this is a derived condition. As
already mentioned, the last plate (sixth plate or postcingulum) is fully separated from the fifth plate. It is made
of two main cusps (right side) or three (left side). Cement fills the interlophs and covers entirely the posterior face of the crown. The most anterior plates are
made of numerous cusps (eight on the second loph of
the left side), while the posterior plates are less derived
(only four cusps). A pretrite column ( conule) is seen
in the first interloph of both right and left M1s, on the
posterior face of the first plate. No other conule is seen.
In occlusal view, the anterior plates (14) are concaveconcave (a derived condition) and the posterior (plates
5 and 6 or postcingulum) are convex-convex.
LT 26327 is the posterior half of a partial M1 or 2
(figure 8.8 [4]). As in LT 23783, the crown is narrow,
not enlarged posteriorly, plate ?4 is straight-concave,
and plate ?5 is convex-convex. The main difference
from LT 23783 is that the last plate is more robust and
made of four cusps.

Discussion
The mandible LT 23783 clearly demonstrates the presence of an elephantid previously unrecognized from
Lothagam. The association of several derived character
states indicates that this elephantid is different from
Stegotetrabelodon orbus present in the Upper Nawata
Formation and from Primelephas gomphotheroides now
recognized only in the Lower Nawata. The new species
is also different from elephantid species found in the
Nachukui Formation at Lothagam and at Kanapoi.
Lack of lower tusks precludes close relationship with
S. orbus. The preserved parts of the mandibular symphysis indicate a brevirostry more pronounced than
that of P. gomphotheroides. The symphyseal rostrum is
more gracile than in Loxodonta adaurora from Kanapoi, and, unlike in L. adaurora, it is not bent downward. This mandible belongs to a young individual
(erupting M1s), while that of L. adaurora is known
from an adult with M3s in wear (Maglio 1973:23, plate
5, figures 23) but possible variation due to different
growth stages can be excluded. Unpublished mandibles
from Kanapoi of L. adaurora of the same individual
age as LT 23783 show the same angulation of the ventral border of the symphysis as that seen on the adult
(observations courtesy of M. Leakey and J. Harris personal communication). Definitely, this young mandible from Nawata is closer to mandibles of the Elephas/
Mammuthus clade, such as M. subplanifrons from
Langebaanweg, South Africa (Maglio 1973:plate 15, figure 2) or Elephas recki shungurensis from Shungura,
Ethiopia (Beden 1983:75, figures 314; plate 3-3, figure
66; 1987a).
The plates of dP4 display a wrinkled linear enamel
band similar to those of Elephas ekorensis and E. recki;
the lack of protruding loop makes them different from
P. gomphotheroides and L. adaurora. The lack of columns in the interlophids posterior to plate 2 of the M1
is another character that distinguishes P. gomphotheroides from L. adaurora. The concave-concave contour
of plates 14 of M1 is a derived trait found in Pliocene
species of Loxodonta and Elephas but not seen in P.
gomphotheroides. The convex-convex structure seen in
figure 8.7 is a more primitive condition than that
found in Pliocene elephantids.
This mandible does not belong to the most primitive species of Elephas previously knownElephas ekorensis from Ekora and Kanapoi (Maglio 1973:3334).
All characters of M1 are more primitive than those of
E. ekorensis. The M1s have only six plates and not eight,
and they are shorter. The laminar frequency is lower
(3.64.4 v. 4.36.2). The enamel is thicker (3.9 mm v.
2.13.0 mm). The crown is lower (74.6 mm v. 92.1
mm): the height index (74.3) is slightly higher than
that of P. gomphotheroides (63.571.1) and much lower

Elephantoidea from Lothagam

345

Figure 8.8 Elephantidae dentitions: 1 KNM-LT 23783, Elephas nawataensis sp. nov., left M1 of holotype mandible (Upper
Nawata), occlusal view; 2 KNM-LT 23783, right M1, occlusal view; 3 KNM-LT 23783, right M1, labial view; 4
KNM-LT 26327, Elephas nawataensis sp. nov., portion of left M1 or 2 (Upper Nawata), occlusal view; 5 KNM-LT 23795B,
Elephas cf. E. ekorensis, partial right M1 (Apak Member), occlusal view; 6 KNM-LT 26323, Elephantidae gen. and sp. incertae
sedis B, portion of right M3 (middle Apak Member), occlusal view. Scales 5 cm.

than in E. ekorensis (174.0). The posterior plates of

M1s are made of four main cusps, which is also a
primitive condition. The cusps of LT 26327 are bigger
and less gracile than those of LT 23783 so that the
primitive morphology of this tooth is more accentuated.
In conclusion, LT 23783 belongs to a previously undescribed species and is attributable to Elephas rather
than Loxodonta. The shape of the mandible is different
from that of S. orbus and P. gomphotheroides. Dental
characters are more derived than those in S. orbus and
P. gomphotheroides, but they are more primitive than
those in E. ekorensis. This mandible is allocated to a

primitive species of the genus Elephas, the first described from Miocene strata, Elephas nawataensis, sp.
nov. Morphologically, this species represents an intermediate stage between P. gomphotheroides and E. ekorensis and can be seen to be a possible forerunner of
E. ekorensis. As the upper member of the Nawata Formation is dated between 6.24 Ma and 5.5 Ma, it is
intermediate in time between P. gomphotheroides
(Lower Nawata) and E. ekorensis (Ekora, Kanapoi).
Yet, as early stages of Mammuthus evolution are still
poorly known in East Africa, only further discoveries
and investigations will yield firm diagnostic data of
early species of Elephas and Mammuthus.

346

Pascal Tassy

Elephantidae gen. and sp. indet.

(Figure 8.5 [45]; table 8.13)

Lothagam Material

Lower Nawata: 342, Rt. and Lt. M2; 26331, anterior
part of Rt. M?2 or 3.

Upper Nawata: 350, partial mandible with Rt. and
Lt. M2.

Horizon indet.: 376, isolated plate of ?Rt. M3.
Most specimens allocated to this taxon are from the
early collection. One fragment of a lower molar from
the Lower Nawata (LT 26331) is new. One was previously described with different interpretations (LT 350),
and the two other specimens had not been published
(LT 342 and LT 376).
Maglio (1973:plate 2, figure 4) described LT 350 under the number LT 342, and this mistake was perpetuated by later authors (Maglio and Ricca 1977; Tassy
1986; Kalb and Mebrate 1993; Froehlich and Kalb
1995). Maglio was aware of both specimens LT 342
(field number 57-67K) and LT 350 (field number 12067K); the fact that they are nearly identical probably
explains Maglios mislabeling. In this contribution I adhere to the original accession numbersthat is, LT 350
is Maglios LT 342, whereas LT 342 is a different specimen. The reader should be aware that Maglios (1973)
figure 4 of plate 2 is the right ramus with M2 of LT 350,
but I suspect that Maglios (1973:18) listing of LT 342,
complete left lower M2, and partial right lower M2 is
indeed LT 342 and not the specimen figured under this
number in his plate 4.
The horizontal rami and M2s comprising LT 350
were previously allocated to Stegotetrabelodon orbus by
Maglio (1973:plate 2, figure 4) and Maglio and Ricca
(1977:plate l, figure 7). Later they were allocated to Primelephas gomphotheroides (Froehlich and Kalb 1995:
390; Kalb and Mebrate 1993:44; Tassy 1986:113), but
this identification is no longer supported here. They
belong to a taxon clearly distinct from these two species.
The mandible LT 350 consists only of portions of
right and left rami; it is missing those parts anterior to
the alveoli of M1 and behind the level of the origin of
the ascending branch. At its posterior edge, the cross
section of the ramus is different from that of the holotype mandible of Stegotetrabelodon orbus; it is both
smaller (height 112, width 135) and less deep (the
height of S. orbus is circa 160180). The rounder cross
section of LT 350 is more characteristic of a brevirostrine mandible.
The M2s of LT 350 (left M2; figure 8.5 [5]) were originally described by Maglio (1970, 1973). The right and
left M2s of LT 342 (left M2; figure 8.5 [4]) are nearly

identical in size, morphology, and state of wearthe

first four plates being worn. There are, in all, five plates
and a large postcingulum that forms the posterior side
of the molars and is completely separated from the fifth
plate. The crown is enlarged posteriorly at the level of
the fifth plate, which is a primitive trait noted by Maglio
(1973:19). Columns ( central conules) are prominent
on the posterior face of plates 13 (LT 350 and right
M2 of LT 342) and of plates 14 (left M2 of LT 342).
The wear facet of the columns of the first (LT 350) and
the first and second interlophid (LT 342) is that of a
strong, rounded loop. There is no column (or even enamel thickening) on the anterior faces of the plates. The
cement fills the interlophids and embeds the posterior
cingulum, except in the right M2 of LT 350. The plates
are made of numerous cusps (apical digitations), and
all are concave-concave in occlusal view.
Allocated to the same taxon are LT 376, an isolated
plate, and LT 26331, the anterior part of a right lower
molar. LT 26331 from the Lower Nawata consists of a
very worn precingulum and first plate, worn second
plate, and part of third plate. The plates are concaveconcave, and the enamel is thick (4.4 mm measured on
the pretrite half of third plate). The wear facet of the
posterior loop of the second plate is confluent with the
third loop; this means that the pretrite half of the third
plate is inflated at its base, a character that cannot be
seen on less worn teeth.

Discussion
The two individuals represented by LT 342 and LT 350
differ from both Stegotetrabelodon orbus and Primelephas gomphotheroides. Characters compatible with their
allocation to S. orbus are primitiveM2s with five plates
and posterior enlargement. The posterior enlargement
is also retained in P. gomphotheroides. Derived characters previously used to exclude LT 350 from the original
hypodigm of S. orbus as conceived by Maglio (1973) are
(1) the numerous digital apications (Tassy 1986) and
(2) concave-concave plates (Kalb and Mebrate 1993).
All plates of these lower M2s are concave-concave, including the last one; in some lower molars of S. orbus
the two anterior plates are concave-concave, as is the
second plate of the only known M2 of P. gomphotheroides. The M2s are much wider than those of P. gomphotheroides, but, unlike in the right M2 LT 358 of P.
gomphotheroides, no anterior columns are present. The
plates are thinner and more slender than in P. gomphotheroides, and their labial and lingual sides are less
conical. The enamel is thinner (4.04.1 vs. 5.46.9 in P.
gomphotheroides). All these traits distinguish these M2s
also from S. orbus. Except for the wide posterior end of
the M2s, all characters displayed by LT 350 and LT 342

Elephantoidea from Lothagam

are more derived than those of P. gomphotheroides.

However, the laminar frequency is comparable to that
of P. gomphotheroides (3.4).
The lower M3 (LT 352) was described earlier in this
contribution as a peculiar example of S. orbus (plates
thinner with more numerous apical digitations). The
association of this M3 with the M2s described here is
unlikely because of the primitive convex-convex shape
of the plates of LT 352.
The morphology of these M2s is seen in no other
species, and the narrow M1s of Elephas nawataensis do
not match with the M2s described here. Because of their
singular morphology, it seems very unlikely that LT 342
and LT 350 are variants of S. orbus, P. gomphotheroides,
or Elephas nawataensis. They appear to belong to a different species, but more precise identification requires
a larger sample and other teeth. The stratigraphic range
of this unidentified species is uncertain. LT 342 is questionably assigned to the Lower Nawata (but could well
be Upper Nawata, according to M. Leakey, personal
communication), and LT 350 is from the Upper Nawata. Only LT 26331 is undoubtedly from the Lower
Nawata, but the allocation of this worn fragment to the
same taxon is tentative. More specimens are needed to
assess the stratigraphic distribution of this elusive species. Although a precise systematic identification is not
possible from the present material, it is sufficient to illustrate the important diversity of elephantids as early
as the Late Miocene.

347

enlarged at the level of plates 5 and 6. The posterior

column of plates 5 and 6 is reduced and nearly merged
in the folded figures of the enamel band. The seventh
plate is narrow and consists of three main cusps. All
plates are concave-concave, except the seventh which is
convex-convex.
The left M1 is not entirely visible. It has at least seven
plates with the probable formula of 7. At least the
first five plates are concave-concave in occlusal view.
The plates are made of five main cusps, and the highest
cusps are the two cusps on each side of the remnant
median sulcus. Pretrite columns are present in the first
four interlophids: the most distinct is posterior to the
second plate, posterior to the third plate, and anterior
to the fourth plate. Although the precise height cannot
be measured, the plates are moderately high and slightly
wider than tall (HI circa 83 calculated on the second
plate). In lateral view, the plates are forwardly inclined
with a concave anterior face.
LT 23795B, a right M1 (figure 8.8 [5]) is tentatively
allocated to the same taxon. It consists of the first three
plates. The shape of the plates is elephantine with a
moderate folding of the enamel band. Each plate possesses a posterior pretrite column. Seven apical digitations are seen on the top of the third plate, which is less
worn than the others. Cement is preserved in the interlophs.

Discussion
Elephas cf. E. ekorensis Maglio, 1970
(Figures 8.8 [5], 8.9 [12]; tables 8.14, 8.15)

Lothagam Material

Apak Member: 23795, partial Rt. M1; 26320, Lt. hemimandible with dP4 and M1.
The left hemimandible LT 26320 belongs to a young
individual with dP4 in wear and unworn M1 erupting
(figure 8.9 [12]). This mandible is brevirostrine, although the symphyseal rostrum is broken. The largest
mandibular foramen is situated at the anterior border
of dP4. The ascending ramus is high, and this morphology conforms to a cranium with a strongly elevated
basicranium. The top of the ascending branch is broken
and lacking both coronoid apophysis and condyle.
The left dP4 has seven plates and a reduced postcingulum (only one cuspule), all worn. Plates 14 possess
a distinct pretrite column ( conule), the wear of
which forms a lanceolate figure. On plates 2 and 3, a
weak anterior column is partly merged in the folded
figures of the enamel band. The crown is moderately

This elephantine mandible is different from that of

Stegotetrabelodon orbus or Primelephas gomphotheroides
from the Nawata Formation. By its brevirostry and
shape of the rami in the symphyseal area, it is closer to
Pliocene elephantines. It does not belong to Loxodonta
because neither the plates of dP4 nor those of M1 exhibit
characteristic loxodont median loops. Relatively derived
elephantine character states, such as the folding of the
enamel band due to wear, make it closer to Elephas. Yet,
the taxon is not Elephas nawataensis, the early Elephas
from the Nawata Formation. Many traits of the molars
of LT 26320 are more derived than those seen on the
holotype of E. nawataensis, the juvenile mandible LT
23783 of the same ontogenetic age. Both dP4 and M1
have more plates, a higher laminar index, and taller
height index. The morphology of M1 in particular reflects more derived traits generally found in Pliocene
species, namely taller plates, more closely spaced, with
cusps less conical and more compressed anteroposteriorly, and with concave anterior sides in lateral
view. All these characters match better with the later
species E. ekorensis and E. recki, although E. recki with
much more derived molars must be excluded. But LT
26320 is not absolutely identical to E. ekorensis, as

348

Pascal Tassy

Figure 8.9 Elephas and Loxodonta dentitions: 1 KNM-LT 26320, Elephas cf. E. ekorensis, left mandible with dP4 and M1

(upper Apak Member), lingual view; 2 KNM-LT 26320, occlusal view; 3 KNM-LT 23794, Loxodonta sp. indet. (?aff. L.
exoptata), right M3 broken anteriorly (Apak Member), occlusal view; 4 KNM-LT 26321, Loxodonta sp. indet. (?aff. L. exoptata), anterior part of right M3, (middle Apak Member), occlusal view; 5 KNM-LT 23786, Loxodonta sp. indet. (?aff. L.
exoptata), posterior part of left M3, (Apak Member). Scales 5 cm.

described from Ekora and Kanapoi, at about 4.2 Ma.

The differences noticed between the Lothagam material
described here and (1) the original hypodigm described
by Maglio (1973:3334) or (2) an unpublished specimen of E. ekorensis from Kanapoi (courtesy M. Leakey
and J. Harris) are all related to more primitive character
states displayed by the material from Apak. The dP4 of

LT 26320 has still a primitive shape (posterior enlargement). The M1 is lower crowned and it is still less high
than wide, unlike the proportions in E. ekorensis. On
the basis of these two traits alone, it is not possible to
accept a priori an individual variation and consider that
LT 26320 is a bona fide E. ekorensis. These differences
can probably be attributed to the more primitive evo-

Elephantoidea from Lothagam

lutionary stage of the Apak material. They demonstrate

that if the material from upper Apak belongs to the
same lineage as E. ekorensis, it is necessarily older than
4.2 Ma.

Loxodonta F. Cuvier, 1825

(anonymous emendation 1827)
Loxodonta sp. indet. (?aff. L. exoptata)
(Figure 8.9 [35]; table 8.16)

Lothagam Material

Apak Member: 23786, posterior Lt. M3; 23794, Rt. M3
fragment; 26321, anterior part Rt. M3.
The preserved portion of LT 23794, a right M3 (figure
8.9 [3]), consists of six convex-convex plates. All plates
except the postcingulum (two cusps) show wear. The
wear figures (the dentine is visible on the first five preserved plates) display an incipient loxodont loop. The
median area of the plates is inflated with a strong anterior column but posterior columns are weak. One
consequence of this asymmetrical development of central columns is the convex shape of the posterior side
of the plates. The laminar frequency is low. The enamel
is rather thick and not much folded.
The preserved posterior portion of LT 23786, left M3
(figure 8.9 [5]), consists of three and a half plates and
the postcingulum made of one cusp. The median area
of the plates is inflated and the wear figures make a
nearly symmetrical loxodont loop. The second preserved plate is concave-concave. More posterior plates
belong to the very end of the tooth; they are narrow
and convex-straight and convex-convex. The enamel is
thick.
The anterior part of LT 26321, right M3 (figure 8.9
[4]), is worn to the base and can only be tentatively
compared to the other M3s described above. The shape
of the plates in occlusal view is nearly rectangular. They
are concave-concave; the anterior side of the third plate
is straight. The median part of the plates is inflated. The
central columns are prominent. In the interlophids, facing anterior and posterior loops are in contact.

Discussion
These three partial molars indicate that a loxodont species very likely existed in the Apak Member of the Nachukui Formation. Its status remains obscure. The M3
is different from either Loxodonta exoptata from Koobi
Fora and Laetoli (Beden 1987b) or Loxodonta sp. indet
(Lukeino stage) from Lukeino and Uganda (Tassy

349

1994). The M3s of both taxa, like the modern L. africana,

have convex-concave plates. The concave shape of the
posterior plates is partly due to the enlargement of the
posterior loop, not realized on LT 23794. Yet, the wear
facets of LT 23794 can be found on some specimens of
the collection of Loxodonta exoptata from Laetoli described by Beden (1987b:plate 8-3, figures 18 and 23:
M2, fourth plate of M3). The M3 described here is also
different from L. adaurora from Kanapoi (Maglio 1970,
1973). L. adaurora is an atypical loxodont because the
plates of the molars are nearly straight, without inflation
medially, although a loxodont sinus can be present
(though not on every tooth). In contrast, the M3 from
Apak exhibits the normal concave-concave shape of
Loxodonta.
All in all, these partial molars seem more closely related to a group formed by Loxodonta sp. Lukeino
stage and L. exoptata than to L. adaurora. However,
three partial molars are not enough for an unequivocal
systematic conclusion. Yet, because these molars are
clearly different from those allocated to the genus Elephas and also found in the Apak Member, it can be
concluded that two contemporaneous species representing the modern elephant genera Loxodonta and Elephas were present at Lothagam between ca. 4 and 5
million years.

Elephantidae gen. and sp. incertae sedis A

(Figure 8.6 [6]; table 8.17)

Lothagam Material

Upper Nawata: 405, partial Rt. dP3.

Apak Member: 23785, portion of Rt. M3.
The two preserved (anterior) plates of LT 405, a right
dP3 (figure 8.6 [6]), are thin and linear. The anterior
cingulum is plate-like. Central columns are present on
the posterior sides of the first and second plate and on
the anterior side of the second plate.
LT 23785, a partial M3, consists of a posterior half
molar with five plates preserved and a postcingulum
made of two cusps. It is an unworn tooth embedded in
cement and only the top of the two anterior plates can
be observed. The most anterior preserved plate shows
six apical cones. This tooth has a low laminar index. It
is moderately tall.

Discussion
There is no conclusive evidence that the dP3 and M3
described here belong to the same taxon, but they share

350

Pascal Tassy

derived traits compared to Stegotetrabelodon orbus and

Primelephas gomphotheroides, and should probably be
allocated to a more evolved elephantid. The dP3 shows
a more elephantine condition compared to the smaller
dP3 (LT 22866) allocated to Stegotetrabelodon or Primelephas (see above): the plates are thinner, more linear;
the anterior cingulum is more plate-like. The M3 is taller
than the M3s of Stegotetrabelodon orbus and Primelephas
gomphotheroides, and belongs to an elephantine with a
low laminar index. Because only a small part of this
specimen is not covered in cement, it is not possible to
allocate it more precisely to an early species of Elephas
or Loxodonta or to what has been described here as
Elephantidae gen. and sp. indet.

General Conclusions
Recent fieldwork at Lothagam by Meave Leakey and
colleagues proves that the elephantoid diversity in the
Lothagam area is greater than previously thought (figure 8.10). The presence of Anancus, Stegotetrabelodon,
and Primelephas is confirmed. In addition to these
classical taxa, a primitive species of the extant genus
ElephasElephas nawataensis sp. nov.is described
from the Late Miocene strata of the Nawata Formation.
Another species, previously erroneously allocated either
to Stegotetrabelodon or to Primelephas and here called
genus and species indet., is also present in the Nawata
Formation and Upper Nawata/?Apak. Loxodonta adaurora appears at Lothagam in the Kaiyumung Member
of the Nachukui Formation, above the Lothagam basalt.

Elephantidae gen. and sp. incertae sedis B

(Figure 8.8 [6]; table 8.18)

Lothagam Material

Apak Member: 26323, portion of Rt. M3.
LT 26323 comprises the rear portion of a partial M3
(figure 8.8 [6]). The three preserved plates are linear
and made up of four or five main cusps. They are
straight-straight. The most anterior preserved plate is
worn, and the wear figure shows the existence of a partly
damaged posterior central column. No other central
column is seen. The postcingulum is bulbous, made of
five cups in a circle.

Discussion
This partial tooth exhibits no distinctive traits associated with any of the different taxa described elsewhere
in this contribution. The plates are too linear and thin
for Stegotetrabelodon orbus; the resemblance to Primelephas gomphotheroides is greater, although the plates
are also thinner. The laminar frequency is also higher
(3.75) than that of the holotype lower M3 of P. gomphotheroides (3.4), and the enamel is thinner (3.9 mm
vs. 4.9 mm). These derived characters make this tooth
closer to elephantines such as Elephas nawataensis, Elephas cf. E. ekorensis, or Elephantidae gen. and sp. indet. But the unlikely association of an M3 with straightstraight plates and an M2 with concave-concave plates
such as these of Elephantidae gen. and sp. indet. (LT
342 and LT 350) still has to be found in a single specimen. This partial tooth could then belong to a primitive member of the Elephas lineage.

Figure 8.10 Stratigraphic extension of the elephantoid taxa

recognized in the Nawata Formation and Apak Member of
Nachukui Formation: LB Lothagam basalt; dash possible but uncertain extension. Stratigraphy taken from Leakey
et al. (1996) and McDougall and Feibel (1999).

Elephantoidea from Lothagam

This provenance equates with that of the skull LT 353

described from Lothagam 3 by Maglio (1973:23) and of
which only one M3 survived transportation from Harvard University to Nairobi after Maglios study. An indeterminate species of the genus Loxodonta, differing
from L. adaurora and compared here to L. exoptata, is
described from the Apak Member of the Nachukui Formation. From the same unit, a species of the genus Elephas intermediate in molar morphology between Elephas nawataensis and Elephas ekorensis is here assigned
to Elephas cf. E. ekorensis.
A consequence of these discoveries is that the differentiation of elephantines is now seen to precede 4 Ma
and to have occurred in the Late Miocene. At Lothagam,
Elephas occurs in the upper part of the Nawata Formation ca. 6.75.2 Ma. Loxodonta occurs in the Apak
Member of the Nachukui Formation, ca. 5.54.2 Ma.
This species of loxodont is likely to be an intermediate
between Loxodonta sp. Lukeino stage (described from
Lukeino [ca. 6 Ma] and in the Nkondo Formation of
Uganda [ca. 64 Ma]) and Loxodonta exoptata described from Laetoli. Thus two loxodont lineages were
contemporaneous in the late Miocene: this taxon and
the L. adaurora lineage. Although the morphological
evidence is scanty, the Apak loxodont fits with the previous scheme that considers Loxodonta adaurora as an
offshoot of the loxodont lineage (Beden 1983; Kalb and
Mebrate 1993).
The taxon here called Elephantidae, gen. and sp.
indet., from the Nawata Formation and Upper Nawata/?Apak (ca. 85 Ma), certainly played a role in the
differentiation of elephantines. Because this taxon is
contemporaneous with both Stegotetrabelodon orbus
and Primelephas gomphotheroides, the hypothesis that P.
gomphotheroides was the stem species of elephantines is
clearly an oversimplification. Despite all the important
fieldwork done in the Lothagam area, with its bounty
of new specimens representing new taxa, our knowledge
today of Primelephas gomphotheroides is nearly the same
as it was in the 1970s. At Lothagam the species is restricted to four individuals only, certainly a frustrating
state of affairs for students of elephantine evolution.
If the origin and differentiation of modern genera of
elephants occurred earlier than previously thought, it is
also worth noticing that trilophodont gomphotheres
persisted in East Africa up to the latest Miocene. An
isolated trilophodont M1 from the uppermost Nawata
Formation (ca. 5.2 Ma) is the only evidence for this
statement but is enough to contrast elephantoid biodiversity in the Late Miocene at Lothagam with that of
Sahabi, which is the only other African locality with a
trilophodont gomphothere ca. 65 Ma.
New discoveries at Lothagam do not include stegodonts. Tassy (1994), who described numerous stegodont molars from Uganda, interpreted the absence of

351

this taxon at Lothagam to emphasize the separation between the biota of the Eastern Rift and Western Rift.
However Sanders (1999) has recently described the discovery of Stegodon at Mpesida in Kenya.
After recent fieldwork by the National Museums of
Kenya expeditions from 1989 to 1993, Lothagam is confirmed as the type area of three elephantoid species:
Stegotetrabelodon orbus, Primelephas gomphotheroides,
and the new Elephas nawataensis. Nevertheless, it
should be remembered that the fossil evidence for elephantid evolution is mainly based on molar morphology. Often when associated character states given by
different organs are found together, we are forced to
alter previous hypotheses. Consequently, we will continue to await new information from future discoveries.

Acknowledgments
I am especially eager to thank Meave Leakey, former
head of paleontology at the National Museums of
Kenya, for having given me the opportunity to study
the Lothagam material. My research in the museum in
1997 benefits from her invaluable help and goodwill. I
also thank her and John M. Harris for letting me look
at the unpublished material from Kanapoi. My stay at
Nairobi was financed by the CNRS (UMR 8569, dir. P.
Janvier). Many thanks to Robert Campbell, who is responsible for the fine photographs of this article and to
F. Pilard, D. Serrette, and D. Visset for their artistic
help.

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Table Abbreviations

L length
LF laminar frequency
P number of plates; x prefix anterior cingulum;
x suffix posterior cingulum; prefix anterior
portion missing; suffix posterior portion missing
po posttrite half loph (po1, 2, etc., first posttrite half
loph, second posttrite half loph, etc.)
pr pretrite half loph (pr1, 2, etc., first pretrite half
loph, second pretrite half loph, etc.)
W width (in parentheses the plate or loph measured)

overestimated measurement
incomplete measurement
[ ] indices and measurements in brackets indicate an
estimation or reconstruction
E enamel thickness
H maximum crown height (in parentheses the plate
or loph measured)
HI height index (100H/W)
I index of robustness (100W/L)
KNM National Museums of Kenya, Nairobi

TABLE 8.1 Measurements (in mm) for Dentition of Anancus kenyensis from Lothagam

HI

34.1 (po3)

(76.8) (3)

91.2

dP4
KNM-LT 28567

e69.0

44.4 (3)

KNM-LT 346

45.2

44.3

52.0 (1)

38

65

(1)

48.1 (pr1)

(48.9)

74.0 (1)

63 (1)

53.4 (pr1)

84.8 (1)

(69.6) (3)

51.7 (pr3)

(74.3) (3)

M1
KNM-LT 383

(po1)

1 or 2

KNM-LT 340
KNM-LT 23790

133.0

KNM-LT 341B
dP4
KNM-LT 23781

84.6

42 (34)

57.5 (4)

KNM-LT 341A

62 (3)

6.8 (pr?3)

KNM-LT 341C

M1
KNM-LT 361
M3
(57)

(5)

TABLE 8.2 Measurements (in mm) of M1 of Gomphotheriidae gen. and sp. indet. from Lothagam

KNM-LT 26324

89.6

e56 (3)

e62.5

3.2 (po3)

TABLE 8.3 Measurements (in mm) of the Cranium of Stegotetrabelodon orbus (KNM-LT 26318) from Lothagam

Parameter
Palatal length, from the anterior alveoli to the choanae

Measurement
250.0

Internal maximal width of the palate

79.0

External maximal width of the palate

270.0

Internal width of the palate taken at the anterior grinding teeth

43.8

Minimal palatal width between the interalveolar cristae ( maxillary ridges)

41.0

Source: Parameters from Tassy (1996b:94).

TABLE 8.4 Measurements (in mm) of Second and Third Molars of Stegotetrabelodon orbus from Lothagam

LF

HI

157.8

92.4 (4)

58.55

3.25

KNM-LT 347

210.0

105.9 (3)

50.4

3.2

KNM-LT 354

103.2 (3)

2.85

73.0 (3)

70.7

(3)

KNM-LT 359 r

107.9 (2)

77.2 (4)

64.0

(2)

M2
KNM-LT 354
M

KNM-LT 366

86.0 (3)

49.7

3.25

KNM-LT 367

173.1
234.6

103.4 (4)

44.1

2.8

KNM-LT 26318r

241.8

110.0 (3)

45.5

2.7

KNM-LT 26318 l

244.7

107.4 (23)

43.4

62.8 (2)

165.0

89.0 (5)

53.9

KNM-LT 344 r

KNM-LT 344l

KNM-LT 349

(?)7

233

78.3 (2)

33.6

64.5 (5)

73.7

(5)

4.0

44.6 (2)

71.0

(2)

3.6

3.7

3.4

M1 or 2
3

KNM-LT 26334
M2
KNM-LT 354
M3

5.26.5

KNM-LT 352

?8

e260.0

102.4 (?4)

e39.4

KNM-LT 354

266.0

87.9 (3)

33.0

KNM-LT 355 r

KNM-LT 355 l

KNM-LT 359 r

263.0

108.5 (3)

41.2

2.9

72.7 (3)

67.0

(3)

KNM-LT 359 l

259.0

108.0 (3)

41.7

3.0

68.0 (3)

62.9

(3)

KNM-LT 23791

211.3

87.8 (4)

41.5

3.65

47.2 (7)

74.4

(7)

83.6

81.0

2.5

60.0 (5)

60.6 (5)

78.7

89.5

(3)

5.0 (pr2)

(3)

5.4 (pr)

5.2 (po)

5.4

TABLE 8.5 Comparative Measurements of M2 of Stegotetrabelodon

LF

ET

HI

S. orbus (Lothagam)

157.8

92.4

3.25

S. orbus (Adu-Asa)

145.0

e85.0

45.0

4.0

54.0

S. syrticus (Sahabi)

45

158.0172.0

96.0

49.0

45

174.4176.9

101.4

50.2

2.93.0

49.5

S. syrticus (Shuwaihat)c
a
b
c

Kalb and Mebrate (1993).

Gaziry (1987).
Tassy (1999).

TABLE 8.6 Comparative Measurements (in mm) of M3 in the Genera Stegotetrabelodon and Primelephas

S. orbus (Lothagam)
S. syrticus (Sahabi)

S. syrticus (Shuwaihat)

LF

ET

HI

67

173.1244.7

86.0110.0

64.577.2

2.73.25

5.57.0

64.073.7

232.0242.0

109.8126.1

73.080.1

2.83.2

6.07.1

65.667.2

P. gomphotheroides (Lothagam)
a

e103

211.6

96.6

59.7

2.9

60.3

52.5 (4)

3.1

57.1

Maglio (1973).
Tassy (1999).

TABLE 8.7 Comparative Measurements (in mm) of M3 in the Genera Stegotetrabelodon and Primelephas

S. orbus (Lothagam)
S. syrticus (Sahabi)

LF

ET

HI

78

211.3266.0

78.3108.4

68.078.7

2.53.7

5.06.5

62.989.5

280.0317.4

115.0123.4

57.074.1

2.63.1

5.86.0

60.1

S. syrticus (Shuwaiat)

109.1

62.0

3.0

6.6

56.4

Stegotetrabelodon sp. (Uganda)c

68

241.6

94.0102.0

81.0

3.0

5.46.8

79.0

P. gomphotheroides (Lothagam)

249.5

92.6

58.8 (5)

3.4

5.6

64.3

a
b
c

277.0

Modified from Maglio (1973).

Tassy (1999).
Tassy (1995).

TABLE 8.8 Measurements (in mm) of ?left P4 of Primelephas gomphotheroides from Lothagam

KNM-LT 23782

HI

43

52.1

121.2

22

42.7

2.6

TABLE 8.9 Measurements (in mm) of Molars of Primelephas gomphotheroides from Lothagam

LF

HI

170.8

87.7 (4)

51.3

3.5

59.5 (4)

67.8 (4)

167.1

85.6 (4)

51.2

3.4

58.4 (4)

68.2 (4)

208.0

96.6 (1)

46.4

3.1

55.4 (4)

60.1 (4)

5.7 (po2)

106.0

60.3 (4)

56.9

4.4

59.0 (4)

M2
KNM-LT 358 r
KNM-LT 358 l
M

KNM-LT 351
M1
KNM-LT 358 r
KNM-LT 358 l

KNM-LT 375

94.0

KNM-LT 358 r

170.0

e80 (5)

e47

KNM-LT 358 l

79.4 (6)

249.0

92.7 (3)

62.6

4.65

66.6

4.0

3.0 (pr2)

3.65

54.7 (4)

74.3 (4)

5.46.2 (pr5-po6)

53.7 (6)

67.6 (6)

6.9 (pre6)

37.2

3.4

67.2 (5)

73.8 (5)

4.9 (pr2)

M2

M3
KNM-LT 351

TABLE 8.10 Measurements (in mm) of Premolars and Deciduous Premolars Allocated to Stegotetrabelodon or

Primelephas

55.5

4
2

HI

39.5 (3)

71.2

25.4 (3)

64.7

41.1 (3)

63.5

17.3

14.7 (2)

85.0

45.7

31.0 (3)

67.8

21.0 (3)

67.7 (3)

KNM-LT 26329

32.8

25.7 (2,3)

78.3

15 (2)

58.4 (2)

KNM-LT 26339

33.8

25.0 (2)

74.0

dP

?2

KNM-LT 26326

22.7

dP3
KNM-LT 22866

64.3 (3)

1.5 (3)

dP

KNM-LT 434
dP2
KNM-LT 26332
dP3
KNM-LT 26332
P3

TABLE 8.11 Measurements (in mm) of the Mandible of Elephas nawataensis sp. nov. (KNM-LT 23783) from Lothagam,

Upper Nawata

Parameter

Measurement

Mandibular width taken at the root of ascending rami

335

Width of the horizontal ramus taken at the root of the ascending branch

113 (left side)

Width of the horizontal ramus, taken at the anterior of dP3

50.3

Width of the horizontal ramus, taken at the anterior of dP4

64

Posterior symphyseal width

159.1

Anterior symphyseal width

45.1

Height of symphyseal rostrum (taken at 60 mm of the tip)

25.8

Internal width between the dP4s

63.2

Height of the horizontal ramus taken at the root of the ascending branch

e103 (left side)

Source: Parameters from Tassy (1996b:95).

TABLE 8.12 Measurements (in mm) of Molars of Elephas nawataensis sp. nov. from Lothagam

KNM-LT 23783 (rt)

KNM-LT 23783 (lt)

KNM-LT 23783 (rt)

KNM-LT 23783 (lt)

LF

HI

111.1

51.9 (4)

46.7

50.3 (5)

2.1 (po4)

145.3

63.0 (2)

43.2

47.0 (2)

74.6 (2)

138.8

61.8 (4)

44.5

45.6 (3)

74.4 (3)

69.5 (?4)

3.4

4.3

dP4

M1

M1 or 2
KNM-LT 26327

TABLE 8.13 Measurements (in mm) of Molars of Elephantidae gen. and sp. indet. from Lothagam

LF

HI

KNM-LT 342 (rt)

95.4 (5)

56.6 (5)

59.3

KNM-LT 342 (lt)

96.1 (5)

3.0

4.1 (pr3)

KNM-LT 350 (rt)

171.1

103.3 (5)

60.4

3.4

4.0 (po2)

KNM-LT 350 (lt)

173.2

101.9 (5)

58.8

3.3

4.0 (po2)

67.4 (2)

4.4 (pr)

M2

M?2 or 3
KNM-LT 26331

TABLE 8.14 Measurements (in mm) of the Hemimandible of Elephas cf. E. ekorensis, KNM-LT 26320

Parameter

Measurement

Width of the horizontal ramus taken at the root of the ascending branch

99.4

Height of the horizontal ramus taken at the level of the fifth interlophid of dP4

113.0

Height of the horizontal ramus taken at the root of the ascending branch

105.2

Source: Parameters from Tassy (1996b:95).

TABLE 8.15 Measurements (in mm) of Elephas cf. E. ekorensis from the Apak Member of the Nachukui Formation

LF

HI

67.0 (3)

4.85

120.4

58.6 (5)

48.7

6.3

(143)

c60 (2)

c41.9

5.3

(53)

(83.3)

KNM-LT 23795B
dP4
KNM-LT 26320
M1
KNM-LT 26320

TABLE 8.16 Measurements (in mm) of Loxodonta sp. indet. (?aff. L. exoptata) from the Nachukui Formation

95.4

LF

HI

4.3

3.64

M3
KNM-LT 23794
M3
KNM-LT 23786

4.9 (po)

KNM-LT 26321

72.4 (3)

3.8 (pr3)

TABLE 8.17 Measurements (in mm) of the Upper Molar of Elephantidae gen. and sp. incertae sedis A from Lothagam

LF

HI

41.2 (2)

20.8 (2)

168.5

85.1

72.8

88.1

dP3
KNM-LT 405
M

KNM-LT 23785

TABLE 8.18 Measurements (in mm) of M3 of Elephantidae gen. and sp. incertae sedis B from Nachukui Formation

KNM-LT 26323

LF

HI

126

e81

3.75

3.9

8.2
Deinotheres from the Lothagam Succession
John M. Harris

Deinotheres were rare components of the Lothagam biota, but deinothere enamel is distinctive and
Deinotherium bozasi is documented from all four terrestrial members of the succession.

Figure 8.11 Restoration of Deinotherium bozasi by Mauricio Anton. Shoulder height estimated at about 3 meters.

360

John M. Harris

Deinotheres were evidently restricted in distribution to

Neogene and Quaternary localities of the Old World
and appear to have originated in Africa as a sister group
of the Elephantoidea. The earliest known representatives of the family, from Early Miocene localities in
Kenya, had attained elephantine size but were characterized by dental and postcranial features that differentiated them unequivocally from the elephantoids.
The dentition, in particular, is fundamentally diagnostic, and the combination of characters that is peculiar
to the deinotheresabsence of upper tusks, downward
curvature of the lower tusks, bifunctional and (mostly)
bilophodont cheek teeth that are retained throughout
the life of the individualreadily distinguish representatives of this family from other proboscideans.
Prodeinotherium hobleyi is characteristic of Early
Miocene faunas from East and North Africa, and similar
deinotheres became established in Europe (P. bavaricum) and Asia (P. pentapotamiae) by the onset of the
Late Miocene. Toward the end of the Miocene, larger
and more progressive forms representing the genus
Deinotherium became established in Eurasia, initially
coexisting with, and eventually replacing the smaller
Prodeinotherium species. In Africa, replacement of P.
hobleyi with D. bozasi reportedly occurred during the
time interval represented by the Namurungule Formation of the Samburu Hills in Kenya (Nakaya et al. 1984).
Deinotheres became extinct in Eurasia during the Pliocene but were still present in East Africa during the
Early Pleistocene, and they occurred as late as Shungura
Member L in the northern Lake Turkana Basin (Beden
1979).
Deinotheres are represented throughout the Lothagam succession. They are not common at any interval,
and few specimens have been collected, but deinothere
enamel is readily recognizable and has been collected
for isotopic analysis, as well as for voucher specimens
for the collections of the National Museums of Kenya.
There is no indication that P. hobleyi might have persisted from the Namurungule Formation into the lowest part of the Nawata sequence.

Systematic Description
Family Deinotheriidae
Deinotherium Kaup, 1829
Deinotherium bozasi Arambourg, 1934
(Figure 8.11; table 8.19)

Diagnosis
Species of Deinotherium with teeth of similar size to
those of D. giganteum. Skull rostrum steeply turned

down anteriorly (as in Prodeinotherium hobleyi), with

narrower external nares and rostral trough than in D.
giganteum; preorbital swelling greatly reduced and sited
just in advance of P3; occiput steeply inclined; nasals
with anterior median projection. Mandibular symphysis flexed at 90. Upper premolars lacking the subsidiary
styles of those of D. giganteum.

Lothagam Material

Lower Nawata: 26346, tooth fragments.

Upper Nawata: 26344, tooth fragment.

Apak Member: 23806, molar fragments; 26345, tooth
fragments.

Kaiyumung Member: 23677, Rt. dP4 fragment.

Horizon indet.: 356, RM1 and molar fragments.
Only two partial teeth were recovered from Lothagama lower deciduous fourth premolar fragment
from the Kaiyumung Member and an incomplete right
upper molar recovered in 1967 and hence of unknown
provenance. The upper molar seemed to be slightly
smaller than a comparable specimen from Koobi Fora
(Harris 1983) but was not complete enough to afford
detailed comparison. Other enamel fragments were
consistent with derivation from D. bozasi rather than P.
hobleyi.

Discussion
The bilophodont brachyodont teeth of deinotheres
were superficially similar to those of tapirs and were
admirably suited for processing soft vegetation. It is interesting to note, therefore, that whereas all the elephantoid proboscideans sampled from the Lothagam
assemblages adopted a C4 grazing diet during the course
of accumulation of the Nawata Formation, the deinotheres persisted as C3 browsers. Indeed, deinotheres and
giraffine giraffes were the only large African mammals
to have retained a diet of C3 browse throughout their
known history. However, whereas the d18O content of
giraffid tooth enamel indicates these mammals derive(d) much of their water from their food, the more
negative d18O values from deinothere tooth enamel suggests that deinotheres drank from local water sources.

Acknowledgments
I thank the government of Kenya and museum trustees
of the National Museums of Kenya for permission to
study the Lothagam deinothere material. I also thank
the curatorial and preparation staff of the palaeontology

Deinotheres from the Lothagam Succession

division of the National Museums of Kenya, Nairobi,

for making the material available for study.

References Cited
Arambourg, C. 1934. Le Dinotherium des gisements de lOmo.
Comptes Rendus de la Societe Geologique de France 1934:
8687.
Beden, M. 1979. Les elephants (Elephas et Loxodonta) dAfrique

361

orientale: Systematique, phylogenie, interet biochronologique. Ph.D. diss., University of Poitiers.

Harris, J. M. 1983. Family Deinotheriidae. In J. M. Harris, ed.,
Koobi Fora Research Project. Vol. 2. The Fossil Ungulates: Proboscidea, Perissodactyla, and Suidae, pp. 2239. Oxford: Clarendon Press.
Kaup, J. J. 1829. Deinotherium giganteum. Isis 4:401404.
Nakaya, H., M. Pickford, Y. Nakano, and H. Nishida. 1984. The
Late Miocene large mammalian fauna from the Namurungule Formation, Samburu Hills, northern Kenya. African
Study Monographs, Supplementary issue 2:87131.

TABLE 8.19 Measurements (in mm) of M1 of

Deinotherium bozasi

LT 356
Length

91.5

Protoloph tr

70.1

Metaloph tr

76.7

Tritoloph tr