Journal of Human Evolution 80 (2015) 179e183

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A 750,000 year old hominin molar from the site of Nadung'a,

West Turkana, Kenya
Scott D. Maddux a, *, Carol V. Ward a, Francis H. Brown b, J. Michael Plavcan c,
Fredrick K. Manthi d
a
Department of Pathology and Anatomical Sciences, University of Missouri, Columbia, MO 65212, USA
b
Department of Geology and Geophysics, University of Utah, Salt Lake City, UT 84112, USA
c
Department of Anthropology, University of Arkansas, Fayetteville, AR 72701, USA
d
Department of Earth Sciences, National Museums of Kenya, P.O. Box 40658-00100, Nairobi, Kenya

a r t i c l e i n f o

Article history: Geological age of Nadung'a

Received 19 October 2013
Accepted 12 November 2014
Available online 5 January 2015
Keywords:
KNM-WT 51261
Nariokotome
Africa
Dentition
Homo
Introduction
Compared to that of Eurasia, the hominin fossil record of Middle
Pleistocene Africa is exceedingly sparse (Manzi, 2004, 2011;
Rightmire, 2008). This is especially true for the earliest stages of
the Middle Pleistocene (~780e600 kya), for which only a handful of
fragmentary African fossil specimens have been recovered (Asfaw
et al., 2002; Manzi, 2004, 2011). In 2010, the West Turkana Pale-
ontology Project from the National Museums of Kenya recovered a
hominin molar, KNM-WT 51261, from the site of Nadung'a on the
west side of Lake Turkana. The molar derives from the level of the
Silbo Tuff dated to 750,000 (þ/20,000) years ago (McDougall and
Brown, 2006), making Nadung'a one of the few early Middle
Pleistocene fossil localities known in Africa. Here, we present
geochronology and comparative analyses to provide an interpretive
context for the KNM-WT 51261 specimen.
* Corresponding author.
E-mail address: madduxs@missouri.edu (S.D. Maddux).
Exposed west of Lake Turkana in northern Kenya, the KNM-WT
51261 molar was discovered at the site of Nadung'a, located within
the upper part of the Nariokotome Member of the Nachukui For-
mation (Fig. 1). As a part of the extensively studied Omo Group of
East African Pliocene-Pleistocene sediments (de Heinzelin, 1983),
the geological and radiometric framework of the entire Nachukui
Formation is known with confidence. Accordingly, the Nariokotome
Member is securely dated to between 1.3 and 0.7 Mya (Brown et al.,
2006; McDougall and Brown, 2008). The KNM-WT 51261 molar
specifically derives from the surface of marginal lacustrine deposits
at the level of the Silbo Tuff dated to 750,000 (þ/20,000) years ago
(McDougall and Brown, 2006) and exhibits the dark mottled col-
oring characteristic of fossils from the Nachukui Formation in this
area (Fig. 2). Exposures at Nadung'a (~4.14 N, 35.84 E) lie between
450 and 460 m and exhibit clear beach features, with most of the
area covered with a thin veneer of deposits left during the recession
of Lake Turkana. These ancient beach deposits are comprised of fine
sandstones, possibly because they developed in an embayment
north of the Nariokotome channel, and suggest that the tooth was
interred on the western shore of Lake Turkana during an ancient
high stand.
Morphological description of KNM-WT 51261
KNM-WT 51261 is a permanent left mandibular molar germ.
More than a third of the total crown is formed, including the entire
buccal groove that extends to what would have likely been the
widest portion of the crown (Fig. 2). As the tooth never erupted,
neither occlusal nor interproximal wear have altered the crown,
resulting in the rare preservation of near perfect occlusal
morphology. The outline of the occlusal surface is subrectangular,
elongated in the mesiodistal dimension with slight distal tapering.
Five distinct cusps are visible, arranged in a Y pattern. The proto-
conid is the largest of the five cusps (25.8 mm2), followed in size by
the metaconid (24.3 mm2), hypoconid (22.8 mm2), entoconid

http://dx.doi.org/10.1016/j.jhevol.2014.11.004
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180 S.D. Maddux et al. / Journal of Human Evolution 80 (2015) 179e183

Figure 1. Stratigraphic column of the Nariokotome Member of the Nachukui Formation at Nachukui, Kaitio (south), Kaitio, and Nadung'a showing the position of the Nadung'a
molar (KNM-WT 51261) site. Detailed discussion in Harris et al. (1988).

(17.4 mm2), and hypoconulid (13.5 mm2). The size of the hypo- positioned groove, while the distal marginal ridge is comparatively
conulid corresponds to a grade 3 in the Arizona State University thick and uninterrupted. The central groove exhibits a clear offset
Dental Anthropology System (ASUDAS: Turner et al., 1991). The between the buccal and lingual grooves, and terminates mesially in
mesial marginal ridge is narrow and intersected by a centrally a moderately expanded anterior fovea (ASUDAS grade 2) and

Figure 2. Photograph (A) and reconstructed 3D laser scan image (B) of the KNM-WT 51261 occlusal surface.
 S.D. Maddux et al. / Journal of Human Evolution 80 (2015) 179e183
distally in a relatively small but deeply excavated posterior fovea.
The anterior fovea of KNM-WT 51261 conforms to the “classic”
morphology of Martino
n-Torres et al. (2008, 2012) in being a
triangular depression that coalesces with the mesial marginal
ridge, rather than a deep pit-like fovea (see also Hrdli cka, 1924;
Scott and Turner, 1997). The posterior fovea presents as a distal
bifurcation of the central groove. The enamel distal to this fovea is
confluent with the distal marginal ridge and is not sufficiently
differentiated to classify as an accessory cusp 6 (entoconulid/
tuberculum sextum). The tooth does not present a cusp 7 (meta-
conulid/tuberculum intermedium). The three distal cusps lack
distinct supplemental grooves and accessory ridges, while the
protoconid and metaconid each exhibit a supplemental groove
delineating a distal accessory ridge on each cusp. These distal
accessory ridges both slope gradually into the central fossa and are
clearly separated by the central groove, and thus do not form a
distal trigonid crest (Scott and Turner, 1997; Bailey et al., 2011). As
the mesial portions of the protoconid and metaconid are similarly
separated by the central groove, no mid-trigonid crest is present
(Bailey et al., 2011). The mesial ridge of the metaconid is straight
and does not deflect distally. No protostylid is present along the
buccal surface of the crown.
The tooth measures 11.4 mm mesiodistally and 10.0 mm buc-
colingually as preserved. However, as KNM-WT 51261 is incom-
pletely formed, its dimensions may have increased slightly with
further development. To assess this possibility, the maximum
mesiodistal and buccolingual crown dimensions of 31 recent Homo
sapiens and 20 fossil hominin first mandibular molars1 were
compared to the same dimensions at 1/3 crown height
(Supplementary Online Material [SOM] Table 1). These compari-
sons reveal that, while the mesiodistal length of KNM-WT 51261
had almost certainly attained maximum adult size, the buccolin-
gual breadth of the tooth may have increased by as much as 0.4 mm
with continued development. Furthermore, as buccolingual di-
ameters of unworn, five-cusped mandibular molars rarely exceed
mesiodistal diameters in Homo (Wood and Abbott, 1983; Wood,
1991; Bermúdez de Castro et al., 1999), it can reasonably be
assumed that the buccolingual dimension of KNM-WT 51261
would not have exceeded 11.4 mm.
Comparative morphology of KNM-WT 51261
As an isolated tooth lacking positional information from inter-
proximal wear, root morphology, or accompanying mandibular
remains, exact identification of KNM-WT 51261 as a first, second, or
third mandibular molar is not conclusively possible. However, the
exceedingly simple occlusal morphology of KNM-WT 51261 likely
precludes classification as an M3, as contemporaneous fossils from
the sites of Ternifine (Algeria), Gran Dolina (Spain), and Zhou-
koudian (China) exhibit highly crenulated M3 occlusal surfaces
(Weidenreich, 1937; Arambourg, 1955; Bermúdez de Castro et al.,
1999, 2007, 2008). Moreover, the M3s of African Early Pleistocene
(e.g., KNM-ER 806, KNM-ER 992, KNM-ER 1812) and later Middle
Pleistocene Homo (e.g., Rabat, Sidi Abderrahman, KNM-BK 67,
KNM-BK 8518) are similarly characterized by complex occlusal
morphology (Arambourg and Biberson, 1956; Thoma and Vallois,
1977; Wood and Van Noten, 1986; Wood, 1991). Comparisons be-
tween well-preserved M1 and M2 teeth in Early and Middle Pleis-
tocene specimens (e.g., KNM-ER 992, KNM-WT 15000, Dmanisi
D211, Ng 8503, Sb 8103, Zhoukoudian G1-6, Gran Dolina ATD6-5,
1
Only molars with minimal occlusal wear (all cusps distinctly visible) were
included to ensure accurate assessment of 1/3 crown height. First mandibular
molars were employed to permit inclusion of younger juvenile specimens.
181
KNM-BK 67) reveal a trend toward the M2 exhibiting slightly
greater occlusal complexity compared to the M1 (Weidenreich,
1937; Wood, 1991, 1999; Walker and Leakey, 1993; Bermúdez de
Castro et al., 1999; Kaifu et al., 2005; Martino
n-Torres et al.,
2008). Further, the similar cusp areas of the protoconid
(25.8 mm2) and metaconid (24.3 mm2) in KNM-WT 51261 may
support classification as an M1, as the metaconid of the M2 is
considerably reduced among early African Homo and the Gran
Dolina fossils (Wood et al., 1983; Bermúdez de Castro et al., 1999).
Regardless of its exact positional identity, the occlusal
morphology of KNM-WT 51261 is broadly consistent with molars of
African Early and Middle Pleistocene Homo. For example, the lack of
a mid-trigonid crest in KNM-WT 51261 is consistent with the
infrequent expression of this feature among African Pleistocene
Homo, contrasting with the prevalence of mid-trigonid crests in
Eurasian Middle and Late Pleistocene Homo (Bailey, 2002a, 2002b;
Martinon-Torres et al., 2007, 2008, 2012; Bailey et al., 2011). Deeply
excavated posterior foveae are also fairly common among African
Homo molars, with specimens such as KNM-WT 15000 (M1), KNM-
ER 806 (M1, M2), and KNM-ER 820 (M1) exhibiting morphology
similar to that seen in KNM-WT 51261 (Wood, 1991; Walker and
Leakey, 1993). Conversely, compared to many Early Pleistocene
molars from East Africa (especially M2 and M3 specimens), KNM-
WT 51261 lacks an accessory cusp 7 (Rightmire, 1980; Wood and
Abbott, 1983; Wood and Van Noten, 1986; Wood, 1991).
Although the exact, fully developed buccolingual dimension of
KNM-WT 51261 cannot be ascertained with certainty, even with
generous allowances for further growth (Fig. 3; SOM Table 1) it
appears that KNM-WT 51261 would have been relatively small
once completely developed. Indeed, when compared to a diverse
sample of hominin mandibular molars,2 even the largest reason-
able size estimate for KNM-WT 51261 (11.4 MD  11.4 BL) is notably
smaller than any M1, M2, or M3 dated to the early Middle Pleisto-
cene of Africa, including contemporary fossils from East Africa (OH
22, OH 51). Moreover, as occlusal morphology argues against clas-
sification as an M3, it appears that KNM-WT 51261 likely represents
a particularly small M1 or M2. This possibility is intriguing, as a lack
of small posterior teeth among African early Middle Pleistocene
Homo has resulted in these hominins being described as exhibiting
dental “gigantism” compared to contemporaneous Eurasian pop-
ulations (Bermúdez de Castro et al., 2007; Martino
n-Torres et al.,
2007). KNM-WT 51261, however, may indicate that the lower
ranges of posterior dental size in African and Eurasian Homo were
actually relatively similar during this time period. Furthermore,
based on dental assessments of sexual dimorphism in Pleistocene
Homo (see Frayer and Wolpoff, 1985; Bermúdez de Castro et al.,
1993, 2001 for reviews), contemporaneous specimens with
smaller posterior teeth from Gran Dolina and Zhoukoudian are
generally regarded as representing females (Weidenreich, 1937;
Bermúdez de Castro et al., 2007). Accordingly, the small size of
KNM-WT 51261 in relation to other African early Middle
2
The comparative sample of hominin molars was derived from published
measurements (SOM Table 1). To maximize the range of variation in the sample,
individual antimeres were included when available, rendering a total sample of 842
specimens (285 M1s, 316 M2s, and 241 M3s). Following generally recognized
paleoanthropological conventions, the specimens are subdivided into the following
groups: Homo habilis (~1.9e1.6 Mya), African Early Pleistocene H. erectus
(~1.8e1.0 Mya), European Early Pleistocene H. erectus (~1.8 Mya), Asian Early
Pleistocene H. erectus (~1.8e1.0 Mya), H. antecessor (~940e780 kya), African early
Middle Pleistocene Homo (~780e600 kya), Zhoukoudian H. erectus (~780e640 kya),
African H. heidelbergensis (~600e200 kya), European H. heidelbergensis
(~600e200 kya), H. neanderthalensis (~200e35 kya), H. sapiens (~200 kya-present),
and H. floresiensis (~13 kya).
182 S.D. Maddux et al. / Journal of Human Evolution 80 (2015) 179e183

Pleistocene specimens may also suggest that the tooth derived from
a female individual.

Summary and conclusion

The KNM-WT 51261 molar represents one of the few fossil

specimens securely dated to the early Middle Pleistocene
(~780e600 kya) of Africa, joining a small number of fragmentary
fossils from Olduvai Gorge in Tanzania, Ternifine (Tighenif) in
Algeria, and Bodo in Ethiopia (Conroy et al., 1978; Rightmire, 1980,
1996; Geraads et al., 1986). The morphology of KNM-WT 51261
suggests classification as an M1, or possibly M2, with its occlusal
topography inconsistent with identification as an M3. Even after
developmental size adjustments, it appears that KNM-WT 51261
would have been relatively small compared to other African early
Middle Pleistocene molars, suggesting a collective African size
distribution more similar to contemporaneous Eurasian pop-
ulations than previously recognized. It is hoped that ongoing
fieldwork by the West Turkana Paleo Project at Nadung'a and
neighboring sites will further contribute to the fossil record of Af-
rican Middle Pleistocene Homo.

Acknowledgments

We thank all team members of the West Turkana Paleo Project,

the directors and staff of the National Museums of Kenya, and the
Government of Kenya. We thank Mark Teaford and the anonymous
reviewers for constructive comments which improved this manu-
script. We thank Lauren Butaric, Jill Scott, and Zachary Cofran for
insightful discussions. We also thank Erik Trinkaus and John Will-
man at Washington University in St. Louis for access and assistance
with skeletal and cast material. Funding was provided by the Lea-
key Foundation, Wenner-Gren Foundation, and the Palae-
ontological Scientific Trust (PAST) of South Africa.

Appendix A. Supplementary data

Supplementary data related to this article can be found at http://

dx.doi.org/10.1016/j.jhevol.2014.11.004

References

Arambourg, C., 1955. A recent discovery in human paleontology: Atlanthropus of

Ternifine (Algeria). Am. J. Phys. Anthropol. 13, 191e201.
Arambourg, C., Biberson, P., 1956. The fossil human remains from the Paleolithic site
of Sidi Abderrahman (Morocco). Am. J. Phys. Anthropol. 14, 467e489.
Asfaw, B., Gilbert, W.H., Beyene, Y., Hart, W.K., Renne, P.R., WoldeGabriel, G.,
Vrba, E.S., White, T.D., 2002. Remains of Homo erectus from Bouri, Middle
Awash, Ethiopia. Nature 416, 317e320.
Bailey, S.E., 2002a. A closer look at Neanderthal postcanine dental morphology: the
mandibular dentition. Anat. Rec. 269, 148e156.
Bailey, S.E., 2002b. Neandertal dental morphology: implications for modern human
origins. Ph.D. Dissertation. Arizona State University.
Bailey, S., Skinner, M., Hublin, J.-J., 2011. What lies beneath? An evaluation of lower
molar trigonid crest patterns based on both dentine and enamel expression.
Am. J. Phys. Anthropol. 145, 505e518.
Bermúdez de Castro, J.-M., Durand, A.I., Ipin ~ a, S.L., 1993. Sexual dimorphism in the
human dental sample from the SH site (Sierra de Atapuerca, Spain): a statistical
approach. J. Hum. Evol. 24, 43e56.

Figure 3. Distribution of individual computed crown areas (mesiodistal  buccolin-

gual dimensions) for Homo first (top), second (middle), and third (bottom) mandibular
molars. Crown area for KNM-WT 51261 (star) is plotted with an estimated develop-
mental increase of 0.4 mm in the buccolingual dimension. Lower whisker extension for
KNM-WT 51261 indicates size based on actual preserved dimensions, while upper
whisker extension indicates size based on maximum possible increase in the bucco-
lingual dimension (see discussion in text). Computed crown area data unavailable for
H. floresiensis third mandibular molars.
 S.D. Maddux et al. / Journal of Human Evolution 80 (2015) 179e183
Bermúdez de Castro, J.-M., Rosas, A., Nicola
s, M.E., 1999. Dental remains from
Atapuerca-TD6 (Gran Dolina site, Burgos, Spain). J. Hum. Evol. 37, 523e566.
Bermúdez de Castro, J.-M., Sarmiento, S., Cunha, E., Rosas, A., Bastir, M., 2001. Dental
size variation in the Atapuerca-SH Middle Pleistocene hominids. J. Hum. Evol.
41, 195e209.
Bermúdez de Castro, J.-M., Martino
n-Torres, M., Go
mez-Robles, A., Prado, L.,
Sarmiento, S., 2007. Comparative analysis of the Gran Dolina-TD6 (Spain) and
Tighennif (Algeria) hominin mandibles. Bull. Me
m. Soc. Anthropol. Paris 19,
149e167.
Bermúdez de Castro, J.-M., Pe
rez-Gonz
alez, A., Martino
n-Torres, M., Go
mez-
Robles, A., Rosell, J., Prado, L., Sarmiento, S., Carbonell, E., 2008. A new early
Pleistocene hominin mandible from Atapuerca-TD6, Spain. J. Hum. Evol. 55,
729e735.
Brown, F.H., Haileab, B., McDougall, I., 2006. Sequence of tuffs between the KBS Tuff
and the Chari Tuff in the Turkana Basin, Kenya and Ethiopia. J. Geol. Soc. 163,
185e204.
Conroy, G.C., Jolly, C.J., Cramer, D., Kalb, J.E., 1978. Newly discovered fossil hominid
skull from the Afar depression, Ethiopia. Nature 275, 67e70.
de Heinzelin, J., 1983. The Omo Group. Geologische Wetenschappen 85.
Frayer, D.W., Wolpoff, M.H., 1985. Sexual dimorphism. A. Rev. Anthropol. 14,
429e473.
Geraads, D., Hublin, J.-J., Jaeger, J.J., Tong, H., Sen, S., Toubeau, P., 1986. The Pleis-
tocene hominid site of Ternifine, Algeria: new results on the environment, age
and human industries. Quatern. Res. 25, 380e386.
Harris, J.M., Brown, F.H., Leakey, M.G., 1988. Stratigraphy and paleontology of the
Nachukui Formation, Lake Turkana Region, Kenya. Contrib. Sci. Los Angeles
County Mus. Nat. Hist. 399, 1e128.
Hrdli
cka, A., 1924. New data on the teeth of early man and certain fossil European
apes. Am. J. Phys. Anthropol. 7, 109e132.
Kaifu, Y., Aziz, F., Baba, H., 2005. Hominid mandibular remains from Sangiran:
1952e1986 collection. Am. J. Phys. Anthropol. 128, 497e519.
Manzi, G., 2004. Human evolution at the Matuyama-Brunhes boundary. Evol.
Anthropol. 13, 11e24.
Manzi, G., 2011. Before the emergence of Homo sapiens: overview on the Early-to-
Middle Pleistocene Fossil Record (with a proposal about Homo heidelbergensis
at the subspecific level). Int. J. Evol. Biol. 2011, e582678.
Martino
n-Torres, M., Bermúdez de Castro, J.-M., Go
mez-Robles, A., Arsuaga, J.L.,
Carbonell, E., Lordkipanidze, D., Manzi, G., Margvelashvili, A., 2007. Dental ev-
idence on the hominin dispersals during the Pleistocene. Proc. Natl. Acad. Sci.
USA 104, 13279e13282.
Martino
n-Torres, M., Bermúdez de Castro, J.-M., Go
mez-Robles, A.,
Margvelashvili, A., Prado, L., Lordkipanidze, D., Vekua, A., 2008. Dental remains
183
from Dmanisi (Republic of Georgia): morphological analysis and comparative
study. J. Hum. Evol. 55, 249e273.
Martino
n-Torres, M., Bermúdez de Castro, J.-M., Go
mez-Robles, A., Prado-Simo

n, L.,
Arsuaga, J.L., 2012. Morphological description and comparison of the dental
remains from Atapuerca-Sima de los Huesos site (Spain). J. Hum. Evol. 62, 7e58.
McDougall, I., Brown, F.H., 2006. Precise 40Ar/39Ar geochronology for the upper
Koobi Fora Formation, Turkana Basin, northern Kenya. J. Geol. Soc. Lond. 163,
205e220.
McDougall, I., Brown, F.H., 2008. Geochronology of the pre-KBS Tuff sequence, Omo
Group, Turkana Basin. J. Geol. Soc. 165, 549e562.
Rightmire, G.P., 1980. Middle Pleistocene hominids from Olduvai Gorge, northern
Tanzania. Am. J. Phys. Anthropol. 53, 225e241.
Rightmire, G.P., 1996. The human cranium from Bodo, Ethiopia: evidence for
speciation in the Middle Pleistocene? J. Hum. Evol. 31, 21e39.
Rightmire, G.P., 2008. Homo in the Middle Pleistocene: hypodigms, variation, and
species recognition. Evol. Anthropol. 17, 8e21.
Scott, G.R., Turner II, C.G., 1997. The Anthropology of Modern Human Teeth: Dental
Morphology and its Variation in Recent Human Populations. Cambridge Uni-
versity Press, Cambridge.
Thoma, A., Vallois, H.V., 1977. Les dents de l'Homme de Rabat. Bull. Me
m. Soc.
Anthropol. Paris 4, 31e58.
Turner, C.G., II, Nichol, C.R., Scott, G.R., 1991. Scoring procedures for key morpho-
logical traits of the permanent dentition: the Arizona State University dental
anthropology system. In: Kelley, M., Larsen, C. (Eds.), Advances in Dental An-
thropology. Wiley-Liss, New York, pp. 13e31.
Walker, A., Leakey, R.E. (Eds.), 1993. The Nariokotome Homo erectus Skeleton.
Harvard University Press, Cambridge.
Weidenreich, F., 1937. The dentition of Sinanthropus pekinensis: a comparative
odontography of the hominids. Palaeont. Sin. New Ser. D 1, 1e180.
Wood, B., 1991. Koobi Fora Research Project, vol. 4. Clarendon Press, Oxford.
Wood, B.A., 1999. Plio-Pleistocene hominins from the Baringo Basin, Kenya. In:
Andrews, P., Banham, P. (Eds.), Late Cenozoic Environments and Hominid
Evolution: A Tribute to Bill Bishop. Geographic Society, London, pp. 113e122.
Wood, B.A., Abbott, S.A., 1983. Analysis of the dental morphology of Plio-Pleistocene
hominids. I. Mandibular molars: crown area measurements and morphological
traits. J. Anat. 136, 197e219.
Wood, B., Van Noten, F., 1986. Preliminary observations on the BK 8518 mandible
from Baringo, Kenya. Am. J. Phys. Anthropol. 69, 117e127.
Wood, B.A., Abbott, S.A., Graham, S.H., 1983. Analysis of the dental morphology of
Plio-Pleistocene hominids. II. Mandibular molarsestudy of cusp areas, fissure
pattern and cross sectional shape of the crown. J. Anat. 137, 287e314.