YEARBOOK OF PHYSICAL ANTHROPOLOGY 159:S37–S78 (2016)

Hominin Taxic Diversity: Fact or Fantasy?

Bernard Wood1* and Eve K. Boyle1,2
1
Center for the Advanced Study of Human Paleobiology, George Washington University, Washington, DC 20052
2
Hominid Paleobiology Graduate Program, George Washington University, Washington, DC 20052

KEY WORDS hominin; diversity; species; taxonomy

ABSTRACT

In this review of the evidence for and against taxic diversity within the hominin clade, we begin by looking at the
logic and the history of simple “ladder-like” interpretations of the hominin fossil record. We then look at the hominin
fossil record in a series of time intervals and use current published evidence about the first and last appearance dates
of each taxon to decide whether a species or specimens should be included in one, or more, of the intervals. Within
each time interval, we consider the strength of the evidence that more than one hominin species is sampled. Deci-
sions about whether taxonomic diversity exists depend on what alpha taxonomic hypotheses are used and although
we use a relatively speciose interpretation of the hominin fossil record, we also explore what impact more inclusive
interpretations of alpha taxonomy would have on assessments of hominin taxic diversity. With the exception of the
oldest (7–5 Ma) all of the other time intervals have in our judgment at least one well-supported example of taxic
diversity and several have multiple examples. In summary, not all new hominin taxa are created equally, but while
taxic diversity may not be as prevalent as some have claimed, it is a feature of the hominin clade from 4 Ma until
c.40 ka years ago. Am J Phys Anthropol 159:S37–S78, 2016. VC 2016 American Association of Physical Anthropologists

INTRODUCTION Should the hominin fossil record be interpreted as sam-

pling a smaller, or a larger, number of taxa? These two
“Diversity” refers to difference and most studies of tax- contrasting interpretations of human evolutionary his-
onomic diversity interpret “difference” to mean looking tory are linked in the sense that those who favor a more
at the factors that influence the numbers and variety of speciose interpretation of the hominin fossil record gen-
species across a given space and time, or the numbers erally also interpret the structure of the hominin twig of
and variety of species sampled through deep time. Most, the TOL as more complex. These very different system-
if not all, can agree with the broad-brush statement that atic philosophies have a substantial impact on judg-
hominin evolution involves a series of time-related grade
ments about extent of taxic diversity within the hominin
shifts. This series of regionally-mosaic grade shifts
clade.
begins with creatures at the base of the hominin clade
Our ultimate aim is to consider the evidence for line-
that share more features with the extant African apes,
age diversity, but lineage diversity cannot exist without
and ends with those closer to the present day that share
fewer features with the extant African apes, and more taxic diversity. Because claims for the latter involve less
features with modern humans. But hominin taxic diver- complex hypotheses than claims about the former, in
sity, in the sense we discuss it here, does not refer to this review we focus on examining the evidence for syn-
diachronic diversity (i.e., through time). Instead, it chronic taxa (i.e., hypotheses about alpha taxonomy) and
focuses on whether there is evidence of synchronic taxo- not on the evidence for synchronic lineages (i.e., hypoth-
nomic diversity (i.e., within the same time interval) at eses about clade structure).
pre-determined temporal intervals between the esti-
mated time of origin of the hominin clade and the pres-
ent. In this review, we assess taxic diversity across the
hominin clade as a whole regardless of whether the spe- Correction added on 5 February 2016, after first online publication.
cies in a time interval are sympatric or allopatric.
One of the most contentious debates in paleoanthro- Grant sponsors: GW Provost’s Signature Program Funding, GW
Provost’s Diversity Graduate Fellowship; Grant sponsor: NSF-
pology is about the structure of the twig of the Tree of
IGERT; Grant number: DGE-0801634.
Life (TOL) that includes modern humans. Is it ladder-
like in the sense that it has a single well-defined stem,
*Correspondence to: Bernard Wood, Center for the Advanced
relatively few branches and a structure that can be Study of Human Paleobiology, George Washington University, Sci-
clearly discerned? Or is a bush a more apt metaphor for ence and Engineering Hall, 800 22nd Street NW, Suite 6000, Wash-
the hominin clade, in that there is not just one stem ington, DC 20052, USA. E-mail: [email protected]
leading up from the base, but several, each with multi-
ple branches that create a branching pattern that is DOI: 10.1002/ajpa.22902
difficult to determine? Another controversial—and Published online in Wiley Online Library (wileyonlinelibrary.
related—debate concerns hominin alpha taxonomy. com).

Ó 2016 AMERICAN ASSOCIATION OF PHYSICAL ANTHROPOLOGISTS

38 B. WOOD AND E. BOYLE

CONTEXT AND HISTORY OF HOMININ TAXIC a single phyletically gradualistic lineage (i.e., an
DIVERSITY ancestor-descendant sequence) connects the hypothetical
common ancestor of modern humans and chimpanzees/
The idea that the evolutionary history of the living bonobos within the TOL to modern humans on its
world can be represented by a tree-like branching struc- branch tip on the surface. In such a scenario at any one
ture was one of Charles Darwin’s many profound time in human evolutionary history there only needs to
insights. All living taxa are on the tips of the branches be one hominin species representing that single lineage,
that reach the surface of the TOL and all of the taxa and each of these time-successive species would be
that lived in the past are on branches within the TOL. ancestors of modern humans. Under this strict ladder-
Not that long ago it was assumed that the extant great like model there would be no hominin taxic diversity
apes—chimpanzees, bonobos, gorillas, and orangutans— (i.e., no evidence of more than one contemporary species
were more closely related to each other than they were within the hominin clade) and the consequent null
to modern humans. This was reflected in the pre- hypothesis is that every early hominin species is a mod-
molecular taxonomy that placed modern humans in their ern human ancestor. At the other end of the complexity
own family, the Hominidae, and the great apes in a dif- spectrum are models of human evolution that expect
ferent family, the Pongidae. To reflect this family-level hominins to be as diverse and speciose as some groups
division, the informal term “hominid” was used to refer of large mammals (e.g., alcelaphins, tragelaphines, ele-
to modern humans and any extinct species considered to phants). In these “bushy” scenarios, close non-ancestral
be more closely related to modern humans than to any relatives would outnumber ancestral relatives with the
of the great apes. The equivalent informal term for the consequent null hypothesis that an extinct hominin spe-
great apes was “pongid.” cies is not a modern human ancestor until proved
In the early days of DNA analyses, both hybridization otherwise.
and sequencing methods were used to generate hypothe-
ses about the relationships among living hominoids, but Single species hypothesis
the latter methods rapidly dominated, and ultimately
The first use of the term was by Milford Wolpoff in a
replaced, the former. The number of sequence-based paper entitled “‘Telanthropus’ and the single species
studies increases each year and there are now good draft
hypothesis,” in which he suggested the single species
sequences of the nuclear genomes of the chimpanzee hypothesis was an interpretive framework derived from
(Consortium, 2005), orangutan (Locke et al., 2011), the principle of competitive exclusion (i.e., that only one
gorilla (Scally et al., 2012), and bonobo (Pr€ ufer et al., species at a time can occupy a specific ecological niche)
2012). Also Das et al. (2014) have studied the differences (Wolpoff, 1968). Three years later Wolpoff published an
between the complete mitochondrial genomes of one article entitled “Competitive exclusion among Lower
Eastern (Gorilla beringei) and one Western gorilla (G. Pleistocene hominids: the single species hypothesis” in
gorilla). The observed DNA differences among the living which he outlined the arguments supporting the hypoth-
apes and modern humans, plus morphological evidence esis (Wolpoff, 1971).
(Shoshani et al., 1996; Gibbs et al., 2002; Lockwood Wolpoff ’s single species hypothesis is a restatement of
et al., 2004; Diogo and Wood, 2011), are consistent with an earlier proposal set out by Ernst Mayr who applied
the hypothesis that chimpanzees/bonobos are more the principle of the “biologically defined polytypic spe-
closely related to modern humans than they are to goril- cies” (Mayr, 1950) to the hominin fossil record. After giv-
las. If these differences are calibrated using paleontologi- ing “due consideration of the many differences between
cal evidence for the split between the apes and the Old Modern man, Java man, and the South African
World Monkeys, and if we make the assumption that apeman,” Mayr stated that he “did not find any morpho-
most of the DNA differences are neutral, this suggests logical characters that would necessitate separating
that the hypothetical ancestor of modern humans and them into several genera” (ibid, p. 110). Instead he sug-
chimpanzees/bonobos lived between about 5 and 8 Ma, gested that within a single hominin genus, Homo, the
and probably closer to 5 than to 8 Ma (Bradley, 2008). only speciation that has occurred “has been phyletic spe-
Estimates based on empirical data about generation ciation resulting in Homo sapiens” (ibid, p. 110), and
times (Langergraber et al., 2012) and mutation rates within what we now call the hominin clade he recog-
(Venn et al., 2014) suggested that the date may closer to nized just three time-successive species: Homo transvaa-
8 and 13 Ma, respectively, but the most recent analysis lensis, Homo erectus, and Homo sapiens. Having
of a larger data set (Prado-Martinez et al., 2013) support accepted that throughout hominin evolution “the known
estimate of an 5 Ma, but recalibration of the molecular diversity of fossil man can be interpreted as being the
clock to take account of the Oligocene catarrhine Rukwa- result of geographic variation within a single species of
pithecus fleaglei, which has been argued to be a basal Homo” (ibid, p. 116) (i.e., there has been no speciation
hominoid (Stevens et al., 2013), could further affect within hominin evolution involving reproductive isola-
these estimates. tion), Mayr then speculated about the reasons for “this
All you need are ancestors puzzling trait of the hominid stock to stop speciating”
(ibid, p. 116). He concluded that the reason for this is
Extant species are on the surface of the TOL. All of “man’s great ecological diversity” (ibid, p. 116), so that if
the species that have ever lived in the past, most of “man occupies successfully all the (ecological) niches
which we only know from morphological evidence, lie that are open for a Homo-like creature, it is obvious that
within it. So strictly speaking the only extinct species he cannot speciate.” Mayr noted that this conforms to
that have to be within the TOL are the ones situated on Gause’s rule (ibid, p. 116), but the principle of competi-
the branches that lead to the species on its surface. tive exclusion developed by Gause (1934) was based on
Thus, because the hominin twig has only one living spe- observations and experiments with organisms (e.g., pro-
cies, modern humans, the minimum requirement is that tozoa and yeast) whose ecology, unlike that of mammals,

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 39
can be replicated and manipulated in a laboratory c.2.5 Ma (Asfaw et al., 1999; Guy et al., 2008), and
setting. c.2.0–1.8 Ma (Leakey et al., 2012; Spoor et al., 2015).
Wolpoff (1971) also referred to Gause’s rule, but he did These interpretations have been challenged by research-
so in a different sense than Mayr (1950). Wolpoff (1968) ers who suggest that those who support hominin taxic
had suggested that human culture is such a specialized diversity in those time periods did not successfully dem-
ecological niche that “no more than one culture-bearing onstrate that the hypodigms of the proposed “new” taxa
hominid could have arisen and been maintained” (ibid, lie outside the range of variation of the existing taxa
p. 477), and that once culture had been acquired there (White, 2003; Smith, 2005; Ackermann and Smith, 2007;
could only ever have been one synchronic hominin spe- White et al., 2009; Lordkipanidze et al., 2013).
cies. Wolpoff (1971) argued that tool use could only have
occurred if a hominin was bipedal and both Washburn EVIDENCE OF HOMININ TAXIC DIVERSITY
(1950) and Mayr (1950) assumed that all hominins, and ACROSS SPACE AND TIME
only hominins, were bipedal, with the latter author sug-
gesting that “when the Homo-line acquired upright pos- We assign the species, site collections, and individual
ture it entered a completely different adaptive zone” specimens listed in Table I to one or more of the time
(ibid, p. 111). intervals we describe below. We exclude hypotheses
As more hominin fossils were discovered during the about the presence of species that we and others con-
late 1960s and early 1970s, especially in East Africa, sider to be idiosyncratic. Researchers use the term first
most researchers interpreted the new evidence as appearance datum, or FAD, to refer to the date of a tax-
strengthening the case for the presence of two forms of on’s first appearance in the fossil record, and last
early hominin, “gracile” and “robust.” Gracile hominin appearance datum, or LAD, to refer to the date of that
species were allocated to Homo, Telanthropus, or Aus- taxon’s last occurrence in the fossil record. The alloca-
tralopithecus, and the robust species were allocated to tions to time intervals are based on conservative ver-
Paranthropus or Zinjanthropus. Wolpoff argued that the sions of the FADs and LADs of each species (Table I)
differences between gracile and robust hominins repre- (Appendix A ) based on the published ages of the fossils.
sented sexual dimorphism in a single species. He ini- The time intervals have to be long enough to capture
tially made these arguments on the basis of evidence several taxa, but not so long that they are uninformative
from southern African sites (Wolpoff, 1968), but later he about diachronic changes in taxic diversity. We are
expanded them to include the early East African discov- acutely aware that our decisions about the length and
eries (Wolpoff, 1971). The discovery and recognition of the registration of the time intervals we use to investi-
Telanthropus provided at a single site, Swartkrans, evi- gate the evidence for hominin diversity, that is evidence
dence of a Homo erectus-like hominin alongside Para- of distinct synchronic hominin taxa, are subjective. We
nthropus robustus (Broom and Robinson, 1950), so it had are also aware that these decisions about their length
been put forward as a refutation of the single species and registration can potentially bias the outcome, but
hypothesis. Wolpoff rejected the refutation on two when we repeated the exercise with the longer intervals
grounds. First, he did not recognize Telanthropus as a (see below) registered 0.5 Ma later, and with the shorter
distinct taxon. Second, he was not convinced the two intervals registered at 0.25 Ma and 0.75 Ma, the out-
taxa were synchronic, to the extent that strict synchrony come was not materially different. So, given this, and
can ever be determined from the paleontological record the various reasons why the observed FADs and LADs
(but see Schroer and Wood, 2015). are likely to underestimate the time span of the species
By dint of persistent advocacy the single species we consider (see Discussion), we suggest that our results
hypothesis as applied to hominin evolution was kept are as robust as the available data allow.
alive for nearly another decade. For many (but see The first time interval includes the fossil evidence
Hunt, 2003) its demise came in 1976 with the demon- between c.7 and 5 Ma. Although one of us is not con-
stration that an early African Homo erectus cranium vinced that these taxa belong in the hominin clade
(KNM-ER 3733) and a Paranthropus boisei cranium (Wood and Harrison, 2011), in this review we consider
(KNM-ER 406) had both been found effectively in situ in all of the species that have been claimed to be early
what was then referred to as the Upper Member of the hominins. The next two time intervals each span one
Koobi Fora Formation dated to c.1.7 Ma (LeaKey and million years (i.e., 5.0–4.0 Ma and 4.0–3.0 Ma), and the
Walkes, 1976). These two specimens are so different in next four consider the fossil evidence in half million-year
morphology that there has been little or no dissent from time intervals (i.e., 3.0–2.5 Ma, 2.5–2.0 Ma, 2.0–1.5 Ma,
the hypothesis that they should be included in separate and 1.5–1.0 Ma). These smaller time intervals reflect the
genera. larger numbers of hominin species post 3 Ma. We then
The falsification of the single species hypothesis as consider evidence for diversity within the hominin clade
applied to Plio-Pleistocene hominins, along with the between 1.0 Ma and 0.25 Ma, and finally in the interval
expansion of the hominin fossil record, the introduction between 0.25 Ma and the present. If a taxon spans more
of new dating methods, and the widespread adoption of than one time interval, we review the evidence for that
phylogenetic methods, has led to the general acceptance taxon in the earliest time interval.
that at several times since the origin of Homo the homi- Decisions about whether taxonomic diversity exists
nin clade has consisted of more than one lineage (e.g., depend on what alpha taxonomic hypotheses are used.
Wood and Baker, 2011). However, debates about hominin For each time interval we consider in order of their pub-
diversity around the time of early Homo, and prior to lication the hominin species, or groups of fossils, or even
the origin of Homo, continue to be polarized. Those on individual fossils, that have been recognized as discrete
the diversity side of the scales cite hominin fossil evi- evolutionary units. We deliberately begin with a speciose
dence consistent with multiple hominin species in three interpretation of the hominin fossil record, but where
time periods, between c.3.5 and 3.3 Ma (Leakey et al., relevant we also explore what impact more inclusive
2001; Spoor et al., 2010; Haile-Selassie et al., 2015), interpretations of alpha taxonomy would have on

Yearbook of Physical Anthropology

40 B. WOOD AND E. BOYLE
TABLE I. For each hominin species, site collection or individual fossil referred to in this review we provide the ages that correspond
to the conservative (column 1) first appearance date (FAD) and FAD that incorporates dating error (column 2), and to the conserva-
tive (column 3) last appearance date (LAD) and LAD that incorporates dating error (column 4)
With dating error With dating error
Conservative Conservative
Taxon FAD FAD LAD LAD
Ardipithecus kadabba 6.3 Ma 6.7 Ma 5.2 Ma 5.11 Ma
Ardipithecus ramidus 4.51 Ma 4.6 Ma 4.3 Ma 4.262 Ma
Australopithecus afarensis 3.7 Ma 3.89 Ma 3.0 Ma 2.9 Ma
Australopithecus africanus 3.0 Ma 4.02 Ma 2.4 Ma 1.9 Ma
Australopithecus anamensis 4.2 Ma 4.37 Ma 3.9 Ma 3.82 Ma
Australopithecus bahrelghazali 3.58 Ma 3.85 Ma 3.58 Ma 3.31 Ma
Australopithecus deyiremeda 3.5 Ma 3.596 Ma 3.3 Ma 3.33 Ma
Australopithecus garhi 2.5 Ma – 2.45 Ma 2.488 Ma
Australopithecus sediba 1.98 Ma 2.05 Ma 1.98 Ma 1.91 Ma
Burtele Foot 3.4 Ma 3.47 Ma 3.4 Ma 3.2 Ma
Denisovans 48.65 ka 50.63 ka 29.2 ka 28.84 ka
Homo antecessor 1.0 Ma 1.2 Ma 0.936 Ma –
Homo erectus 1.81 Ma 1.85 Ma 27 ka –
Homo ergaster 1.7 Ma 2.27 Ma 1.4 Ma 0.87 Ma
Homo floresiensis 74 ka 108 ka 17 ka 16 ka
Homo georgicus 1.85 Ma – 1.77 Ma –
Homo habilis sensu lato 2.35 Ma 2.6 Ma 1.65 Ma –
Homo heidelbergensis 700 ka – 100 ka –
Homo helmei 260 ka – 80 ka –
Homo neanderthalensis 130 ka 197 ka 40 ka 39.22 ka
Homo rhodesiensis 600 ka – 300 ka –
Homo rudolfensis 2.0 Ma 2.09 Ma 1.95 Ma 1.78 Ma
Homo sapiens 195 ka 200 ka Present –
Kenyanthropus platyops 3.54 Ma 3.65 Ma 3.35 Ma –
Ledi-Geraru 2.80 Ma 2.85 Ma 2.75 Ma 2.65 Ma
Orrorin tugenensis 6.0 Ma 6.14 Ma 5.7 Ma 5.52 Ma
Paranthropus aethiopicus 2.66 Ma 2.73 Ma 2.3 Ma 2.23 Ma
Paranthropus boisei 2.3 Ma 2.5 Ma 1.3 Ma 1.15 Ma
Paranthropus robustus 2.0 Ma 2.27 Ma 1.0 Ma 0.87 Ma
Sahelanthropus tchadensis 7.2 Ma 7.43 Ma 6.8 Ma 6.38 Ma
Sima de los Huesos (SH) 780 ka – 427 ka 415 ka

assessments of hominin diversity. Howell (1999) recom- fossil evidence in the two hypodigms samples the same
mended that it might be less controversial to use p- anatomical regions, and the taxonomic valency of any
demes (an informal category used for a geographically shared anatomical regions. Other factors that affect our
localized population of closely related individuals) rather ability to make these judgments include how well the
than species for discussions about the taxonomy of the species were initially described and diagnosed, and
hominin fossil record. Presumably one or more p-demes whether any published species comparisons were quanti-
may correspond to regional variants within a single spe- tative or just qualitative. We readily concede that others
cies (i.e., equivalent to a subspecies), or they could corre- may well reach different conclusions, but we hope our
spond to the hypodigm of a species. But to our mind attempt to rank each species difference will stimulate
although the use of p-demes may be a useful way to dis- and encourage others to explore alternative methods
cuss the different parts of the fossil record prior to mak- and approaches.
ing decisions about its taxonomy, their use does not
solve the problem of how many species to recognize in c.7.0–5.0 Ma
the fossil record.
After we have considered the hominin species, groups Orrorin tugenensis. The first putative hominin spe-
of fossils, or even individual fossils, that have been rec- cies we consider in the c.7.0–5.0 Ma time interval is
ognized as discrete evolutionary units, we informally Orrorin tugenensis (Senut et al., 2001), the genus and
review the evidence for species diversity within each species established to accommodate cranial and postcra-
time interval as pairwise comparisons. Our attempt to nial remains recovered from c.6.0 Ma Lukeino Forma-
rank each comparison in terms of our degree of confi- tion sediments exposed at Aragai, Cheboit, Kapcheberek,
dence that each species pair really is different (see Table and Kapsomin in the Baringo District, Kenya. The femo-
2) is based on making judgments about how the morpho- ral morphology has been interpreted to mean that O.
logical differences between the hypodigms compare with tugenensis was at least a facultative biped (Pickford
the variation observed within and among closely-related et al., 2002; Richmond and Jungers, 2008; Almecija
extant taxa (e.g., Wood et al., 1991). But such judgments et al., 2013), but other researchers interpret the internal
are greatly influenced by factors over which we have no structure of the femoral neck as indicating a mix of
control. These factors include the multiple ways the bipedal and non-bipedal locomotion (Galik et al., 2004;
hypodigms we are comparing may differ. Such differen- Ohman et al., 2005). Almecija et al. (2010) suggested
ces include, but are not limited to, their state of preser- that the palmar surface of the pollical distal phalanx of
vation, their sample size, to what extent the preserved O. tugenensis (BAR 1901’01) shows evidence of a flexor

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 41
TABLE 2. For each of the nine time intervals relevant to the Low confidence
hominin fossil record, we assess the strength of the evidence for Homo erectus vs. Homo ergaster
taxic diversity among hominin species by allocating each pair- Homo erectus vs. Homo georgicus
wise comparison to one of three categories of confidence, high, 1.5–1.0 Ma
moderate or low High confidence
Homo erectus vs. Paranthropus robustus
7–5 Ma Homo erectus vs. Paranthropus boisei
High confidence Paranthropus robustus vs. Homo habilis sensu lato
N/A Paranthropus robustus vs. Homo antecessora
Moderate confidence Paranthropus boisei vs. Homo antecessora
Orrorin tugenensis vs. Sahelanthropus tchadensis Homo habilis sensu lato vs. Homo antecessora
Orrorin tugenensis vs. Ardipithecus kadabba Moderate confidence
Sahelanthropus tchadensis vs. Ardipithecus kadabba Homo erectus vs. Homo habilis sensu lato
Low confidence Paranthropus robustus vs. Paranthropus boisei
N/A Low confidence
5–4 Ma Homo erectus vs. Homo ergaster
High confidence Homo erectus vs. Homo antecessor
Ardipithecus ramidus vs. Australopithecus anamensis 1.0–0.25 Ma
Moderate confidence High confidence
N/A Homo erectus vs. Homo heidelbergensis
Low confidence Homo erectus vs. Homo rhodesiensis
N/A Homo erectus vs. Homo helmei
4–3 Ma Homo erectus vs. Sima de los Huesos
High confidence Homo heidelbergensis vs. Sima de los Huesos
Australopithecus afarensis vs. Burtele foot Homo rhodesiensis vs. Sima de los Huesos
Moderate confidence Homo helmei vs. Sima de los Huesos
Australopithecus afarensis vs. Kenyanthropus platyops Moderate confidence
Low confidence N/A
Australopithecus afarensis vs. Low confidence
Australopithecus bahrelghazali Homo heidelbergensis vs. Homo rhodesiensis
Australopithecus afarensis vs. Australopithecus deyiremeda Homo heidelbergensis vs. Homo helmei
Australopithecus bahrelghazali vs. 0.25 Ma to present
Australopithecus deyiremeda High confidence
Australopithecus bahrelghazali vs. Kenyanthropus platyops Homo sapiens vs. Homo neanderthalensis
Australopithecus bahrelghazali vs. Burtele foot Homo sapiens vs. Homo erectus
Kenyanthropus platyops vs. Australopithecus deyiremeda Homo sapiens vs. Homo heidelbergensis
Kenyanthropus platyops vs. Burtele foot Homo sapiens vs. Homo floresiensis
Burtele foot vs. Australopithecus deyiremeda Homo sapiens vs. Denisovans
3.0–2.5 Ma Homo erectus vs. Homo heidelbergensis
High confidence Homo erectus vs. Homo floresiensis
Australopithecus africanus vs. Paranthropus aethiopicus Homo neanderthalensis vs. Homo erectus
Australopithecus africanus vs. LD 350-1 Homo neanderthalensis vs. Homo floresiensis
Paranthropus aethiopicus vs. LD 350-1 Homo heidelbergensis vs. Homo floresiensis
Moderate confidence Moderate confidence
N/A Homo neanderthalensis vs. Denisovans
Low confidence Homo erectus vs. Denisovans
N/A Low confidence
2.5–2.0 Ma Homo neanderthalensis vs. Homo heidelbergensis
High confidence Homo heidelbergensis vs. Denisovans
Australopithecus africanus vs. Paranthropus boisei a
Paranthropus boisei vs. Homo habilis sensu lato If the partial mandible (ATE9-1) from the Sima del Elefante
Australopithecus garhi vs. Homo habilis sensu lato does not belong to H. antecessor then these “high confidence”
Moderate confidence pairwise comparisons would fall away.
Australopithecus africanus vs. Homo habilis sensu lato
Australopithecus africanus vs. Australopithecus garhi pollicis longus insertion that is modern human-like. Other-
Paranthropus boisei vs. Australopithecus garhi wise, the discoverers admit that much of the taxonomically
Homo habilis stricto stricto vs. Homo rudolfensis critical dental morphology is “ape-like” (Senut et al., 2001,
Low confidence
N/A p. 6). O. tugenensis may prove to be a hominin, but it is
2.0–1.5 Ma equally, and perhaps more, likely that it belongs to an
High confidence extinct clade closely related to panins and hominins.
Homo erectus vs. Paranthropus robustus
Homo erectus vs. Paranthropus boisei
Homo erectus vs. Australopithecus sediba Sahelanthropus tchadensis. The second putative
Paranthropus robustus vs. Homo habilis sensu lato
Paranthropus robustus vs. Australopithecus sediba
hominin species to be described in the c.7.0–5.0 Ma time
Paranthropus boisei vs. Homo habilis sensu lato interval is Sahelanthropus tchadensis (Brunet et al.,
Paranthropus boisei vs. Australopithecus sediba 2002). This species was established to accommodate a
Homo habilis sensu lato vs. Australopithecus sediba partial cranium, two partial mandibles, and some iso-
Moderate confidence lated teeth recovered in 2001 and 2002 from the c.7 Ma
Homo erectus vs. Homo habilis sensu lato Anthracotheriid unit at Toros-Menalla in the Chad
Paranthropus robustus vs. Paranthropus boisei Basin. Two more partial mandibles and an isolated P3,
Homo habilis sensu stricto vs. Homo rudolfensis
also recovered in 2001 and 2002, were later added to the
hypodigm (Brunet et al., 2005). Apparently, there is also

Yearbook of Physical Anthropology

42 B. WOOD AND E. BOYLE

Fig. 1. A conservative estimate of the temporal ranges of the hominin species, site collections, and individual fossils referred to
in this review. The bottoms and tops of the continuous columns represent, respectively, the published first and last appearances
dates. The sources of the data are set out in the Appendices A and B.

postcranial evidence (Beauvilain and Watte, 2009), but no between Ar. kadabba and Ar. ramidus (see below) are
details have been published. The only reference to differen- that the apical crests of the upper canine crown of the
ces between O. tugenensis and S. tchadensis in Brunet former taxon are longer and the P3 crown outline of Ar.
et al. (2002) is where the latter authors note that S. tcha- kadabba more asymmetrical than in Ar. ramidus. Haile-
densis is distinct from the former taxon because it has Selassie et al. (2004) suggest there is a morphocline in
upper I1s “with multiple tubercles on the lingual fossa” upper canine morphology, with Ar. kadabba exhibiting
and because the latter’s upper canines are “non chimp- the most ape-like morphology (see Fig. 1D in Haile-
like” and show “extensive apical wear” (ibid, p. 146). Selassie et al., 2004). Haile-Selassie (2001) also suggest
that there are differences in upper canine morphology
between Ar. kadabba and O. tugenensis, with the later
Ardipithecus kadabba. The third putative hominin having more “primitive” upper canine crowns that “lack
species in this time interval is Ardipithecus kadabba. the elevated crown shoulders” of “Ardipithecus and all
Initially, fossils recovered from four localities (Saitune other hominids” (ibid, p. 180). The formal diagnosis of Ar.
Dora, Alayla, Asa Koma, and Digiba Dora) in the West- kadabba (Haile-Selassie et al., 2004) refers to differences
ern Margin region of the Middle Awash study area were between the upper canines of Ar. kadabba and O. tuge-
assigned to Ar. ramidus as a separate subspecies, Ar. nensis, with the former having “a more circular upper
ramidus kadabba (Haile-Selassie, 2001), but subse- canine outline” (ibid, p. 1504), whereas the latter has an
quently that initial hypodigm, plus some additional upper canine crown that is “relatively more elongate
specimens, were elevated to species rank (Haile-Selassie mesiodistally” (ibid, p. 1505). It is also claimed that there
et al., 2004, 2009). The hypodigm of the new species con- are differences with respect to the morphology of the lin-
sists of the original cranial and postcranial remains gual face of the crown of the upper canine with that of Ar.
(partial mandible, four isolated teeth, fragments of a left kadabba being “relatively flat,” whereas there is
upper limb, and a proximal foot phalanx) announced in “stronger hollowing” on the only upper canine of O. tuge-
2001, plus six more dental specimens that were recov- nensis (ibid, p. 1504). The postcranial evidence of Ar.
ered from five c.5.8–5.2 Ma localities, in the Middle kadabba is generally ape-like, with the proximal foot pha-
Awash study area, Ethiopia. The main differences lanx (AME-VP-1/71) claimed as evidence of bipedalism

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 HOMININ TAXIC DIVERSITY 43
based on the proximal joint surface (Haile-Selassie, challenged, given the small size of the hypodigms of the
2001). However, both the orientation and the degree of three species, the lack of overlap in the parts of the skel-
curvature of this joint are like those seen in Australopi- eton represented in those hypodigms, and the lack of
thecus afarensis (i.e., African ape-like). any current evidence of temporal overlap, perhaps the
hypothesis that the fossils in this time interval sample
Evidence of diversity three separate species should also be challenged. In our
White et al. (2009) suggest that the morphological dif- opinion species differences in this time interval have at
ferences between Ar. ramidus and both O. tugenensis best only moderate support.
and S. tchadensis do not justify either of the latter spe- 5.0–4.0 Ma
cies being assigned to their own genus, so they proposed
they should be transferred to the genus with priority Ardipithecus ramidus. The first putative hominin
(i.e., Ardipithecus) as Ardipithecus tugenensis (Senut species in the 5.0–4.0 Ma time interval is Ar. ramidus
et al., 2001; White et al., 2009) and Ardipithecus tcha- (White et al., 1994, 1995), a species established to
densis (Brunet et al., 2002; White et al., 2009), respec- accommodate cranial and postcranial fossils recovered
tively. However, because White et al. (2009) did not from c.4.5–4.4 Ma localities at Aramis on the northeast-
question the decision to recognize O. tugenensis and S. ern flank of the Central Awash Complex in the Middle
tchadensis as separate species, their proposal does not Awash study area of Ethiopia. The authors claimed Ar.
affect taxic diversity. ramidus shares some features with living species of Pan,
Since its initial description, publications about S. tcha- others with the African apes in general and, crucially,
densis (Guy et al., 2005; Zollikofer et al., 2005) have several dental and cranial features that are shared only
mainly focused on defending its status as a hominin with later hominins such as Au. afarensis. The taxon
rather than on defending the decision to recognize it as was initially included within the genus Australopithecus,
a new species distinct from O. tugenensis. Zollikofer but it was subsequently transferred to a new genus,
et al. (2005) and Guy et al. (2005) each emphasize the Ardipithecus (White et al., 1995). Fossils recovered from
substantial (2,500 km) distance separating the localities four localities (Saitune Dora, Alayla, Asa Koma, and Dig-
where the fossil evidence of the two taxa has been iden- iba Dora) in the Western Margin region of the Middle
tified, but there is no a priori reason why the same Awash study area were initially assigned to Ar. ramidus
taxon could not have existed in the two regions (e.g., as a separate subspecies, Ardipithecus ramidus kadabba
later discussion about taxonomic diversity between Au. (Haile-Selassie, 2001), but those specimens, plus additional
afarensis and Australopithecus bahrelghazali). When specimens found since the initial publication, were later
Brunet et al. (2005) described two new mandibles transferred to a separate species, Ar. kadabba (Haile-Selas-
belonging to S. tchadensis they made no reference to sie et al., 2004) (see above). The first reported additions to
how they compare with what is preserved of the mandib- the Ar. ramidus hypodigm came from the Gona study area
ular morphology of O. tugenensis. However, apart from (Semaw et al., 2005), but subsequently more fossils, includ-
the mandible, and some positions along the maxillary ing the ARA-VP-6/500 associated skeleton recovered from
tooth row, there is little overlap in the parts of the skele- the Aramis locality (White et al., 2009), as well as fossils
ton preserved in the two hypodigms. from two other localities, Kuseralee Dora and Sagantole,
In the article establishing Ar. kadabba as a separate in the Central Awash Complex, have been added to the
species, Haile-Selassie et al. (2004) drew attention to dif- hypodigm (White et al., 2009).
ferences between the crown morphology of the upper The chewing teeth of Ar. ramidus are relatively small
canines of O. tugenensis and Ar. kadabba, but in the and the form of the reconstructed pelvis and the mor-
summary they suggest that with respect to the dentition phology of the lateral side of the foot have been cited as
“Sahelanthropus and Orrorin”. . .“are very similar to A. evidence that the posture and gait of Ar. ramidus were,
kadabba” (ibid, p. 1503). Also, when Haile-Selassie et al. respectively, more upright and bipedal than is the case
(2009) reviewed the fossil evidence for Ar. kadabba they in the living apes. The enamel covering on the teeth is
concluded that the mandibular morphology of Sahelan- not as thin as that of chimpanzees/bonobos, but it is not
thropus is “broadly compatible with that exhibited by as thick as that seen in archaic (sensu Wood, 2010)
the ALA-VP-2/10 mandible (of Ar. kadabba) (ibid, p. hominins such as Au. afarensis (Suwa et al., 2009), and
208), but no evaluation of any similarities or differences according to Rak et al. (2007) the morphology of the
between the mandibular morphology of O. tugenensis ramus of the mandible is similar to that of Pan. Initial
and Ar. kadabba was offered “because of the lack of estimates based on the size of the shoulder joint sug-
detailed information” about O. tugenensis (ibid, p. 208). gested that Ar. ramidus weighed about 40 kg, but
With respect to the postcanine dentition, Haile-Selassie researchers claim the enlarged hypodigm indicates an
et al. (2009) concluded that “the mandibular postcanine estimated mean body mass of c.50 kg (Lovejoy et al.,
dentition of Sahelanthropus (TM266-02-154-2) closely 2009), although an alternative (lower) estimate of
matches the ALA-VP-2/10 equivalent in the available P4 c.36 kg is obtained based on hip joint size (Almecija
to M3 metrics” (ibid, p. 218), and once again they offer et al., 2015) using a modern human baseline. Grabow-
no comparison with O. tugenensis because the “teeth are ski et al. (2015) suggest an even smaller c.32 Kg,
not described or illustrated in sufficient detail” (ibid, p. estimate.
218). Yet, despite these acknowledged similarities, Haile-
Selassie et al. (2009) do not advocate any change to the
conventional taxonomy that recognizes species- and Australopithecus anamensis. The second putative
genus-level differences between O. tugenensis, S. tcha- hominin species to be announced from the 5.0 to 4.0 Ma
densis, and Ar. kadabba. time interval, Australopithecus anamensis (Leakey
Thus, although the hypothesis that the fossils in this et al., 1995), was established to accommodate a left dis-
time interval sample three separate genera has been tal humeral fragment (KNM-KP 271) recovered by

Yearbook of Physical Anthropology

44 B. WOOD AND E. BOYLE

Bryan Patterson at Kanapoi, Kenya, in 1965 (Patterson time interval, the substantial overlap in the parts of the
and Howells, 1967), plus cranial remains recovered from skeleton represented by the hypodigms of Ar. ramidus
c.3.9–4.2 Ma localities at Allia Bay and Kanapoi. Addi- and Au. anamensis strengthens the evidence for taxic
tional fossils from Allia Bay and Kanapoi were described diversity in this time interval.
three years later (Leakey et al., 1998). Leakey et al.
(1995) claimed that the dental morphology (e.g., mandib- 4.0–3.0 Ma
ular canine morphology, the asymmetry of the premolar Australopithecus afarensis. The hominin species
crowns, and the relatively simple crowns of the decidu- with historical priority in the 4.0–3.0 Ma time interval,
ous first mandibular molars) of the fossils collected at Australopithecus afarensis (Johanson et al., 1978, but
Allia Bay and Kanapoi are more primitive than those of see ICZN, 1999), was established to accommodate the
Au. afarensis, but in other respects (e.g., the low cross- c.3.7–3.0 Ma cranial and postcranial remains recovered
sectional profiles and bulging sides of the molar crowns) from Laetoli, Tanzania, and Hadar, Ethiopia. White
the postcanine teeth of Au. anamensis show similarities et al. (1981) compared the hypodigms of Au. afarensis
to Paranthropus (see below). Upper limb remains were and Australopithecus africanus and made a compelling
also recovered and were described as being australopith- case for recognizing Au. afarensis as a distinct species
like (Leakey et al., 1998), and although a tibia (KNM- with generally more primitive craniodental anatomy
KP 29283) with features associated with obligate biped- than Au. africanus. Kimbel et al. (1984, 2004) and Kim-
alism was attributed to Au. anamensis it was not bel and Delezene (2009) provide further information
included in the list of paratypes. White et al. (2006) about the morphological differences between Au. afaren-
attributed 31 fossils from the Middle Awash study area sis and Au. africanus.
in Ethiopia to Au. anamensis. One, a maxilla, was found To judge from the evidence of the associated skeleton
at Aramis, with the remainder coming from three local- A.L. 288-1, the hind limbs of Au. afarensis are substan-
ities at Asa Issie, a collecting area 10 km/6 miles west of tially shorter than those of a modern human of similar
Aramis. Biostratigraphic dating suggests an age of stature, and the appearance of the pelvis and the rela-
c.4.2–4.1 Ma for both sets of Middle Awash fossils. Cra- tively short lower limb suggests that while Au. afarensis
niodentally (e.g., anterior tooth and postcanine relative was capable of bipedal walking it was not adapted for
tooth size, crown morphology, enamel thickness, etc.), long-range bipedalism (e.g., Jungers, 1982). The upper
the new material from the Middle Awash is consistent limb, especially the hand and the shoulder girdle,
with an attribution to Au. anamensis. Upper limb retains morphology that some workers suggest reflects a
remains have been recovered and were described as significant element of arboreal locomotion (e.g., Stern
being australopith-like (Leakey et al., 1998), but a tibia and Susman, 1983), yet its upper limb morphology, and
(KNM-KP 29283) with features associated with obligate especially its hand, suggests that Au. afarensis was
bipedalism that was attributed to Au. anamensis was capable of the type of modern human-like manipulation
not included in the list of paratypes. A specimen pre-
(Alba et al., 2003, Drapeau, 2012; Almecija and Alba,
serving the proximal three-quarters of a right femur
2014) which is not surprising given that human hand
shaft, ASI-VP-5/154, looks like a slightly more primitive
proportions are largely primitive (Almecija et al., 2015).
version of the Au. afarensis femoral morphology.
The upper limb, especially the hand and the shoulder
Evidence of diversity girdle, retains morphology that some workers suggest
reflects a significant element of arboreal locomotion, yet
When they prepared their diagnosis of Ar. ramidus it Drapeau (2012) suggests that the Au. afarensis upper
was natural that White et al. (1994) focused on the differ- limb morphology was capable of the type of manipula-
ences between Ar. ramidus and Au. afarensis, but since tion needed for the manufacture of crude artefacts.
1994 Au. anamensis has superseded Au. afarensis as the Recent body mass estimates for Au. afarensis range
oldest known East African Australopithecus species. Thus, from c.30 to c.65 kg (Grabowski et al., 2015), known
evidence of taxic diversity within the 5.0–4.0 Ma time endocranial volumes range between 385 and 550 cm3,
interval is dependent on being able to demonstrate signifi- and estimates of the standing height of adult individuals
cant differences between Ar. ramidus and Au. anamensis, range between 1.0 and 1.5 m. Reno et al. (2003) sug-
yet most comparisons of Ardipithecus and Australopithecus gested that skeletal size sexual dimorphism in Au. afar-
by White et al. (2009) and Suwa et al. (2009) were made ensis is at a relatively low level, but most researchers
between Ar. ramidus and Au. afarensis. (e.g., Gordon et al., 2008) have concluded that this taxon
In their diagnosis of Au. anamensis, Leakey et al. shows a substantial level of sexual dimorphism.
(1995) suggest that the new taxon “can be distinguished
from Ardipithecus” because it has “absolutely and rela-
tively thicker tooth enamel; upper canine buccal enamel Australopithecus bahrelghazali. The second homi-
thickened apically; molars more buccolingually nin species to be announced from the 4.0–3.0 Ma time
expanded; first and second lower molars not markedly interval, Australopithecus bahrelghazali (Brunet et al.,
different in size; tympanic tube extends only to the 1996), was established to accommodate a tooth-bearing
medial edge of the postglenoid process rather than to midline mandible fragment plus an upper premolar
the lateral edge or beyond it; lateral trochlear ridge of tooth, both recovered from c.3.5 to 3.0 Ma sediments in
humerus weak” (ibid, p. 565). These distinguishing fea- the Bahr el Ghazal region, Koro Toro, Chad. The man-
tures also occur in the list of inferred shared-derived dibular fragment had previously been assigned to Aus-
characters that White et al. (2009) claim distinguish Ar. tralopithecus aff. Australopithecus afarensis (Brunet
ramidus from Au. anamensis (ibid, Table I, pp. 82–83). et al., 1995), but Brunet et al. (1996) assigned it to a
In that list 7 out of 11 cranial and mandibular charac- new species because they claimed it had thicker enamel
ters, and 18 out of 34 dental characters, differ between than Ar. ramidus, a more vertically oriented and more
the two taxa. Thus, unlike the situation in the previous gracile symphysis than Au. anamensis, more complex

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 45
premolar roots than Au. afarensis, and larger incisors which, while obviously distinguishing it from the pedal
and canines and more complex premolar roots than Au. remains of Au. afarensis (Haile-Selassie et al., 2010a,b;
africanus. Ward et al., 2011; Haile-Selassie et al., 2015), also aligns
it with the type of foot seen in Ar. ramidus (Lovejoy
et al., 2009). Haile-Selassie et al. (2015) admit that they
Kenyanthropus platyops. The third hominin species cannot reject the null hypothesis “that BRT-VP-2/73
in the 4.0–3.0 Ma time interval, Kenyanthropus platyops belongs to Au. deyiremeda” (ibid, Supporting Informa-
(Leakey et al., 2001), was established to accommodate tion Note 1), but the authors demur from assigning the
cranial remains recovered from the c.3.5 Ma Kataboi foot to the new taxon because none of the fossil evidence
Member at Lomekwi, West Turkana, Kenya. The initial for the latter “is clearly associated with BRT-VP-2/73”
report lists the holotype cranium and the paratype max- (ibid, p. 483).
illa, plus 34 other craniodental specimens including
three mandible fragments, a maxilla fragment, and iso- Evidence of diversity
lated teeth. Some of these may also belong to the hypo-
digm, but the researchers reserved their judgment about One of the signal achievements in paleoanthropology
the taxonomy of most of these remains (Leakey et al., in the past several decades has been the assembly of the
2001) several of which had only recently been referred substantial collection of fossils (now >400 specimens)
to Au. afarensis (Brown et al., 2001). The main reasons that forms the hypodigm of Au. afarensis (Kimbel and
Leakey et al. (2001) did not assign the Lomekwi mate- Delezene, 2009; Haile-Selassie et al., 2010a,b; Ward
rial to Au. afarensis are its reduced subnasal progna- et al., 2012). Although there have been suggestions that
thism, anteriorly-situated zygomatic root, the flatter and the Au. afarensis hypodigm samples more than one
more vertically orientated malar region, the generally taxon (Johanson and Taieb, 1976; Olson, 1981,
relatively small but thick-enameled molars, and the 1985a,1985b; White et al., 1981; Senut, 1983; Tardieu,
unusually small M1 when compared to the size of the P4 1983), none of them have received consistent support
and M3. Some of the morphology of the new genus is from the researchers most familiar with the accumu-
Paranthropus-like, yet K. platyops lacks the postcanine lated fossil evidence. Thus, any proposal for a new homi-
megadontia that characterizes Paranthropus. The nin species in the 4.0–3.0 Ma time interval must
authors note the face of the new material resembles that demonstrate that the morphology of the new fossil evi-
of Homo rudolfensis (see below), but they point out that dence lies outside the envelope of morphological varia-
the postcanine tooth row of the latter is substantially tion expected within the species sampled by the existing
longer than that of KNM-WT 40000. fossil evidence for Au. afarensis.
When this test is applied to Au. bahrelghazali, while
Guy et al. (2008) claim that the symphyseal outline of
Australopithecus deyiremeda. The most recent addi- Au. bahrelghazali distinguishes it from Au. afarensis,
tion to the hominin species recognized in the 4.0–3.0 Ma others, including the authors, are more inclined to inter-
time interval is Australopithecus deyiremeda (Haile- pret the differences between the modest-sized hypodigm
Selassie et al., 2015). It was established to accommodate of Au. bahrelghazali and Au. afarensis as evidence of
several c.3.4 Ma cranial fossils including a left maxilla, geographical variation within the latter taxon.
part of the right maxilla from what might be the same The proposal to establish a new species and genus, K.
individual, and two mandibles, all recovered from the platyops, for fossil hominins discovered at West Turkana in
Burtele (BRT) and Waytaleyta (WYT) collection areas in 1998 and 1999 was based on the hypothesis that two speci-
the Woranso-Mille study area in Ethiopia. The authors mens, the type, KNM-WT 40000, a c.3.5 Ma cranium, and
claim that the fossils assigned to Au. deyiremeda have the paratype, KNM-WT 38350, a c.3.3 Ma partial maxilla
thicker enamel, more complex P4 roots and a more (Leakey et al., 2001), passed the test set out above. The
robust mandibular corpus than Ar. ramidus, and when case for rejecting Au. afarensis as the appropriate taxon for
compared to Au. anamensis they have a mandibular this material is complicated because while much of the face
symphysis that is more vertical, and a mandibular cor- and vault of the KNM-WT 40000 cranium are preserved,
pus that is more robust. Haile-Selassie et al. (2015) the cranium is plastically deformed and permeated by
claim that the mandibular corpus of Au. deyiremeda matrix-filled cracks. White (2003) took the view that these
lacks the lateral hollowing seen in Au. afarensis, the factors are responsible for the unusual facial morphology of
roots of the mandibular ramus and the zygomatic pro- KNM-WT 40000 and he interprets it as a taphonomically
cess of the maxilla are more anteriorly-located, and the altered Au. afarensis cranium. Spoor et al. (2010)
M1 crown is smaller than is the case in Au. afarensis. responded by making what, to us, is a convincing case that
They also suggest that the mandibular corpus of Au. the matrix-filled cracking and deformation are not respon-
deyiremeda is more robust, and the root of the mandibu- sible for the differences between the face of KNM-WT
lar ramus more anteriorly-located, than is the case in 40000 and Au. afarensis. In any event, the taphonomic
most of the mandibles attributed to early Homo. Den- alteration of the bone cannot explain the dental differences.
tally, Haile-Selassie et al. (2015) claim that Au. deyire- The case made by Haile-Selassie et al. (2015) that Au.
meda differs from K. platyops in having a larger canine, deyiremeda lies outside of the envelope of Au. afarensis
and from Australopithecus garhi because it has smaller has elements in common with the argument made by
canines and smaller postcanine tooth crowns. Leakey et al. (2001) for K. platyops. The authors claim
that the mandibular corpus of Au. deyiremeda is more
robust and lacks the lateral hollowing seen in Au. afar-
Burtele foot. The final piece of evidence from this ensis. Haile-Selassie et al. (2015) also suggest that the
time interval is a partial foot, BRT-VP-2/73, recovered in roots of the mandibular ramus and the zygomatic pro-
2009, also from Burtele (Haile-Selassie et al., 2012). cess of the maxilla are more anteriorly located than in
What is notable about the foot is its opposable hallux, Au. afarensis. With respect to the dentition, they suggest

Yearbook of Physical Anthropology

46 B. WOOD AND E. BOYLE

that the postcanine tooth crowns are smaller, and as 2003) belong to the Au. africanus hypodigm, or to a dif-
was the case for K. platyops, it is the M1 crown that is ferent species (Clarke, 2008).
particularly small. However, for many of these differen- The hypodigm of Au. africanus is numerically one of
ces the condition in Au. deyiremeda is close to, or at the the better fossil records of an early hominin species. The
edge of the range of, the hypodigm of Au. afarensis. To cranium, mandible, and dentition are well sampled, and
us, it seems that the claim of distinctiveness rests very although the postcranial and the axial skeleton are less
largely on what Haile-Selassie et al. (2015) declined to well represented, there is at least one specimen of each
do, which is to link Au. deyiremeda with the foot from long bone. However, many of the fossils have been
Burtele (see below). crushed and deformed before they were fully hardened
Haile-Selassie et al. (2012) note that the short, oppos- by fossilization. The chewing teeth of Au. africanus are
able, hallux, and overall morphology of the partial foot relatively large and apart from the reduced canines the
from Burtele are good evidence that it “does not belong skull is relatively ape-like. Its mean endocranial volume
to the contemporaneous species Au. afarensis.” They is c.460 cm3. Morphological and functional analyses sug-
suggest that BRT-VP-2/73 retains “a grasping capacity gests that although Au. africanus was capable of walk-
that would allow it to exploit arboreal settings” (ibid, p. ing bipedally, it was probably more arboreal than most
568), making it unlike the foot of Au. afarensis that has other archaic hominin taxa. The Sterkfontein evidence
been described as “functionally like that of modern suggests that males and females of Au. africanus dif-
humans” and the foot of a “committed terrestrial biped” fered substantially in body size, but probably not to the
(Ward et al., 2011, p. 750). Thus, although the Burtele degree they did in Au. afarensis.
foot has not been formally assigned to a taxon, we sug- Several researchers have commented on the unusual
gest that the profound structural, and inferred func- nature and degree of variation within Au. africanus
tional, differences between BRT-VP-2/73 and the foot of (Lockwood and Tobias, 2002), but there has been scant
Au. afarensis provide perhaps the strongest evidence for agreement on how the specimens that make up the
hominin taxic diversity in the 4.0–3.0 Ma time interval. hypodigm should be partitioned. Clarke (1988, 1999,
No meaningful comparison can be made between the 2008) has consistently argued that the Sterkfontein
Burtele foot and K. platyops, Au. bahrelghazali and Au. Member 4 and Makapansgat hypodigm of Au. africanus
deyiremeda because the hypodigms of the latter three samples a second, more Paranthropus-like, taxon. He
taxa include no pedal remains, plus we believe it is not included Sts 1 and 71, StW 183, 252, 384, 498 and 505,
presently possible to falsify the hypothesis that the two and MLD 2 in a second species he refers to as Australo-
latter taxa are junior synonyms of Au. afarensis, and pithecus prometheus (Granger et al., 2015), thus reviving
this assessment also weakens the likelihood that a com- the species name Dart (1948) used for the hominin fos-
parison between Au. bahrelghazali and Au. deyiremeda sils from Makapansgat. The differences between the sec-
is going to provide evidence of hominin taxic diversity. ond taxon and Au. africanus mainly relate to
Moreover, any comparison between Au. bahrelghazali craniofacial structure and dental size (Clarke, 2008).
and K. platyops suffers from the twin problems of small
sample sizes and little or no overlap in the regional mor-
phology represented in the two hypodigms. Paranthropus aethiopicus. The second species to be
Several researchers (e.g., Ward et al., 1999; Kimbel considered in the 3.0–2.5 Ma time interval is Paraustra-
et al., 2006) have drawn attention to dental, facial, and lopithecus aethiopicus (Arambourg and Coppens, 1968),
mandibular differences between the early component of now Paranthropus aethiopicus (Chamberlain and Wood,
the Au. afarensis hypodigm from Laetoli and the geologi- 1987). The latter is the species name used by research-
cally younger part of the hypodigm from Hadar, and to ers who do not recognize Paraustralopithecus as a sepa-
similarities between the Laetoli remains and those of Au. rate genus, but who do consider that >2.3 Ma hyper-
anamensis. It remains to be seen whether these similar- megadont hominins from the Omo-Turkana Basin belong
ities are sufficient evidence to sustain the hypothesis that to a species that is distinct from P. boisei. As presently
Au. anamensis evolved via anagenesis into Au. afarensis interpreted, the hypodigm of such a species would
(e.g., Haile-Selassie et al., 2010). If the taxa are related in include a well-preserved adult cranium from West Tur-
this way it would have no effect on any claims about kana (KNM-WT 17000) together with mandibles (e.g.,
hominin taxic diversity, but it would if new evidence sug- KNM-WT 16005) and isolated teeth from the Shungura
gests that their fossil records are synchronic. Formation. Some also assign the juvenile braincase, L.
338y-6, to this taxon. The only postcranial fossil consid-
3.0–2.5 Ma ered part of the hypodigm of P. aethiopicus is a proximal
tibia from Laetoli. The cranial evidence for P. aethiopi-
Australopithecus africanus. The first of the two spe- cus resembles P. boisei except that the face of the former
cies in the 3.0–2.5 Ma time interval, Au. africanus taxon is more prognathic, the cranial base is less flexed,
(Dart, 1925), was established to accommodate an imma- the incisors judging from their preserved alveoli are
ture skull recovered from the limeworks at Taungs (now inferred to be larger, and the postcanine teeth, especially
called Taung) in 1924. In addition to the type specimen the mandibular premolars, are less morphologically spe-
from Taung the Au. africanus hypodigm as presently cialized (Suwa, 1988).
interpreted includes fossils from Member 4 at Sterkfon-
tein, fossils from Members 3 and 4 at Makapansgat, and
fossils recovered from lime worker’s dumps and LD 350-1. The only other hominin known from the
extracted in situ from the breccia exposed at Gladysvale, 3.0–2.5 Ma time interval is represented by a single spec-
all of these sites are located in South Africa. It remains imen, LD 350-1, the left side of a 2.8–2.75 Ma adult
to be seen whether the associated skeleton StW 573 hominin mandible found in the Lee Adoyta region of the
from Sterkfontein Member 2, and 12 hominin fossils Ledi-Geraru research area in the Afar Regional State in
recovered from the Jakovec Cavern (Partridge et al., Ethiopia (Villmoare et al., 2015). LD 350-1 preserves the

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 HOMININ TAXIC DIVERSITY 47
crowns and roots of the canine through to the M3. Before size dimorphism, presumed to be sex-related. The fossil
concluding that it was most parsimonious to allocate the record of P. boisei extends across about one million years
mandible to Homo sp., the researchers compared it with of time during which there is little evidence of any sub-
Au. afarensis. Their judgment was that although it was stantial change in the size or shape of the cranium,
within the size range of Au. afarensis and shared with it mandible, and dentition (Wood and Constantino, 2007).
mandibular features such as a sloping symphysis and a
robust inferior transverse torus, it differed from Au.
afarensis by having a less robust corpus that lacked lat-
eral hollowing, plus the height of the corpus was consist- Homo habilis sensu lato. The third species in this
ent along the tooth row. Dentally, the P3 is more time interval, Homo habilis (Leakey et al., 1964), was
symmetrical, the M1/M2 hypoconid is reduced, the M1 established to accommodate non-megadont fossil homi-
has a C7, and the P3/molar wear pattern is reversed nins (OH 4, 6, 7, 8, 13, 14, and 16) recovered between
(Villmoare et al., 2015). Villmoare et al. (2015) suggest 1959 and 1963 from Beds I and II at Olduvai Gorge in
that the isolated left and right P3-M2 crowns of a single Tanzania. The authors claimed that its cranial and den-
individual (KNM-ER 5431) from the upper Tulu Bor tal morphology and endocranial volume, plus inferences
Member (i.e., between 3.0 and 2.7 Ma) at Koobi Fora, about its dexterity and locomotion, both distinguished
Kenya and non-robust teeth from Mbs B-C of the Shun- the new taxon from Au. africanus and justified its inclu-
gura Formation, may belong to the same taxon as LD sion in Homo. Subsequent discoveries at Olduvai (e.g.,
350-1. OH 24, 62, and 65) and from other sites (e.g., Koobi
Fora: KNM-ER 1470, 1802, 1805, 1813, 3735; Sterkfon-
tein: Stw 53; Swartkrans: SK 847; Hadar: A.L. 666-1)
Evidence for diversity. There is an excellent case for have also been assigned to, or affiliated with, H. habilis
distinguishing between P. aethiopicus and Au. africanus, sensu lato. The hypodigm set out above has a relatively
and the taxonomic distinctions between both of these wide range of cranial and dental morphology (e.g., endo-
taxa and the LD 350-1 mandible from Ledi-Geraru are cranial volumes range from just less than 500 cm3 to
also impressive. Villmoare et al. (2015) understandably c.800 cm3). All the crania in this group are wider across
focus on whether LD 350-1 differs from the existing the base of the cranium than across the vault. Facial
hypodigm of Au. afarensis, and although the mandible and mandibular morphology varies (e.g., KNM-ER 1470
does not currently overlap temporally with that hypo- has a flat and wide midface, whereas KNM-ER 1813 is
digm, for reasons we set out below, we believe it was a broadest across the upper face). Postcanine teeth also
sensible strategy. The case for distinguishing between differ in size and crown morphology, with some mandib-
LD 350-1 and Au. afarensis is weaker than the case for
ular premolars and molars being narrow buccolingually
distinguishing between the former and P. aethiopicus
(e.g., OH 7), whereas other mandibular premolar teeth
and Au. africanus, but all of these distinctions are more
within the hypodigm (e.g., KNM-ER 1802) have bucco-
impressive than the claims for taxic diversity within the
lingually broader crowns (P3s), larger talonids, and more
conventional hypodigm of Au. africanus.
complex root systems (P4s). Some researchers consider
2.5–2.0 Ma the cranial variation within the hypodigm set out above
to be excessive in scale and unlike the pattern of intra-
Australopithecus africanus. The first hominin spe- specific variation seen in the African ape clade, and they
cies with historical priority in this time interval, Au. suggest H. habilis sensu lato subsumes two taxa, Homo
africanus, has already been reviewed. habilis sensu stricto and Homo rudolfensis (e.g., Wood,
1991; Spoor et al., 2015).
Paranthropus boisei. The second species in the 2.5–
2.0 Ma time interval is Zinjanthropus boisei (Leakey,
1959), now Paranthropus boisei (Robinson, 1960). It has Paranthropus aethiopicus. The fourth species
a comprehensive craniodental fossil record that includes sampled in this time interval, P. aethiopicus, has already
an especially complete and well-preserved skull from been reviewed.
Konso in Ethiopia (Suwa et al., 1997), several well-
preserved crania, and many mandibles and isolated
teeth. P. boisei is the only hominin to combine a massive,
wide, flat, face, large-crowned premolars and molars, Australopithecus garhi. The fifth species to be con-
and small-crowned anterior teeth. The face of P. boisei is sidered in the 2.5–2.0 Ma time interval, Australopithe-
wider than that of P. robustus (see below), yet their cus garhi (Asfaw et al., 1999), was established to
endocranial brain volumes are similar. The mandible of accommodate a fragmented cranium recovered from the
P. boisei has a large and wide body or corpus and the c.2.5 Ma Hatayae Member of the Bouri Formation at the
tooth crowns apparently grow at a fast rate. Apart from Bouri, Gamedah, and Matabaietu collection areas, Mid-
a partial upper limb from Olduvai Gorge and two very dle Awash study area, Ethiopia. It combines a primitive
fragmentary possible P. boisei partial skeletons from cranium with large postcanine teeth, and especially
Koobi Fora, no other postcranial evidence can be attrib- large P3s. However, unlike other hyper-megadont species
uted to P. boisei with any certainty. Indeed, some of the such as P. aethiopicus and P. boisei, the incisors and can-
postcranial fossils from Bed I at Olduvai Gorge tradi- ines of Au. garhi are also large and the enamel appa-
tionally linked with Homo habilis (see below) may rently lacks the extreme thickness seen in those taxa. A
belong to P. boisei (Wood, 1974; Wood and Constantino, partial skeleton combining a long femur with a long
2007). There is evidence of both large- and small-bodied forearm was found nearby, but it is not associated with
individuals in the P. boisei hypodigm, and the range of the type cranium (Asfaw et al., 1999) and these fossils
the size difference suggests a substantial degree of body- have not been formally assigned to Au. garhi.

Yearbook of Physical Anthropology

48 B. WOOD AND E. BOYLE

Homo rudolfensis. If the fossils assigned to Homo between it and the cranial hypodigm of P. boisei suggest
habilis sensu lato (see above) sample not one, but two that the claim for a distinction between those taxa is a
species, and if that second species includes KNM-ER strong one, and the differences between Au. garhi and
1470, then the final species with historical priority in Au. africanus provide additional evidence for taxic diver-
this time interval is Pithecanthropus rudolfensis (Alex- sity in this time interval.
eev, 1986), later transferred to Homo (Groves, 1989) as
Homo rudolfensis. Leakey et al. (2012) described a face 2.0–1.5 Ma
(KNM-ER 62000) and two mandibles (KNM-ER 1482 Homo erectus. The first hominin species in the 2.0–
and 60000) that match KNM-ER 1470, and Spoor et al. 1.5 Ma time interval is Homo erectus (Dubois, 1893). In
(2015) make the case that the dental arcade of the his initial publication of the Trinil remains, Dubois
enlarged hypodigm of H. rudolfensis is distinctively dif- referred the skullcap to Anthropopithecus erectus, a
ferent (e.g., more divergent tooth rows, flatter anterior choice that reflected his initial conviction that he had
dental arch) from the more primitively-shaped dental discovered the remains of a fossil ape, but a year later
arcade of H. habilis sensu stricto. he convinced himself that he had discovered a human
Evidence of diversity ancestor and transferred the new species to a novel
genus, Pithecanthropus. The discovery of the Trinil calo-
From the time of the introduction of Homo habilis in tte was significant because of its small cranial capacity
1964, researchers have questioned the wisdom of estab- (c.940 cm3), low brain case, and sharply angulated occi-
lishing a new taxon to accommodate the non-Paranthro- pital region. Most of the discoveries made by Ralph von
pus fossils from Beds I and II of Olduvai Gorge. Initially, Koenigswald at Sangiran, also in Indonesia, were added
the main criticism was that within the linear, anagenetic to the hypodigm of P. erectus, and later fossils that had
model of evolution prevailing when the taxon was intro- been recovered from what was then called Choukoutien
duced there was insufficient “morphological space” (now called Zhoukoudian) and initially assigned to
between Au. africanus and Homo erectus (see below) for Sinanthropus pekinensis (Black, 1927), were compared
another hominin taxon. Critics claimed that the H. with the P. erectus hypodigm. The researchers responsi-
habilis hypodigm from Olduvai Gorge was an amalgam ble for analyzing the two collections suggested that the
of geologically older “advanced” Au. africanus fossils and Indonesian and Chinese hypodigms were “related to
geologically younger “primitive” H. erectus remains (Rob- each other . . . in the same way as two different races of
inson, 1965). This criticism was effectively countered by present mankind” (von Koenigswald and Weidenreich,
the demonstration that one of the most morphologically 1939, p. 928) and a year later Weidenreich proposed that
advanced H. habilis specimens (OH 24) was also geologi- the two hypodigms should be formally merged within a
cally among the oldest fossils in the hypodigm, coming single genus and species, as Homo erectus pekinensis
as it did from close to the base of Bed I. Researchers and Homo erectus javanensis, respectively (Weidenreich,
have also demonstrated that the distinguishing features 1940). Subsequently, the hypodigms of Meganthropus
of H. habilis are not simply a mix of the characteristics palaeojavanicus (Mayr, 1944, p. 14; Le Gros Clark, 1955,
of archaic hominins, such as Au. africanus and Au. afar- pp. 86–87), Atlanthropus (Le Gros Clark, 1964, p. 112),
ensis, and H. erectus, but they constitute a distinctive and Telanthropus (Robinson, 1961) were transferred to
morphological combination (Tobias, 1966; Wood, 1991). Homo erectus. Some researchers (e.g., Kaifu et al., 2008)
In our opinion, the hypothesis that Au. africanus is taxo- interpret H. erectus as a chronospecies that evolves
nomically distinct from H. habilis sensu lato enjoys at through time, whereas others (e.g., Widianto and Zei-
the least a moderate level of confidence. toun, 2003) argue there are potential species-level differ-
The effective synchronicity of P. boisei and H. habilis ences between the Sangiran/Trinil hypodigms and the
sensu lato (Leakey and Walker, 1976; Wood, 1991) is per- more recent evidence from Sambungmacan and
haps the strongest evidence of taxic diversity in the Ngandong.
hominin fossil record. Even researchers who fiercely
criticize, and even deride, the notion of hominin taxic
diversity (e.g., White, 2003) accept this as an example of Paranthropus robustus. The second species in this
the phenomenon. However, the other claims for hominin time interval, Paranthropus robustus (Broom, 1938),
taxic diversity in the 2.5–2.0 Ma time interval (e.g., H. was established to accommodate fossil hominins recov-
habilis sensu stricto vs. H. rudolfensis) are of a different ered in 1938 from what was then referred to as the
order in the sense that for various reasons, including “Phase II Breccia” (now called Member 3) at Kromdraai
sample size, the evidence is less compelling. That does B, in Gauteng Province, South Africa. Most of the cur-
not mean there is no case, just that it is modest in com- rent hypodigm comes from Swartkrans (Members 1, 2,
parison with the number and scale of the craniodental and 3), with other fossil evidence coming from Cooper’s,
differences between P. boisei and H. habilis sensu lato Drimolen, and Gondolin. Research at Drimolen was only
(see Wood, 1991 for details) that do not conform with the initiated in 1992, but since then more than 80 hominin
sort of intraspecific variation that one would predict specimens have been recovered and it promises to be a
based on a comparative analysis of the African ape rich source of evidence about P. robustus. Although the
clade. The case for distinguishing between P. aethiopicus dentition is well represented in the hypodigm of P.
and P. boisei is also much less strong than the case for robustus, most of the mandibles are crushed or distorted.
distinguishing the latter from H. habilis sensu lato. The brain, face, and chewing teeth of P. robustus are
Given that the fossil records of P. aethiopicus and P. boi- larger than those of Au. africanus, yet the incisors and
sei do not overlap in geological time, any decision to rec- canines are smaller, and whereas P. robustus includes
ognize these two taxa as separate species has no impact crania with ectocranial crests, there are no Au. africanus
on the issue of hominin taxic diversity. There is only one crania with unambiguous crests. What little is known
fragmented cranium for Au. garhi, but the differences about the postcranial skeleton of P. robustus suggests

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 HOMININ TAXIC DIVERSITY 49
that the morphology of the pelvis and the hip joint is the 2–1.5 Ma time interval provides the first evidence of
much like that of Au. africanus. hominin species in Asia, both on the mainland and in at
least one of the islands of Southeast Asia.
There is little doubt that the fossil evidence assigned
Paranthropus boisei and Homo habilis sensu to H. erectus, P. robustus, and Au. sediba, respectively,
lato. The third and fourth hominin species in the 2.0– sample three different species. There is also strong, but
1.5 Ma time interval, P. boisei and H. habilis sensu lato, not quite as compelling, evidence to suggest that P. boi-
have already been reviewed. sei is distinct from P. robustus (Tobias, 1967; Rak, 1983;
Wood and Constantino, 2007), H. habilis sensu stricto is
distinct from H. rudolfensis, and H. habilis sensu lato is
Homo ergaster. The fifth hominin species, Homo
distinct from H. erectus (Tobias, 1991; Wood, 1991, 1992;
ergaster (Groves and Maz ak, 1975), was established to
Spoor et al., 2007; Leakey et al., 2012, 2015; but see
accommodate fossil hominins recovered from Koobi Fora
Lordkipanidze et al., 2013 for a contrary interpretation),
that, in the judgment of the authors, did not belong in
but the evidence for those distinctions, as convincing as
the taxa identified up to that time. Wood (1994) used the
we believe it is, is not as clear-cut as the case for recog-
taxon name H. ergaster for hominin remains (e.g., KNM-
nizing H. erectus, P. robustus, and Au. sediba as sepa-
ER 730, 820, and 992) that are generally more primitive
rate species.
and lack the more extreme expressions of some of the
The case for making specific distinctions among H.
derived features (e.g., thick inner and outer table, sagit-
erectus, H. ergaster, and H. georgicus is weaker still.
tal keeling, etc.) seen in Asian Homo erectus. The best
Two categories of features are claimed to distinguish H.
postcranial evidence of H. ergaster comes from KNM-WT
ergaster from H. erectus. The first, which comprises fea-
15000 (Brown et al., 1985; Walker and Leakey, 1993).
tures for which H. ergaster is more primitive than H.
erectus, includes details of the mandibular dentition,
Homo georgicus. The sixth species we consider in and in particular the mandibular premolars. For exam-
this time interval, Homo georgicus (Gabounia et al., ple, the crowns of H. ergaster mandibular premolars are
2002), was established for hominins recovered from generally more complex than those of H. erectus, and
Dmanisi. The holotype is the mandible D2600. No para- the P3 roots of H. ergaster tend to have an accessory
types were formally designated, but the mandible D211, mesiobuccal root (and are thus more like those of the
the calvaria D2280, the cranium D2282, and the skull hypothetical common ancestor of the hominins) than are
and associated skeleton D2700 were referred to as evi- the simpler P3 roots of the majority of H. erectus speci-
dence that “will complete the characteristics of the new mens. The second category includes features in which
species” (Gabounia et al., 2002, p. 244). Although several H. ergaster is less derived in its cranial vault and cranial
of the authors of the original publication (e.g., Lordkipa- base morphology than is H. erectus. For example, it is
nidze et al., 2013) no longer support a separate taxon for claimed that H. ergaster lacks some of the more derived
this material, we treat it separately in our initial discus- features of H. erectus (e.g., thickened inner and
sion of the evidence in this time interval because the cir- outer tables and prominent sagittal and angular tori)
cumstances of the site are that the hominin evidence (Wood, 1984, 1991), but other researchers dispute the
from Dmanisi provides a sense of the range of variation distinctiveness of this material, and Spoor et al. (2007)
within what is likely to be a relatively short interval of found that the expression of some features is related to
time. Overall, the fossils are most similar to H. erectus, the overall size of the cranium such that larger H. erec-
and where there are differences they involve morphology tus crania are more likely to show the derived
inferred to be more primitive than that seen in H. morphology.
erectus.
1.5–1.0 Ma
Homo erectus, Paranthropus robustus, Paranthro-
Australopithecus sediba. The final species in the
pus boisei, Homo habilis sensu lato, and Homo
2.0–1.5 Ma time interval, Australopithecus sediba (Berger
ergaster. All of these species were reviewed in earlier
et al., 2010), was established to accommodate two associ-
time intervals.
ated skeletons, MH1, a sub-adult presumed male, and
MH2, an adult presumed female, recovered from Malapa,
Gauteng Province, South Africa. Berger et al. (2010) Homo antecessor. The sixth species, Homo antecessor
authors suggested that Au. sediba has cranial (e.g., more (Berm udez de Castro et al., 1997; Carbonell et al.,
globular neurocranium, gracile face), mandibular (e.g., 2008), was established to accommodate hominins recov-
more vertical symphyseal profile, a weak mentum ered from Level 6 of the Gran (or Trinchera) Dolina,
osseum), dental (e.g., simple canine crown, small anterior one of a complex of caves in the Atapuerca hills near
and postcanine tooth crowns), and pelvic (e.g., acetabuloc- Burgos, Spain. The hypodigm presently includes at
ristal buttress, expanded ilium and short ischium) mor- least 11 individuals, all but one of which are infants,
phology that departs from that seen in Au. africanus, and children, or juveniles. The modern human-like mor-
which is only shared with early and later Homo taxa. phology of the face and the apparent lack of derived
(Carlson et al., 2011; Kivell et al., 2011; Zipfel et al., Homo neanderthalensis features, combined with differ-
2011) make similar claims for the endocranial, hand, and ences between the Gran Dolina hominins and H. erec-
foot morphology, respectively, of Au. sediba. tus, led Berm udez de Castro et al. (1997) to propose
Evidence of diversity that the former fossils should be assigned to a new spe-
cies, H. antecessor, which they suggest is probably the
Just as the 4–3 Ma time interval witnessed evidence most recent common ancestor of H. neanderthalensis
of hominin species beyond eastern and central Africa, and H. sapiens.

Yearbook of Physical Anthropology

50 B. WOOD AND E. BOYLE

Evidence of diversity neanderthalensis. Other researchers (e.g., Dean et al.,

1998; Carbonell et al., 2005; Hublin, 2009) argue, contra
The evidence for the following species pairs, H. erectus Mounier et al. (2009), that H. heidelbergensis is an early
and P. boisei, H. erectus and P. robustus, and P. boisei stage in the accretion model for the origin of H. neander-
and H. habilis sensu stricto, being distinct is exception- thalensis (i.e., it should be included in H. neanderthalen-
ally strong. The case for distinguishing P. boisei and P. sis if that taxon is interpreted inclusively).
robustus is strong, but the scale of the morphological dif-
ferences between them is less than the comparisons
listed above. Homo rhodesiensis. The third hominin species in the
Despite the claims of Lordkipanidze et al. (2013), the 1.0–0.25 Ma time interval, Homo rhodesiensis (Wood-
case for distinguishing between H. erectus and H. habilis ward, 1921), is a species of pre-modern Homo introduced
is a strong one that involves detailed, and not so to accommodate the cranium and limb bones (Kabwe 1
detailed, cranial, mandibular, dental differences (Wood, or E 686) recovered from the Broken Hill lead mine at
1991) as well as some postcranial differences (Brown Kabwe, in what then was the British protectorate of
et al. 1985). It is also a substantially stronger case than Northern Rhodesia, now Zambia. Woodward reasoned
the one for distinguishing between H. erectus and H. that a new species was needed for the specimen because
ergaster (Wood, 1991). it did not fit within any of the existing Homo taxa. It
Assessments of the distinctiveness of H. antecessor are was not as primitive as what we now refer to as H. erec-
hindered because most of its hypodigm consists of indi- tus, nor as derived as either H. sapiens or H. neander-
viduals that have not stopped growing, so their morphol- thalensis. Morphologically, similar remains include
ogy cannot be easily compared with the mostly adult fossils from Hopefield/Elandsfontein in southern Africa,
remains belonging to other taxa in this time interval. Ndutu in East Africa, Sale in North Africa, and Bodo in
But nonetheless comparisons between H. antecessor and Ethiopia. The taxon H. rhodesiensis is used by research-
P. robustus, P. boisei and H. habilis sensu stricto, respec- ers who see H. heidelbergensis as an exclusively Euro-
tively, all carry a higher level of confidence than the pean pre-modern Homo taxon (see above).
comparisons between H. antecessor and H. erectus, and
H. antecessor and H. ergaster. But H. antecessor could
equally well be considered in the 1.0–0.25 Ma time inter- Homo helmei. The last formal species in the 1.0–0.25
val because (Berm udez de Castro et al., 2011) concluded Ma time interval, Homo helmei (Dreyer, 1935), was
there is not enough evidence to assign the Sima del Ele- established for the Florisbad 1 partial cranium discov-
fante mandible to H. antecessor. ered in 1932 in Florisbad, South Africa. Some research-
ers interpret the cranium as being intermediate in
1.0–0.25 Ma morphology between H. heidelbergensis and H. sapiens,
Homo erectus. The hominin species with historical with its face, brow ridges, and steeper inclination of the
priority in this time interval, H. erectus, was reviewed frontal bone, distinguishing it from the former, and
in an earlier time interval. retained primitive features (e.g., large brow ridge, more
receding frontal and low greatest breadth in the vault)
distinguishing it from anatomically modern humans.
Homo heidelbergensis. The second species in this Others have suggested that Jebel Irhoud, Ngaloba (aka
time interval, Homo heidelbergensis (Schoetensack, LH 18), and Omo II belong to the same hypodigm as the
1908), was created to accommodate a hominin mandible Florisbad 1 cranium.
found in 1907 in a commercial sandpit at Mauer, near
Heidelberg, Germany. Schoetensack concluded that the
Mauer mandible’s mosaic of primitive (no chin, robust Sima de los Huesos. The penultimate group in this
corpus, broad ramus, and an anterior–posteriorly deep time interval is not a formal species, but the unusually
mandibular symphysis) and derived (reduced canines complete and well-preserved collection of hominins
and modern human-like dental proportions) features recovered from the Sima de los Huesos, one of the many
was sufficient to distinguish it from Homo sapiens, H. breccia-filled cave systems that make up the Cueva
neanderthalensis, and what was then called Pithecan- Mayor-Cueva del Silo within the Sierra de Atapuerca,
thropus erectus. But H. heidelbergensis attracted little near Burgos in northern Spain. To date, more than
interest until it was suggested that it might be the most 6,500 hominin specimens belonging to at least 28 indi-
appropriate species name for a group of Afro-European viduals (Berm udez de Castro et al., 2003) have been
hominin fossils (e.g., Arago, Bodo, Kabwe, Mauer, recovered from excavations conducted both in the main
Ndutu, and Petralona) that had traditionally been cave itself (SH-A, SH-B, etc.) and in three pits (SRA,
labeled as “archaic” Homo sapiens (Rightmire, 1995). SRM and SRB) in the ramp that leads down to the cave.
Mounier et al. (2009) set out the morphological grounds The hominin remains include numerous crania, mandi-
for recognizing H. heidelbergensis as a taxon separate bles, hundreds of teeth, a nearly complete pelvis, verte-
from H. neanderthalensis, H. sapiens, and H. erectus as brae, ribs, hand and foot bones, and multiple specimens
well as providing a definition and a differential diagnosis of long bones. The cranial and mandibular sample as a
(ibid, pp. 243–244, also see Rightmire, 2008). whole shows a number of clearly derived features of H.
Mounier et al. (2009) discriminate between the “Afro- neanderthalensis including pronounced mid-facial prog-
European” and “European” hypotheses about H. heidel- nathism, the form of the brow ridge, a flat articular emi-
bergensis. The former interprets H. heidelbergensis as a nence of the glenoid fossa, a retromolar space, and an
geographically widely dispersed species that gave rise to asymmetrical configuration of the ramus of the mandi-
H. neanderthalensis in Eurasia and H. sapiens in Africa, ble. In contrast, the cranial vault is generally more ple-
whereas the latter interpretation sees H. heidelbergensis siomorphic (e.g., large, projecting, mastoid processes,
restricted to Europe where it is only ancestral to H. rounded neurocranium), with some incipient derived

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 HOMININ TAXIC DIVERSITY 51
traits of H. neanderthalensis (e.g., weak expression of be older than 150 ka, and most are probably younger
the suprainiac fossa). The dental remains show all of the than 100 ka.
traits that have been defined as typical of Neanderthals,
including a pronounced buccolingual expansion of the
anterior dentition and a reduction in the size of the post-
canine tooth crowns (Gomez-Robles et al., 2015). Homo neanderthalensis. The next species for consid-
eration in this time interval, Homo neanderthalensis
(King, 1864), was established for the partial skeleton
Homo antecessor. The final species to be considered recovered in 1856 from the Kleine Feldhofer Grotte in
in the 1.0–0.25 Ma time interval, H. antecessor, was
the part of the D€ussel valley named after Joachim Nean-
reviewed earlier.
der. Excavations of the same sediments in 1997 and
Evidence of diversity thereafter resulted in the recovery of fauna, artifacts,
and some 80 hominin fragments from at least two indi-
The best evidence for taxic diversity in the 1.0–0.5 Ma viduals were recovered. Discoveries made before 1856,
time interval is the co-existence in time, if not in space, such as the infant’s cranium from Engis (1828) and the
of H. erectus in China and Southeast Asia and H. heidel- partial cranium from Forbes’ Quarry, Gibraltar (1848),
bergensis in Europe. As for hominin taxonomic diversity were subsequently recognized as belonging to H. nean-
within Europe, researchers familiar with the evidence derthalensis. In the following half-century, remains
from the Sima de los Huesos make a distinction between attributed to H. neanderthalensis were discovered at
that site sample and the hypodigm of H. neanderthalen- other European sites, including La Naulette and Spy in
sis (e.g., Perez-Perez et al., 1999; Berm
udez de Castro 
Belgium, Sipka in Moravia, Krapina in Croatia, and
et al., 2003; Arsuaga et al., 2014; Arsuaga et al., 2015). Malarnaud, La Chapelle-aux-Saints, Le Moustier (lower
But even though the researchers working at the site shelter), La Ferrassie, and La Quina, among others, in
assigned the Sima de los Huesos hominins to H. heidel- France. In 1924–1926 the first H. neanderthalensis
bergensis, they acknowledged that H. heidelbergensis is remains were found outside of Western Europe at Kiik-
related to the Neanderthals in much the same way that Koba in the Crimea, and thereafter came discoveries at
Au. anamensis is related to Au. afarensis (see above). Tabun cave on Mount Carmel in the Levant, at Teshik-
Others more inclined to treat the fossils from the Sima Tash in central Asia, and at Saccopastore and Monte
de los Huesos as evidence of an early stage of a H. nean- Circeo in Italy. Further evidence was added after WWII,
derthalensis chronospecies include them within that first from Shanidar in Iraq, then from Amud and Kebara
taxon (Hublin, 2009). As for H. heidelbergensis and H. in Israel, and from Dederiyeh in Syria. New fossiliferous
rhodesiensis, the jury is still out on whether the latter is localities continue to be discovered in Europe (e.g.,
a junior synonym of the former, or a distinct taxon that Saint-Cesaire and Moula-Guercy in France, Zafarraya in
was the common ancestor of both H. neanderthalensis Spain, Vindija in Croatia, and Lakonis in Greece) and
and modern humans (see Fig. 1 in Hublin, 2009). Western Asia (e.g., Mezmaiskaya and Denisova in Rus-
While some researchers suggest that Florisbad 1 could sia). To date, Neanderthal remains have been found
serve as the holotype of a distinct species, H. helmei, throughout much of Europe below 558N, in the Near
most take the view there is no satisfactory diagnosis for East and in Western Asia.
such a taxon that separates it from H. heidelbergensis The earliest fossils that most researchers would accept
(or from H. rhodesiensis if the distinction between H. as H. neanderthalensis are from OIS 5 (i.e., c.130 ka),
heidelbergensis and H. rhodesiensis is accepted), on the but beyond that date there is no consensus. For exam-
one hand, and H. sapiens, on the other. ple, some (e.g., Hublin, 2009) interpret the fossil evi-
0.25 Ma to the present dence from Swanscombe (OIS 11 i.e., c.425–375 ka) and
the Sima de los Huesos (possibly as early as OIS 12 i.e.,
Homo sapiens. The species with historical priority in c.475–425 ka) as showing enough H. neanderthalensis-
this most recent time interval is Homo sapiens (Lin- like morphology to justify inclusion in H. neanderthalen-
naeus, 1758). The first widely accepted evidence that sis (Arsuaga et al., 2014), whereas others (e.g., Rosas
modern humans were ancient enough to have fossilized et al., 2006a) see a distinction between these specimens,
representatives came when a series of skeletal remains which they would include in H. heidelbergensis, and
were discovered by workmen at the Cro-Magnon rock- later “true” Neanderthals they claim do not appear until
shelter at Les Eyzies de Tayac, France, in 1868. Before OIS 6.
long discoveries of H. sapiens-like fossils were made Nuclear and mtDNA have been used to generate esti-
elsewhere in Europe (e.g., Mladec, Predmosti, and Brno), mates of the date of divergence of H. sapiens and H.
Asia and South-East Asia (e.g., Wadjak, Zhoukoudian neanderthalensis. An estimate using mtDNA and based
Upper Cave, and Niah Cave), the Near East (e.g., Skhul on an assumed divergence time of c.6–7 Ma for the mod-
and Djebel Qafzeh) and Australia (e.g., Willandra ern human and chimpanzees/bonobo lineages suggests a
Lakes). The first African fossil evidence for H. sapiens coalescence age of 660 6 140 ka (Green et al., 2008).
came in 1924 from Singa in the Sudan, and thereafter Estimates of the divergence time of the ancestral mod-
comparable evidence has come from Border Cave and ern human and Neanderthal populations based on the
Klasies River Mouth in South Africa, Dar es Soltane in nuclear genome range from 440 ka (this assumes an ear-
Morocco, and Dire-Dawa, Herto, and Omo-Kibish in lier, 8.3 Ma, divergence date for chimpanzees/bonobos
Ethiopia. With the exception of the c.190 ka date for and modern humans) to 270 ka (assuming a later, 5.6
Omo-Kibish (McDougall et al., 2005) and the c.170 ka Ma, divergence date) (Green et al., 2010). The most
date for Herto (Clark et al., 2003), there is no firm evi- recent estimates of the timing of the split between mod-
dence to suggest that any of the above sites is likely to ern humans and Neanderthals suggest it was c.600 ka).

Yearbook of Physical Anthropology

52 B. WOOD AND E. BOYLE

Homo erectus and Homo heidelbergensis. The next nuclear genome sequence showed that Denisovans and
two species in this time interval, H. erectus and H. hei- Neanderthals split from each other after their common
delbergensis, were reviewed in earlier time intervals. ancestor had separated from the line leading to modern
humans. A recent study of mitochondrial DNA of an
individual from Sima de los Huesos (Meyer et al., 2014)
Homo floresiensis. After H. erectus and H. heidelber- has shown a closer affinity between the mtDNA of Deni-
gensis, the next hominin species in the interval between sovans and Sima de los Huesos than between Deniso-
0.25 Ma and the present, Homo floresiensis (Brown vans and Neanderthals or modern humans. The
et al., 2004), was established to accommodate LB1, a implications of this finding remain to be tested with
partial adult hominin skeleton, and LB2, an isolated left nuclear DNA data from Sima de los Huesos.
P3, recovered in 2003 from the Liang Bua cave on the
island of Flores, Indonesia. More material belonging to Evidence of diversity
LB1 and evidence allocated to individuals LB4–9, includ-
ing LB6, a partial skeleton lacking a cranium, was Any proposal for a hominin species other than H. sapi-
recovered in 2004 (Morwood et al., 2005). The hypodigm ens in the 0.25 Ma to the present time interval must rise
now includes close to 100 individually numbered speci- to a simple challenge. Does the morphology of the fossil
mens that are estimated to represent fewer than 10 evidence lie outside the envelope of the morphological
individuals. variation documented for H. sapiens? And, if more than
The new species was immediately controversial for at one species meets that challenge, can those species be
least two reasons. First, its estimated geological age of distinguished from one another?
between c.17 and c.74 ka (it is probably closer to the lat- Those who claim that H. neanderthalensis is specifi-
ter age) substantially overlapped with the estimated cally distinct from H. sapiens recognize morphological
ages of evidence of the presence of modern humans in autapomorphies that distinguish the former from both
the region. Second, its discoverers and describers inter- earlier (e.g., H. erectus and H. heidelbergensis) and con-
preted its small overall size (the stature of LB1 is esti- temporary (i.e., H. sapiens) Euro-Asian hominin taxa.
mated to be c.105 cm and its body mass to be roughly These features are found in the cranium (e.g., large,
between 25 and 30 kg), small brain (c.420 cm3) and rounded discrete brow ridges, projecting mid-face,
primitive morphology, as evidence of a novel endemically angled cheeks, small mastoid process, suprainiac fossa,
dwarfed species of Homo. Initially it was suggested that and occipital bun), mandible (long corpus, retro-molar
H. floresiensis was a dwarfed H. erectus, but the burden space, and asymmetric mandibular notch), dentition
of subsequent analyses suggests that it may be more (e.g., large shovel-shaped incisors, distinctive occlusal
closely related to a more primitive hominin such as H. morphology of molars and premolars, a high incidence of
habilis sensu stricto (Tocheri et al., 2007; Argue et al., taurodontism), and the postcranial skeleton (e.g., long
2009; Brown and Maeda, 2009; Morwood and Jungers, clavicle, teres minor groove extending onto the dorsal
2009). Other researchers have suggested that no new surface of the scapula, large infraspinous fossa, long,
taxon needs to be erected because they claim the “Homo thin pubic ramus, and large joints). Especially signifi-
floresiensis hypodigm” has been sampled from a popula- cant are reports that the ontogeny of H. neanderthalen-
tion of Homo sapiens—most likely related to the small- sis differed in several ways from that of modern humans
statured Rampasasa people who live on Flores today— (Tillier, 1982; Nelson and Thompson, 2005; Coqueugniot
afflicted by either an endocrine disorder or a range of and Hublin, 2007; Ponce de Leon et al., 2008; Gunz
syndromes that include microcephaly, or who retain et al., 2010; Smith et al., 2010).
primitive morphology (see Baab et al., 2013 and West- The taxon H. neanderthalensis is currently the only
away et al., 2015 for refutations of these hypotheses). extinct hominin species for which there is ancient DNA
evidence from many individuals and sites. The draft
Denisovans. The final evidence to be considered in sequence of the Neanderthal nuclear genome (Green
this time interval is based on the analysis of ancient et al., 2010), which focused on three individuals
DNA recovered from the distal phalanx of the fifth digit (Vi33.16, 25, and 26) from Vindija, was compared with
(little finger) of a hominin hand (Denisova 3) recovered smaller amounts of sequence data from Neanderthal
from the c.48–30 ka layer 11 in Denisova Cave in the specimens from El Sidron, Kleine Feldhofer Grotte, and
Altai Mountains in Russia. When mtDNA from the dis- Mezmaiskaya, as well as the sequenced nuclear genomes
tal phalanx was compared with the mtDNA of 54 mod- of five modern humans. The results show that whereas
ern humans, one chimpanzee and one bonobo, six the modern humans from sub-Saharan Africa contained
Neanderthals, and a single fossil H. sapiens from Kos- no evidence of Neanderthal DNA, the three modern
tenki (Krause et al., 2010) it was concluded that it came humans from outside of Africa showed similar, low
from a hominin that, while distinct from both modern amounts (between 1 and 4%) of DNA shared with Nean-
humans and Neanderthals, shared a common ancestor derthals. These results are compatible with either a
with both species c.1.0 million years ago. Reich et al. deep split within Africa between the population that
(2010) sequenced the nuclear genome of the Denisova gave rise to modern Africans and a second one that gave
hominin phalanx plus mtDNA from a large-crowned rise to present-day non-Africans plus Neanderthals, or
maxillary molar (Denisova 8) found in the cave whose with the hypothesis that there was hybridization
crown morphology is distinct from both modern humans between Neanderthals and modern humans soon after
and Neanderthals. Meyer et al. (2015) recently published the latter left Africa, perhaps in western Asia. In sum-
a high-coverage sequence from the same individual. Sub- mary, the morphological differences between modern
sequently, a deciduous molar (Denisova 2) and another humans and Neanderthals are consistent with a species-
maxillary molar (Denisova 4) have been identified as level distinction, and the genetic evidence is also consist-
Denisovans (Sawyer et al., 2015; Slon et al., 2015). The ent with such a conclusion.

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 53

Fig. 2. A more realistic estimate of the temporal ranges of the hominin species, site collections, and individual fossils referred
to in this review. The bottoms and tops of the continuous columns represent, respectively, the published first and last appearances
dates. The tiles below and above many of the columns reflect the various sources of error that should be added to the mean ages
that were used to locate the bottom and the top of each continuous column. The sources of the data are set out in the Appendices A
and B.

With respect to the alpha taxonomy of H. floresiensis, scribe to the interpretation that the fossil evidence from
views are sharply polarized. The consensus is that if you Liang Bua is a dwarfed early Homo species that is dis-
take the hypodigm as a whole it is most parsimoniously tinct from all of the other taxa in this time interval.
interpreted as evidence of a novel endemically dwarfed Differences between the ancient mtDNA and nuclear
pre-modern Homo, or early Homo, species (Brown et al., DNA extracted from the distal phalanx of the fifth (little
2004; Morwood et al., 2005; Tocheri et al., 2007; Mor- finger) digit of a hominin hand, and the mtDNA
wood and Jungers, 2009). Less than a handful of extracted from a large-crowned maxillary molar, both
researchers, literally, cling to the view that the “Homo found in Denisova Cave, and the DNA of modern
floresiensis hypodigm” samples a H. sapiens popula- humans and Neanderthals, are consistent with Deniso-
tion—most likely related to the small-statured Rampa- vans and modern humans belonging to different species
sasa people who live on Flores today—all of which are (Reich et al., 2011; Patterson et al., 2012; Pr€ ufer et al.,
afflicted by either an endocrine disorder (see Obendorf 2014). Denisovans and Neanderthals were also likely to
et al. (2008) and a rebuttal by Brown (2012)) or one, or have been separate species given that DNA recovered
more, of a range of syndromes that include microcephaly. from Neanderthals across Europe was consistently more
Both explanations, a novel dwarfed early hominin spe- similar to one another than any individual Neanderthal
cies, or a pathological population of modern humans, are was to the DNA of the Denisovans.
exotic, but those who espouse a pathological explanation So within the time interval between 0.25 Ma to the
for the individuals represented by LB1–15 need to present time, there seems sound evidence for hominin
explain what pathology results in a phenotype that taxic diversity in the form of species-level differences
resembles an early Homo-like cranial vault, primitive among H. sapiens, H. neanderthalensis, H. erectus, and
mandibular, dental, carpal, and pedal morphology, and a H. floresiensis. There is also less strong, but still poten-
brain that, while very small, apparently has none of the tial, evidence for two more taxa, H. heidelbergensis and
morphological features associated with the majority of the Denisovans. It is possible that the H. heidelbergensis
types of microcephaly (Vannucci et al., 2011). We sub- hypodigm and the Denisovan DNA sample the same

Yearbook of Physical Anthropology

54 B. WOOD AND E. BOYLE

taxon (Meyer et al., 2014), especially if mtDNA results Smith (2005) exposes with forensic effectiveness just
were to be confirmed by the results of future nuclear how small the sample sizes of some early hominin hypo-
DNA analyses, but the substantial morphological and digms are, and what this means for our ability to dem-
metric differences between at least one of the teeth from onstrate that a new sample exceeds the variation within
Denisova and equivalent teeth from the Sima de los an existing species. That being said, if new fossil evi-
Huesos need be taken into account. dence happens to preserve a region in which the ratio of
inter- to intra-specific variation is favorable (see below),
and if the specimens are relatively complete, it is possi-
DISCUSSION ble to demonstrate taxic diversity even with relatively
The assessment of taxic diversity depends on pro- small sample sizes. Indeed, there are several instances
posals about (A) the existence of species, and (B) how in the hominin fossil record where just one new fossil
long-lived those species are (Figs. 1 and 2). If these pro- (e.g., OH 7) is so different from the fossil evidence of a
posals are incorrect, the assessment will be flawed. Both synchronic species (e.g., OH 5) that the case for taxo-
types of proposal are bedeviled by the adequacy of the nomic distinctiveness is obvious. But, as Smith (2005)
respective samples that are available. Namely, how close points out, not all claims for new hominin taxa are as
are the data we have to the data one would need in securely based as this example.
order to determine whether the hypothesis that a given How can the situation be improved? The obvious solu-
sample of fossils includes more than one species is a reli- tion is to find (and publish!) additional fossil evidence from
able one (i.e., it resists attempts at falsification)? And, new and existing sites and localities, and then provide, in
even if a new species is justified, how close are the avail- a timely fashion, detailed and reliable information about
able data to the evidence one would need in order to the new evidence. This will serve to expand our knowledge
generate reliable hypotheses about its temporal span? of existing fossil hominin taxa. But while we await the
Turning first to alpha taxonomy, the criteria that should assembly of larger hypodigms is there anything else we
be used to determine whether a newly discovered fossil can do? One strategy is to expand our understanding of
sample falls outside the range of variation of an existing, variation within closely-related extant taxa and apply any
lessons learned to interpreting the hominin fossil record.
reference, species are the same as the criteria that are
Comparative studies that collect data from large samples
used to determine if a newly discovered living animal jus-
of the great apes for a particular region, such as the stud-
tifies the erection of a new species. Its phenotype is com-
ies of dental morphology by Uchida (2004) and Pilbrow
pared with museum collections that sample the species’ it
(2010), are especially helpful because they allow us to con-
resembles, and if researchers are convinced the new speci-
duct the thought experiment of imagining what variation
men falls comfortably outside that range of variation then
in a large sample of an early hominin might look like.
there are grounds for erecting a new species.
Another strategy is to look at patterns of inter- and intra-
Museum collections of most living animals comprise
specific variation within and among the extant taxa most
complete specimens numbered in the hundreds, thus they closely-related to fossil hominins. If there are any common
relatively faithfully reflect the population (i.e., the extant patterns, and if we make the reasonable working assump-
species) from which they were drawn. The equivalent exer- tion that the hominin clade shares the same pattern (i.e.,
cise involving the hominin fossil record would be to use the principle of the phylogenetic bracket, see Witmer,
the observed range of variation for the relevant variables 1995), parsimony suggests that the same pattern should
of the reference taxon (e.g., Au. afarensis) in the time also apply to fossil hominin taxa. One of us has attempted
interval occupied by the newly discovered fossils (e.g., c.3.4 to do this in the past (Wood, 1975; Wood et al., 1991) as
Ma), and if the newly discovered fossils fall comfortably have others (e.g., Lieberman et al., 1988; Kramer, 1993;
outside that range of variation then there are grounds for Shea et al., 1993; Lockwood et al., 1996; Miller, 2000; Har-
erecting a new species. The problem is that we do not vati et al., 2005). Sound comparative evidence about inter-
have the equivalent of comprehensive museum collections and intra-specific variation could then be used to generate
to estimate the parameters of the reference fossil hominin additional criteria to help researchers make judgments
species, just the fossils that make up its hypodigm. In the about the taxonomic significance of any observed morpho-
case of Au. afarensis, and thanks to decades of field work, logical differences between small samples of fossil
researchers have access to what in relative terms is a hominins.
respectable sample from Hadar, with important but much Until we have larger hypodigms for fossil hominin
smaller samples from other sites. But that becomes a taxa, the factors we have reviewed above suggest to us
much smaller sample when you restrict it to the parts of that the procedures traditionally used in paleoanthropol-
the hypodigm known from c.3.4 Ma, and even if one ogy are likely to lead to the over-reporting of new taxa,
relaxes the time constraint and considers the regions of with the inevitable result that taxic diversity will also
the skeleton (e.g., dentition and mandible) that are best be over-reported (i.e., Type I error).
represented in the Au. afarensis hypodigm, we are still The assessment of taxic diversity also depends on the
talking about sample sizes that would be considered unac- ability to determine how long-lived hominin taxa are.
ceptably low in studies of extant primates. One strategy is to use comparative evidence about spe-
But this is a “best case” scenario, for there many cies longevity from other mammalian groups. Two stud-
examples of species in the hominin fossil record (e.g., P. ies that looked at the time span of mammalian taxa in
aethiopicus, O. tugenensis, and Au. garhi) where the ref- the Cenozoic resulted in very different estimates of how
erence hypodigm is both smaller and less comprehensive long species persist. Foote and Raup (1996) suggest the
than that of Au. afarensis in terms of coverage of the median duration of a species is 1.7 Ma, whereas the
regions of the skeleton. This means that the existing estimate of Barnosky et al. (2011) of 0.6 Ma is substan-
hypodigms of many early hominin taxa fail to realisti- tially shorter. In the early hominin fossil record, two
cally capture the nature and range of variation of the species, namely Au. afarensis and P. boisei, have a good
parent species population. In a much under-cited review, fossil record with well-dated samples from several sites,

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 55
and their hypodigms span c.0.7 Ma and c.1.0 Ma, CONCLUSIONS
respectively. These data, meager as they are, would
suggest it is a reasonable working hypothesis to assume It is evident from this review that not all proposals to
that the median duration of an early hominin species recognize new hominin taxa are equally compelling, but
was in the order of one million years, plus or minus the strength of the case for establishing a new species
0.25 Ma years. can change because our perspective is constantly shift-
A second strategy is to work with the data we have ing as new discoveries expand hypodigms and thus
for each hominin species. To assign species, site sam- change our understanding of the context of such
ples, and individual fossils to time intervals we used proposals.
conservative versions of the FAD and LAD of each spe- In Table 2, we present our assessment of the strength
cies based on their published ages (Table I) (Fig. 1) of the existing evidence for species distinctions in the
(Appendix A). We also assembled a second version that, form of pairwise comparisons of species, or species equiv-
where applicable, incorporates the age, and the pub- alents, for each of the time intervals. In each time inter-
lished error of the age, of the nearest underlying dated val we list the species, or species equivalents, in order of
horizon in the case of the FAD, and the age, and the their proposal or discovery and score the case for each of
published error of the age, of the nearest overlying the comparisons into one of three confidence categories,
dated horizon in the case of the LAD (Table I) (Fig. 2) high, medium, or low. In Appendix B we set out some of
(Appendix A). For various reasons even this observed the factors we took into account when making these
FAD of a taxon is almost certainly later than the time decision, but we want to emphasize that these are sub-
the taxon actually originated in, or migrated into, that jective assessments others can challenge. But at the
region, just as the equivalent observed LAD of a taxon very least these proposals for taxonomic diversity are
is almost certainly earlier than the time the taxon “on the table” for debate.
actually became extinct, or emigrated from that region. As can be seen from Table 2, apart from the period
How much earlier than the observed FAD the actual prior between 7 and 5 Ma, in all of the other time inter-
origination or migration occurred, and just how much vals we sampled there is at least one example of taxic
later than the observed LAD of a taxon the actual diversity that enjoys what we judge to be a high level of
extinction, or emigration occurred, is determined by confidence. Two time intervals (5–4 Ma, 4–3 Ma) have
many factors. These include technical considerations one example, two (3.0–2.5 Ma and 2.5–2.0 Ma) have
such as dating errors (see above), but a much more three examples, one (1.5–1.0 Ma has six examples 1.0–
potent factor is the nature of the relevant mammalian 0.25 Ma) have seven examples, one (2.0–1.5 Ma) has
fossil record before and after the current observed FAD eight examples and one (0.25 Ma to the present) has ten
and LAD. The problem is the old adage “absence of evi- examples of species diversity. All of these numbers, espe-
dence is not evidence of absence” – in other words it is cially those for the earlier time intervals, are likely to
yet another sampling problem. At most sites early hom- increase as hypodigms expand and new sites extend the
inins are such a rare component of the mammalian fau- time ranges of the existing species. There is also the
nal record (c.1–2%) that researchers need to find a potential for fossil evidence of additional species. There
substantial number of non-hominin mammalian fossils is also the potential for finding fossil evidence of addi-
(at least several hundred) without finding any evidence tional species such as Homo naledi (Berger et al., 2015).
of a particular hominin species before it can be reason- Our preliminary assessment of the fossil hominin evi-
ably assumed that species was not part of the faunal dence recovered from the Dinaledi chamber of the Rising
assemblage being sampled. So, for example, whereas Star system suggests the claim that this represents a
researchers can be reasonably confident there were no new species needs to be taken seriously. However, even
hominins at Omo-Shungura in Ethiopia prior to their if we assume that the discoveries in the Dinaledi Cham-
FAD, at nearby Koobi Fora, in the same lake basin, ber add to the evidence for taxonomic diversity, until
there is a major break in sedimentation that spans researchers can determine its age (Dirks et al., 2015) we
several hundred thousand years prior to the FAD of will not know which time interval, or intervals, would be
several hominin species (i.e., there is no fossil record affected by the new evidence.
during that time). So we have little basis for concluding As for the future, researchers who find new fossil evi-
there were no hominins at Koobi Fora prior to their dence that lies outside the envelope of the existing hypo-
FAD at that site (Bobe and Leakey, 2009). We know digms of closely-related fossil species should at least
even less about the time span of the early hominin convince themselves that the new material is unlikely to
species that are presently found only at southern Africa be sampled from the population (i.e., the species) from
sites. which the samples (i.e., the existing hypodigm) of those
For all of these reasons there are uncertainties about taxa has been drawn. There is also a need to guard
the temporal span of all of the fossil hominin taxa we against the tendency in all of us to “focus on the
reviewed above, especially those with small hypodigms strength or extremeness of the available evidence” and
sampled at one or two sites. However, because it is impos- to have “insufficient regard for its weight or credence”
sible for a FAD to occur before a species actually origi- (Griffin and Tversky, 1992, p. 411). This may explain
nated, or for an LAD to occur after a species actually went why people in general, and perhaps paleoanthropologists
extinct, all of the species we reviewed will have had longer in particular, are “often more confident in their judg-
temporal spans than the ones we used. So, any species rec- ments than is warranted by the facts” (ibid, p. 411).
ognized in the hominin fossil record is likely to extend into Devising a formal strategy to take account of this tend-
more, rather than fewer, time intervals, however they are ency for over-confidence is a challenge that faces all
defined. This source of error, unlike the problems that researchers involved in fossil discovery. However, if
bedevil the identification of new taxa, will almost certainly forced to choose between systematic over-reporting of
result in the systematic under-estimation of hominin taxic hominin species and systematic under-reporting, we
diversity (i.e., Type II error). would opt for the former. When you compare the history

Yearbook of Physical Anthropology

56 B. WOOD AND E. BOYLE

of H. erectus and H. habilis, it is clear that having for- just above the Bodele Tuff. Samples from the Botele Tuff
mal labels (i.e., Pithecanthropus erectus and Sinanthro- yield dates of 6.24 6 0.19 Ma and 6.48 6 0.22 Ma, leading
pus pekinensis) for the units that now comprise H. Simpson et al. (2015) to suggest a date of c.6.3 Ma for
erectus made the process of taxonomic revision of that the ABD1 specimen. The material from Amba East is
taxon a lot easier to manage than past and current younger, coming from the 5.55 6 0.09–5.18 6 0.07 Ma
efforts to split the H. habilis sensu lato hypodigm. Kuseralee Member of the Sagantole Formation (Renne
What this evidence for taxic diversity within the homi- et al., 1999).
nin clade means for lineage diversity within that clade FAD: 6.3/6.7 Ma; LAD: 5.2/5.11 Ma
is outside the scope of this review, but while lineage
diversity cannot exist without taxonomic diversity, taxo-
nomic diversity does not always mean there is lineage Ardipithecus ramidus
diversity. For example, although we suggest there is evi-
Key publications: White et al., 1994, 1995, 2009;
dence of species-level distinctions between Au. anamen-
Semaw et al., 2005
sis and Au. afarensis and between P. aethiopicus and P.
Main sites: Localities in Ethiopia, and possibly also in
boisei, there is also good evidence that at least one of
Kenya (see below). In the Central Awash Complex in the
these pairs of taxa (Kimbel et al., 2006) belong to the
Middle Awash study area: Aramis, Kuseralee Dora,
same lineage. In addition, we are not as convinced as
Sagantole. Also, As Duma in the Gona Western Margin.
some of our colleagues that current phylogenetic hypoth-
Temporal range: Magnetostratigraphy and argon dat-
eses are reliable enough to generate sound hypotheses
ing have yielded an age range of c.4.5–4.3 Ma for Ar.
about lineage diversity.
In conclusion, we suggest that there is no need for ramidus (WoldeGabriel et al., 1994, 2009; Renne et al.,
Ernst Mayr, or anyone else, to seek a solution to the 1999; Kleinsasser et al., 2008; Quade et al., 2008;). Most
puzzle of why the “hominid stock” stopped speciating of the Ar. ramidus material from the Middle Awash
(Mayr, 1950, p. 116). It is apparent that hominins did study areas is from the Lower Aramis Member of the
speciate on a scale commensurate with that of some Sagantole Formation, which is underlain by the
other large mammal clades. So, to return to the question 4.419 6 0.068 Ma G aala Vitric Tuff Complex (GATC) and
posed in the title, hominin taxic diversity exists. overlain by the 4.416 6 0.031 Ma Daam Aatu Basaltic
Tuff (DABT) (WoldeGabriel et al., 2009). Some specimens
ACKNOWLEDGMENTS found above the DABT are overlain by the Wodara
Basaltic Tuff (WOBT) (White et al. 2009). WOBT, which
Authors would like to thank Andrew Barr, Aida Gomez- is not dated, lies underneath the 4.317 6 0.055 Ma Kul-
Robles, Bill Kimbel, and Richard Smith for their insightful lunta Basaltic Tuff (KUBT), and Gani and Gani (2011)
comments on earlier versions. A later version of the manu-
argue that an average sedimentation rate of 0.5
script was improved by comments from Shara Bailey,
mm yr21 between DABT and KUBT indicates that
Katharine Balolia, Andrew Du, Fred Smith, and an anony-
WOBT was likely deposited within 5 ky of the deposition
mous reviewer. BW would like to thank the Provost of
of the DABT. Until an age for WOBT is determined, we
George Washington University for his continuing support
use 4.262 Ma for the minimum LAD of Ar. ramidus. The
for the GW University Professorship of Human Origins
Ar. ramidus specimens from Gona are found in the
and via the GW Signature Program for his support of
CASHP. <5.2–>4.6 Ma as Duma Member and the 4.6–4.2 Ma
Segala Noumou Member of the Sagantole Formation,
APPENDIX A: TEMPORAL EVIDENCE FOR though the member affiliation for remains from GWM-
SPECIES, SITE COLLECTIONS, AND SINGLE 10 is uncertain (Quade et al., 2008). These dates there-
SPECIMENS fore call into question the original 4.51–4.32 Ma tempo-
ral span estimation for the material from this area
(Semaw et al., 2005). The specimens from GWM-5
Ardipithecus kadabba series,-9n,-16 are aged to >4.6 Ma. However, since
Quade et al. (2008) describe the dating of the As Duma
Key publications: Haile-Selassie, 2001; Haile-Selassie Member as a “work in progress,” we use a maximum
et al., 2004 FAD of 4.6 Ma for these specimens until the results of
Main sites: Localities in Ethiopia. In the Middle
further research are published. The specimens from
Awash study area: Saitune Dora, Alayla, Asa Koma, and
GWM-3,-3w have a more reliable estimated age range of
Digiba Dora in the Western Margin, and Amba East in
4.4–4.3 Ma.
the Central Awash Complex. Localities in the Gona Pale-
Wood (2013) and Kissel and Hawks (2015) suggest
oanthropological Research Project study area are in the
that the morphology of the 4.42 Ma Tabarin mandible
Western Ethiopian Escarpment and Asbole Dora.
(KNM-TH 13150) has affinities with Ar. ramidus, and
Temporal range: The Ar. kadabba specimens from the
Western Margin and Gona derive from the Adu-Asa For- tentatively assign it to that taxon, but it would not alter
mation, which has been dated via magnetostratigraphy, the estimated temporal range of this taxon.
tephrostratigraphy, and argon dating to >6.4–5.2 Ma FAD: 4.51/4.6 Ma; LAD: 4.3/4.262 Ma
(Kleinsasser et al., 2008; Quade et al., 2008). The fossil-
iferous layers from the Western Margin are confined to Australopithecus afarensis
the Asa Koma Member and have an estimated age range
of 5.77–5.54 Ma (WoldeGabriel et al., 2001). In addition Key publications: Leakey, 1976; Johanson et al.,
to describing Ar. kadabba remains from c.5.4 Ma layers 1978,1982a,b,; White et al., 1981; Clark et al., 1984;
of the Adu-Asa Formation at Gona, Simpson et al. (2015) Kimbel et al., 1984, 1994, 2004; Kimbel, 1988; Suwa,
tentatively attribute one specimen (ABD1/P1) to this 1990; Alemseged et al., 2005, 2006; Kimbel and Dele-
taxon that comes from a layer of this formation that lies zene, 2009; Haile-Selassie et al., 2010.

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 57
Main sites: Localities in Ethiopia in the Middle Awash Australopithecus anamensis
study area (Belohdelie [tentative], Dikika, Hadar, Maka,
and Omo [Usno]), and the Korsi Dora in the Woronso- Key publications: Fleagle et al., 1991; Coffing et al.,
Mille study area. Also, localities in Koobi Fora, Kenya, 1994; Leakey et al., 1995, 1998; Ward et al., 1999, 2001;
and Laetoli, Tanzania. White, 2002; White et al., 2006
Temporal range: Radiometric dating, biostratigraphy, Main sites: Localities in the Omo-Turkana Basin in
magnetostratigraphy have established a temporal span Kenya (Allia Bay and Kanapoi) and Ethiopia (Fejej), in
of c.3.7–2.9 for Au. afarensis (Kimbel and Delezene, 2009). the Middle Awash Study Area (Aramis and Asa Issie)
The Belohdelie frontal bone remains were found above the and in the Woronso-Mille study area of Ethiopia.
3.97 6 0.03 Ma Tuff VT-1 (Moiti tuff), and have an esti- Temporal range: The specimens from Allia Bay come
mated age of c.3.89–3.86 Ma (Clark et al., 1984; McDougall from within and just beneath the 3.97 6 0.03 Ma Moiti
and Brown, 2008; McDougall et al., 2012). Confirmation of Tuff, and have an estimated age of c.3.9 Ma (Coffing
these fossils as Au. afarensis would make them the earliest et al., 1994; Leakey et al., 1995; McDougall and Brown,
evidence for that taxon and push back the FAD to c.3.89 2008; McDougall et al., 2012). The material from Kanapoi
Ma. The Dikika skeleton derives from the c.3.42–3.24 Ma is bracketed by the 4.2 6 0.033 Ma Lower pumiceous tuff
Sidi Hakoma Member of the Hadar Formation and has an and the 4.11 6 0.03 Ma Kanapoi Tuff (McDougall et al.,
estimated age range of c.3.37–3.34 (Alemseged et al., 2012). Geomagnetic and radiogenic dating yield a mini-
2006). The DIK-2-1 mandible comes from the basal mem- mum age estimate for the Fejej material of c.4.18–4.0 Ma
ber of the Hadar Formation and thus has a minimum age (Kappelman et al., 1996). If these specimens do belong to
of c.3.42 Ma (Alemseged et al., 2005). The fossils from Au. afarensis as originally argued, this pushes back the
Hadar span the entirety of the Hadar Formation, ranging FAD of that taxon to 4.18 Ma, but if these specimens are
in age from c.3.42 to 2.9 Ma (Kimbel and Delezene, 2009). attributed to Au. anamensis (White, 2002; Kimbel and Del-
The specimens from Maka come from the “Maka sands” ezene, 2009), then they fall well within the estimated tem-
horizon, which is dated via tephrocorrelation to c.3.4 Ma poral span for that taxon. The Aramis maxilla was found
(Renne et al., 1999). The teeth from the Usno Formation just above sediments that appear to preserve the base of
in the Omo-Turkana basin have an approximate age of chron C2Ar, yielding an age of c.4.21 Ma (White et al.,
c.3.0 Ma (Feibel et al., 1989; Suwa, 1990). The skeleton 2006). The locality where this specimen was found (ARA-
from Korsi Dora was from just above a 3.6 6 0.03 Ma tuff, VP-14) is bracketed by the 4.317 6 0.055 Ma Kullunta
and has an age of c.3.6–3.58 Ma (Haile-Selassie et al., Basaltic Tuff of the Sagantole Formation and the
2010). The calvaria from Koobi Fora is from the Tulu Bor 4.041 6 0.06 Ma MA 94-55C volcanic layer (Renne et al.,
Member of the Koobi Fora Formation, and has an age of 1999; White et al., 2006). We therefore use an age of c.4.37
c.3.3 Ma (Kimbel and Delezene, 2009). The fossils and foot- Ma as the FAD for this taxon. Cumulative evidence from
prints from the Upper Laetolil Beds at Laetoli span the magnetostratigraphy, argon dating, and biostratigraphy
age range from c.3.7 to 3.5 Ma (Kimbel and Delezene, from Asa Issie led White et al. (2006) to provide an esti-
2009). mated age range of c.4.12–3.77 Ma for the specimens at
FAD: 3.7/3.89 Ma; LAD: 3.0/2.9 Ma. this site, though the authors note the older date is likely
more accurate. The Au. anamensis specimens from
Woronso-Mille that have been argued as evidence for ana-
Australopithecus africanus genetic evolution between this taxon and Au. afarensis
span the period from c.3.82 to 3.57 Ma (Deino et al., 2010).
Key publications: Dart, 1925, 1948; Berger, 1992; We will thus use the older of these ages as the less con-
Clarke and Tobias, 1995; Lockwood and Tobias, 1999; servative LAD estimate for Au. anamensis. However, if the
Partridge et al., 2003; Toussaint et al., 2003; Moggi- FAD for Au. afarensis is actually 3.89 Ma (see below), then
Cecchi et al., 2006 a c.3.82 LAD for Au. anamensis is problematic if these
Main sites: Several localities in Gauteng Province, taxa are related anagenetically.
South Africa including Gladysvale, Makapansgat (Mem- FAD: 4.2/4.37 Ma; LAD: 3.9/3.82 Ma.
bers 3 and 4), Sterkfontein (Member 4, possibly Member
2), and possibly Jacovec cavern. The type locality; Taung
in the North West Province, South Africa. Australopithecus bahrelghazali
Temporal range: Curnoe (1999) suggests that the Au.
africanus remains at Gladysvale may derive from inter- Key publications: Brunet et al., 1995, 1996
nal deposits at the cave and are likely 2.4–1.9 Ma in Main site: Bahr el Ghazal, Koro Toro, Chad.
age. Herries (2003) and Herries et al. (2010) provide an Temporal range: The Au. bahrelghazali specimens
age range of c.3.0–3.6 Ma for the fossiliferous deposits at from the KT 12 locality have an estimated age of
Makapansgat Members 3 and 4. Magnetostratigraphy 3.58 6 0.27 Ma based on biostratigraphy and cosmogenic
and U-series dating at Sterkfontein Member 4 yield an nuclide dating (Lebatard et al., 2008).
age range of 2.65 6 0.30 Ma to 2.01 6 0.05 Ma for the fos- FAD: 3.58/3.85 Ma; LAD: 3.58/3.31 Ma
sils from this layer (Pickering and Kramers, 2010). Bio-
stratigraphy suggests an estimated age range of 3.0–2.6 Australopithecus deyiremeda
Ma for the Taung child (Herries et al. 2013). If StW 573
from Sterkfontein Member 2 belongs within Au. africa- Key publication: Haile-Selassie et al., 2015
nus and if the newly estimated age for this deposit is Main sites: The Burtele and Waytaleyta collection areas
accepted, then the FAD of Au. africanus would be c.3.67 in the Woranso-Mille study area, central Afar region,
Ma (Granger et al., 2015). If the Jacovec cavern speci- Ethiopia.
mens belong to this taxon, then the FAD of Au. africa- Temporal range: Magnetostratigraphy and argon dat-
nus would be c.4.02 Ma (Partridge et al., 2003). ing provide an estimated age range of c.3.5–3.3 Ma for
FAD: 3.0/4.02 Ma; LAD: 2.4/1.9 Ma. the Au. deyiremeda specimens (Haile-Selassie et al.,

Yearbook of Physical Anthropology

58 B. WOOD AND E. BOYLE

2015). Specifically, the localities fall within the 3.596– Temporal range: Thermoluminesence, electron spin
3.330 Ma C2An.3n magnetozone. resonance spectroscopy, biostratigraphy, and magneto-
FAD: 3.5/3.596 Ma; LAD: 3.3/3.33 Ma stratigraphy have yielded an age of c.1 Ma for the H.
antecessor remains from the Sierra de Atapuerca sites
(Pares and Perez-Gonz alez, 1995, 1999; Cuenca-Bescos
Australopithecus garhi et al., 1999; Falguères et al., 1999; Garcıa and Arsuaga,
Key publication: Asfaw et al., 1999 1999; Berger et al., 2008; Pares et al., 2013; Moreno
Main site: Bouri, Ethiopia in the Middle Awash study et al., in press). Paleomagnetic sampling of younger lev-
area. els provided a minimum age of 0.936 Ma for the fossils
Temporal range: The Au. garhi hypodigm derives from and suggests they were likely deposited in the TD6 level
the c.2.5 Ma Hatayae Member of the Bouri Formation during OIS 25 (Pares et al., 2013). The age of the fossils
that has been dated with argon dating, lithostratigraphy, from TD6 has been further refined to c.800–900 ka
and tephrostratigraphy (de Heinzelin et al., 1999). The based on electric spin resonance applied on quartz (Mor-
holotype was recovered from just above the 2.496 6 0.008 eno et al., in press). If the Sima del Elefante hominin is
Ma Maoleem Vitric Tuff. de Heinzelin et al. (1999) sug- included, the FAD date would extend to 1.2–1.1 Ma
gest that the hominin remains are no younger than 2.45 (Rosas et al., 2006ab).
Ma based on predicted sedimentation accumulation rates. FAD: 1.0/1.2 Ma; LAD: 0.936 Ma
FAD: 2.5 Ma; LAD: 2.45/2.488 Ma
Homo erectus
Australopithecus sediba
Key publications: Dubois, 1893; Black, 1927, von Koenigs-
Key publication: Berger et al., 2010 wald, 1936, 1968, 1975; Weidenreich, 1936a, 1936b, 1937,
Main site: Malapa, Gauteng, South Africa 1944, 1951; Leakey, 1961; Jacob, 1973; Rightmire, 1979;
Temporal range: An age range of 1.95–1.78 Ma was Santa Luca, 1980; Widianto and Grimaud-Herve, 1993;
originally given for the Au. sediba hypodigm based on Anton, 1999; Kaifu et al., 2008; Zaim et al., 2011
biostratigraphy, uranium-lead dating, and magnetostra- Main sites: Localities in Asia including Java, Indone-
tigraphy (Dirks et al. 2010). A flowstone layer with an sia (Kedung Brubus, Mojokerto, Ngandong, Ngawi, Sam-
age of 2.026 6 0.021 Ma underlies the breccia that con- bungmacan, Sangiran, and Trinil) and Zhoukoudian,
tains the fossils. Uranium-lead dating of a 2.048 6 0.140 China. Also Bouri from the Middle Awash study area in
Ma flowstone that caps the hominin remains and magne- Ethiopia and Olduvai Gorge, Tanzania.
tostratigraphy provide a refined age of 1.977 6 0.002 Ma Temporal range: The Mojokerto calvaria from the Dje-
for the fossils, which correlates to the pre-Olduvai geo- tis Beds has an approximate age of 1.81 6 0.04 Ma
magnetic event (Pickering et al., 2011). (Swisher et al., 1994). Dating of Ngandong has been con-
FAD: 1.98/2.05 Ma; LAD: 1.98/1.91 Ma troversial, with U-series dating yielding an age range of
c.101–31 ka for the H. erectus remains and electron spin
Burtele foot resonance spectroscopy dating providing an age range of
c.46.4–27.3 ka (Orchiston and Siesser 1982; Bartstra
Key publication: Haile-Selassie et al., 2012 et al., 1988; Gr€
un et al., 1997). We will use c.27 ka as the
Main sites: Burtele collection area in the Woranso- LAD of H. erectus, even though that date is controversial.
Mille study area, central Afar region, Ethiopia. The Sambungmacan cranial remains have an estimated
Temporal range: The 3.469 6 0.008 Ma Burtele tuff is age of c.53–27 ka based on electron spin resonance spec-
27 m below the pedal remains of BRT-VP-2/73, yielding troscopy dating and U-series dating (Swisher et al.,
a maximum age of c.3.47 (Haile-Selassie et al., 2012). 1996). The H. erectus fossils from the Sangiran Dome
Sediment accumulation rate suggests an approximate deposits span the Sangiran and Bapang Formations, and
age range of c.3.4–3.2 Ma for the foot. have an age range of 1.51 6 0.08 to c.0.9 Ma (Larick
FAD: 3.4/3.47 Ma; LAD: 3.4/3.2 Ma et al., 2001; Ciochon et al., 2005). Biostratigraphy provide
ages of c.1 Ma for the Trinil remains and c.800 ka for the
Kedung Brubus fossils (Weinand, 2005). Cosmogenic
Denisovans nuclide dating, U-series dating, and correlation of OIS,
Key publication: Krause et al., 2010 and loess sequences suggest an age of c.780–400 ka for
Main site: Denisova Cave, Siberia, Russia the material from Zhoukoudian (Shen et al., 2009). The
Temporal range: Although biostratigraphy suggests a Bouri (Daka) material has an age of c.1.0 Ma (Asfaw
maximum age of c.125 ka for the occupation levels at the et al., 2002). The specimens from Olduvai Gorge are from
cave, the Denisova phalanx comes from a level that has an 3.6 to 3 m below Bed III in Bed II, and above the
uncalibrated radiocarbon age range of 48.65 6 2.38 ka to 1.72 6 0.03 Ma Tuff IIA, giving them an estimated age
29.2 6 0.36 ka (Derevianko et al., 2008; Krause et al., 2010). range of c.1.7–1.2 Ma (Manega, 1993).
Sequence differences suggest a much old age for teeth FAD: 1.81/1.85 Ma; LAD: 27 ka
recovered from deepes levels in the cave (Slon et al., 2015).
FAD: 48.65/50.63 ka; LAD: 29.2/28.84 ka
Homo ergaster

Homo antecessor Key publications: Leakey, 1974; Groves and Maz ak,
1975; Leakey and Walker, 1985; Wood, 1994
Key publications: Berm udez de Castro et al., 1997, Main sites: Localities in Kenya (Koobi Fora and West
2008; Carbonell et al., 2005 Turkana), and perhaps also Swartkrans, Gauteng Prov-
Main site: Gran Dolina, Sierra de Atapuerca, Spain ince, South Africa.

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 59
Temporal range: Recent argon dating by McDougall et al. (Spoor et al., 2015). It was found in sediments just below
(2012) provides ages of 1.47 6 0.03 Ma and 1.44 6 0.03 Ma the KBS Tuff of the Koobi Fora Formation, thus having a
for the specimens from the Nachukui Formation in West Tur- minimum age of 1.87 6 0.02 Ma (McDougall et al., 2012).
kana, an age range of 1.65 6 0.05 Ma to 1.43 6 0.04 Ma for KNM-ER 1813 is from Area 123 in the Upper Burgi Mem-
the material from the Karari Ridge at Koobi Fora, and an ber of the Koobi Fora Formation and has an approximate
age of 1.50 6 0.04 Ma for the remains from the Ileret region age of c.1.90–1.88 Ma (Feibel et al., 1989), though McDou-
of Koobi Fora. We will therefore use 1.7 Ma and 1.4 Ma as the gall et al. (2012) note that the stratigraphy of this locality
conservative FAD and LAD ages, respectively, for H. ergaster. is still debated. KNM-ER 3735 is also from this horizon.
The specimens from Swartkrans attributed to H. ergaster Spoor et al. (2015) express uncertainty about whether the
come from Members 1, 2, and potentially Member 3 (postcra- c.1.44 Ma KNM-ER 42703 specimen belongs to H. habilis
sensu stricto, so we will use 1.65 Ma, the age of OH 13, as
nial remains) of the Swartkrans Formation, yielding an esti-
the LAD. The same authors also suggest that A.L. 666-1
mated temporal range of 2.19 6 0.08 to 0.96 6 0.09 Ma
from Hadar, which derives from sediments just under
(Gibbon et al., 2014). These less conservative dates make H. Bouroukie Tuff 3 in the Busidima Formation and thus has
ergaster contemporaneous with P. robustus in South Africa. a minimum age of c.2.35 Ma (Campisano, 2012), may not
FAD: 1.7/2.27 Ma; LAD:1.4/0.87 Ma belong to H. habilis, but even if it is removed from the
hypodigm it does not affect the FAD of this taxon (see
Homo floresiensis below). The cranial fragments from the Shungura Forma-
tion come from Member G, and are c.2.0 Ma (McDougall
Key publications: Brown et al., 2004; Morwood et al., et al., 2012). The H. habilis material from Olduvai Gorge
2005 come from Bed I and Lower Bed II, with an estimated
Main site: Liang Bua, Flores, Indonesia temporal span of 1.877 6 0.013 Ma to c.1.65 Ma (Deino,
Temporal range: Thermoluminesence and carbon dat- 2012). If the c.2.6–2.0 Ma craniodental remains (Stw 53)
ing provide an age range for the occupation deposits at from Sterkfontein belongs within H. habilis, it becomes
Liang Bua of 95 6 13 ka to c.18–16 ka (Morwood and the oldest evidence of this taxon (Herries et al., 2010). SK
Jungers et al., 2009; Roberts et al., 2009; Westaway 847 comes from Member 1 of the Swartkrans Formation
et al., 2009). Homo floresiensis remains in the cave are and thus has a maximum age of 2.19 6 0.08 Ma (Gibbon
et al., 2014). Based on previously noted similarities with
found in excavation levels spanning c.74–17 ka.
KNM-ER 1802, the mandible from Uraha is likely also H.
FAD: 74/108 ka; LAD: 17/16 ka
habilis, and has an estimated age of c.2.5–2.3 Ma based on
biostratigraphy (Kullmer, 2008).
Homo georgicus FAD: 2.35/2.6 Ma; LAD: 1.65 Ma

Key publications: Gabounia et al., 2002; Vekua et al.,

2002; Lordkipanidze et al., 2007, 2013
Homo heidelbergensis
Main site: Dmanisi, Georgia. Key publications: Schoetensack, 1908; Kokkoros and
Temporal range: Magnetostratigraphy has aged the occu- Kanellis 1960; de Lumley and de Lumley 1973; Rightmire,
pation at Dmanisi to the interval from c.1.85 to 1.77 Ma, 1995; Manzi et al., 2001; Mounier et al., 2009
spanning the last half of the Olduvai Subchron through the Main sites: Several localities in Europe: Mauer,
earliest Upper Matuyama chron (Ferring et al., 2011). Germany, Caune de I’Arago, France, Petralona, Greece,
FAD: 1.85 Ma; LAD: 1.77 Ma1 Montmaurin, France, plus Tighenif, Algeria.
Temporal range: The Mauer mandible has been dated to
OIS 15 based on biostratigraphy, magnetostratigraphy, elec-
Homo habilis sensu stricto tron spin resonance spectroscopy, and U-series dating, with
Key publications: Leakey et al., 1964, 1971, 1989; an approximate age of 609 6 40 ka (Wagner et al., 2010). The
Leakey, 1974; White et al., 1981; Leakey and Walker, material from Arago has a minimum age of c.350 ka, based
on U-series dating of an overlying stalagmitic formation,
1985; Johanson et al., 1987; Tobias, 1991; Wood, 1991;
which is consistent with an age of c.400 ka for the Arago XXI
Grine et al., 1993; Schrenk et al., 1993; Kimbel et al., facial remains (Falguères et al., 2004). Several dating meth-
1996; Grine, 2001; Blumenschine et al., 2003; Spoor ods have been used to approximate an age for the Petralona
et al., 2015 cranium, but a reliable age has not yet been specified
Main sites: Localities in Kenya (Koobi Fora and West Tur- (reviewed by Poulianos, 2005). Age estimates for this cra-
kana), Ethiopia (Hadar [tentative] and Omo-Shungura), nium range from the lower Middle to late Middle Pliestocene,
Tanzania (Olduvai), Gauteng Province, South Africa (Sterk- from c.700 to 160 ka. The Montmaurin mandible has an esti-
fontein and Swartkrans), and Malawi (Uraha). mated age range of c.390–100 ka (Girard, 1973). The speci-
Temporal range: Argon dating by McDougall et al. mens from Tighenif have an estimated age of c.700 ka based
(2012) provides estimated ages of 1.75 6 0.05 Ma for KNM- on magnetostratigraphy (Geraads et al., 1986).
ER 1805 and 2.09 6 0.05 Ma for the H. habilis specimen FAD: 700 ka; LAD: 100 ka
from Kangaki, West Turkana. Based on similarities with
OH 7, KNM-ER 1802 is likely to belong to H. habilis
Homo helmei
1 Key publications: Dreyer, 1935; Ennouchi, 1962; Leakey
In our review, pairwise comparisons are made between Homo habilis
sensu lato and other taxa in the relevant time intervals. The temporal
et al., 1969; Day et al., 1980
range of H. habilis sensu lato is the combination of the temporal ranges Main sites: Several sites in Africa including Florisbad,
of H. habilis sensu stricto and Homo rudolfensis. But because the tem- South Africa, Jebel Irhoud, Morocco, Ngaloba Beds, Laetoli,
poral range of H. rudolfensis is narrower and within that of H. habilis Tanzania, Omo-Kibish, Ethiopia and Singa, Sudan.
sensu stricto (Fig. 2), the temporal range of H. habilis sensu lato is Temporal range: Electron spin resonance spectroscopy
effectively that of H. habilis sensu stricto. dating suggests that the Florisbad cranium has an age

Yearbook of Physical Anthropology

60 B. WOOD AND E. BOYLE

of c.260 ka (Gr€ un et al., 1996). Electron spin resonance Temporal range: KNM-ER 1470 has an age of 2.03 6 0.05
spectroscopy and U-series dating yield an estimated age Ma based on argon dating by McDougall et al. (2012). This
range of c.190–90 ka Jebel Irhoud remains (Hublin, date was subsequently adjusted to 2.03–2.09 Ma by Joordens
1992; Gr€ un and Stringer, 1991; Smith et al., 2007). The et al. (2013). We will thus use c. 2.0 Ma as the conservative
LH 18 cranium from Laetoli has an age of c.200 ka, as FAD for this taxon, and 2.09 Ma as the less conservative esti-
determined by amino acid racemization (Manega, 1993). mate. The recently described facial and mandibular remains
The Omo II calvaria is contemporaneous with the Omo I with affinities to H. rudolfensis have an estimated age range
H. sapiens material, with an estimated age of 195 6 ka of c.1.95–1.78 Ma (Leakey et al., 2012; Spoor et al., 2015).
(McDougall et al., 2005; Feibel, 2008). U-series dating and FAD: 2.0/2.09 Ma; LAD: 1.95/1.78 Ma
electron spin resonsance spectroscopy provide an estimated
age range of 145.5 6 7.5 to 89 6 9.3 ka for the Singa 1 cal-
varia (Gr€un and Stringer, 1991; McDermott et al., 1996). Homo sapiens
FAD: 260 ka; LAD: 80 ka
Key publications: Keith, 1912; Day, 1967; White et al.,
2003; Schwartz and Tattersall, 2010
Homo neanderthalensis Main sites: Many sites in the Old World and some in
the New World.
Key publications: King, 1864; Schmitz et al., 2002; Temporal range: Argon dating has yielded ages for the ear-
White et al., 2014 liest anatomically modern humans, with ages of 195 6 5 ka
Main sites: Many sites in Europe, Asia, and the Near for the Omo I material from Omo-Kibish, Ethiopia (McDougall
East.
et al., 2005) and an estimated age range of c.160–154 ka for
Temporal range: An age range of c.200 ka–40 ka for
the material from Herto, Ethiopia (Clark et al., 2003).
Neanderthals has been established via various dating meth-
FAD: 195/200 ka; LAD: present
ods, including radiocarbon, thermoluminesence, electron
spin resonance spectroscopy, and uranium-series dating
techniques. Though H. neanderthalensis-like morphologies Kenyanthropus platyops
appear earlier, the fossils that most researchers would accept
as H. neanderthalensis sensu stricto are in OIS 5 c.130 ka Key publications: Leakey et al., 2001
(e.g. Bruner and Manzi, 2006). The inclusion of some conten- Main site: Lomekwi,West Turkana, Kenya.
tious specimens like the c.170 ka Lazaret material, the c.240 Temporal range: The K. platyops material originally
ka Biache-Saint-Vaast specimens, or the c.400 ka Swan- had an estimated age range of c.3.5–3.3 Ma, based on
scombe skull would significantly push back the first appear- argon dating and magnetostratigraphy (Leakey et al.,
ance date of this taxon (Hublin 2009; Michel et al., 2009; 2001). Recent argon dating for the deposition of the Omo
Bahain et al., in press). Recently, a Neanderthal skeleton Group beds yields slightly older ages for these specimens
from Altamura, Italy was dated to 172 6 15 to 130.1 6 1.9 ka (McDougall et al., 2012). The type specimen is from the
(Lari et al., 2015) via U-series dating, so we will use the Kataboi Member of the Nachukui Formation, found 12 m
upper age from this analysis as this taxon’s less conservative above the 3.60 6 0.05 Ma Lokochot Tuff and 8 m below the
FAD. The disappearance of Neanderthals happened at differ- 3.44 6 0.02 Ma Tulu Bor Tuff, and thus has an estimated
ent times across sites (Higham et al., 2014). Using improved age of c.3.54 Ma. Other specimens are from the lower
carbon dating techniques on samples from 40 sites and
Lomekwi Member, and have estimated ages of c.3.35 Ma.
Bayesian age modeling, Higham et al. (2014) estimate a last
FAD: 3.54/3.65 Ma; LAD: 3.35 Ma
appearance date of 40.89–39.22 ka for Neanderthals in
Europe, assuming this taxon was responsible for both the
Mousterian and Chatelperronian traditions. The authors LD 350-1
note that it is possible that some populations, such as those
in southern Iberia, may have persisted after this date. Key publication: Villmoare et al., 2015
FAD: 130/197 ka; LAD: 40/39 ka Main site: Lee Adoyta region of the Ledi-Geraru study
area, Ethiopia
Temporal range: Based on magnetostratigraphy and
Homo rhodesiensis stratigraphic scaling, the Ledi-Geraru mandible has an esti-
Key publications: Woodward, 1921; Conroy et al., 1978 mated age range of 2.80–2.75 Ma (Villmoare et al., 2015).
Main sites: Kabwe (Broken Hill), Zambia and Bodo, The fossil is bracketed by the 2.842 6 0.007 Ma Gurumaha
Ethiopia. Tuff and by the 2.665 6 0.016 Ma Lee Adoyta Tuff.
Temporal range: Dating attempts involving amino acid FAD: 2.80/2.85 Ma; LAD: 2.75/2.65 Ma
racemization and biostratigraphy for the Broken Hill remains
have been problematic, but it is generally accepted that the H. Orrorin tugenensis
rhodesiensis specimens from this site date to the Middle Pleis-
tocene, most likely c.300 ka (Barham et al., 2002; Trinkaus, Key publication: Senut et al., 2001
2009; but see McBrearty and Brooks, 2000). Based on biostra- Main sites: Localities in the Lukeino Formation at
tigraphy and argon dating, the Bodo hominins have an esti- Aragai, Cheboit, Kapcheberek, and Kapsomin, Tugen
mated age of c.600 ka (Kalb et al., 1980; Clark et al., 1994). Hills, Baringo District, Kenya.
FAD: 600 ka; LAD: 300 ka Temporal range: The Orrorin hypodigm has been dated
to c.6.0 Ma by magnetostratigraphy and potassium-argon
Homo rudolfensis radiometric dating (Deino et al. 2002, Sawada et al., 2002)
that approximates a 6.0–5.7 Ma accumulation range for the
Key publications: Alexeev, 1986; Groves, 1989; Wood, Lukeino Formation. The fossils from Aragai and Cheboit are
1992; Leakey et al., 2012; Spoor et al., 2015 from the Kapgoywa Member, which is bounded by two zones
Main sites: Koobi Fora, Kenya. of reverse polarity that have ages of 6.14–6.05 Ma and 5.89–

Yearbook of Physical Anthropology

 HOMININ TAXIC DIVERSITY 61
5.83 Ma (Hill, 1985; Baksi, 1993; Cande and Kent 1995; Wei, FAD: 2.3/2.5; LAD: 1.3/1.15 Ma
1995). Most of the O. tugenensis specimens are from the Kap-
somin Member, and have an estimated age range of 5.9–5.8
Ma (Sawada et al., 2002). The single specimen from the Kap- Paranthropus robustus
cheberek member is the youngest and lies near the top of the Key publications: Broom, 1938, 1949; Grine, 1989;
Lukeino Formation, with minimum age estimates of Berger et al., 1995; Menter et al., 1999; Keyser, 2000;
5.66 6 0.14 Ma (Sawada et al., 2002) and 5.73 6 0.05 Ma Steininger and Berger, 2000; Thackeray et al., 2001; de
(Deino et al., 2002). Ruiter et al., 2009
FAD: 6.0/6.14 Ma; LAD: 5.7/5.52 Ma Main sites: Localities in Gauteng, South Africa: Coop-
er’s, Drimolen, Kromdraai (Member 3), Sterkfontein
Paranthropus aethiopicus (Member 5b) and Swartkrans (Members 1, 2, and 3).
Gondolin in the North West Province, South Africa.
Key publications: Arambourg and Coppens, 1968; Temporal range: Uranium-lead dating of a basal speleo-
Walker et al., 1986; Wood and Chamberlain, 1987; Suwa, them (CDD1) at Cooper’s D has yielded a maximum age of
1988 1.526 6 0.088 Ma for the fossil material in this infill (de
Main sites: Omo, Ethiopia, West Turkana, Kenya, and Ruiter et al., 2009). A flowstone layer above the basal spe-
Laetoli, Tanzania. leothem (CDD3) has been aged to 1.526–1.413 Ma. Some
Temporal range: KNM-WT-17000 is underlain by the fossils are preserved above this layer, yielding an approxi-
2.53 6 0.02 Ma Lokalalei Tuff and has an estimated age of mate age range of c.1.5–<1.4 Ma for the Cooper’s D fossils.
c.2.5 Ma (Feibel et al., 1989; McDougall et al., 2012). Keyser et al. (2000) suggest an age range of 2.0–1.5 Ma for
KNM-WT 16005 is from the Upper Lomekwi Member and the Drimolen material based on biostratigraphy, though
is thus slightly older, with an age of c.2.55 Ma (Feibel O’Regan and Menter (2009) note that the Drimolen felids
et al., 1989; McDougall et al., 2012). If the KNM-WT 16002 resemble those from Cooper’s D, implying a younger 1.6–
femur belongs within P. aethiopicus, then the FAD of this <1.4 Ma age range for Drimolen (Herries and Adams,
taxon extends back to 2.7 6 0.03 Ma (Brown et al., 2001). 2013). Magneto- and biostratigraphy yield a likely age
The specimens from Members C through F of the Shun- range of c.1.8–1.6 Ma for the Kromdraai material (Herries
gura Formation at Omo span the estimated age range from et al., 2009; Herries and Adams, 2013). P. robustus at Gon-
c.2.6 to 2.27 6 0.04 Ma (McDougall et al., 2012). Potassium- dolin has an age range of 1.95–1.78 Ma based on magneto-
argon and argon dating of the Upper Ndolanya Beds pro- and biostratigraphy, though Herries and Adams (2013)
vide an age estimate of c.2.66 Ma for the putative P. aethio- note that the younger age is a better estimate. If P. robus-
picus remains at Laetoli (Deino, 2011). tus is represented at Sterkfontein Member 5, Herries and
FAD: 2.66/2.73 Ma; LAD: 2.3/2.23 Ma Shaw (2011) suggest an age range of c.1.4–1.2 Ma for this
material based on electron spin resonance spectroscopy,
Paranthropus boisei uranium-lead dating, and magnetostratigraphy. Members
1, 2, and 3 at Swartkrans preserve P. robustus specimens
Key publications: Leakey, 1959; Robinson, 1960; Lea- that span the entire age range of this taxon (Herries and
key and Leakey, 1964; Tobias, 1967; Day, 1969; Carney Adams 2013). Cosmogenic nuclide burial dating of these
et al., 1971; Leakey and Walker, 1988; Suwa, 1988; members by Gibbon et al. (2014) refines the temporal span
Suwa et al., 1997; Kullmer et al., 1999; Wood and Con- estimate for P. robustus at Swartkrans, and yields a maxi-
stantino, 2007 mum age of 2.19 6 0.08 Ma (an average of 1.99 6 0.19 Ma)
Main sites: Localities in Ethiopia (Konso and Omo), for Member 1, and an age of 0.96 6 0.09 Ma for Member 3.
Kenya (Chesowanja, Koobi Fora, and West Turkana), FAD: 2.0/2.27 Ma; LAD: 1.0/0.87 Ma
Malawi (Malema), and Tanzania (Olduvai Gorge and Peninj)
Temporal range: The specimens from Konso have an age
range of c.1.45–1.3 Ma based on biostratigraphy and argon Sahelanthropus tchadensis
dating (Suwa et al., 1997). Argon dating provides an esti-
Key publication: Brunet et al., 2002
mated age range of 2.27 6 0.04 to c.1.2 Ma for the specimens
Main site: TM 266 locality, Toros-Menalla, Chad
from Member G and above of the Shungura Formation at
Temporal range: Toros-Menalla has been dated by bio-
Omo (McDougall et al., 2012). The fossils from the Chemoi-
stratigraphy and cosmogenic nuclide dating to c.7.0 Ma,
gut Formation at Chesowanja have an age of c.1.43 Ma
with an estimated range from 7.2 to 6.8 Ma (Lebatard
(Hooker and Miller, 1979). McDougall et al. (2012) yield an
age range of 1.59 6 0.05 to 1.41 6 0.05 Ma for the P. boisei et al., 2008). Specifically, section TM 266 is bracketed by
specimens from the Ileret region of Koobi Fora, and an age ash tuffs aged 6.83 6 0.45 Ma and 7.12 6 0.31 Ma, yield-
range of 1.75 6 0.05 to 1.54 6 0.05 Ma for the specimens from ing an age of 7.04 6 0.18 Ma for the S. tchadensis
the Karari Ridge at Koobi Fora. P. boisei from the Kalochoro remains (Lebatard et al., 2008).
and Kaitio Members of the Nachukui Formation span the FAD: 7.2/7.43 Ma; LAD: 6.8/6.38 Ma
range from c.2.3 to 1.6 Ma at West Turkana (Feibel et al., Sima de los Huesos hominins
1989). Kullmer (2008) uses biostratigraphy to generate an
age range of c.2.5–2.3 Ma for a maxilla assigned to P. boisei Key publications: Arsuaga et al., 1993, 1997, 1999
from Malema. Based on argon dating and magnetostratigra- Main site: Sima de los Huesos, Sierra de Atapuerca, Spain
phy, the specimens from Bed I and Bed II at Olduvai Gorge Temporal range: The remains from Sima de los Huesos
span the range from c.1.92 to 1.15 Ma, with OH 3, OH 36, have been most recently dated via luminescence dating
and possibly OH 38 from Upper Bed II as the youngest speci- and magnetostratigraphy (Arnold et al., 2014). The
mens (Tamrat et al., 1995; Deino, 2012). The Peninj 1 mandi- updated minimum age estimate is 427 6 12 ka, and the
ble has an age range of c.1.5–1.3 Ma based on potassium- maximum age is c.780 ka.
argon and argon dating (Domınguez-Rodrigo et al., 2009). FAD: 780 ka; LAD: 427/415 ka

Yearbook of Physical Anthropology

 APPENDIX B: SUMMARY JUSTIFICATION FOR ALLOCATING LEVELS OF CONFIDENCE TO THE MAIN PAIRWISE COMPARISONS SPECIES 62

Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatricb differentiation
0
7.0 Ma–5.0 Ma Orrorin tugenensis vs. Sahelan- 10 Mandible, dentition, femur, Yes Yes No Moderate
thropus tchadensis humerus, phalanx vs. Cranium,
mandible, dentition
Orrorin tugenensis vs. Ardipithe- 100 Mandible, dentition, femur, Yes No No Moderate

Yearbook of Physical Anthropology

cus kadabba humerus, phalanx vs. Mandible,
isolated teeth, upper limb long
bones & phalanges, pedal
phalanx
Sahelanthropus tchadensis vs. 100 Cranium, mandible, dentition vs. Yes Yes No Moderate
Ardipithecus kadabba Mandible, isolated teeth, upper
limb long bones & phalanges,
pedal phalanx
5.0 Ma–4.0 Ma Ardipithecus ramidus vs. Australo- 102 Dentition and fragmented remains Yes Yes Yes High
pithecus anamensis of the cranium, mandible, verte-
brae, pelvis, upper and lower
limb long bones, carpals and tar-
sals, metacarpals and metatar-
sals, hand and pedal phalanges
vs. Maxilla, mandible, dentition,
humerus, femur
4.0 Ma–3.0 Ma Australopithecus afarensis vs. Bur- 102 Nearly the entire skeleton vs. Yes No Yes High
tele foot Metatarsals and phalanges
B. WOOD AND E. BOYLE

Australopithecus afarensis vs. Ken- 102 Nearly the entire skeleton vs. Cra- Yes Yes No Moderate
yanthropus platyops nium and dentition
2
Australopithecus afarensis vs. Aus- 10 Nearly the entire skeleton vs. Yes Yes No Low
tralopithecus bahrelghazali Mandible and teeth
Australopithecus afarensis vs. Aus- 102 Nearly the entire skeleton vs. Yes Yes Yes Low
tralopithecus deyiremeda Maxilla, mandible, and dentition
0
Australopithecus bahrelghazali vs. 10 Mandible and teeth vs. Maxilla, No No No Low
Australopithecus deyiremeda mandible, and dentition
Australopithecus bahrelghazali vs. 100 Mandible and teeth vs. Cranium No No No Low
Kenyanthropus platyops and dentition
0
Australopithecus bahrelghazali vs. 10 Mandible and teeth vs. Metatar- No No No Low
Burtele foot sals and phalanges
Kenyanthropus platyops vs. Aus- 100 Cranium and dentition vs. Maxilla, Yes Yes No Low
tralopithecus deyiremeda mandible, and dentition
0
Kenyanthropus platyops vs. Bur- 10 Cranium and dentition vs. Meta- No No No Low
tele foot tarsals and phalanges
Burtele foot vs. Australopithecus 100 Metatarsals and phalanges vs. Yes No Yes Low
deyiremeda Maxilla, mandible, and dentition
 APPENDIX B Continued

Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatricb differentiation
3 Ma–2.5 Ma Australopithecus africanus vs. Par- 101 Cranium, mandible, dentition and No No No High
anthropus aethiopicus some postcranial evidence vs.
Cranium, mandible, dentition
Australopithecus africanus vs. LD 102 Cranium, mandible, dentition and Yes Yes No High
350-1 some postcranial evidence vs.
Mandible and dentition
Paranthropus aethiopicus vs. LD 101 Cranium, mandible, dentition vs. Yes Yes No High
350-1 Mandible and dentition
2.5 Ma–2.0 Ma Australopithecus africanus vs. Par- 100 Cranium, mandible, dentition and Yes Yes No High
anthropus boisei some postcranial evidence vs.
Cranium, mandible, dentition,
distal humerus, radius, femur,
and tibia fragment
Australopithecus africanus vs. 100 Cranium, mandible, dentition and Yes Yes Potentially Moderate
Homo habilis sensu lato some postcranial evidence vs.
Cranium, mandible, dentition,
hand phalanges, tarsals, meta-
tarsals, and other fragmented
postcranial remains including
upper and lower long bone
shafts
Australopithecus africanus vs. Aus- 102 Cranium, mandible, dentition and Yes Yes No Moderate
tralopithecus garhi some postcranial evidence vs.
Cranial fragments, maxilla,
dentition
Paranthropus boisei vs. Australopi- 102 Cranium, mandible, dentition, dis- Yes Yes No Moderate
HOMININ TAXIC DIVERSITY

thecus garhi tal humerus, radius, femur, and

tibia fragment vs. Cranial frag-
ments, maxilla, dentition
Paranthropus boisei vs. Homo 100 Cranium, mandible, dentition, dis- Yes Yes Yes High
habilis sensu lato tal humerus, radius, femur, and
tibia fragment vs. Cranium,
mandible, dentition, hand pha-
langes, tarsals, metatarsals, and
other fragmented postcranial
remains including upper and
lower long bone shafts
Australopithecus garhi vs. Homo 102 Cranial fragments, maxilla, denti- Yes Yes No High
habilis sensu lato tion vs. Cranium, mandible, den-
tition, hand phalanges, tarsals,
metatarsals, and other frag-
mented postcranial remains
including upper and lower long
bone shafts
63

Yearbook of Physical Anthropology

 APPENDIX B Continued
64
Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
b
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatric differentiation
1
Homo habilis stricto stricto vs. 10 Cranium, mandible, dentition, Yes Yes Yes Moderate
Homo rudolfensis hand phalanges, tarsals, meta-
tarsals, and other fragmented
postcranial remains including
upper and lower long bone
shafts vs. Cranium, dentition,
mandibles
2.0 Ma–1.5 Ma Homo erectus vs. Paranthropus 100 Nearly the entire skeleton except Yes Yes Potentially High
robustus for most hand and foot bones vs.

Yearbook of Physical Anthropology

Cranium, mandible, dentition,
hand bones, vertebrae, femur,
talus, and other fragmentary
postcranial remains
Homo erectus vs. Paranthropus 100 Nearly the entire skeleton except Yes Yes Yes High
boisei for most hand and foot bones vs.
Cranium, mandible, dentition,
distal humerus, radius, femur,
and tibia fragment
Homo erectus vs. Homo habilis 100 Nearly the entire skeleton except Yes Yes Yes High
sensu lato for most hand and foot bones vs.
Cranium, mandible, dentition,
hand phalanges, tarsals, meta-
tarsals, and other fragmented
postcranial remains including
upper and lower long bone
shafts
B. WOOD AND E. BOYLE

Homo erectus sensu stricto vs. 101 Cranium, mandible, dentition, Yes Yes Yes Low
Homo ergaster humerus, clavicle, vertebra,
lunate, pelvis, lower limb long
bones vs. Nearly the entire skel-
eton except for most hand and
foot bones
Homo erectus vs. Homo georgicus 101 Nearly the entire skeleton except Yes Yes Potentially Low
for most hand and foot bones vs.
Cranium, mandible, dentition,
vertebrae, clavicle, scapular
fragment, ribs, humerus, hand
phalanges, femur, patella, tibia,
some tarsals and metatarsals,
foot phalanges
Homo erectus vs. Australopithecus 100 Nearly the entire skeleton except Yes Yes Yes High
sediba for most hand and foot bones vs.
Cranium, mandible, dentition,
upper limb long bones, complete
hand, pelvis fragments, lower
limb long bones, partial foot
 APPENDIX B Continued

Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatricb differentiation
2.0 Ma–1.5 Ma Paranthropus robustus vs. Homo 100 Cranium, mandible, dentition, Yes Yes Yes High
habilis sensu lato hand bones, vertebrae, femur,
talus, and other fragmentary
postcranial remains vs. Cra-
nium, mandible, dentition, hand
phalanges, tarsals, metatarsals,
and other fragmented postcra-
nial remains including upper
and lower long bone shafts
Paranthropus robustus vs. Para- 100 Cranium, mandible, dentition, Yes Yes No High
nthropus boisei hand bones, vertebrae, femur,
talus, and other fragmentary
postcranial remains vs. Cra-
nium, mandible, dentition, distal
humerus, radius, femur, and
tibia fragment
Paranthropus robustus vs. Austral- 101 Cranium, mandible, dentition, Yes Yes No High
opithecus sediba hand bones, vertebrae, femur,
talus, and other fragmentary
postcranial remains vs. Cra-
nium, mandible, dentition, upper
limb long bones, complete hand,
pelvis fragments, lower limb
long bones, partial foot
Paranthropus boisei vs. Homo See above See above See above See above See above See above
habilis sensu lato
Paranthropus boisei vs. Australopi- 101 Cranium, mandible, dentition, dis- Yes Yes No High
HOMININ TAXIC DIVERSITY

thecus sediba tal humerus, radius, femur, and

tibia fragment vs. Cranium,
mandible, dentition, upper limb
long bones, complete hand, pel-
vis fragments, lower limb long
bones, partial foot
Homo habilis sensu lato vs. Aus- 100 Cranium, mandible, dentition, Yes Yes No High
tralopithecus sediba hand phalanges, tarsals, meta-
tarsals, and other fragmented
postcranial remains including
upper and lower long bone
shafts vs. Cranium, mandible,
dentition, upper limb long bones,
complete hand, pelvis fragments,
lower limb long bones, partial
foot
Homo habilis sensu stricto vs. See above See above See above See above See above See above
Homo rudolfensis
65

Yearbook of Physical Anthropology

 APPENDIX B Continued
66
Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
b
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatric differentiation
1.5 Ma–1.0 Ma Homo erectus vs. Paranthropus See above See above See above See above See above See above
robustus
Homo erectus vs. Paranthropus See above See above See above See above See above See above
boisei
Homo erectus vs. Homo habilis See above See above See above See above See above See above
sensu lato
Homo erectus vs. Homo ergaster See above See above See above See above See above See above
Homo erectus vs. Homo antecessor 100 Nearly the entire skeleton except Yes Yes No Low
for most hand and foot bones vs.

Yearbook of Physical Anthropology

Cranium, mandible, dentition,
upper limb long bones, clavicle,
ribs, carpals, metacarpals, hand
phalanges, tarsals, metatarsals,
foot phalanges
Paranthropus robustus vs. Para- See above See above See above See above See above See above
nthropus boisei
Paranthropus robustus vs. Homo See above See above See above See above See above See above
habilis sensu lato
Paranthropus robustus vs. Homo 100 Cranium, mandible, dentition, No No No Highc
antecessorc hand bones, vertebrae, femur,
talus, and other fragmentary
postcranial remains vs. Cra-
nium, mandible, dentition, upper
limb long bones, clavicle, ribs,
carpals, metacarpals, hand pha-
langes, tarsals, metatarsals, foot
B. WOOD AND E. BOYLE

phalanges
Paranthropus boisei vs. Homo 100 Cranium, mandible, dentition, dis- No No No Highc
antecessorc tal humerus, radius, femur, and
tibia fragment vs. Cranium,
mandible, dentition, upper limb
long bones, clavicle, ribs, car-
pals, metacarpals, hand pha-
langes, tarsals, metatarsals, foot
phalanges
Homo habilis sensu lato vs. Homo 100 Cranium, mandible, dentition, Yes Yes No Highc
antecessorc upper limb long bones, complete
hand, pelvis fragments, lower
limb long bones, partial foot vs.
Cranium, mandible, dentition,
upper limb long bones, clavicle,
ribs, carpals, metacarpals, hand
phalanges, tarsals, metatarsals,
foot phalanges
1.0 Ma20.25 Ma Homo erectus vs. Homo 100 Nearly the entire skeleton except Yes Yes Yes High
heidelbergensis for most hand and foot bones vs.
Cranium, mandible, dentition
and some isolated postcranial
evidence
 APPENDIX B Continued

Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatricb differentiation
0
Homo erectus vs. Homo 10 Nearly the entire skeleton except Yes Yes Yes High
rhodesiensis for most hand and foot bones vs.
Cranium, mandible, dentition,
pelvis, sacrum, humerus, femur,
tibia
Homo erectus vs. Homo helmei 101 Nearly the entire skeleton except Yes Yes Yes High
for most hand and foot bones vs.
Cranium, mandible, dentition,
fragmentary postcrania
Homo erectus vs. Sima de los 100 Nearly the entire skeleton except Yes Yes Potentially High
Huesos for most hand and foot bones vs.
Nearly the entire skeleton
except for some axial skeleton
and upper limb bones, DNA
Homo rhodesiensis vs. Sima de los 102 Cranium, mandible, dentition, pel- Yes Yes Yes High
Huesos vis, sacrum, humerus, femur,
tibia vs. Nearly the entire skele-
ton except for some axial skele-
ton and upper limb bones, DNA
Homo helmei vs. Sima de los 102 Cranium, mandible, dentition, Yes Yes Yes High
Huesos fragmentary postcrania vs.
Nearly the entire skeleton
except for some axial skeleton
and upper limb bones, DNA
Homo heidelbergensis vs. Homo 100 Cranium, mandible, dentition and Yes Yes Potentially Low
rhodesiensis some isolated postcranial evi-
dence vs. Cranium, mandible,
HOMININ TAXIC DIVERSITY

dentition, pelvis, sacrum,

humerus, femur, tibia
Homo heidelbergensis vs. Homo 100 Cranium, mandible, dentition and Yes Yes Potentially Low
helmei some isolated postcranial evi-
dence vs. Cranium, mandible,
dentition, fragmentary
postcrania
Homo heidelbergensis vs. Sima de 102 Cranium, mandible, dentition and Yes Yes No Low
los Huesos some isolated postcranial evi-
dence vs. Nearly the entire skel-
eton except for some axial
skeleton and upper limb bones,
DNA
0.25 Ma–present Homo sapiens vs. Homo 100 Entire skeleton, DNA vs. Entire Yes Yes Yes High
neanderthalensis skeleton, DNA
Homo sapiens vs. Homo erectus 102 Entire skeleton, DNA vs. Nearly Yes Yes Yes High
the entire skeleton except for
most hand and foot bones
67

Yearbook of Physical Anthropology

 68

APPENDIX B Continued

Difference in
abundance
of material Qualitatively Quantitatively Confidence
(orders of compared in compared in in species
Time interval Species etc., compared magnitude)a Nature of the evidence the literature the literature Sympatricb differentiation
c
0.25 Ma–present Homo sapiens vs. Homo 10 Entire skeleton, DNA vs. Yes Yes Yes High
heidelbergensis Cranium, mandible, dentition
and some isolated postcranial
evidence
Homo sapiens vs. Homoi 102 Entire skeleton, DNA vs. Nearly Yes Yes Yes High

Yearbook of Physical Anthropology

floresiensis the entire skeleton except for
some axial skeleton, upper limb,
hand and tarsal bones
Homo sapiens vs. Denisovans 10c Entire skeleton, DNA vs. Isolated Yes Yes Yes High
teeth, hand phalanx, DNA
Homo neanderthalensis vs. Homo 101 Entire skeleton, DNA vs. Nearly Yes Yes Potentially High
erectus the entire skeleton except for
most hand and foot bones
Homo neanderthalensis vs. Homo 102 Entire skeleton, DNA vs. Cranium, Yes Yes Yes Low
heidelbergensis mandible, dentition and some
isolated postcranial evidence
Homo neanderthalensis vs. Homo 10c Entire skeleton, DNA vs. Nearly Yes Yes No High
floresiensis the entire skeleton except for
some axial skeleton, upper limb,
hand and tarsal bones
Homo neanderthalensis vs. 10c Entire skeleton, DNA vs. Isolated Yes Yes No Moderate
Denisovans teeth, hand phalanx, DNA
Homo erectus vs. Denisovans 102 Nearly the entire skeleton except Yes Yes Yes Moderate
B. WOOD AND E. BOYLE

for most hand and foot bones vs.

Isolated teeth, hand phalanx,
DNA
Homo heidelbergensis vs. Homo 100 Cranium, mandible, dentition and Yes Yes No High
floresiensis some isolated postcranial evi-
dence vs. Nearly the entire skel-
eton except for some axial
skeleton, upper limb, hand and
tarsal bones
Homo heidelbergensis vs. 101 Cranium, mandible, dentition and Yes Yes No Low
Denisovans some isolated postcranial evi-
dence vs. Isolated teeth, hand
phalanx, DNA
a
Approximate orders of difference in the abundance of specimens in the two hypodigms. 100 5 less than a ten-fold difference; 101 5 ten-fold difference or more; 102 5 hundred-fold
difference or more; 103 5 thousand fold difference or more.
b
All assessments are based on the current fossil evidence. “Potentially” indicates that the species could be sympatric depending on how the fossil evidence is interpreted.
c
If the partial mandible (ATE9-1) from the Sima del Elefante does not belong to H. antecessor then these “high confidence” pairwise comparisons would fall away.
 HOMININ TAXIC DIVERSITY 69
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