Evolutionary Anthropology 1
NEWS
Systematics of “Humankind”
A
n international colloquium,
“Taxonomy and Systematics of
the Human Clade,” was held
February 6 –9, 2000, in the tranquil
surroundings of Palma de Mallorca,
Spain. This colloquium was conceived
and orchestrated as a workshop with
round-table discussions by the philos-
opher of science Camilo Cela-Conde
of La Universidad de las Islas
Baleares, who strove to collect, in a
locale unassociated with the fossilized
trappings of human evolution, a vari-
ety of individuals of different back-
grounds and perspectives. Cela-
Conde’s goal was to see if such a
diverse group of variably opinionated
researchers could hammer out the ba-
sis for a consensus on the matter of
what constitutes the human clade.
Following a format that differed
from the typical meeting schedule of
presentations, Cela-Conde organized
the colloquium around a series of
half-hour “target” papers and ten-
minute contributions. The longer pre-
sentations were meant literally to be
targets for debate, comment, and re-
buttal, and were expected to raise for
discussion a broader suite of poten-
tially relevant issues. The shorter pre-
sentations helped focus the group on
specific issues within the larger frame-
work and kept discussions directed to-
ward the intended goal of the collo-
quium.
Although they did not give formal
presentations, Francisco Ayala (Uni-
versity of California, Irvine), F. Clark
Howell (University of California,
Berkeley), Phillip V. Tobias (Univer-
sity of the Witwatersrand), Daniel
Turbón (Universidad de Barcelona),
Alejandro Pérez-Pérez (representing
the International Association of Hu-
man Paleontology), and Jordi Serra-
Evolutionary Anthropology 10:1–3 (2001)
llonga (coordinator of Hominids,
Spanish Association for the Study for
Hominid Evolution) were prominent
in debate and discussion throughout
the scheduled sessions of the collo-
quium. All in attendance were actively
involved in the final hours, when Cela-
Conde challenged the assembled
group to reach a meaningful and ac-
ceptable consensus on what consti-
tutes the human clade.
After the opening comments and
greetings, Jeffrey H. Schwartz (Uni-
versity of Pittsburgh) gave the first of
the target papers (“The Relevance of
Fossils and Other Considerations to
the Concept and Representation of
Hominoidea”), in which he reviewed
the parallel histories of the study of
fossil “hominoids” versus “hominids”
as a way of illustrating that any deci-
sion regarding taxonomic inclusivity
or exclusivity must take into account
fossil as well as extant taxa, and the
picture of diversity that they repre-
sent. Schwartz also argued that sys-
tematists must try to integrate in-
sights from developmental genetics
into their approaches to phylogenetic
reconstruction, recognizing, for ex-
ample, that clues to phylogenetic re-
latedness may be more appropriately
appreciated at the level of regulatory
homology, with more closely related
taxa being “similar” through the re-
cruitment in similar signal transduc-
tion pathways of similar regulatory
molecules.
Morris Goodman (Wayne State Uni-
versity) then followed with the latest
version of his and his colleagues’ molec-
ular reconstruction of the relationships
of extant primates (“An Objective Time-
Based Phylogenetic Classification of
Primates That Places Chimpanzees and
Humans in the Genus Homo”), which
represents decades of laborious and
dedicated research. Harking back to a
position he developed in the 1960s,
Goodman made his strongest plea yet
for equating overall molecular similar-
ity with phylogenetic propinquity. He
argued that, especially in the case of the
resultant hierarchical arrangement of
the large-bodied hominoids, this should
be directly translated into a classifica-
tion wherein Pan, if not also Gorilla, is
subsumed in the genus Homo. Conse-
quently, all fossil taxa would have to be
accommodated within this framework:
one genus, a plethora of species.
The two diametrically opposed posi-
tions represented initially by Goodman
and Schwartz continued as major
themes throughout the colloquium:
How much or how little diversity
should be reflected in perceptions of,
and operational procedures regarding,
the systematics and classification of
Homo sapiens and its relatives?
Thus, Bernard Wood (George Wash-
ington University) summarized his re-
cent efforts with Mark Collard (Univer-
sity College, London) (“The Human
Genus”) in rethinking the stranglehold
history has had on the genus Homo,
which has a priori dictated just how
much (actually very little) taxonomic
diversity should be “allowed” to exist.
Their argument was that the genus
Homo should be recognized, and poten-
tial species allocated to it, on the basis
of an identifiably distinctive shift in
adaptive strategies and associated mor-
phological and behavioral novelties.
For Wood and Collard, the final cut for
inclusion in the genus was with H. er-
gaster (that is, “African H. erectus”). By
implication, more genera would be
needed to receive the taxonomically or-
phaned non-Australopithecus species.
In sharp contrast, Milford Wolpoff
(University of Michigan) presented his
interpretation of the human fossil
record (“Is There a Phylogeny of
Homo?”), in which he has found little
convincing evidence for diversity within
the genus Homo: variation within a spe-
cies, yes, but diversity among an array
of species, no. From the perspective
2 Evolutionary Anthropology
that the presence of similar features in
chronologically successive specimens
of fossil hominid indicates a continuity
that is artificially segmented by im-
posed species names, Wolpoff argued
in favor of recognizing H. sapiens at
least as far back as the middle Pleisto-
cene.
Using a different approach to the
issue of conceptualizing and opera-
tionalizing “the human clade,” Colin
Groves (Australian National Univer-
sity) (“Morphological, Molecular; Cla-
distics; Unranked Taxonomies; Link-
ing Time to Taxonomic Rank”) looked
at the human fossil record in terms of
how many genera and species have
typically been recognized in other
mammalian groups. Having earlier
been a co-author with Goodman in
promoting the idea that at least Pan
should be subsumed in the genus
Homo, Groves here argued that the
relatively short time depth of the hu-
man fossil record is consistent with
classifications of other taxa in which
the genus and some number of species
were sufficient to accommodate the
lot. As Goodman had proposed in his
presentation, so too did Groves offer
suggestions for inserting below-ge-
nus-level taxonomic categories in rec-
ognition of some degree of diversity
represented in the fossil record.
The shorter contributions were
thoughtfully interspersed between the
target papers. David S. Strait (New
York College of Osteopathic Medi-
cine) presented the cladistic analysis
he had carried out with Frederick E.
Grine (SUNY, Stony Brook) concern-
ing the question, “Is Australopithecus
garhi the First Human?” From among
the array of equally defensible phylo-
genetic hypotheses thus produced
came the answer: no. In the only mor-
phometrically oriented presentation,
Leslie Aiello (University College, Lon-
don) pointed to pitfalls in “Recogniz-
ing Species and Species Diversity in
the Fossil Record,” demonstrating
how the choice and manipulation of
postcranial morphometric characters
could yield totally different interpreta-
tions and blur the distinction between
variation and diversity. Shifting the
focus to a different philosophical
realm in “Classification and Phylog-
eny in Human Evolution,” Ian Tatter-
sall (American Museum of Natural
History) reminded the participants
that classification and phylogeny are
not the same in either intellectual pur-
suit or consequence. He pointed to
problems inherent in the particular
history of human evolutionary stud-
ies, which has emphasized the search
for continuity of variation from the
present into the past and the con-
straining of the picture of human evo-
lution by the classifications of Mayr
and Dobzhansky. On the matter of the
philosophical and methodological as-
pects of phylogenetic reconstruction
and the reconstruction of evolution-
ary events, Frederick S. Szalay
(CUNY, Hunter College) discussed
“The Issues of Operational Taxonomic
Units (OTUs), the Morphological Spe-
cies Concept, and the Tests for Specia-
tion in the Fossil Record.” Drawing on
his recent studies on marsupial post-
cranial comparative anatomy, Szalay
discussed the degrees to which these
pursuits were viable and results
thereby attainable. As the titles of
these presentations clearly indicate,
the contributions importantly punctu-
ated sometimes lengthy discussion
with necessary fresh air and focus.
The final target paper, by Camilo
Cela-Conde (“Can a Cladogram Be
Falsified?”), sought to bring the dis-
cussion into the realm of practical and
theoretical systematics. After briefly
reviewing the historical and philo-
sophical aspects of the debates on fal-
sification, Cela-Conde reminded the
participants of the inherently “fragile”
nature of phylogenetic reconstruction
and the problems in delineating
clades, as well as the need to bear
these matters in mind when extrapo-
lating from an hypothesized cla-
dogram to arenas of greater specula-
tion, such as scenarios of phylogenetic
history, or greater constraint, such as
classifications. The final short contri-
bution, by Emiliano Aguirre (Muséo
de Ciencias Naturales, Madrid), “Cri-
teria to Apply Taxonomic Categories
to Human Fossils,” brought the wis-
dom and experience of a seasoned pa-
leontologist to bear not only on the
specific topics discussed by individual
participants, but also on the matter
that still stood before the participants:
attempting to reach a consensus on
the question of what constitutes the
human clade.
NEWS
In an unparalleled act of rising
above the details of individual prefer-
ences, and within the allotted number
of hours, this assembled group con-
structed a proposal that all hoped will
serve as a starting point for more effi-
cient and mutually understandable
discourse among the incredibly di-
verse lot of students involved in re-
search on human evolution. The text
of the statement is offered here, not as
the final word on the subject, but as
the beginning of international collab-
oration on the phylogenetic history
and classification of our own group.
General Considerations
1. Humans have evolved from non-
human ancestors and continue to
evolve. Since their emergence from
their ape ancestors, humans diversi-
fied into different lineages, of which
only one has survived to the present
day. All evidence indicates that the hu-
man species is not evolving toward
increasingly diverse groups, but
rather the opposite.
2. Current human beings, their an-
cestors not shared with any extant
nonhuman species, and the collateral
descendants of these ancestors, will
be henceforward referred to as the hu-
man “clade.”
3. Scholars engaged in studying the
human clade use a variety of ap-
proaches and methods, all of which
have a place in human systematics.
Different methods may yield different
outcomes concerning human system-
atics. Principles and methods origi-
nally devised for classifying living
forms often encounter difficulties
when applied to fossils.
4. Organisms were originally classi-
fied on the basis of morphology (in-
cluding ontogeny). Molecular, biogeo-
graphical, behavioral, and ecological
approaches have added significant
new lines of evidence. Morphology re-
tains a preeminent role in systematics,
particularly so in the study of fossils.
The time dimension is notably signif-
icant in the assessment of phyloge-
netic relationships.
5. Phyletics, phenetics, and cladis-
tics are all used in systematics. Taxo-
nomic classification should conform,
to the extent possible, to phylogenetic
relationships. Since the second half of
the twentieth century, cladistics has
NEWS
made particularly valuable contribu-
tions in the field of classification, but
we note that cladistics identifies sister
groups, not ancestor/descendant rela-
tionships.
6. Molecular considerations are
valuable in determining relationships
between lineages that have extant de-
scendants. In the case of fossil lin-
eages having no extant descendents,
the application of molecular studies is
limited at present, but advancing
techniques may reduce this limita-
tion.
Classificatory Principles
7. Taxonomic practice should be
consistent with the principle of mono-
phyly. On this ground, Simpson’s tra-
ditional concept of a family Pongidae
should be rejected because it em-
braces paraphyly.
The family Hominidae has been
widely used as the taxon assigned to
the human clade. It is advisable to
Fossils, Footprints, and Foragers:
Bipedalism Evolving
The origin of bipedalism in early
hominids is a focus of paleoanthropo-
logical research to such an extent that
the evolution of bipedalism after that
point is often disregarded. At the 2000
annual meeting of the American Asso-
ciation of Physical Anthropologists,
Jeff Meldrum (Idaho State University)
and Chuck Hilton (University of New
Mexico) presented a symposium of re-
search aimed at remedying that.
David Begun (Toronto) began the
symposium with a perspective on the
implications of recent phylogenetic
analyses with respect to the precur-
sors of human bipedalism. Given that
genetic data support a chimp-human
clade, knuckle-walking becomes ei-
ther a primitive state for African apes
and humans or else evolved indepen-
dently in gorillas and chimps. Begun
Evolutionary Anthropology 10:3– 4 (2001)
treat taxonomic rankings conserva-
tively.
Several alternative ranks have been
proposed for the human clade, such
as subfamily (Homininae), tribe
(Hominini), subtribe (Hominina), ge-
nus (Homo), or subgenus (H. Homo).
None of them has achieved a substan-
tial degree of consensus among the
world community of scholars, though
the use of tribe has received increas-
ing support.
We recommend that active steps be
taken to elicit a consensus viewpoint
on this subject. To this end, we pro-
pose that a Workshop or Study Group
be convened to meet at the V Interna-
tional Congress of Human Paleontol-
ogy, to be held in Barcelona (Spain) in
2003.
8. The transfer of a species taxon
from one superspecific taxon to an-
other should be avoided if it increases
paraphyly in the recipient taxon.
There are practical difficulties in fol-
analyzed the carpal anatomy of homi-
noids and hominids, and demon-
strated a series of characters shared
between modern humans and African
apes. These characters imply either
significant homoplasy in the wrist of
the chimp-human clade or a period of
knuckle-walking in hominin ancestry.
Begun reasoned that humans most
likely evolved from a knuckle-walking
ancestor.
Ron Clarke (Germany and South
Africa) described the new 3.3 million
year old Australopithecus skeleton
(StW 573) recovered from Sterkfon-
tein. The foot skeleton reported five
years ago has now been supplemented
with additional elements, which con-
firm an ape-like architecture. Clarke
recovered a complete distal upper
limb, including a hand skeleton for
StW 573, which seems to display no
derived traits for knuckle-walking.
Clarke maintains that hominids retain
a primitive upper limb design that is
consistent with a locomotor ancestry
Evolutionary Anthropology 3
lowing this principle. We recommend
that the above mentioned Workshop
address this issue.
9. No consensual means of identi-
fying biological species in the fossil
record exists at the present time.
Jeffrey H. Schwartz
Departments of Anthropology and History
and Philosophy of Science
University of Pittsburgh
Pittsburgh, PA 15260 (USA)
jhs⫹@pitt.edu
Mark Collard
Department of Anthropology and
The AHRB Centre for the Evolutionary
Analysis of Culture,
University College London,
Gower Street,
London WC1E 6BT (UK)
m.collard@ucl.ac.uk
Camilo J. Cela-Conde
Departamento de Filosofiá
Universidad de las Islas Baleares
07071 Palma de Mallorca (Spain)
cela@atlas-iap.es
© 2001 Wiley-Liss, Inc.
based on an upright posture and fore-
limb-assisted locomotion in the trees.
Peter Schmid (Switzerland) re-
cently reanalyzed the actual Laetoli
footprints. This experience convinced
him that the casts of these footprints
can be misleading. He observed key
modern features in the footprints,
such as a medial longitudinal arch,
but found no evidence in any of the
prints of abducted big toes. He en-
couraged us to remember the great
range of variation in gait patterns in
humans and apes when considering
the very few available ancient impres-
sions.
Jeff Meldrum reported on modern
human footprints preserved in the
Kilauea volcanic ash in Hawaii, which
may be the best possible database for
studying the effects of ash on human
foot impressions. Compared to the
Hawaiian footprints, the Laetoli casts
longer lateral toe impressions, lack of
a prominent ball impression, and am-
biguous evidence of a medial longitu-
4 Evolutionary Anthropology
dinal arch. Meldrum reiterated his
proposal that the australopithecine foot
retained a degree of midtarsal flexibil-
ity and even suggested that this archi-
tecture persisted in Homo ergaster.
Roshna Wunderlich (James Madi-
son University) and Dan Schmitt
(Duke University) detailed hindlimb
anatomical adaptations to the unique
nature of heel-strike plantigrady in-
voked by apes and humans walking on
the ground, testing the hypotheses
that a heel-striking quadrupedal ape
should have flexed hips, extended
knees, and protracted hindlimbs.
These behavioral traits during terres-
trial bipedalism in chimpanzees result
from a kinematic pattern expected in
an animal with a hindlimb adapted to
climbing on vertical supports.
Patricia Kramer (University of Wash-
ington) approached bipedalism from a
life-history and ecological perspective.
Trade-offs among walking velocity,
range, and energetic efficiency may
have shifted during the evolution of bi-
pedalism from australopithecines to
modern humans. For example, the high
carrying loads common to modern
humans would have compromised the
energetic efficiency of early hominid lo-
comotion. Moreover, the differing ca-
pacities of male and female anatomies
for parceling out these trade-offs must
be accounted for in models of how bi-
pedalism evolves. Kramer’s work sug-
gests that changes in the locomotor
niche between modern humans and
early hominids involved increased ca-
pacities for faster speeds or longer dis-
tances, heavier carrying burdens for fe-
males, and better energetic efficiency in
males at high speeds.
D. R. Carrier (University of Utah)
modeled hypotheses of intrasexual
competition and specializations for
locomotor speed that may have influ-
enced selection on forelimbs versus
hindlimbs in the evolution of Homo.
In the hindlimb, Homo shows changes
toward improved speed and efficiency
coincident with a lack of anatomy for
competition or aggression. By analogy
to hoofed mammals that have adapted
a “new” weapon system that preserves
their need for locomotor speed, Car-
rier hypothesized that the advent of
nonbiological weaponry by humans
uncoupled adaptations for cursorial
specialization and competition in the
locomotor skeleton.
Chris Ruff (Johns Hopkins) con-
NEWS
cluded the symposium by reviewing
themes relating to how bipedalism is
manifest in the human fossil record.
He stressed the anatomical mosaicism
apparent in early hominid skeletons
and reminded the audience to study
closely the relationship between ge-
netically determined versus environ-
mentally plastic traits in the locomo-
tor skeleton. The Laetoli footprints,
hominid fossils, and laboratory stud-
ies of ape locomotion will always be
part of the investigative landscape,
but should not be studied in isolation
or outside of a paradigm that consid-
ers the effects of nonlocomotor behav-
iors on locomotor anatomy and mo-
bility strategies. These integrated
approaches do not necessarily com-
promise the rigor of formalized, cla-
distic-based, analytical methods, but
rather require increased collaboration
and creativity in experimental de-
signs.
Walter Carl Hartwig
Department of Basic Sciences
832 Walnut Avenue
Touro University College of
Osteopathic Medicine
Mare Island, CA 94592
© 2001 Wiley-Liss, Inc.