Vous êtes sur la page 1sur 15

Anthropological Review • Vol.

72, 66–80 (2009)

Two interpretations of human evolution:


Essentialism and Darwinism

Maciej Henneberg
Biological Anthropology and Comparative Anatomy Research Unit, University of Adelaide,
Adelaide 5005, and Department of Archaeology, Flinders University Australia;
E-mail: Maciej.Henneberg@adelaide.edu.au

ABSTRACT Despite intensive studies of a large number of fossils discovered during the
20th century there is no consensus as to the interpretation of the process of hominin evolution.
Some authors see as many as six genera and some 17 species, while others argue for a single
lineage from Plio/Pleistocene until today. Such diversity of interpretations of the same facts
indicates lack of a uniform theoretical basis underlying studies of human evolution. Debates
can be resolved using basic principles of scientific inquiry – parsimony and falsification of null
hypotheses. Hypothesis testing is now possible with respect to the evolution of basic hominin
characteristics such as brain size, body size and the size of the dentition that have sample sizes
of a few hundred individual data points each. These characters display a continuous change
with time. Analyses of variance do not falsify the null hypothesis of the existence of only one
species at any time – variances around regression lines on time do not differ from the variance
observed in the single species of Homo sapiens – distributions of residuals are normal. Thus,
splitting of the hominin lineage into coeval species can only be based on descriptive character-
istics that are liable to errors of subjective judgment.
KEY WORDS: hominin, hominid, Australopithecus, brain, stature

Understanding of the evolutionary pro- how our major traits – such as mental ca-
cess that produced modern humans is of pacities and erect bipedal locomotion –
crucial importance for the way people emerged are still ongoing [Carruthers and
perceive themselves and their lives. It in- Chamberlain 2000, Cela-Conde and Ayala
forms ideologies underlying political and 2007]. The number of validly named hu-
economic decisions. Thus it is important man species by the end of the 20th century
to have this process documented and inter- exceeded 50 [Meikle and Parker 1994,
preted in accordance with the best rules of Henneberg 1997], while it has been pro-
scientific practice. Despite over 150 years posed that an even larger number should
of studies of human evolution, debates on be identified [Tattersall 1987] and actually
Received 10 June; accepted 8 December 2009
DOI: 10.2478/v10044-008-0016-2
© 2009 Polish Anthropological Society
Two interpretations of human evolution 67

has recently [Brown et al. 2004]. It does the ideal pattern of the world of living
not add, though, to the clarity of the pic- things, the “pattern of creation”. Similari-
ture of our origins. ties among individuals were used to distil
Over the last half-century several au- essences of ideal types of their respective
thors proposed that the parsimony and fal- kinds and those types were then organized
sification of hypotheses should be applied into nested hierarchies of species, genera,
to studies of human evolution postulating families, classes, orders, phyla, etc. [Lin-
a “single species hypothesis” [Brace 1967; naeus 1758]. Thus, conceptually, the small-
Wolpoff 1968, 1971]. According to Hunt est unit of life was a species, not an indi-
[2003], Frank Livingstone proclaimed vidual. An individual was but an example
himself to be the proponent of single spe- of an ideal species type. Individual varia-
cies hypothesis still at the beginning of the tion was consciously ignored for the sake
21st century. Kevin Hunt himself argues of clarity. Once described, taxa became
strongly in support of the single species hy- units of study for biologists. Discussions
pothesis using data on taxonomic diversity about the complexity of life in a particular
of large-bodied genera of mammals. Re- region or the entire world were based on the
cently Holliday [2003] proposed an inter- number of species and higher taxa present
mediate interpretation of hominid diversity. in various environments, the interactions
He used the notion of syngameon which is still being studied as those between par-
a network of separate cohesive taxa that ex- ticular taxa, while descriptions of evolution
change genes among themselves. He sup- consist of biblical lists of who is a descend-
ported this interpretation with observations ant of whom and what the family tree looks
of gene flow among several species of the like [Henry and Wood 2007, Stanford et al.
two genera of baboons – Theropithecus 2009].
and Papio – [Jolly 1993, 2001; Jolly et al. As an example, following a suggestion
1997]. Quintyn [2009] summarized current by Robert Eckhardt:
discussions concerning hominin diversity And unto Enoch was born Irad: and Irad be-
and concluded that a temporary ban on cre- gat Mehujael: and Mehujael begat Methush-
ating new hominin taxa should be imposed ael; and Methushael begat Lamech [Genesis
until a clearer understanding of hominin 4:18, 1611].
variation can be reached. Homo erctus is descended from Homo habi-
lis, which in turn descended from Australo-
pithecus garhi … [Larsen 2008].
Taxonomy, essentialism and the
concept of species Figure 1 is an example of the classic rep-
resentation of a hominin family tree. Such
To make sense of the variety of forms representation may be as well reflecting
that surround us we must be able to reduce a pattern of creation. No explanations of
it to a manageable number of generalizing how and why particular branches arose are
categories that can then be used in various given. No uncertainty as to possible mix-
thought processes. Thus, the human ten- ing of categories is indicated, nor is there
dency to categorize: From the time platonic uncertainty indicated in separating some
idealism arose in classical antiquity to the categories from others.
early enlightenment, the task of natural- Taxa are perceived as real entities and
ists was perceived as that of discovering they can only have two states: existence or
68 M. Henneberg

Australopithecus afarensis
Australopithecus aethiopicus
Australopithecus boisei
Australopithecus robustus
Australopithecus africanus
Homo habilis
Homo erectus
“Archaic” H. sapiens
Neandertals
Homo sapiens sapiens

Fig. 1. Scheme originally published on page xxix of the Encyclopedia of Human Evolution and Prehisto-
ry [Tattersall et al. 1988] and repeated with modification in the second edition [Delson et al. 2000], author’s
own drawing. This is not evolution, this is systematics. It may as well be a “pattern of creation”.

extinction; categorically yes or no. Such [Lee 2003] in the face of the overwhelming
an approach, though producing voluminous understanding that life is variable and no
descriptions, cannot provide explanations of stable categories actually exist. Following
the process of evolution since evolutionary Lee [2003], current concepts of species can
mechanisms are based on individual varia- be subdivided into (1) similarity concepts
tion and transmission of this variation from using phenetic resemblance of organisms as
one generation of individuals to the next. the sole criterion, (2) cohesion concepts in
It can be argued that before we can analyze which organisms are purported to share fea-
anything and search for processual explana- tures allowing them to remain coherent from
tions, we must first provide a solid descrip- generation to generation, (3) monophyly
tion. Such description, however, must be in concepts stressing membership of a single
terms that lend themselves to further pro- nonreticulating lineage, and (4) interbreed-
ductive analysis. In biology, the use of ar- ing concepts in which members of a species
bitrarily and sharply delimited categories exchange genes among themselves while
is not conducive to productive analysis. It being reproductively isolated from others.
leads rather to “catastrophic” explanations Concepts belonging to the first three groups
in the style of George Cuvier [Brace 1981]. are valid for any taxonomic rank, not be-
Taxonomic descriptions may be useful in ing specific to species. Depending on how
certain applications, such as ecology, but strictly one defines phenetic similarity, level
the fact that they ignore individuals and the of cohesion or monophyletic inclusiveness,
flow of time from generation to generation one can define supraspecific taxa (like ge-
is always a limiting factor. nus) or infraspecific groupings (like races).
Currently there exist some 23 definitions The interbreeding concepts emphasize that
of species [Mayden 1997, Quintyn 2009]. actual exchange of genetic material oc-
They all attempt to “salvage a Linnean rank” curs only within a species [Mayr 1969,
Two interpretations of human evolution 69

1995], a phenomenon which is unique to whether a Neandertal man and a modern


this taxonomic rank and making it the only woman would produce fertile offspring.
“real” supraindividual biological unit. For a test falsifying a hypothesis of their
This is the unit of evolution because it is conspecificity we must rely on secondary
in the process of differential reproduction inferences derived from comparisons of
of populations isolated genetically from morphology and partly reconstructed DNA
other populations that the adaptive change sequences.
in gene pool can occur. The phylogenetic
species concept stresses the same phenom- Individual variation and
enon of reproductive continuity, but from Darwinian view of life
a point of view of change through time
– all descendants of a particular ances- In reality, only individuals exist at a par-
tor are a taxon [Velasco 2009]. A specific ticular point in time. Some of them are
variety of this approach is the genomic- more similar to each other than to many
phylogenetic species concept [Staley other individuals. Such similar individu-
2009] inferring evolution of an organism als can be grouped together into a similum
from sequences of its genes. [Henneberg and Brush 1994, Henneberg
In all definitions, however, boundaries 1997] without any prejudice as to their for-
of species are imprecise because reproduc- mal taxonomic status. Simila are defined
tive isolation is often incomplete with in- solely by mutual similarities of individuals
ter-species gene flow occurring at various to each other, not by a similarity of indi-
rates, while it takes a minimum of three- viduals to some idealized, immutable type.
generations for a monophyletic lineage Simila are thus capable of changing con-
(phylogenetic species) to separate from tinuously through time, while taxa are not
others [Samadi and Barberousse 2009]. In because their change requires abrupt ap-
this situation, an audacious solution to the pearance of a new ideal type. In most cases
removal of the ambiguity inherent in the of metazoans, observation of individuals
biological species concept has been pro- over a period of their lives reveals that they
posed. Gonzalez-Forero [2009] postulates mate with other members of the similum
that in situations of relaxed reproductive to produce new individuals who undergo
isolation, some populations should be con- a process of ontogenetic development and
sidered members of more than one species. growth. During this process individuals
This should be reflected in their trinomial change their physical form, physiology and
nomenclature. An example relating to ho- behavior, sometimes to a very large extent.
minins would be Homo neanderthalensis- Depending on their sex, individuals may
sapiens. differ so substantially in their characteris-
Despite a multitude of theoretical means tics that they may be mistaken for different
of reconciling a descriptive platonic cat- species [Manning and Dawkins 1998]. In
egory with the current understanding of extant organisms it is easy to study ontoge-
biology, when dealing with fossil material, netic processes and assess sexual dimor-
only morphological similarity, and mor- phism, although it may happen that various
phological variation, are available for study ontogenetic stages, or different sexes, may
and must be used for testing taxonomic hy- mistakenly be described as separate species
potheses. It is impossible to test directly [Eckhardt 2000]. Even when ontogenetic
70 M. Henneberg

and sexual variations are known, there re- have a range of variation so wide as to be
mains substantial interindividual variation almost worthless for any diagnoses. Over
due to both genetic polymorphism and vari- centuries, the variation of brain size is fur-
ous responses to environmental stimuli dur- ther added to by microevolutionary trends.
ing ontogeny. This variation can hardly be During the Holocene Homo sapiens cranial
categorized. capacity decreased in size by about 10%
Brain size, especially when related to body from the beginning of this period to its end
size, is amongst the most prominent defin- [Henneberg 1988, 1998] while in the Anthro-
ing characteristics of Homo sapiens. Yet its pocene (considered to start in the late 18th
variation is very substantial – CV approach- century), due to general secular increase in
es 12% [Henneberg 1990] compared with body size, it increased again by about 6%
about 4% for body height and 10% for arm in some countries [Henneberg et al. 1985].
circumference (approximate values calcu- Such changes would be undetectable during
lated from Frisancho [1990] data of anthro- the Plio/Pleistocene due to technical errors
pometric standards). In absolute figures, the of dating methods that may extend to a few
range for modern human cranial capacity ex- thousand years [Eckhardt 2000].
tends from 3 standard deviations (SD) below In paleoanthropology, when individual
the mean at 879 ml to 3 SD above the mean fossils are found, in most instances it is dif-
at 1821 ml [Henneberg 1990]. This is more ficult to be sure of their sex, and their devel-
than double the minimum value. Amongst opmental age is sometimes debatable (see
adults only about one quarter of this variation uncertainties in the developmental ages of
is explained by sexual dimorphism, another the Taung Child and Nariokotome Boy,
quarter or so can be attributed to differences Lacruz et al. [2005], Walker and Leakey
among populations, while the remaining 50 [1993]) and their membership of a partic-
percent occurs among same-sex individu- ular local population, as opposed to some
als in the same population (Fig. 2, Henne- other one, unknown. Dating methods of
berg [1990]). Were we to add variation re- Pliocene and Pleistocene fossils have wide
sulting from ontogenetic doubling in brain error ranges, easily incorporating several
size, what seems to be an obvious species- hundreds of years during which microevo-
specific “big brain” characteristic would lutionary changes could occur [Eckhardt
2000]. The situation is further complicated
by the fragmentary nature of fossils. In this
situation, researchers attempt to perform
a multivariate assessment based on all traits
that can be estimated from a given fossil. It
usually takes form of a descriptive and cat-
egorical, rather than metrical, analysis and
leads to attributing a fossil to an idealized
taxon, usually a species. When the multi-
variate “appearance” of a particular fossil
is somewhat different from previously de-
scribed fossils it is left to an arbitrary deci-
Fig. 2. Components of variance in the human
sion of researchers whether to assign it to
brain size. one of the already established “species” or
Two interpretations of human evolution 71

to a new one. The amount of variation that not explain processes of evolution but pro-
can be tolerated within one species is still vides only a static description of its course.
debated [Cope and Lacy 1992, 1996; Al- A phylogeny is only as good a description of
brecht and Miller 1993; Kelley 1993; Plav- the course of evolution as are the definitions
can 1993; Rose and Bown 1993; Plavcan of species and other taxa that are included
and Cope 2001; Ackermann et al. 2006]. with it. Were the taxa incorrectly identified,
Uncertainties abound, especially because or incorrectly characterized, the descrip-
there are so many different definitions of tion of the process of evolution would be
species. The Latin word specere means faulty. Thus, any attempts at explanations
simply “a kind” or “a form”, thus anything of evolutionary processes operating dur-
looking like a new kind or new form of or- ing such phylogeny would be incorrect. In
ganism can be designated as a new species. common descriptions of hominin evolution
Since there are no two organisms that look many homoplasies are identified, mean-
precisely alike in all details – even identi- ing that supposedly the same adaptations
cal twins differ somewhat on closer inspe- evolved separately in various hominin taxa
ction – researchers must judge whether the [Holliday 2003, Lieberman et al. 1996].
form of this particular fossil they are try- This might be true, accepting correctness of
ing to classify differs from other similar the standard phylogeny, but it might not be
forms more than expected for members of true if individuals assigned to separate taxa
the same species, or less. Some definitions actually belonged to a single polymorphic
of species, especially those akin to the species that was evolving as a whole. Then,
Biological Species Concept [Mayr 1969, various individuals of such species are ex-
1995], allow for the existence of fairly pected to have same adaptations.
wide ranges of intraspecific variation, Construction of phylogenies based on-
while others, such as especially the Phylo- species that are considered ideal static
genetic Species Concept or morphological descriptions of organisms of a particular
species concepts, may consider any “diag- “kind”, imposes on the explanations of tran-
nosable” difference as defining a new spe- sitions from one taxon to the next abrupt,
cies, thus narrowing substantially allowa- largely unexplainable events in the form
ble ranges of intraspecific variation [Hen- of “punctuations” [Eldredge and Gould
nig 1966, Platnick 1977a,b, Eldredge and 1972] rather than adaptive processes based
Cracraft 1980, Nelson and Platnick 1981, on generation-by-generation reproductive
Mishler and Donoghue 1982, Mishler and success of some variants over others. The
Brandon 1987, De Queiroz and Donoghue use of platonic ideal entities – immutable
1988, Nixon and Wheeler 1990, Kimbel species – prevents an understanding of the
and Rak 1993, Mishler and Theriot 2000]. actual mechanism of evolution and factors
This increases the arbitrariness of deci- that play a role in evolutionary changes.
sions assigning particular specimens to As Charles Darwin [1859: 485] himself re-
previously known species or to new ones. marked “… we shall at least be freed from
Once a new species is created, it replaces the vain search for the undiscovered and
a particular specimen, or sample of speci- undiscoverable essence of the term spe-
mens in consideration of phylogenies, that cies.” He also stated that he does not see any
is, in descriptions of the process of macro- substantial difference between the notion
evolution. Production of phylogenies does of variety and a species. His explanation
72 M. Henneberg

of the way life evolves rests on the obser- earliest human ancestors. For later periods
vation that individuals are variable, that that have more abundant fossil representa-
they are adapted to their living conditions tion, greater number of individual fossils
only to a certain degree and that those who necessarily produces greater variability of
happen to be best adapted at a given mo- morphological characters. This, coupled
ment have greater reproductive success with the tendency to categorize, and with
than others. Life consists of assemblages the common belief that evolution proceeds
of individuals who, though clustering into by emergence of new species, leads most
groups based on mutual similarity, are researchers to splitting of the Pliocene and
variable and reproduce with different suc- Pleistocene fossil record into numerous
cess that is a measure of their adaptation to species while also recognizing that they
momentary conditions. Those conditions can be organized into chronological grades
change through time and thus midpoints of [Henry and Wood 2007]. We are therefore
ranges of variation of individuals shift with left with a picture of human evolution in
the passage of time. which the necessary use of similarities to
identify our ancestors and the natural in-
Identification of the human lineage dividual, regional, microevolutionary and
sexual variation act in opposite directions
With regard to human ancestry, research- – reducing and increasing postulated di-
ers are faced with a difficult task that au- versity. Disputes regarding detailed mor-
tomatically produces a conflict between phological, chronological and geographi-
identification of taxonomic diversity and cal characteristics of particular fossils can
lineage continuity. The only way to identify continue interminably, unless some gen-
human ancestors in the fossil record is to erally accepted method of distinguishing
find among assemblages of primate fossils intra-specific variation from inter-specific
from the Miocene, Pliocene and Pleistocene diversity can be applied.
those skeletal remains that resemble humans
more than other primates. As one goes back Quantitative testing of the
in time, having already identified human- hypothesis of human lineage
like fossils at a later date, one can then try to
compare even earlier fossils with those lat- The 20th century produced a large, tem-
er ones and so on. Eventually, one enters porally and geographically widespread fos-
the epoch that contains no fossils whose sil record of hominins. Thus, detailed de-
similarity to later putative human ancestors bates regarding small number of individual
can convincingly be shown. At present, fossils can be replaced by a generalizing
this process has reached back some 6-7 overview using statistical methods of hy-
Ma, identifying forms such as Sahelan- potheses testing to resolve the perennial
thropus chadensis and Orrorin tugenensis dispute between “lumpers” and “splitters”.
as possible earliest hominins (Stanford et The approach to testing hypotheses of hu-
al. 2009). The necessity to identify fossils man evolution should conform to the ba-
somehow similar to those that are similar to sic rules of scientific methodology: par-
humans, while the density of fossil record simony and null hypothesis falsification.
decreases on going back in time, reduces We should use the commonly accepted ap-
the reliability of the identification of the proach in which a null hypothesis must be
Two interpretations of human evolution 73

falsified before alternative hypotheses can be hominin species [Wood and Richmond
entertained. In the case of fossil documenta- 2000]. One can add to those generalizing
tion of hominin lineage, the null hypothesis characteristics the direct metric character-
that it consisted of one species at each point istics of dentition since teeth preserve well
in time can be reasonably built based on the in the fossil record.
general consideration of the speciosity of The study of variation in some 210 ho-
genera of large eurytopic mammals [Conroy minin brain sizes, some 205 body size es-
2002, Hunt 2003]. Such genera usually com- timates and 915 sizes of dentition [Hen-
prise just one or two species. This is true of neberg and Thackeray 1995; Henneberg
the hominid genera Pan, Gorilla and Pongo. and de Miguel 2004; Henneberg 1997,
Although the number of hominin fossils 2006, 2008] invariably shows that there is
is of the order of thousands of specimens, a gradual change in those characteristics
these are fragmentary. Thus sample sizes through geological time, that distributions
for determination of their various detailed of individual values around regressions
characteristics can be small. It is practical describing temporal changes are unimo-
to analyze for the largest possible number dal, normal and of the magnitude not dif-
of fossils general characteristics that can be ferent from that of a single modern species
uniformly reconstructed from various parts Homo sapiens. The scattergrams of body
of skeletal elements. These are: body size height, mass and cranial capacity (Fig. 3)
and cranial capacity. The latter is still exten- illustrate temporal continuity without
sively used to characterize various putative abrupt changes and branching.

Fig. 3. Body height, mass and cranial capacity of hominins from Pliocene to the end of Pleistocene. All
scales logarithmic. Note continuity of distributions and lack of branching. All available in the literature:
cranial capacities of adult hominins are used (N = 209, as reported by de Miguel and Henneberg [2001],
supplemented from more recent literature). Reconstructed body heights and weights of hominins from
Mathers and Henneberg [1995] and de Miguel and Henneberg [1999].
74 M. Henneberg

Fig. 4. Average total tooth areas for various hominin taxa compared with several averages and 2
standard deviation ranges for various modern human groups. Note that differences between hominin
species are of the same order as those between modern human populations. The only exception are robust
australopithecines. Data for all australopithecines, H. habilis, H. erectus and H. sapiens (modern) from
Tobias [1988], for Neandertals from Brace and Mahler [1971], for Nubians from Calcagno [1989], for
Early Upper Palaeolitic and Mesolitic Europe from Frayer [1978], for Australian Aborigines and for
French from Brace [1995].

The averages of the “total dental mate- through time in the hominin lineage [Hen-
rial” [Tobias 1988], that is, the sum of hori- neberg 2007, Martin 2007]. It is the single,
zontal crown areas of all teeth, of various unfossilized skeleton labeled LB1 found,
hominin taxa do not differ more than av- together with isolated bone elements and
erages of various modern human groups teeth of a few other humans on the island of
(Fig. 4) with exception of the robust aus- Flores, Indonesia in 2003–2004 [Brown et
tralopithecines. Since total dental mate- al. 2004, Morwood et al. 2004, Brumm et
rial is a composite variable and difficult to al. 2006]. Both its body size (1.06 – 1.35 m)
calculate for individuals, its variation can and cranial capacity (about 430 ml) fall 3 to
only be estimated. Here we use an SD es- 5 SD below the distributions for other ho-
timate of approximately 100 mm2 derived minins of similar dating (Late Pleistocene)
from Brace et al. [1991]. When intervals of while size of the dentition is close to the av-
2 SD are built around averages for various erage. The find has been, and still is, hotly
groups, most of the variation among hom- debated as a number of authors proposed
inin taxa averages is included in those in- that it is a pathologically deformed individ-
tervals (see Fig. 4). ual, not a representative of a new species
There is only one exception from the [Henneberg and Thorne 2004, Jacob et al.
pattern of brain and body size change 2006, Weber et al. 2005, Hershkovitz et al.
Two interpretations of human evolution 75

2007, Martin et al. 2006a,b, Obendorf et who, responding to the pressures of nat-
al. 2008]. It has been suggested that its dat- ural selection, and later to the forces of
ing is incorrect and it is a modern human artificial selection resulting from self-am-
[Henneberg and Schofield 2008]. Until this plifying feedbacks between human biolo-
debate is resolved, we can assume that this gy, technology and culture [Bielicki 1969,
exception is a result of doubtful attribution Strzałko and Henneberg 1982], change to-
rather than a challenge to the general pic- wards modern human form. The present-
ture of hominin evolution. day human similum comprises a variety
The recent description of a large as- of individuals bearing signs of their an-
semblage of 4.4 Ma old fossils of the Ar- cestral adaptations to the same wide range
dipithecus ramidus [White et al. 2009] fits of climatic and economic conditions as
the pattern well. The cranial capacity of it was in the past, except that over time
300–350 ml and body height of 1.2 m could the external conditions to which adapta-
be predicted by extrapolating backwards in tions occurred were different and those
time the trends already documented (see conditions changed over time due both to
Fig. 3). natural forces and human activities. Such
an interpretation of human evolution is
Conclusion: A single variable, simpler than the long litany of fossil “spe-
adapting lineage cies” changing abruptly and inexplicably
from one to another. Most importantly, it
It can be concluded that at no point in allows us to focus attention on the proc-
time since the late Miocene in the distribu- esses of adaptation, explainable in terms
tions of basic metric characteristics of ho- of Darwinian mechanisms, that led from
minins are there to be found any significant sparse populations of hunter/gatherer ho-
discontinuities, deviations from normal- minids to the widespread, very dense and
ity or ranges expanded beyond what is ob- numerically dominant form of large mam-
served in the only living hominin species. mal.
Although arguments can be made that upon
a study of descriptive characteristics some Notes
categorical discontinuities can be detected,
the subjective nature of the descriptions This paper was presented (in Polish) at the
42nd biannual meeting of the Polish Anthropo-
makes it difficult to objectively test such logical Society in Łódź in September 2009.
arguments. Thus the null hypothesis that
hominins are a single lineage cannot be fal-
sified and must be accepted at the present References
time. This is the most parsimonious and
Ackermann R.R., J. Rogers, J.M. Cheverud,
simplest interpretation able to be made. It 2006, Identifying the morphological signa-
is also compatible with the way by which tures of hybridization in primate and human
hominin ancestors are identified among evolution, J. Hum. Evol., 51, 632–45
fossils. Albrecht G.H., J.M. Miller, 1993, Geographic
variation in Primates: A review with impli-
Since the Miocene, we are dealing with cations for interpreting fossils, [in:] Species,
populations of interbreeding, geographi- Species Concepts, and Primate Evolution,
cally widespread ecologically eurytopic W.H. Kimbel & L. Martin (eds.), Plenum
[Henneberg 2001] human individuals Press, New York, pp. 123–61
76 M. Henneberg

Bielicki T., 1969, Niektóre związki zwrotne Darwin C., 1859 [1952], The origin of species
w procesie ewolucji Hominidae, Mat. i Prace by means of natural selection, Encyclopaedia
Antrop., 77, 3–60 Britannica, Chicago
Brace C.L., 1967, The Stages of Human Evolu- Delson E., I. Tattersall, J.A. van Couvering,
tion, Prentice-Hall, Englewood Cliffs, NJ A.S. Brooks, 2000, Brief introduction to hu-
Brace C.L., 1981, Tales of the phylogenetic man evolution and prehistory, [in:] Encyclo-
woods: The evolution and significance of pedia of Human Evolution and Prehistory,
evolution trees, Am. J. of Phys. Anthropol., 2nd ed., E. Delson, I. Tattersall, J.A. Couver-
56, 411–29 ing, A.S. Brooks (eds.), Garland Publishing,
Brace C.L., 1995, Trends in the evolution of hu- New York, pp. xvii–xxii
man tooth size, [in:] Aspects of Dental Biol- De Miguel C., M. Henneberg, 1999, Variation
ogy: Palaeontology, Anthropology, and Evo- in hominid body size estimates: Do we know
lution, J. Moggi-Cecchi (ed.), Int. Inst. for the how big our ancestors were?, Persp. Hum.
Study of Man, Florence, pp. 437–46 Biol., 4(1), 65–80
Brace C.L., P.E. Mahler, 1971, Post-Pleisto- De Miguel C., M. Henneberg, 2001, Variation
cene changes in the human dentition, Am. in hominid brain size: How much is due to
J. Phys. Anthropol., 34, 191–204 method?, Homo, 52, 2–56
Brace C.L., S.L. Smith, K.D. Hunt, 1991, De Queiroz K., M.J. Donoghue, 1988, Phylo-
What big teeth you had grandma! Human genetic systematics and the species problem,
tooth size, past and present, [in:] Advances Cladistics, 6, 61–75
in Dental Anthropology, M.A. Kelley & Eckhardt R.B., 2000, Human paleobiology,
C.S. Larsen (eds.),  Wiley-Liss, New York, Cambridge Univ. Press, Cambridge
pp. 33–57 Eldredge N., S.J. Gould, 1972, Speciation
Brown P., T. Sutkina, M.J. Morwood, R.P. Soe- and punctuated equilibria: An alterna-
jono, E. Jatmiko, et al., 2004, A new small- tive to phyletic gradualism, [in:] Models in
bodied hominin from the Late Pleistocene of paleobiology,T. Schopf (ed.), Freeman Coo-
Flores, Indonesia, Nature, 431, 1055–61 per, San Francisco, pp. 85–120
Brumm A., F. Aziz, G.D. van den Burgh, M.J. Eldredge N., J. Cracraft, 1980, Phylogenetic
Morwood, M.W. Moore, et al., 2006, Early Patterns and the Evolutionary Process, Co-
stone technology on Flores and its implica- lumbia Univ. Press, New York
tions for Homo floresiensis, Nature, 441, Frayer D., 1978, The Evolution of the Dentition
624–28 in Upper Paleolithic and Mesolithic Europe,
Calcagno J., 1989, Mechanisms of Human Univ. of Kansas Publ. in Anthropology, 10,
Dental Reduction, A Case Study from Post- Lawrence, Kansas
Pleistocene Nubia, Univ. of Kansas Publ. in Frisancho A.R., 1990, Anthropometric stan-
Anthropology, 18, Lawrence, Kansas dards for the assessment of growth and nu-
Conroy G.C., 2002, Speciosity in the early tritional status, Univ. of Michigan Press, Ann
Homo lineage: Too many, too few, or just Arbor
about right? J. Hum. Evol., 43, 759–66 Genesis, 1611, The Holy Bible, Robert Barker,
Carruthers P., A. Chamberlain (eds.), 2000, London
Evolution and the human mind, Cambridge Gonzalez-Forero M., 2009, Removing ambi-
Univ. Press, Cambridge guity from the biological species concept,
Cela-Conde C.J., F.J. Ayala, 2007, Human J. Theor. Biol., 256, 76–80
evolution. Trails from the past, Oxford Univ. Henneberg M., 1988, Decrease of human skull
Press, Oxford size in the Holocene, Hum. Biol., 60, 395–
Cope D.A., M.G. Lacy, 1992, Falsification of 405
a single species hypothesis using the coef- Henneberg M., 1990, Brain size/body weight
ficient of variation: A simulation approach, variability in Homo sapiens: Consequences
Am. J. Phys. Anthropol., 89, 359–78 for interpreting hominid evolution, Homo,
Cope, D.A., M.G. Lacy, 1996, The CV as a tool 39, 121–30
for assessing the taxonomic composition of Henneberg M., 1997, The problem of species
fossil sample, Am. J. Phys. Anthropol., Sup- in hominid evolution, Persp. Hum. Biol., 3,
pl. 22, 89 21–31
Two interpretations of human evolution 77

Henneberg M., 1998, Evolution of the human floresiensis and patients with primary growth
brain: Is bigger better?, Experim. Clin. hormone insensitivity (Laron Syndrome), Am.
Physiol. Pharmacol., 25, 745–49 J. Phys. Anthropol., 134, 198–208
Henneberg M., 2001, The gradual eurytopic Holliday T.W., 2003, Species concepts, re-
evolution of humans: Not from Africa alone, ticulation, and human evolution, Curr. An-
[in:] Man: Past, present, and future, E. In- thropol., 44, 653–60
driati (ed.), Bigraf Publishing, Yogyakarta, Hunt K.D., 2003, The single species hypoth-
Indonesia, pp. 42–52 esis: Truly dead and pushing up bushes, or
Henneberg M., 2006, The rate of human mor- still twitching and ripe for resuscitation?,
phological microevolution and taxonomic Hum. Biol., 75, 485–502
diversity of hominids, Studies in Historical Jacob T., E. Indriati, R.P. Soejono, K. Hsü, D.
Anthropology, 4 [2004], 49–59 Frayer, et al., 2006, Pygmoid Australomela-
Henneberg M., 2007, The mode and rate of nesian Homo sapiens skeletal remains from
human evolution and the recent Liang Bua Liang Bua, Flores: Population affinities and
finds, [in:] Recent advances of Southeast Asia pathological abnormalities, Proc. Natl. Acad.
paleoanthropology and archaeology, E. In- Sci., 103, 13421–26
driati (ed.), Gadjah Mada University, Yogya- Jolly C.J., 1993, Species, subspecies, and
karta, pp. 24–29 baboon systematics, [in:] W.H. Kimbel &
Henneberg M., 2008, Darwinian interpretation L. Martin (eds.), Species, Species Concepts,
of human evolution based on the evolution and Primate Evolution, Plenum Press, New
of body size and brain size, [in:] Ksiega ju- York, pp. 67–107
bileuszowa, stulecie Zakladu Antropologii Jolly C.J., 2001, A proper study for mankind
Uniwersytetu Jagiellonskiego 1908–2008, analogies from the papionin monkeys and
K. Kaczanowski (ed.), Plus Publishers, Kra- their implications for human evolution,
ków, pp. 71–85 Yearb. Phys. Anthropol., 44, 177–204
Henneberg M., G. Brush, 1994, Similum, a con- Jolly C.J., T. Woolley-Barker, S. Beyene, T.
cept of flexible synchronous classification re- Distotell, J. Phillips-Conroy, 1997, Inter-
placing rigid species in evolutionary think- generic hybrid baboons, Int. J. Primatol., 18,
ing, Evol. Theor., 10, 278 597–628
Henneberg M., A. Budnik, M. Pezacka, A.E. Jones S., R.D. Martin, D.R. Pilbeam, 2000, The
Puch, 1985, Head size body size and intel- Cambridge Encyclopedia of Human Evolu-
ligence intraspecific correlation in Homo sa- tion, Cambridge Univ. Press, Cambridge
piens species, Homo, 36, 207–18 Kelley J., 1993, Taxonomic implications of sex-
Henneberg M., C. de Miguel, 2004, Hominins ual dimorphism in Lufengpithecus, [in:] Spe-
are a single lineage: Brain and body size vari- cies, species concepts, and primate evolution,
ability does not reflect postulated taxonomic W.H. Kimbel & L.B. Martin (eds.), Plenum
diversity of hominins, Homo, 55, 21–37 Press, New York, pp. 429–58
Henneberg M., J. Schofield, 2008, The Hobbit Kimbel W., Y. Rak, 1993, The importance of spe-
Trap, Wakefield Press, Adelaide cies taxa in paleoanthropology and an argu-
Henneberg M., J.F. Thackeray, 1995, A single- ment for the phylogenetic concept of the species
lineage hypothesis of hominid evolution, category, [in:] Species, species concepts, and
Evol. Theor., 13, 31–38 primate evolution, W.H. Kimbel & L.B. Martin
Henneberg M., A. Thorne, 2004, Flores human (eds.), Plenum Press, New York, pp. 461–84
may be pathological Homo sapiens, Before Lacruz R.S., F.R. Rozzib, T.G. Bromage, 2005,
Farming, 4, 2–4 Dental enamel hypoplasia, age at death, and
Hennig W., 1966, Phylogenetic Systematics, weaning in the Taung child, S. Afr. J. Sci.,
Univ. of Illinois Press, Illinois 101, 567–69
Henry A.G., B. Wood, 2007, Whose diet? An in- Larsen C.S., 2008, Our Origins: Discovering
troduction to the hominin fossil record, [in:] Physical Anthropology, W.W. Norton, New
Evolution of the human diet, P.S. Ungar (ed.), York
Oxford Univ. Press, Oxford, pp. 11–28 Lee M.S.Y., 2003, Species concepts and species
Hershkovitz I., L. Kornreich, Z. Laron, 2007, reality: Salvaging a Linnean rank, J. Evol.
Comparative skeletal features between Homo Biol., 16, 179–88
78 M. Henneberg

Lieberman D.E., D.R. Pilbeam, B.A. Wood, & R. Meier (eds.), Columbia Univ., pp. 44–
1996, Homoplasy and early Homo: An analy- 54
sis of the evolutionary relationship of H. ha- Morwood M.J., R.P. Soejono, R.G. Roberts, T.
bilis sensu stricto and H. rudolfensis, J. Hum. Sutikna, C.S.M. Turney, et al., 2004, Ar-
Evol., 30, 97–120 chaeology and age of a new hominin from
Linnaeus C., 1758, Systema Naturae, Editio de- Flores in eastern Indonesia, Nature, 431,
cimal, Laur Salvius, Holmiae 1087–91
Manning A., M.S. Dawkins, 1998, An introduc- Nelson G.J., N.I. Platnick, 1981, Systematics
tion to animal behaviour, 5th ed., Cambridge and biogeography: Cladistics and vicari-
Univ. Press, Cambridge ance, Columbia Univ. Press, New York
Martin R.D., 2007, Problems with the tiny brain Nixon, K.C., Q.D. Wheeler, 1990, An ampli-
of the Flores hominid, [in:] Recent advances fication of the phylogenetic species concept,
of Southeast Asia paleoanthropology and Cladistics, 6, 211–23
archaeology, E. Indriati (ed.), Gadjah Mada Obendorf P., C.E. Oxnard, B.J. Kefford, 2008,
University, Yogyakarta, pp. 9 –23 Are the small human–like fossils found on
Martin R.D., A.M. MacLarnon, J.L. Phillips, Flores human endemic cretins?, Proc. R. Soc.
L. Dussubieux, P.R. Williams, W.B. Dobyns, B, 275, 1287–96
2006a, Comment on “The brain of LB1, Platnick N.I., 1977a, Cladogram, phylogenetic
Homo floresiensis”, Science, 312, 999 trees, and hypothesis testing, Syst. Zool., 26,
Martin R.D., A.M. MacLarnon, J.L. Phillips, 438–42
W.B. Dobyns, 2006b, Flores hominid: New Platnick N.I., 1977b, Monophyly and the origin
species or microcephalic dwarf?, Anat. Rec., of higher taxa: A reply to E.O. Wiley, Syst.
288A, 1123–45 Zool., 26, 355–57
Mayr E., 1969, Principles of systematic zool- Plavcan J.M., 1993, Catarrhine dental variabil-
ogy, McGraw-Hill, New York ity and species recognition in the fossil record,
Mayr E., 1995, Species, classification, and [in:] Species, species concepts, and primate
evolution, [in:] Biodiversity and Evolution. evolution, W.H. Kimbel & L.B. Martin (eds.),
R. Arai, M. Kato, Y. Doi, (eds.), National Plenum Press, New York, pp. 239–93
Science Museum Foundation, Tokyo, pp. Plavcan J.M., D.A. Cope, 2001, Metric varia-
3–12 tion and species recognition in the fossil re-
Mathers K., M. Henneberg, 1995, Were we ever cord, Evol. Anthropol., 10, 204–22
that big ? Gradual increase in hominid body Quintyn C., 2009, The naming of new species
size over time, Homo, 46, 141–73 in hominin evolution: A radical proposal
Mayden R.L., 1997, A hierarchy of species con- – a temporary cessation in assigning new
cepts: the denouement in the saga of the spe- names, Homo, 60, 307–41
cies problem, [in:] Species, the Units of Bio- Rose K.D., T. Bown, 1993, Species concepts
diversity, M.F. Claridge, H.A. Dawah, M.R. and species recognition in Eocene primates,
Wilson (eds). Chapman and Hall, London, [in:] Species, species concepts, and primate
pp. 381–424 evolution, W.H. Kimbel & L.B. Martin (eds.),
Meikle W.E., S.T. Parker, 1994, Naming our Plenum Press, New York, pp. 299–330
ancestors. An anthology of hominid Taxono- Samadi S., Barberouse A., 2009, Species: To-
my, Waveland Press, Prospect Heights, Ill. wards new, well-grounded practices, Biol.
Mishler B.D., M.J. Donoghue, 1982, Species J. Linn. Soc., 97, 217–22
concepts: a case for pluralism, Syst. Zool., Staley J.T., 2009, Universal species concept:
31, 491–503 Pipe dream or a step toward unifying biol-
Mishler B.D., R.N. Brandon, 1987, Individu- ogy?, J. Ind. Microbiol. Biotechnol., 36,
ality, pluralism, and the phylogenetic species 1331–36
concept, Biol. & Philos., 2, 397–414 Stanford C., J.S. Allen, C.S. Antón, 2009,
Mishler B.D., E.C. Theriot, 2000, The phylo- Biological Anthropology, Pearson Education,
genetic species (sensu Mishler and Theriot): Upper Saddle River, NJ
Monophyly, apomorphy, and phylogenetic Strzałko J., M. Henneberg, 1982, Hominization
species concepts, [in:] Species concepts and as a necessary effect of evolution of a non-
phylogenetic theory: A debate, Q.D. Wheeler genetic mode of hereditary transmission,
Two interpretations of human evolution 79

[in:] Evolution and environment, V. Novak, Weber J., A. Czarnetski, C.M. Pusch, 2005,
J. Milkovsky (eds.), CSAV Press, Prague, pp. Comment on “The brain of LB1, Homo flore-
367–76 siensis”, Science, 310, 236
Tattersall I., 1987, Species recognition in hu- White T.D., B. Asfaw, Y. Beyene, J. Haile-
man palaeontology, J. Hum. Evol. 15, 165–76 Selassie, C.O. Lovejoy, et al., 2009, Ardipi-
Tattersall I., E. Delson, J.A. van Couvering thecus ramidus and the paleobiology of early
(eds.), 1988, Encyclopedia of Human Evolution hominins, Science, 326, 64
and Prehistory, Garland Publishing, New York Wolpoff M.H., 1968, “Telanthropus” and the
Tobias P.V., 1988, Tooth material in the homin- single species hypothesis, Am. Anthrop., 72,
idae, J. Dent. Assoc. S. Afr., 43, 557–60 447–93
Velasco J.D. 2009, When monophyly is not Wolpoff M.H., 1971, Competitive exlusion
enough: Exclusivity as the key to defining among Lower Pleistocene hominids: The sin-
a phylogenetic species concept, Biol. Philos., gle species hypothesis, Man, 6, 601–14
24, 473–86 Wood B., B.G. Richmond, 2000, Human evolu-
Walker A., R.E. Leakey, 1993, The Nariokotome tion: taxonomy and paleobiology, J. Anat.,
Homo erectus skeleton, Springer, New York 197, 19–60

Streszczenie
Mimo bogactwa materiałów zgromadzonych w ubiegłym stuleciu, wśród autorów
zajmujących się antropogenezą brakuje jednomyślności w rozumieniu tego procesu. Nie-
którzy z nich utrzymują, że w ciągu ostatnich 5 milionów lat istniało 17, lub nawet więcej
gatunków homininów, inni twierdzą, że było ich znacznie mniej – może zaledwie 2, a nawet,
że ewolucja człowieka przebiegła w obrębie jednej, niedzielącej się, linii rozwojowej.
Różne są także interpretacje pojawienia się dwunożności i przyczyn wzrostu względnych
rozmiarów mózgowia. Nadal toczą się spory o rolę neandertalczyków i pochodzenie ludzi
o współczesnych cechach morfologicznych. Taka różnorodność interpretacji tej samej bazy
faktograficznej wskazuje na słabość podstaw teoretycznych, jakimi posługują się zwolennicy
rozmaitych koncepcji. Słabość ta wynika najprawdopodobniej z tego, że badacze antropo-
genezy rekrutują się spośród absolwentów wielu dyscyplin, zarówno humanistycznych jak
i przyrodniczych (rzadko biologii) oraz stosują heterogeniczne paradygmaty i niejednolite
definicje podstawowych pojęć, takich jak gatunek, gen czy specjacja. Przy tym, ze względu
na światopoglądowe znaczenie wyjaśnienia naszego pochodzenia, istnieją uwarunkowania
pozanaukowe – ideologiczne, którym różni autorzy ulegają świadomie lub podświadomie
(ryc. 1).
W tej sytuacji trzeba zmierzać do rozstrzygnięcia istniejących sporów stosując podstawowe
zasady postępowania naukowego – falsyfikację hipotez w oparciu o odpowiednio liczne, jed-
nolicie mierzone lub opisywane, materiały. Takie podejście jest możliwe w stosunku do licz-
nych już obserwacji dotyczących podstawowych cech hominidów, takich jak wielkość mózgu,
rozmiary ciała i uzębienia. Cechy te zmieniały się w ciągu ostatnich 3-4 milionów lat w sposób
ciągły, nie wykazując rozszczepiania w procesach specjacji. Analiza wariancji rozkładów tych
cech w poszczególnych okresach chronologicznych nie pozwala odrzucić hipotezy zerowej,
stanowiącej, że rozkłady te nie różnią się wielkością wariancji wewnątrzgrupowej od rozkładu
charakteryzującego pojedynczy, i jedyny żyjący obecnie, gatunek Homo sapiens (ryc. 2).
Rozkład indywidualnych rozmiarów mózgów 210 homininów wokół linii ich regresji na czas
geologiczny nie różni się istotnie od rozkładu normalnego. Podobnie, rozkłady wielkości
ciała po usunięciu efektu czasu (datowania) nie odbiegają od normalności (ryc. 3). Wielkość
80 M. Henneberg

uzębienia rozmaitych postulowanych gatunków homininów, z wyjątkiem masywnych australo-


piteków, waha się w granicach nie przekraczających różnic pomiędzy rozmaitymi populacjami
człowieka współczesnego (ryc. 4).
Nie da się zatem, w oparciu o te podstawowe cechy, sfalsyfikować hipotezy, że w ciągu os-
tatnich 3-4 milionów lat istniał w każdym momencie tylko jeden gatunek człowieka (lub jego
przodka). Akceptacja hipotezy alternatywnej, o wielogatunkowości, może być rozważana tylko
w oparciu o cechy bardziej szczegółowe, mierzone (opisywane) mniej obiektywnie, których
próby mają mniejszą liczebność, a interpretacja wartości przystosowawczej nie zawsze jest
jasna. Badania antropogenezy winny w tej sytuacji skoncentrować się na wyjaśnianiu ewolucji
przystosowań ludzkich, a nie na formalnej opisowej klasyfikacji poszczególnych znalezisk.

Vous aimerez peut-être aussi