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himpanzees, bonobos, and presume a chimpanzee- or gorilla-like
gorillas are our closest living BIPEDALITY ancestor to explain habitual upright
relatives. The most popular walking. Ar. ramidus was fully capable
reconstructions of human evolution Pan
of bipedality and had evolved a sub-
during the past century rested on the stantially modified pelvis and foot with
A
n essential goal of human evolutionary To many, these fossils were consistent with their occupation of forest refugia (32). Even their
studies is to account for human unique- chimpanzee-based referential scenarios. Bipe- gut structure differs substantially from that of
ness, most notably our bipedality, marked dality had long been argued to have occurred humans (33).
demographic success, unusual reproductive phys- when early hominids ventured onto the expand- How and why did such profound differences
iology, and unparalleled cerebral and technolog- ing savannas and grasslands of the Pliocene between hominids and African apes evolve?
ical abilities. During the past several decades, it (8, 9). More recently, bipedality is seen to have Why did early hominids become the only primate
has been routinely argued that these hominid char- emerged from African ape behaviors, including to completely eliminate the SCC? Why did they
acters have evolved by simple modifications of feeding postures (10, 11), gorilla dominance dis- become bipedal, a form of locomotion with vir-
homologs shared with our nearest living rela- plays (12), and even vertical climbing (13). Many tually no measurable mechanical advantage (34)?
tives, the chimpanzee and bonobo. This method mechanical/behavioral models have been pro- Why did body-size dimorphism increase in their
is termed referential modeling (1). A central tenet posed to explain the evolution of hominid bipe- likely descendants? These are now among the
has been the presumption (sometimes clearly dality, but most have presumed it to have evolved ultimate questions of human evolution. We can,
stated but more often simply sub rosa) that Gorilla from a chimpanzee-like ancestor (4, 14, 15). A of course, only hypothesize their answers. Never-
and Pan are so unusual and so similar to each primary problem with these scenarios has been the theless, by illuminating the likely morphological
other that they cannot have evolved in parallel; remarkably advanced postcranium of early Austra- structure and potential social behavior of the CLCA,
therefore, the earliest hominids must have also lopithecus, which exhibits particularly advanced Ar. ramidus now confirms that extant African ape–
resembled these African apes (2, 3). Without a adaptations to upright walking (16–18). based models are no longer appropriate.
true early hominid fossil record, the de facto null Ardipithecus ramidus. Ardipithecus ramidus Adaptive suites. An alternative to referential
hypothesis has been that Australopithecus was now reveals that the early hominid evolutionary modeling is the adaptive suite, an extrapolation
largely a bipedal manifestation of an African ape trajectory differed profoundly from those of our from optimization theory (35). Adaptive suites
(especially the chimpanzee). Such proxy-based ape relatives from our clade’s very beginning. are semiformal, largely inductive algorithms that
scenarios have been elevated to common wisdom Ar. ramidus was already well-adapted to bipe- causally interrelate fundamental characters
by genomic comparisons, progressively estab- dality, even though it retained arboreal capa- that may have contributed to an organism’s total
lishing the phylogenetic relationships of Gorilla, bilities (19–25). Its postcranial anatomy reveals adaptive pattern. One for the horned lizard
Pan, and Homo (4). that locomotion in the chimpanzee/human last (Phyrnosoma platyrhinos) of the southwesten
Early Australopithecus. Although Australo- common ancestor (hereafter the CLCA) must U.S. serves as an excellent example (Fig. 1)
pithecus was first encountered early in the last have retained generalized above-branch quad- (36, 37). For this species, the interrelation be-
century (5), its biology was only slowly revealed. rupedality, never relying sufficiently on suspen- tween a dietary concentration on ants and its
In the 1970s, abundant earlier Australopithecus sion, vertical climbing, or knuckle walking to impact on body form imply, at first counter-
fossils began to emerge in eastern Africa. These have elicited any musculoskeletal adaptations to intuitively, that elevation of clutch size and inten-
samples broadened our understanding of the ge- these behaviors (26–28). sification of “r” strategy (maximize the number of
nus and included partial skeletons (6) and even Moreover, Ardipithecus was neither a ripe- offspring by minimizing paternal care) are the ul-
footprint trails [the latter extending our knowledge fruit specialist like Pan, nor a folivorous browser timate consequences of this specialization (35–37).
to 3.75 million years ago (Ma)] (7). like Gorilla, but rather a more generalized omni- Such character and behavioral interdependen-
vore (19, 25). It had already abandoned entirely cies can have profound consequences on evolu-
the otherwise universal sectorial canine complex tionary trajectory, as demonstrated by the equally
Department of Anthropology, School of Biomedical Sciences,
(SCC), in which the larger, projecting upper ca- notable differences in clutch size in the common
Kent State University, Kent, OH 44242–0001, USA. E-mail: nine is constantly honed by occlusion against the leopard frog (3500 to 6500 eggs) versus those of
olovejoy@aol.com lower third molar of anthropoid primates (25), numerous species of so-called poison dart frogs
The primitive nature of the craniodental and was the CLCA’s socio-reproductive structure Possible alternative vicars are extant, K-selected
postcranial anatomies of Ar. ramidus suggest that before these events? Whereas African apes have, atelines (New World forms such as spider and
the CLCA, unlike extant African apes, was pre- in the past, almost invariably been selected as woolly monkeys), which exhibit many of the
dominantly arboreal. However, all of its descend- CLCA vicars, Ar. ramidus now allows us to infer CLCA’s likely socio-reproductive characters,
ants have since developed relatively sophisticated that they have undergone far too many pronounced including male philopatry, minimal-to-moderate
adaptations to terrestrial locomotion (23). What and divergent specializations to occupy such a role. canine dimorphism, moderate–to–possibly intense