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Evolution of Early Primates
The proliferation of mammalian forms from
the end of the Cretaceous period (ca. 65 Mya) coincided with the
extinction of many of the other life forms which roamed over the earth at
that time (including dinosaurs). The depopulation of the planet likely
opened up many new niches, which accounts for the rapid increase in
mammalian species following the Cretaceous-Tertiary boundary. Primates -
or their tree shrewlike progenitors - were one of these opportunistic,
niche-filling mammals. However, given that a great many mammalian types
emerged at this time, how is it that primates acquired the peculiar
battery characteristics which would prove so instrumental in the later
success of the order?
Three Hypotheses
An early theory of primate evolution,
proposed by F. W. Jones (1916), relates the primate characteristics of
grasping hands and feet, orbital frontation (increased binocularity of
vision), and enhanced cognitive processing capacity to the challenge of
arboreal life. Generally known as the Arboreal Theory of primate
evolution, this theory explains the enhanced visual-motor systems and
cognitive elaboration in primates as products of arboreal life.
However, one major weakness in the argument
for the Arboreal Theory is the fact that arboreal mammals exist which do
not possess any of these traits, and yet are very successful. An example
of this would be tree squirrels, which have very little visual overlap,
non-opposable digits, and smallish brains; however, as any walk through a
park may attest, squirrels are quite successful in their own right. As
response to this, in 1974, Duke University primatologist Matt Cartmill
proposed a new theory - the Visual Predation Theory - to account
for the evolution of primate characteristics. Instead, he suggested that
arboreal predatory behavior accounted for the grasping hands and feet, and
particularly the increased visual overlap and brain size (both
characteristics observed in terrestrial predators, such as large cats).
However, a weakness in the Visual Predation
Hypothesis lies in the fact that prosimians, considered to be closer to
the ancestral form of all primates, exhibit lower reliance on visual
information for locomotion and predation; rather, they emphasize olfactory
and auditory cues in the pursuit of prey. In light of this fact,
Washington University primatologist Robert Sussman proposed the Mixed
Diet Theory - namely, that an increased exploitation of angiosperms
(flowering plants) selected for modern primate characteristics. Enhanced
visual acuity, color vision, and characteristics amenable to exploiting
terminal branch resources all allowed for efficient acquisition of a
resource with an angiosperm-like distribution. Additionally, the emergence
of flowering plants in the Paleocene roughly coincides with the emergence
of the earliest primate ancestors.
The First Anthropoids
While the earliest potential anthropoid
primates may have emerged close to 54 million years ago, the first
unambiguous remains date back to approximately 36 Mya, from a region
called the Fayum in Egypt. One of the more complete fossils from
the Fayum is a monkey named Catopithecus brownii, a diurnal (active
during the day), arboreal quadrupeds, and probably fed primarily on
insects. One characteristic linking C. brownii with modern day
anthropoids is the shared dental formula between it and all modern Old
World monkeys (2.1.2.3). Since New World monkeys have different dentitions
(2.1.3.3 or 2.1.3.2), C. brownii is not likely to be ancestral to
any of the New World species. Later fossil primates from the Fayum -
including Aegyptopithecus zeuxis, pictured at left - are grouped
into parapithecoids and propliopithecoids. The propliopithecoids (which
includes A. zeuxis) are considered to be possible ! progenitors of
modern anthropoid primates.
The New World Monkey Enigma
The puzzling ancestry of New World monkeys
derives from a low abundance of fossil material, as well as limitations
based on knowledge of continental drift. Similar biological diversity and
geology suggest that the continent of South America was once a part of
Africa. South America eventually broke off of the African mainland
(through a geologic process called continental drift), and over the
course of millions of years, migrated to its present position.
Unfortunately, the separation between South America and Africa occurred
over 100 Mya ago, which is much earlier than the first primate fossils on
either continent. This leads to several possible scenarios for the
ancestry of New World primates:
- African anthropoids crossed the Atlantic
somehow, and radiated into new habitats upon reaching South America
- North American primate forms gave rise
to the current New World species
- African anthropoids emerged earlier than
fossils suggest, and rafted across the Atlantic when it was much
smaller
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The problem with the first interpretation is
the tremendous distance involved in a trans-Atlantic journey (by the late
Oligocene it was already over 2000 miles). Large chunks of land mass have
been known to break off and essentially "float" across large
bodies of water as living cargo vessels, although evidence has yet to
emerge in support for this hypothesis on the origin of New World monkeys.
The second interpretation is problematic in the sense that North America
does not have any evidence of anthropoid species. Thus, descent
from a prosimian-like common ancestor would require a remarkable number of
evolutionary convergences between New World and Old World anthropoids, a
concession many systematists find difficult to accept. The third
possibility provides a compromise of sorts between the first two
explanations, suggesting that an ancestral primate form might have rafted
across the Atlantic, but when it was much smaller. This would push back
the divergence dates! for anthropoid primates beyond those supported by
the fossil record, but as an old anthropological adage goes, "absence
of evidence is not evidence of absence". Estimations of error in
fossil sampling suggest that anthropoids may have actually emerged as
early as 52 Mya. An earlier emergence of anthropoid primates might allow
for a transoceanic voyage before the distance between South America and
Africa became too large.
Emergence of the Hominoids
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The hominoids (ancestors of apes and humans)
first emerge in the late Oligocene (ca. 27 Mya) in Africa. These early
forms are represented by the genus Proconsul (pictured at right).
Elements of their dentition, cranium, and postcranial anatomy suggest they
were quadrupedal and frugivorous, not unlike earlier fossils. The proconsulids,
however, lacked tails, and exhibit limb proportions closer to those of
modern apes. By the middle Miocene (ca. 17 Mya) a large number of distinct
hominoid species emerged, including Dryopithecus, Kenyapithecus,
Oreopithecus, Ouranopithecus, and Sivapithecus; these
forms were distributed throughout Africa, Europe, and Asia, suggesting an
adaptive radiation of hominoid forms occurred during the warmer Miocene
epoch. While some of these species have been closely associated with
modern forms (for example, see a comparison between Sivapithecus
and modern orang! utan), none of them appear to have any clear link
with the earliest hominids. With the aridification of the late Miocene,
the majority of hominoid forms went extinct, leaving behind a few modern
day forms (Pan, Gorilla, and Pongo). Of more
relevance to the study of human origins, the apparent pressure of a
shrinking forest habitat may have driven an ancestral ape to adopt a
peculiar form of locomotion; this adaptation, which had profound effects
on the trajectory of hominid evolution, is manifested in its earliest
members, the Australopithecines. |