BIO223

Evolutionary Transition from Primates to Humans

• Primates (order Primata) are basically herbivores, but are mostly not so large and
mainly forest-dwelling.
• They are represented by a dazzling diversity of species throughout tropical forests
where fruits are produced and trees retain foliage almost year-round.
• The platyrrhine (‘flat-nosed’) monkeys found in South America split off from the
haplorhine (‘simple-nosed’) suborder inhabiting the ‘Old World’ tropics early in
the Eocene ~50 Ma.
• Prosimians (bushbabies, lemurs and allies) diverged even earlier during the late
Cretaceous 74 Ma, based on genetic evidence, and are most diverse in
Madagascar.
• The monkeys inhabiting Africa and Asia (Cercopithecoidea) are separated from
the apes (Hominoidea), which lack tails, at superfamily level. Among apes, the
gibbons, found solely in Asia, are placed in a different family (Hylobatidae) from
the remainder (Hominidae).
• Humans are allied with chimpanzees (Pan troglodytes and P. paniscus) and
gorillas (Gorilla gorilla) in the latter family, but allocated to a distinct subfamily,
the Homininae.
• Ecologically, we have diverged from other apes in habitat, locomotion and diet.
• Evolution takes the form of changes in lineages through time.
• These changes are captured by names given to species, traditionally defined
morphologically with reference to type specimens.
• A conceptual structure has been developed for identifying features that have
been derived from some ancestral form, perhaps shared with sister species, and
those that are novel, distinguishing the species.
• Supporting this approach is the biological species concept, based on whether
individuals can interbreed and produce fertile offspring..
• If they can, they share a gene pool and their populations follow a common
evolutionary trajectory.
• Populations may become isolated if males and females fail to recognise one
another as mates or if the offspring produced are not viable, for whatever reason.
• Genetic divergence, through the accumulation of mutations over time, however
neutral in their effects, can be used to infer that populations have been isolated
for sufficiently long to be assumed distinct at the species level.
• Of course, one cannot be sure that they would not interbreed if they came into
contact later.
• A similar caveat applies spatially. Should populations separated geographically be
regarded as distinct species, or merely subspecies, morphologically different in
some features but without barriers to mating if connected?
• Ecologically, species can coexist only if they differ sufficiently in niche occupation
as defined by resources exploited, physiological tolerance or mechanisms for
evading predation
• The earliest fossils placed in the genus Homo come from Ledi-Geraru in the
Afar region of north-eastern Ethiopia, in the form of a mandible with teeth dated
to almost 2.8 Ma.
• They exhibit features transitional between Au. afarensis and later specimens from
Ethiopia assigned more firmly to early Homo, dated to around 2.4 Ma.
• Vertebrate fossils suggest that the habitat had become prevalently more open
with a mixture of grassland and riparian forest and expanded C4 component
around that time.
• Around one-third of the bovid taxa identified at the site made their first
appearance then, including the earliest wildebeest (Connochaetes sp.).
•.
• The Ledi-Geraru mandible is followed after an interval of 400 kyr by fossils
assigned to early Homo recorded not only in Ethiopia, but also further south in
Kenya and even Malawi.
• This gap in time happens to span the climatic transition into the start of the
recurrent ice ages typifying the Pleistocene. Further specimens assigned more
firmly to H. habilis, including a nearly complete skull, come from both the Omo–
Turkana Basin and Olduvai Gorge, dated ~1.9 Ma.
• This name, meaning ‘handy man’, was given because the fossils were associated
with stone tools defining the Oldowan culture at Olduvai.
• Features distinguishing this species from earlier australopiths include a larger
body size and cranial capacity, less-protruding face, longer legs, smaller jaws, and
smaller molar teeth with thinner enamel.
• Hominins somewhat larger in size discovered to the west of Lake Turkana,
likewise dated to ~2.0 Ma, were named H. rudolfensis, but may not be sufficiently
distinct from H. habilis (or H. ergaster) to justify the species distinction.
• Fossils from the South African cave sites allied with early Homo are generally too
fragmented to be assigned reliably to any particular species.
• These very early humans were rare components of local faunas, making up less
than 0.1 percent of all large mammal fossils, which is less than contributed
by Paranthropus spp. and similar to that formed by large carnivores.
• The scarcity of fossils representing early Homo in South African cave sites
suggests that early humans were either rare in this Highveld plateau region, or
much less susceptible to falling (or being dragged) into cavities than Paranthropus
• The earliest fossils assigned to H. ergaster come from the Omo–Turkana Basin,
dated to ~1.9 Ma, but became common elsewhere only after 1.7 Ma. H.
ergaster is commonly merged with H. erectus, named from fossils found in Java
and elsewhere in south-east Asia.
• It made its appearance during the time when glacial oscillations defining the
Pleistocene, coupled with local earth movements, further accentuated aridity in
eastern Africa.
• These proto-humans exhibited a cranial volume a little larger than that of H.
habilis, stood as tall as modern humans, and walked as competently.
• Their molar teeth were relatively smaller than those of the gracile australopiths,
and incisors somewhat larger, thereby reverting towards dental features of the
great apes. Cranially, they retained a sloping forehead and prominent brow
ridges.
• Surprisingly, a juvenile cranium recently recovered in Drimolen Cave in the Cradle
region of South Africa, ascribed to H. ergaster, has been dated as early as 2.0 Ma,
preceding the first appearance of this species in eastern Africa.
• The next earliest record of H. ergaster in the Cradle region is a specimen from
Swartkrans dated shortly after 1.8 Ma.
• Specimens allied with H. habilis apparently persisted alongside H. ergaster until
~1.4 Ma. Whether these two forms segregated ecologically or represent a single
polytypic species remains debatable.
• Distinctions between them are based mainly on skull shape.
• H. ergaster showed a slow expansion in brain size and progressive reduction in
molar surface area through time, although rather poorly documented.
• It was replaced by H. heidelbergensis, clearly a descendent, around 800 ka, both
in Europe and in Africa.
• This was around the time when the period between glacial extremes lengthened
to around 100 kyr, defining the transition from early to middle Pleistocene when
seasonal extremes of aridity intensified in Africa.
• By that stage the cranial volume of these early humans was almost equal to that
of modern humans. H. heidelbergensis was clearly ancestral to the European
Neanderthals (H. neanderthalensis), but how it connected with early humans
inhabiting Africa remains obscure.
• Genetic evidence suggests that the Neanderthals separated from the human
lineage in Africa around 500 ka.
• They differed from modern humans in their stocky build, prominent brow ridges,
and sloping rather than domed foreheads.
• The earliest fossils sufficiently similar to modern humans to be assigned to the
species Homo sapiens come from a cave in Morocco dated to between 300 and
350 ka.
• These craniums show a reduction in face size, like later humans, but retain fairly
prominent brow ridges and lack the globular shape of the braincase characteristic
of modern humans.
• The Kabwe skull, found in a limestone cave in Zambia, has also been dated to
~300 ka.
• It has been affiliated with H. heidelbergensis due to its lack of more modern
features.
• A partial cranium from the Lake Ndutu region near Olduvai also represents this
time period.
• Slightly more recent is the Florisbad skull from interior South Africa, dated
approximately to ~260 ka and regarded as representing early H. sapiens.
• Partial human skulls from Herto and Omo in Ethiopia and Turkana in Kenya, dated
• Strangely out on an evolutionary limb are the remains of diminutive humans with
small but complexly shaped brains named Homo naledi, which were found deep
in a cave system in South Africa.
• Although their features resemble those of early humans living around 2 Ma, their
hardly fossilised bones date from as recently as ~280 ka.
• While their leg and foot bones were adapted for efficient walking, their hands
retained long fingers, which would have facilitated climbing trees.
• They had smaller but higher-crowned molars with greater wear-resistance than
the dentition of the australopiths, suggesting that abrasive particles were
prominent in their diet.
• They apparently coexisted alongside H. ergaster for well over a million years
without showing up elsewhere in fossil deposits.
• Around 130 ka, humans with modern features appeared in the Levant region of
the Middle East, but did not persist there for very long.
• This was around the beginning of the relatively moist interglacial period that
defines the transition into the late Pleistocene.
• The major dispersal of modern humans beyond Africa through the Middle East
and onwards occurred ~60 ka.
• These people evidently followed a coastal route through tropical Asia, arriving in
Australia ~55 ka. By 47 ka they had spread westward into Europe, displacing the
Neanderthals within a few thousand years.
• Fossils document the dental trend that australopiths showed towards enlarged
molars of low relief with thickened enamel, enabling them to chew and chew,
while their canine teeth became reduced so as not to obstruct grinding.
• Au. anamensis had initiated this trend by 4 Ma and it became accentuated in
later australopiths.
• Molar enlargement reached its epitome in the Paranthropus forms, which
possessed massive jaws with huge molars.
• These dental features seem adapted especially for processing the ‘fallback’ foods
providing subsistence through the dry season, rather than the staples forming the
bulk of the diet during the wet season, which were probably still fruits, seeds and
leaves.
• Nevertheless, greater chewing capabilities might also have been helpful for
breaking open fruits with hard shells, like monkey oranges and baobab pods.
• Eight major adaptive transitions can be recognised in the lineage linking some forest-
dwelling ape to modern humans:
• 1. The emergence of the distinct genus Ardipithecus with features facilitating
bipedal locomotion, around 5.7 Ma.
• 2. The transformation of Ardipithecus into Australopithecus with greater bipedal
competence, by 3.5 Ma.
• 3. The separation of Homo, with more generalised dentition supported by a stone tool kit,
from Paranthropus with dental specialisations for chewing tough foods, around 2.8 Ma.
• 4. The appearance of H. erectus/ergaster with fully competent walking and substantially
bigger brain, accompanied by shaped stone implements, around 1.7 Ma.
• 5. The surge in brain volume towards that of modern humans via H. heidelbergensis,
beginning around 0.8 Ma.
• 6. The establishment of a finely crafted tool kit, plus other cultural artefacts, by big-brained
‘modern’ H. sapiens, between 600 and 200 ka.
• 7. The development of projectile points launched from bows and daubed with poison in
finely elaborated tool kits around 70 ka, predisposing the expansion of humans beyond
Africa.
• It is apparent that these adaptive shifts tended to be coupled with times of major
environmental transitions:
• (1) establishment of seasonally dry savannas between 12 and 5 Ma,
• (2) expansion of C4 grasslands during this period,
• (3) establishment of recurrent glacial advances bringing greater aridity around 2.6
Ma,
• (4) tectonic disruptions accentuating local aridity ~1.8 Ma,
• (5) widening climatic oscillations after 0.8 Ma,
• (6) extremes of aridity during the last-but-one glacial maximum 140 ka, and
• (7) amelioration of local aridity during the Holocene after 20 ka.
• It is intriguing that there are blanks in the fossil record just around the times when
some of these transitions took place.
• Unconformities in sedimentary layers indicate conditions either too arid to form
deposits or so wet as to wash away sediments.
• Puzzlingly, some of these adaptive advances seem to precede, rather than follow