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The Origin and Evolution of Mammals - Moodle

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272 THE ORIGIN AND EVOLUTION OF MAMMALS<br />

<strong>The</strong> other modern anthropoid group are the<br />

Catarrhini, the Old World monkeys <strong>and</strong> the apes.<br />

Several primitive members occur in the Late<br />

Eocene levels <strong>of</strong> the Fayum deposits, such as<br />

Catopithecus, for which there are several skulls preserved<br />

(Simons <strong>and</strong> Rasmussen 1996). It combines<br />

derived catarrhine characters <strong>of</strong> the dentition,<br />

including a reduction from three to two premolars<br />

<strong>and</strong> fused symphysis, with primitive features <strong>of</strong> the<br />

skull such as the relatively small brain <strong>and</strong> elongated<br />

snout. Aegyptopithecus is the most familiar<br />

basal catarrhine, coming from the slightly younger<br />

Early Oligocene level in the Fayum sequence. It<br />

was the size <strong>of</strong> a typical monkey, well adapted for<br />

arboreal life, <strong>and</strong> the rounded cusps <strong>of</strong> the molar<br />

teeth indicate a diet <strong>of</strong> fruit. <strong>The</strong>se basal catarrhines<br />

are conveniently grouped as the Propliopithecoidea,<br />

despite presumably being a paraphyletic group<br />

defined only by primitive characters, <strong>and</strong> containing<br />

the ancestors <strong>of</strong> the later anthropoid taxa<br />

(Rasmussen 2002). As with the platyrrhines, there is<br />

no agreement about precisely what the relationship<br />

is between the known basal anthropoids <strong>and</strong> the<br />

catarrhines.<br />

<strong>The</strong>re is a considerable gap in the fossil record<br />

between the Early Oligocene basal catarrhines such<br />

as Aegyptopithecus, <strong>and</strong> the first occurrences <strong>of</strong> the<br />

derived lineages. <strong>The</strong> Old World monkeys,<br />

Cercopithecoidea, are unknown until the Early<br />

Miocene <strong>of</strong> Africa, when the Kenyan Victoriapithecus<br />

is found. <strong>The</strong> first <strong>of</strong> the apes, Hominoidea, may be<br />

slightly earlier if the tooth-bearing maxillary fragment<br />

referred to as Kamoyapithecus (Leakey et al.<br />

1995) is correctly identified <strong>and</strong> correctly dated as<br />

Late Oligocene. Otherwise, hominoids too commence<br />

their radiation in the Early Miocene <strong>of</strong> Africa<br />

in the form <strong>of</strong> the well-known genus Proconsul<br />

(Harrison 2002). By the Middle Miocene, radiations<br />

<strong>of</strong> hominoids were also well under way in Asia <strong>and</strong><br />

Europe. <strong>The</strong> third derived catarrhine group are the<br />

Pliopithecoidea, an exclusively Middle <strong>and</strong> Late<br />

Miocene group restricted to Europe <strong>and</strong> Asia.<br />

During their brief radiation, a range <strong>of</strong> size <strong>and</strong><br />

adaptations comparable to that <strong>of</strong> the Old World<br />

monkeys in Africa evolved (Begun 2002).<br />

<strong>The</strong> story <strong>of</strong> the origin <strong>and</strong> evolution <strong>of</strong> the<br />

Hominidae, the australopithecines <strong>and</strong> hominines<br />

<strong>of</strong> Africa <strong>and</strong> their subsequent worldwide dispersal<br />

has been told so many times that it does not need<br />

repeating here. As far as mammalian radiation is<br />

concerned, the hominids were a very minor group<br />

<strong>of</strong> trivial significance, prior to the ecological effect<br />

<strong>of</strong> one species, Homo sapiens, since the Late<br />

Pleistocene a few thous<strong>and</strong> years ago.<br />

Dermoptera<br />

<strong>The</strong> sole living genus <strong>of</strong> the order Dermoptera is<br />

the colugo or flying lemur Cynocephalus, although<br />

despite this paucity, there is fossil evidence for a<br />

very long, if not very diverse history <strong>of</strong> the group.<br />

<strong>The</strong> dentition is particularly distinctive. <strong>The</strong>re is a<br />

gap between the front <strong>of</strong> the upper tooth rows, <strong>and</strong><br />

the two pairs <strong>of</strong> lower incisors have a unique,<br />

comb-like construction, consisting <strong>of</strong> many separate<br />

‘tines’. <strong>The</strong> cheek teeth are wide <strong>and</strong> have a<br />

curiously wrinkled enamel that creates a series <strong>of</strong><br />

sharp edges after a certain amount <strong>of</strong> wear. <strong>The</strong><br />

postcranial skeleton is also uniquely designed for<br />

supporting a gliding membrane stretched between<br />

all four limbs. On the basis <strong>of</strong> these characters, a<br />

fully evolved fossil dermopteran, Dermotherium, is<br />

recognisable in the Lower Eocene <strong>of</strong> Thail<strong>and</strong><br />

(Ducrocq et al. 1992). <strong>The</strong>re are much earlier specimens<br />

sometimes regarded as members <strong>of</strong> the order,<br />

namely the plagiomenids that occurred from the<br />

Early Palaeocene to Middle Eocene <strong>of</strong> North<br />

America. <strong>The</strong> lower dentition <strong>of</strong> a form such as<br />

Plagiomene certainly resembles that <strong>of</strong> Cynocephalus<br />

(Rose <strong>and</strong> Simons 1977), although others doubt the<br />

relationship (MacPhee et al. 1989).<br />

Dermoptera have been linked by several authors to<br />

the plesiadapiforms, the group <strong>of</strong> mainly Palaeocene<br />

forms sometimes regarded as basal primates. Kay et<br />

al. (1990) described similarities in the ear region, <strong>and</strong><br />

Szalay <strong>and</strong> Lucas (1993) detected homologies in the<br />

postcranial skeleton <strong>of</strong> the two groups. Other authors<br />

have, however, disputed the association <strong>of</strong> any, let<br />

alone all the plesiadapiforms with the Dermoptera,<br />

regarding the similarities as merely superficial,<br />

or primitive for the orders (Wible 1993; Bloch <strong>and</strong><br />

Silcox 2001).<br />

Sc<strong>and</strong>entia<br />

<strong>The</strong> tree shrews, at various times confidently classified<br />

as Insectivora, <strong>and</strong> as basal, virtually ancestral<br />

primates, are a distinct mammalian order, but one

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