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The Origin and Evolution of Mammals - Moodle

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etween Primates, Dermoptera (colugos), <strong>and</strong><br />

Sc<strong>and</strong>entia (tree shrews), which formed part <strong>of</strong> the<br />

old supraordinal taxon Archonta. However, the<br />

anatomy <strong>of</strong> neither fossil nor living forms has ever<br />

<strong>of</strong>fered support for either excluding the bats, or<br />

including the rodents <strong>and</strong> lagomorphs with them,<br />

as is now indicated by molecular evidence.<br />

Primates<br />

<strong>The</strong> fossil record <strong>of</strong> primates (Fig. 7.23) is notoriously<br />

very poor because they are largely arboreal,<br />

forest-dwelling animals, <strong>and</strong> the early members at<br />

least were all small in size (Taveré et al. 2002). On<br />

the other h<strong>and</strong>, the degree <strong>of</strong> effort devoted to finding<br />

them <strong>and</strong> the level <strong>of</strong> attention given to their<br />

interpretation tends to mitigate their sparseness.<br />

<strong>The</strong> Plesiadapiformes (page 232) used to be<br />

regarded as the most primitive primates, indicating<br />

a very early, immediately post-Cretaceous origin<br />

for the order, followed by a radiation through the<br />

Palaeocene resulting in at least five plesiadapiform<br />

families, distributed in North America <strong>and</strong> Europe.<br />

However, the lack <strong>of</strong> such fundamental primate<br />

characters as a postorbital bar, shortened snout, or<br />

opposable hallux <strong>and</strong> pollex eventually led most<br />

authors to exclude Plesiadapiformes from the<br />

order, interpreting them instead as basal archontans,<br />

with the possible relationship to Dermoptera<br />

mentioned later (p272). <strong>The</strong> issue has re-emerged<br />

recently with the description by Block <strong>and</strong> Boyer<br />

(2003a,b) <strong>of</strong> a more or less complete skeleton <strong>of</strong> the<br />

carpolestid plesiadapiform Carpolestes (Fig. 7.5(b)),<br />

showing it does in fact possess opposable, nailbearing<br />

first digits. <strong>The</strong>ir phylogenetic analysis has<br />

carpolestids as the sister-group <strong>of</strong> what they term<br />

the Euprimates, <strong>and</strong> they conclude that plesiadapiforms<br />

ought to be included in the order Primates<br />

after all. Kirk et al. (2003) are unconvinced <strong>and</strong><br />

believe that the postcranial similarities are convergences<br />

in two separate lineages <strong>of</strong> arboreal euarchontan<br />

mammals.<br />

Living primates fall into two informal categories.<br />

<strong>The</strong> ‘prosimians’ are the primitive primates <strong>and</strong><br />

include lemurs, galagos, lorises, <strong>and</strong> tarsiers. <strong>The</strong>y<br />

retain a series <strong>of</strong> ancestral characters, including a<br />

small brain, a long snout, <strong>and</strong> a postorbital bar that<br />

is not exp<strong>and</strong>ed to enclose the back <strong>of</strong> the orbit. <strong>The</strong><br />

incisors are rounded rather than spatulate <strong>and</strong><br />

LIVING AND FOSSIL PLACENTALS 269<br />

the symphysis <strong>of</strong> the m<strong>and</strong>ible is unfused. <strong>The</strong><br />

anthropoids are advanced primates <strong>and</strong> consist <strong>of</strong><br />

the monkeys <strong>and</strong> apes. ‘Prosimii’ is a paraphyletic<br />

group because it is based entirely on ancestral characters;<br />

using derived characters demonstrates that<br />

some prosimians are basal to the anthropoids. Two<br />

monophyletic groups are now recognised. <strong>The</strong><br />

lemurs, galagos, <strong>and</strong> lorises are the Strepsirhini,<br />

characterised by the presence <strong>of</strong> a grooming toothcomb<br />

formed from the compressed lower incisors<br />

<strong>and</strong> canines. <strong>The</strong> tarsiers <strong>and</strong> anthropoids are the<br />

Haplorhini, characterised, amongst other features,<br />

by loss <strong>of</strong> the moist rhinarium on the nose <strong>and</strong><br />

cleft upper lip, loss <strong>of</strong> the tapetum lucidum, <strong>and</strong><br />

presence <strong>of</strong> a haemochorial placenta.<br />

‘Prosimian’ grade primates first appear in the<br />

earliest Eocene, <strong>and</strong> there followed immediately<br />

an Eocene radiation throughout the northern<br />

continents, that produced several distinct lineages.<br />

<strong>The</strong> relationships <strong>of</strong> the Eocene to the modern<br />

‘prosimian’ groups are far from clear, but most<br />

authors accept that both the strepsirhines <strong>and</strong> the<br />

haplorhines were represented, <strong>and</strong> that the latter<br />

had already divided into the tarsier <strong>and</strong> the anthropoid<br />

lineages (Covert 2002). All were relatively<br />

small, <strong>and</strong> arboreal, <strong>and</strong> variously had insectivorous<br />

<strong>and</strong> frugivorous diets.<br />

Adapiformes are first represented by dentitions<br />

<strong>and</strong> partial skulls <strong>of</strong> Cantius (Fig. 7.23b), from the<br />

earliest Eocene <strong>of</strong> Europe <strong>and</strong> North America. <strong>The</strong>y<br />

had unspecialised incisors <strong>and</strong> non-shearing<br />

molars, suggesting a primarily frugivorous diet.<br />

Later forms were generally quite large for Eocene<br />

primates. Notharctus, for example, had an estimated<br />

weight <strong>of</strong> 7 kg (Gebo 2002) <strong>and</strong> proportions resembling<br />

a lemur. A number <strong>of</strong> characters suggest that<br />

adapiforms are strepsirhines (Kay et al. 1997; Ross<br />

et al. 1998). <strong>The</strong>se include the lemuriform-like structure<br />

<strong>of</strong> the ear region, with a ring-shaped ectotympanic<br />

bone within the tympanic cavity, <strong>and</strong><br />

similarities in the bones <strong>of</strong> the h<strong>and</strong> <strong>and</strong> the foot.<br />

<strong>The</strong> Omomyoidea are the second group <strong>of</strong> primates<br />

to occur in the Early Eocene <strong>of</strong> Europe <strong>and</strong><br />

North America. Teilhardina was only about 70 g in<br />

estimated body weight, <strong>and</strong> no omomyoid was<br />

larger than a squirrel. <strong>The</strong>ir unspecialised dentition<br />

suggests that insects formed an important part <strong>of</strong><br />

the diet, <strong>and</strong> at least some members <strong>of</strong> the family

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