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The Origin and Evolution of Mammals - Moodle

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<strong>The</strong> salient feature <strong>of</strong> Widanelfasia is the presence <strong>of</strong><br />

incipiently zalambdodont molar teeth. <strong>The</strong> trigonid<br />

cusps, especially the protoconid are very high <strong>and</strong><br />

form a narrowly acute triangle, <strong>and</strong> the talonid is<br />

much lower. <strong>The</strong> fully zalambdodont molar <strong>of</strong><br />

solenodontids is presumed to be convergent, but<br />

that <strong>of</strong> the tenrecs <strong>and</strong> the rather specialised version<br />

found in chrysochloridans may well be homologous.<br />

It is therefore possible, though by no means<br />

positively demonstrable yet, that Widanelfasia is a<br />

stem afrosoricid. Seiffert <strong>and</strong> Simons (2000) also<br />

point to the upper teeth <strong>of</strong> an Early Eocene primitive<br />

‘insectivore’ from Tunisia, Chambilestes,<br />

described by Gheerbrant <strong>and</strong> Hartenberger (1999)<br />

<strong>and</strong> note that they have a structure compatible with<br />

the lower teeth <strong>of</strong> their Fayum mammal.<br />

Chrysochlorida. <strong>The</strong> golden moles have a very poor<br />

fossil record. It is entirely restricted to Africa, dates<br />

from no earlier than the Early Miocene, <strong>and</strong> consists<br />

<strong>of</strong> specimens that can be placed in the same family<br />

as the nine living genera.<br />

Tenrecida. <strong>The</strong> tenrecs <strong>of</strong> Madagascar along with the<br />

otter shrews <strong>of</strong> sub-Saharan Africa have the poorest<br />

fossil record <strong>of</strong> all the afrotherian orders, consisting<br />

solely <strong>of</strong> zalambdodont teeth <strong>of</strong> an Early Miocene<br />

African form Parageogale (Butler 1984), prior to subfossils<br />

<strong>of</strong> the Pleistocene.<br />

Laurasiatheria<br />

Carnivora<br />

<strong>The</strong> very early appearance <strong>of</strong> members <strong>of</strong> the order<br />

Carnivora has already been described, <strong>and</strong> within the<br />

Early Palaeocene differentiation <strong>of</strong> the two primary<br />

subgroups <strong>of</strong> the order had apparently also occurred,<br />

for one <strong>of</strong> them, Feliformes, is represented by the<br />

viverravids <strong>of</strong> that time (Fig. 7.14(f)). <strong>The</strong> miacids<br />

(Fig. 7.14(e)), which appear in the Late Palaeocene,<br />

are basal members <strong>of</strong> the other group, Caniformes<br />

(Flynn 1998; Janis et al. 1998b). Both <strong>of</strong> these primitive<br />

groups, consisting <strong>of</strong> relatively small animals, radiated<br />

through the first half <strong>of</strong> the Eocene, during<br />

which period it was the creodonts that provided the<br />

larger carnivorous mammals.<br />

<strong>The</strong> family Nimravidae were the earliest group <strong>of</strong><br />

modern-style Feliformes, a group specialised for<br />

LIVING AND FOSSIL PLACENTALS 259<br />

pure carnivory <strong>and</strong> convergent in many features on<br />

the true cats, the Felidae (Martin 1998a). <strong>The</strong>y<br />

appeared in the Late Eocene <strong>of</strong> North America <strong>and</strong><br />

Eurasia, <strong>and</strong> radiated primarily during the<br />

Oligocene; the last survivor, Barbour<strong>of</strong>elis (Fig.<br />

7.18(d)), disappeared no later than the Late<br />

Miocene <strong>of</strong> North America. Nimravids, like felids,<br />

had short, powerful skulls, highly developed carnassial<br />

teeth <strong>and</strong> reduction <strong>of</strong> the more posterior<br />

molars. <strong>The</strong>ir postcranial skeleton indicates that<br />

they were long-legged, cursorial predators with<br />

feline-like claws. Even more markedly than in the<br />

felids, nimravids had a tendency to evolve sabretoothed<br />

forms. <strong>The</strong> early, leopard-sized<br />

Hoplophoneus (Fig. 7.18(c)) already had evolved<br />

enlarged canines. <strong>The</strong> trend is most dramatically<br />

expressed in the lion-sized Barbour<strong>of</strong>elis (Turner <strong>and</strong><br />

Antón 1997).<br />

<strong>The</strong> first true cat, the felid Proailurus (Fig.<br />

7.18(b)), is well known from the Oligocene <strong>and</strong><br />

Early Miocene <strong>of</strong> Europe. It was the size <strong>of</strong> a modern<br />

ocelot <strong>and</strong> its relatively short, robust limbs <strong>and</strong><br />

retractile claws indicate an accomplished arboreal<br />

animal (Agustí <strong>and</strong> Antón 2002). <strong>The</strong> major radiation<br />

<strong>of</strong> felids commenced in the Late Miocene<br />

(Martin 1998b), including their entry into North<br />

America, where they rapidly replaced the nimravids<br />

as the top carnivores. A number <strong>of</strong> genera<br />

independently evolved into sabre-toothed cats,<br />

most famously Smilodon, huge numbers <strong>of</strong> which<br />

are preserved in the Rancho La Brea asphalt pits <strong>of</strong><br />

Los Angeles <strong>and</strong> which survived until the end-<br />

Pleistocene about 10,000 years ago.<br />

<strong>The</strong> other feliform families, Viverridae (civets),<br />

Herpestidae (mongooses), <strong>and</strong> Hyaenidae also<br />

commenced their radiations during the Miocene,<br />

<strong>and</strong> apart from a single Pliocene hyaenid, were<br />

restricted to Eurasia <strong>and</strong> Africa. This is in inexplicable<br />

contrast to the Felidae, which were abundant<br />

in North America.<br />

<strong>The</strong> second branch <strong>of</strong> the Carnivora are the<br />

Caniformes, all the families <strong>of</strong> which do occur in<br />

the fossil record <strong>of</strong> North America, as well as<br />

Eurasia <strong>and</strong> Africa. As mentioned, the earliest<br />

caniniforms are believed to be the Late Palaeocene<br />

miacids <strong>of</strong> North America <strong>and</strong> Europe. <strong>The</strong> modern<br />

caniform families appeared at various times after<br />

this, in a complex biogeographic pattern. Canids

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