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The Origin and Evolution of Mammals - Moodle

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Extinct Notoryctemorphia<br />

At present the sole fossil relatives <strong>of</strong> the marsupial<br />

mole are teeth <strong>and</strong> isolated skeletal elements from<br />

the Riversleigh deposits. <strong>The</strong> molar teeth are not as<br />

extremely modified as those <strong>of</strong> Notoryctes<br />

(Fig. 6.1(g)), <strong>and</strong> when a description is published it<br />

may indicate how the notoryctid-like version <strong>of</strong> the<br />

zalambdodont condition evolved from a basic marsupial<br />

tooth (Archer et al. 1991).<br />

Extinct Diprotodontia<br />

As the principal herbivorous Order <strong>of</strong> Australian<br />

marsupials, diprotodonts are, as expected, the most<br />

abundant <strong>and</strong> diverse component <strong>of</strong> the fossil, as<br />

well as <strong>of</strong> the living, fauna. In fact, the family level<br />

diversity <strong>of</strong> the group at their first appearance in<br />

the Late Oligocene <strong>and</strong> through the Early Miocene<br />

exceeds that <strong>of</strong> the present day. Of the total <strong>of</strong> 16<br />

diprotodont families recognised by McKenna <strong>and</strong><br />

Bell (1997), only two are not known during this<br />

period, the monospecific Tarsipedidae (honey<br />

possum), <strong>and</strong> dispecific Acrobatidae (feather-tailed<br />

glider <strong>and</strong> feather-tailed possum). On the other<br />

h<strong>and</strong>, eight <strong>of</strong> the mid-Cenozoic families have<br />

subsequently disappeared. <strong>The</strong>se extinct families<br />

include some very peculiar <strong>and</strong> highly specialised<br />

diprotodonts.<br />

Wynyardiidae. Members <strong>of</strong> this family only occur<br />

in the Late Oligocene/Early Miocene, although they<br />

were fairly abundant during this time. Two complete<br />

skeletons <strong>of</strong> Muramura (Fig. 6.12(a)) from the<br />

Etadunna Formation <strong>of</strong> the Lake Eyre Basin have<br />

been found (Pledge 1987), so it is a particularly<br />

well-known group. Muramura was dog-sized <strong>and</strong> its<br />

unspecialised syndactyl hindfeet indicate a simple,<br />

terrestrial quadrupedal locomotion. <strong>The</strong> molars had<br />

a primitive degree <strong>of</strong> development <strong>of</strong> transverse<br />

lophs, <strong>and</strong> the stylar cusps on the uppers are still<br />

distinguishable. As far as the details <strong>of</strong> the wynyardiid<br />

braincase are concerned, Aplin (1987)<br />

believed it to be the most plesiomorphic <strong>of</strong> all<br />

diprotodonts. <strong>The</strong>refore, on dental, cranial, <strong>and</strong><br />

postcranial structure, the wynardiids appear to be<br />

close to the base <strong>of</strong> the diprotodont radiation.<br />

Diprotodontidae. <strong>The</strong> diprotodontids (Fig. 6.12(i)<br />

<strong>and</strong> (j)) include the largest marsupials ever to<br />

LIVING AND FOSSIL MARSUPIALS 213<br />

evolve, ranging in size from that <strong>of</strong> a sheep to giants<br />

larger than hippos. <strong>The</strong> Late Pleistocene Diprotodon<br />

optatum was almost 3 m long, stood 2.6 m high at the<br />

shoulder, <strong>and</strong> weighed as much as 2.78 tonnes<br />

(Wroe et al. 2003). <strong>The</strong> diprotodontid dentition was<br />

modified for browsing on relatively s<strong>of</strong>t vegetation,<br />

with enlarged, bilophodont molar teeth. Unlike the<br />

other major herbivore family, the macropodids,<br />

diprotodontids remained fully quadripedal. <strong>The</strong><br />

group finally became extinct in the Late Pleistocene,<br />

as part <strong>of</strong> the general megafaunal mass extinction <strong>of</strong><br />

that time.<br />

Palorchestidae. This family consisted <strong>of</strong> forms<br />

quite similar to diprotodontids <strong>and</strong> they were at<br />

one time actually included in the latter family. Also<br />

like the diprotodontids, they survived until the<br />

Late Pleistocene, when the best-known genus<br />

Palorchestes was the size <strong>of</strong> a horse. However,<br />

palorchestids do differ from diprotodontids in<br />

details <strong>of</strong> the dentition, <strong>and</strong> also in the structure <strong>of</strong><br />

the ear region, in which they more closely resemble<br />

the wombats. Very large, powerful claws tended to<br />

develop on the forelimbs suggesting a diet <strong>of</strong> tubers<br />

<strong>and</strong> roots, <strong>and</strong> in the later forms such as Palorchestes<br />

the snout was elongated in a manner that suggests<br />

that it bore a mobile proboscis or short trunk.<br />

Thylacoleonidae. Famously the most extraordinary<br />

diprotodonts <strong>of</strong> all were the marsupial lions, which<br />

are also one <strong>of</strong> the comparatively rare examples <strong>of</strong><br />

the evolution <strong>of</strong> a carnivore from a specialised<br />

herbivorous ancestor. <strong>The</strong> Late Oligocene–Early<br />

Miocene thylacoleonid Priscileo was a relatively<br />

small, cat-sized animal (Rauscher 1987), <strong>and</strong><br />

Wakaleo (Murray et al. 1987) had a 15-cm skull<br />

(Fig. 6.12(f)). By the Plio-Pleistocene, Thylacoleo<br />

(Fig. 6.12(g)) had evolved into the size <strong>and</strong> general<br />

form <strong>of</strong> a leopard. <strong>The</strong> dentition is modified for<br />

extreme carnivory (Nedin 1991), or perhaps a<br />

hyaena-like scavenging that included dealing with<br />

large bones. <strong>The</strong> first upper <strong>and</strong> sole lower incisors<br />

are enlarged <strong>and</strong> caniniform. <strong>The</strong> upper third<br />

premolar (Fig. 6.12(h)) is an enormous shearing<br />

blade that worked against the almost equally large<br />

lower third premolar plus the first molar. <strong>The</strong> rest<br />

<strong>of</strong> the molars are reduced in both number <strong>and</strong><br />

relative size.

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