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The Origin and Evolution of Mammals - Moodle

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urrowing <strong>and</strong> subsisting on a diet <strong>of</strong> subterranean<br />

roots <strong>and</strong> tubers. <strong>The</strong> other surviving therapsid taxa<br />

were small in size, <strong>and</strong> may have existed initially in<br />

less harsh habitats such as upl<strong>and</strong>s <strong>and</strong> fed primarily<br />

on insects. Of particular interest, the one known<br />

<strong>and</strong> the one inferred lineages <strong>of</strong> cynodonts that<br />

lived through the end-Permian event may perhaps<br />

have owed their survival to a higher level <strong>of</strong> homeostasis<br />

combined with small body size, <strong>and</strong> possibly<br />

burrowing (Damiani et al. 2003).<br />

<strong>The</strong> Triassic decline <strong>of</strong> the <strong>The</strong>rapsida<br />

<strong>The</strong> vegetation soon recovered from the effects <strong>of</strong><br />

the extinction <strong>and</strong> during the Lower Triassic an<br />

ecosystem basically structured like that <strong>of</strong> the<br />

Permian had been re-established. <strong>The</strong>re was an<br />

increase in diversity <strong>of</strong> various primitive conifer<br />

groups, but seed ferns remained abundant. On the<br />

whole, the flora reflected a warm, seasonal climate<br />

(Behrensmeyer et al. 1992). Of the therapsid survivors<br />

<strong>of</strong> the end-Permian mass extinction, only<br />

two lineages were <strong>of</strong> much significance. One is the<br />

kannemeyeriid dicynodonts <strong>and</strong> the other the<br />

eucynodontian cynodonts. Both groups radiated<br />

through the Lower <strong>and</strong> Middle Triassic worldwide,<br />

but therapsids never again dominated as they had<br />

in the Late Permian. <strong>The</strong>y shared the terrestrial<br />

amniote world from the Lower Triassic onwards<br />

with increasing numbers <strong>of</strong> diapsid reptiles,<br />

notably the herbivorous rhynchosaurs <strong>and</strong> a variety<br />

<strong>of</strong> thecodontan archosaurs. <strong>The</strong>n during the<br />

Upper Triassic, they disappeared. Correlation <strong>of</strong> the<br />

ages <strong>of</strong> terrestrial Triassic strata is notoriously difficult,<br />

but the main extinction appears to have<br />

occurred during or at the end <strong>of</strong> the Carnian stage,<br />

by about 220 Ma. This included the disappearance<br />

<strong>of</strong> the kannemeyeriids <strong>and</strong> the chiniquodontid<br />

EVOLUTION OF MAMMAL-LIKE REPTILES 87<br />

cynodonts, while the herbivorous traversodontid<br />

cynodonts were greatly reduced in diversity,<br />

although a few occur in the following Norian Stage<br />

(Benton 1994).<br />

<strong>The</strong>re has been a long dispute about whether the<br />

disappearance <strong>of</strong> the therapsids <strong>and</strong> the radiation<br />

<strong>of</strong> the dinosaurs in the Upper Triassic resulted from<br />

competitive interactions in which the dinosaurs<br />

were somehow superior, or from a change in the<br />

environment that caused a decline in therapsids <strong>and</strong><br />

allowed an opportunistic radiation <strong>of</strong> dinosaurs into<br />

the large terrestrial tetrapod niches (Benton 1986).<br />

<strong>The</strong> traditional view was the competitive one, <strong>and</strong> a<br />

number <strong>of</strong> authors speculated on what might have<br />

been the basis for the dinosaurs’s better general<br />

adaptation. Charig (1984), for example, believed<br />

that dinosaurs simply evolved a more effective<br />

locomotory ability, while Bakker (1968) argued that<br />

dinosaurs evolved endothermy before the mammallike<br />

reptiles. Benton (1986, 1994) argued forcefully<br />

that there was actually a significant mass extinction<br />

in the late Carnian, correlated with an environmental<br />

change. <strong>The</strong>re is evidence for an increase in<br />

the extinction rate <strong>of</strong> plants at least approximately at<br />

the same time (Boulter et al. 1988; Simms et al. 1994).<br />

From this perspective, the decline <strong>of</strong> the therapsids<br />

resulted from the environmental change associated<br />

with this extinction event, <strong>and</strong> the increase in<br />

dinosaur diversity from the Norian onwards<br />

occurred as the latter opportunistically invaded the<br />

habitats vacated by the former.<br />

Whatever the truth <strong>of</strong> the matter, the last part <strong>of</strong> the<br />

Triassic record <strong>of</strong> therapsids consisted only <strong>of</strong> the<br />

highly specialised tritylodontids, the tritheledontans<br />

<strong>and</strong> the small, relatively rare, <strong>and</strong> insignificant mammals<br />

(Lucas <strong>and</strong> Hunt 1994). <strong>The</strong> phase <strong>of</strong> synapsid<br />

evolution represented by the Mesozoic mammals<br />

was about to commence.

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