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The Origin and Evolution of Mammals - Moodle

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186 THE ORIGIN AND EVOLUTION OF MAMMALS<br />

Like insects these also tend to occur as small, discrete<br />

items requiring individual discovery <strong>and</strong> collection.<br />

Small body size is again a prerequisite for<br />

surviving on this diet, but with the complication<br />

that teeth adapted to some degree <strong>of</strong> grinding, <strong>and</strong><br />

hence subjected to a greater degree <strong>of</strong> wear must<br />

evolve. A number <strong>of</strong> Mesozoic mammals occupied<br />

this role, probably the haramiyids <strong>and</strong> the docodontans,<br />

certainly the multituberculates, <strong>and</strong> also the<br />

Late Cretaceous placental zhelestids.<br />

Predaceous carnivory is a mode <strong>of</strong> life that readily<br />

provides enough food, provided there is a source <strong>of</strong><br />

suitable sized prey. During the Mesozoic this would<br />

have been a problem for a large mammal. <strong>The</strong>re<br />

were no medium-sized to large herbivorous mammals,<br />

<strong>and</strong> the dominant herbivores, the dinosaurs,<br />

were mostly too large in body size to be prey items<br />

for all except very large predators indeed. Of the<br />

other terrestrial animals present <strong>and</strong> forming a<br />

potential food source, some tended to be too large<br />

such as crocodiles, <strong>and</strong> others such as lepidosaurs<br />

too small to provide an adequate diet for a middle to<br />

large mammal carnivore. <strong>The</strong>refore, no large carnivorous<br />

mammals evolved. As it happens, the largest<br />

Mesozoic mammals that did exist were the predaceous<br />

members <strong>of</strong> the Eutriconodonta, <strong>and</strong> their<br />

body size <strong>of</strong> up to 1.5 kg is consistent with a diet <strong>of</strong><br />

amphibians, small lizard, <strong>and</strong> other mammals.<br />

<strong>The</strong> fourth basic diet to consider is leaf-eating,<br />

browsing herbivory, the diet followed by the majority<br />

<strong>of</strong> large herbivorous mammals today. This mode<br />

<strong>of</strong> nourishment creates special problems because <strong>of</strong><br />

the relatively low energy content <strong>of</strong> foliage, <strong>and</strong> its<br />

low rate <strong>of</strong> assimilation due to the need for a gut<br />

fermentation flora. Large body size is necessary so<br />

that the basal metabolic rate is relatively less, the<br />

storage capacity <strong>of</strong> the gut is larger, <strong>and</strong> the potential<br />

foraging area is greater. Small mammals cannot<br />

collect or assimilate this kind <strong>of</strong> food at a rate that<br />

is high enough to satisfy their relatively higher<br />

metabolic dem<strong>and</strong>s, a problem exacerbated in a<br />

female committed to lactation.<br />

From these comments, simple as they are, it follows<br />

that the fundamental point was the inability <strong>of</strong><br />

Mesozoic mammals to adapt to the large herbivore<br />

habitat because <strong>of</strong> the absence <strong>of</strong>, or inability to<br />

acquire food at a high enough rate for the endothermic<br />

metabolism. A possible reason for this is a den-<br />

tition unable to cope with the abrasion involved.<br />

As it happens, the first appearance <strong>of</strong> the form <strong>of</strong><br />

enamel found in modern chewing mammals was<br />

in the Early Palaeocene herbivorous mammals.<br />

Koenigswald et al. (1987) showed that the arrangement<br />

<strong>of</strong> the enamel prisms into what are termed<br />

Hunter–Schreger b<strong>and</strong>s was present in certain<br />

condylarths. <strong>The</strong> apparent function <strong>of</strong> this organisation<br />

<strong>of</strong> the enamel is to reduce the tendency <strong>of</strong><br />

teeth to split when actively masticating plant food.<br />

Since similar enamel is found in most large herbivore<br />

mammal taxa thereafter, it must have evolved<br />

convergently several times, implying that it was a<br />

morphological adaptation in response to a common<br />

ecological opportunity in different lineages. Precisely<br />

what that opportunity consisted <strong>of</strong> is unclear.<br />

To conclude, there are two seriously proposed<br />

physiological constraints that might have been<br />

responsible for the persistent small body size <strong>of</strong><br />

Mesozoic mammals: over-heating in the absence<br />

<strong>of</strong> adequate cooling adaptations, <strong>and</strong> undernourishment<br />

in the absence <strong>of</strong> adequate folivorous<br />

adaptations.<br />

End <strong>of</strong> the era: the K/T mass extinction<br />

<strong>and</strong> its aftermath<br />

Sixty-five million years ago, an episode <strong>of</strong> massextinction<br />

affected the world’s biota to the extent<br />

that some 65–75% <strong>of</strong> the species disappeared in,<br />

geologically speaking, an instant. Such events have<br />

occurred every few tens <strong>of</strong> millions <strong>of</strong> years<br />

throughout the history <strong>of</strong> life <strong>and</strong> this one was not<br />

even the largest. That accolade goes to the end-<br />

Permian mass extinction which was experienced by<br />

the therapsid ancestors <strong>of</strong> mammals 250 Ma while<br />

even the Late Triassic mass extinction <strong>of</strong> 210 Ma,<br />

around the time <strong>of</strong> the origin <strong>of</strong> mammals <strong>and</strong><br />

dinosaurs, was at least <strong>of</strong> comparable magnitude<br />

(page 87). Nevertheless, the end-Cretaceous, or<br />

K/T event was the mass-extinction most significant<br />

in the story <strong>of</strong> the origin <strong>and</strong> evolution <strong>of</strong> mammals.<br />

It marks the transition from the ecologically<br />

very limited Mesozoic mammal radiation to the<br />

Tertiary radiation that culminated in today’s mammalian<br />

fauna with its huge variation <strong>of</strong> body size<br />

<strong>and</strong> habitat. Whether the mass extinction acted<br />

by removing the dinosaur competitors <strong>of</strong>

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