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The Origin and Evolution of Mammals - Moodle

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form <strong>of</strong> the tooth, must reflect subtle differences in<br />

niche amongst this new, spectacular richness <strong>of</strong><br />

entomological resources, comparable to what is<br />

seen in the insectivorous mammals <strong>of</strong> today.<br />

Thus the Late Cretaceous became dominated by<br />

two mammalian groups, the multituberculates <strong>and</strong><br />

tribosphenidans. However, the vast preponderance<br />

<strong>of</strong> information about Mesozoic mammal evolution<br />

comes from the Laurasian continents <strong>of</strong> North<br />

America, Europe, <strong>and</strong> Asia, <strong>and</strong> to a small degree<br />

North Africa. It would be misleading to assume<br />

that the events outlined above also involved the<br />

southern continents. Indeed, evidence is accruing<br />

that it would be just plain wrong. Prior to the Late<br />

Cretaceous, close to nothing is known at present,<br />

apart from the pitifully few australosphenidans.<br />

Late Cretaceous mammals occur in the Los Alamitos<br />

Formation <strong>of</strong> Argentina, <strong>and</strong> are composed entirely<br />

<strong>of</strong> non-tribosphenidan mammals. <strong>The</strong>re are indigenous<br />

groups <strong>of</strong> eutriconodontans, symmetrodontans,<br />

dryolestidans eupantotheres, <strong>and</strong> the strange,<br />

hypsodont teeth <strong>of</strong> the uniquely Gondwanan group<br />

Gondwanatheria. Presumably this mammalian<br />

fauna evolved in complete isolation from the familiar<br />

northern fauna, <strong>and</strong> no southern tribosphenidan<br />

is known for certain prior to the Cenozoic.<br />

<strong>The</strong> general biology <strong>of</strong> the Mesozoic<br />

mammals<br />

<strong>The</strong> cladistic relationships <strong>of</strong> the non-mammalian<br />

cynodonts indicates quite clearly that the origin <strong>of</strong><br />

mammals during the Upper Triassic involved<br />

amongst other things fairly extreme miniaturisation.<br />

Sinoconodon was the largest <strong>of</strong> the early forms, with<br />

a skull length up to 6 cm, which is roughly the size<br />

<strong>of</strong> the skull <strong>of</strong> the European hedgehog. <strong>The</strong> postcranial<br />

skeleton is unknown, but, continuing the<br />

comparison with the hedgehog, it would be<br />

expected to have been about 25 cm in presacral<br />

length, <strong>and</strong> to have weighed about 800g. It may<br />

also be <strong>of</strong> significance that Sinoconodon retained the<br />

primitive condition <strong>of</strong> continuous growth, rather<br />

than the typically mammalian pattern where growth<br />

is restricted to a relatively brief juvenile phase. <strong>The</strong><br />

great majority <strong>of</strong> the rest <strong>of</strong> the Mesozoic mammals<br />

were considerably smaller, being <strong>of</strong> the general<br />

body size <strong>of</strong> soricid shrews <strong>and</strong> murid mice <strong>and</strong><br />

THE MESOZOIC MAMMALS 183<br />

rats, with body weights in the range <strong>of</strong> 200 g down<br />

to the 3 g <strong>of</strong> a minute modern shrew such as<br />

Microsorex hoyi (Eisenberg 1981). <strong>The</strong> docodontan<br />

Henkelotherium is typical. It had a skull length <strong>of</strong><br />

4 cm, <strong>and</strong> a presacral body length <strong>of</strong> 11 cm, which<br />

is roughly equivalent to a body weight <strong>of</strong> about 20 g.<br />

<strong>The</strong> smallest Mesozoic mammal so far described is<br />

the sub-adult specimen <strong>of</strong> Hadrocodium, which has<br />

a skull length <strong>of</strong> only 1.5 cm <strong>and</strong> an estimated body<br />

weight <strong>of</strong> 2 g (Luo et al. 2001b). <strong>The</strong>re were also a<br />

few relative ‘giants’. <strong>The</strong> largest are the Early<br />

Cretaceous eutriconodontans such as Gobiconodon<br />

ostromi. This had a 10 cm skull <strong>and</strong> a 35 cm presacral<br />

body length, giving it a body size roughly<br />

comparable to Didelphis virginiana, a mammal<br />

whose body weight is around 1.5 kg. A few <strong>of</strong> the<br />

advanced tribosphenidan mammals <strong>of</strong> the Late<br />

Cretaceous also achieved somewhat increased<br />

body size. <strong>The</strong> largest is probably Zalambdalestes<br />

lechei, but even this was reconstructed by Kielan-<br />

Jaworowska (1978) with a skull length <strong>of</strong> no more<br />

than 4.6 cm, <strong>and</strong> a presacral body length <strong>of</strong> 14.9 cm.<br />

<strong>The</strong> hypothetical ancestral mammal, as well as<br />

being small, is universally agreed to have been<br />

insectivorous, endothermic, <strong>and</strong> nocturnal. <strong>The</strong><br />

inference that its primary diet consisted <strong>of</strong> small,<br />

terrestrial invertebrates is based on the sharpcusped,<br />

shearing nature <strong>of</strong> the postcanine teeth,<br />

which were unsuitable for dealing with any significant<br />

volume <strong>of</strong> plant material. <strong>The</strong> hypothesis that<br />

it was endothermic has been considered at length<br />

earlier, <strong>and</strong> it is difficult to imagine that the whole<br />

integrated suite <strong>of</strong> mammalian morphological characters<br />

was not associated with insulation <strong>and</strong> a<br />

high metabolic rate. Preservation <strong>of</strong> the impression<br />

<strong>of</strong> the pelt <strong>of</strong> the Early Cretaceous placental Eomaia,<br />

<strong>and</strong> <strong>of</strong> hair <strong>of</strong> a Late Palaeocene multituberculate,<br />

powerfully confirms this belief. Nocturnality is<br />

more speculative, but is implied by the form <strong>of</strong> the<br />

brain <strong>and</strong> sense organs. <strong>The</strong> enlargement <strong>of</strong> the<br />

cerebrum to become the primary control centre in<br />

mammals, at the expense <strong>of</strong> the optic lobes, suggests<br />

that the importance <strong>of</strong> the sense <strong>of</strong> olfaction<br />

found in modern nocturnal insectivores was<br />

already present. <strong>The</strong> increased sensitivity <strong>of</strong> the<br />

sense <strong>of</strong> hearing, as witnessed by the evolution <strong>of</strong><br />

the ear ossicle system may also point to nocturnality,<br />

although it has to be said that amongst other

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