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The Origin and Evolution of Mammals - Moodle

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transversely occluding molar teeth that functioned<br />

in the unique manner <strong>of</strong> basic living mammals, <strong>and</strong><br />

there is a new jaw hinge between the dentary bone<br />

<strong>of</strong> the lower jaw <strong>and</strong> the squamosal <strong>of</strong> the skull that<br />

permitted a much stronger bite. Such animals were<br />

undoubtedly mammalian in the biological sense<br />

although not strictly members <strong>of</strong> the crown-group<br />

Mammalia. <strong>The</strong> most primitive <strong>of</strong> these forms is<br />

called Sinoconodon, <strong>and</strong> most palaeobiologists believe<br />

that they should formally be members <strong>of</strong> the clade<br />

designated Mammalia (e.g. Crompton <strong>and</strong> Sun<br />

1985; Kielan-Jaworowska et al. 2004). <strong>The</strong> definition<br />

<strong>of</strong> Mammalia thus becomes: synapsids that possess<br />

a dentary-squamosal jaw articulation <strong>and</strong> occlusion<br />

between lower <strong>and</strong> upper molars with a transverse<br />

component to the movement. This has exactly the<br />

same membership as the clade that includes the<br />

common ancestor <strong>of</strong> Sinoconodon, living mammals,<br />

<strong>and</strong> all its descendants.<br />

This is the concept <strong>of</strong> a mammal that is used in<br />

the chapters that follow, in the belief that conceding<br />

a degree <strong>of</strong> subjectivity in the choice <strong>of</strong> what is a<br />

mammal is a small price to pay for allowing the<br />

focus <strong>of</strong> the work to be on the origin <strong>and</strong> evolution<br />

<strong>of</strong> the quality <strong>of</strong> ‘mammalness’ just as much as on<br />

the origin <strong>and</strong> evolution <strong>of</strong> the taxon Mammalia.<br />

A sketch <strong>of</strong> the plot<br />

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

as told by the fossil record falls into three distinct<br />

phases, the first <strong>of</strong> which led to the origin <strong>of</strong> mammals<br />

as such. <strong>The</strong> lineage <strong>of</strong> amniotes that culminated<br />

in the mammals made its first appearance in<br />

the fossil record <strong>of</strong> the Pennsylvanian (Upper<br />

Carboniferous) about 305 Ma. <strong>The</strong> best-known<br />

form from this time is Archaeothyris, which existed<br />

very soon after the initial appearance <strong>of</strong> the terrestrially<br />

adapted amniote animals. From a hypothetical<br />

ancestor comparable to Archaeothyris, the<br />

radiation <strong>of</strong> mammal-like reptiles commenced <strong>and</strong><br />

occupied the succeeding 100 Ma. Taxa appeared,<br />

flourished, <strong>and</strong> went extinct in a complex kaleidoscopic<br />

pattern <strong>of</strong> successive groups <strong>of</strong> small-,<br />

medium-, <strong>and</strong> large-sized carnivores <strong>and</strong> herbivorous<br />

groups, which over time exhibited increasing<br />

numbers <strong>of</strong> mammalian characters superimposed<br />

on their particular specialisations. In the latest<br />

INTRODUCTION 3<br />

Triassic Period, 210 Ma, one particular lineage <strong>of</strong><br />

small carnivores culminated in the first mammal.<br />

<strong>The</strong> second phase <strong>of</strong> the unfolding story consists<br />

<strong>of</strong> what are referred to as the Mesozoic mammals.<br />

<strong>The</strong>se were the numerous subgroups that radiated<br />

from the mammalian ancestor during the 140 Ma<br />

duration <strong>of</strong> the Jurassic <strong>and</strong> Cretaceous Periods, the<br />

time during which the world’s terrestrial fauna was<br />

dominated by the dinosaurs. A good deal <strong>of</strong> evolution,<br />

especially dental evolution occurred, but none<br />

<strong>of</strong> them ever evolved the medium or large body<br />

size found in so many modern mammals. For twothirds<br />

<strong>of</strong> the whole <strong>of</strong> their history, mammals<br />

remained small animals with the largest being<br />

barely larger than a cat, <strong>and</strong> the vast majority <strong>of</strong> the<br />

size <strong>of</strong> living shrews, mice, <strong>and</strong> rats. With hindsight,<br />

the most important evolutionary event was<br />

the origin <strong>of</strong> the modern mammalian kind <strong>of</strong> molar<br />

tooth, known as the tribosphenic tooth, <strong>and</strong> with it<br />

the roots <strong>of</strong> the two major modern taxa, marsupials<br />

<strong>and</strong> placentals.<br />

Sixty-five million years ago, the mass extinction<br />

marking the close <strong>of</strong> the Mesozoic Era saw the end<br />

<strong>of</strong> the dinosaurs along with the extinction or severe<br />

depletion <strong>of</strong> many other taxa. Several mammal lineages<br />

survived this event, <strong>and</strong> within a mere 2–3 Ma<br />

had radiated explosively to produce a plethora <strong>of</strong><br />

new small forms but also, for the first time, mammals<br />

<strong>of</strong> middle to large body size. This was the<br />

commencement <strong>of</strong> the great Tertiary radiation <strong>of</strong><br />

placental <strong>and</strong>, to a lesser but equally interesting<br />

degree, marsupial mammals during which many<br />

unfamiliar, <strong>of</strong>ten quite bizarre kinds evolved, flourished,<br />

<strong>and</strong> disappeared. <strong>The</strong> Tertiary world has<br />

been a period <strong>of</strong> dramatic biogeographical <strong>and</strong> climatic<br />

change, against which is set this extraordinary<br />

evolutionary pageant. <strong>The</strong> old, single supercontinent<br />

<strong>of</strong> Pangaea occupied by the mammal-like reptiles<br />

had long since broken into Laurasia in the<br />

north <strong>and</strong> Gondwana in the south, <strong>and</strong> by the early<br />

Tertiary, Gondwana itself was breaking up. Africa<br />

<strong>and</strong> India for a short while, <strong>and</strong> South America <strong>and</strong><br />

Australia for most <strong>of</strong> the era were isl<strong>and</strong> continents<br />

with extremely limited biotic contact with each<br />

other <strong>and</strong> with the northern l<strong>and</strong> masses. Even<br />

amongst the Laurasian continents <strong>of</strong> North America,<br />

Europe, <strong>and</strong> Asia, interconnections formed <strong>and</strong><br />

broke from time to time as l<strong>and</strong> masses shifted, sea

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