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The Origin and Evolution of Mammals - Moodle

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fossil mammals, which might help clarify this<br />

question, are exceedingly scarce in North America<br />

(Cifelli <strong>and</strong> Davis 2003).<br />

At any event, during the Late Cretaceous the<br />

North American marsupials radiated into 4 families,<br />

<strong>and</strong> about 19 genera, all <strong>of</strong> them relatively<br />

small forms, but variously adapted, some for insectivory,<br />

some for a rather more carnivorous habit,<br />

<strong>and</strong> some for a more cosmopolitan or omnivorous<br />

diet. Incidentally, it should be remembered in this<br />

context that the Tribosphenida as a whole were less<br />

abundant <strong>and</strong> diverse than the contemporary multituberculate<br />

mammals. It needs also to be borne in<br />

mind that so very little is known about mammals<br />

<strong>of</strong> the Cretaceous in the Gondwanan continents,<br />

including Africa, that a major surprise is always<br />

possible. <strong>The</strong> reported, though disputed presence<br />

<strong>of</strong> a Late Cretaceous marsupial in Madagascar is<br />

a reminder <strong>of</strong> this.<br />

<strong>The</strong> K/T mass extinction event has been described.<br />

As far as the marsupial mammals are concerned,<br />

the thriving North American radiation was all but<br />

eliminated, <strong>and</strong> the only marsupials represented on<br />

that continent in the Palaeocene were the didelphid<br />

Peradectes, <strong>and</strong> one or two closely related genera.<br />

Possibly, as discussed earlier in the chapter, the<br />

mass extinction <strong>of</strong> the marsupials was due to competition<br />

from similarly adapted placental immigrants<br />

from Asia. Whatever the reason, didelphids<br />

did nevertheless persist <strong>and</strong> indeed radiated in a<br />

small way through the Eocene <strong>and</strong> possibly into the<br />

Miocene. Members <strong>of</strong> the family also appeared<br />

sporadically in Europe, Africa <strong>and</strong> Asia during this<br />

time, presumably by dispersal from North America.<br />

Although important as small carnivorous members<br />

<strong>of</strong> certain Old World mammalian faunas, they<br />

never achieved significant taxonomic diversity, <strong>and</strong><br />

have not been recorded in these regions since the<br />

Miocene (McKenna <strong>and</strong> Bell 1997).<br />

South America<br />

<strong>The</strong> next major point in known marsupial history<br />

occurred around the time <strong>of</strong> the K/T boundary.<br />

Having all but disappeared from North America,<br />

they left the Laurasian <strong>and</strong> African world virtually<br />

exclusively to the placentals. However, at about the<br />

same time, marsupials first appear in the South<br />

American fossil record. No marsupial, or placental,<br />

LIVING AND FOSSIL MARSUPIALS 217<br />

mammals have been found in undisputed Cretaceous<br />

rocks anywhere in South America. <strong>The</strong> mammalian<br />

fauna that was present, as most richly represented<br />

by the Campanian or Maastrichtian Los Alamitos<br />

Formation <strong>of</strong> Argentina, consisted solely <strong>of</strong> nontribosphenidan<br />

Mesozoic mammal groups that are<br />

more typical <strong>of</strong> Jurassic <strong>and</strong> Early Cretaceous<br />

deposits elsewhere, <strong>and</strong> which are represented here<br />

mostly by subgroups endemic to South America.<br />

Whether the very earliest South American marsupials<br />

are strictly latest Cretaceous or Early Palaeocene<br />

is not certain, but what is evident is that the fauna<br />

included possible members <strong>of</strong> two groups characteristic<br />

<strong>of</strong> the Late Cretaceous <strong>of</strong> North America,<br />

peradectids <strong>and</strong> pediomyids. Add to these the<br />

very slightly later appearance in South America <strong>of</strong><br />

possible relatives <strong>of</strong> the specialised North<br />

American genus Glasbius, <strong>and</strong> the conclusion seems<br />

inescapable that the South American fauna was<br />

derived by immigration from North America close<br />

to 65 Ma. During the Late Cretaceous, the Caribbean<br />

Archipelago was forming, <strong>and</strong> provided a potential<br />

dispersal route by isl<strong>and</strong> hopping. Dispersal <strong>of</strong> several<br />

other groups <strong>of</strong> terrestrial vertebrates, including<br />

dinosaurs, had indeed been in progress during the<br />

latest Cretaceous (Rage 1988; Gayet et al. 1992).<br />

Within a very short space <strong>of</strong> time, less than five<br />

million years, a truly indigenous South American<br />

marsupial fauna had evolved from the presumed<br />

immigrant ancestors. <strong>The</strong>re was a radiation <strong>of</strong><br />

primarily insectivorous <strong>and</strong> omnivorous didelphids,<br />

that retained the primitive tooth form. Simultaneously,<br />

the medium <strong>and</strong> large carnivore mammal<br />

niches started to be filled by the sparassodonts<br />

(borhyaenoids), a group which was also probably<br />

derived from the immigrant didelphid ancestor.<br />

<strong>The</strong> paucituberculates may be related to the original<br />

Glasbius-like immigrant, <strong>and</strong> they evolved into<br />

a series <strong>of</strong> small- <strong>and</strong> medium-sized omnivore, <strong>and</strong><br />

specialist grain, fruit, <strong>and</strong> tuber-eating herbivores.<br />

However, the South American marsupial radiation<br />

was limited by them having to share their continent<br />

with certain placental mammal groups, which had<br />

also arrived at the start <strong>of</strong> the Palaeocene. <strong>The</strong>se<br />

contributed the medium- to large-sized ungulate<br />

herbivores, <strong>and</strong> also certain specialists, such as<br />

anteaters <strong>and</strong> sloths, that constitute the Xenarthra.<br />

Later in the Cenozoic, marsupials had to contend

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