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The Origin and Evolution of Mammals - Moodle

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(a)<br />

Leptictidium<br />

Palaeoryctidans were also common in the North<br />

American Palaeocene <strong>and</strong> Eocene, <strong>and</strong> present in<br />

Europe. None have yet been found in China,<br />

although there are teeth <strong>of</strong> a family <strong>of</strong> primitive<br />

insectivores, Micropternodontidae, which Butler<br />

(1988) regarded as derived from palaeoryctids.<br />

Others dispute this relationship (e.g. McKenna <strong>and</strong><br />

Bell 1997). <strong>The</strong> shrew-like Eoryctes, described from<br />

a complete Early Eocene skull (<strong>The</strong>wissen <strong>and</strong><br />

Gingerich 1989), is a typical micropternodontid.<br />

Of the other Cretaceous placental groups, a few<br />

zhelestid teeth have been found in the Early<br />

Palaeocene <strong>of</strong> North America, but neither asioryctidans<br />

nor zalambdalestids appear to have survived.<br />

In addition to pre-existing groups, there were<br />

several new lineages that were related to, but taxonomically<br />

distinct from the Cretaceous groups. <strong>The</strong><br />

majority <strong>of</strong> these still consisted <strong>of</strong> relatively small,<br />

basically insectivorous or omnivorous mammals.<br />

However, within as little as 2 or 3 million years<br />

after the start <strong>of</strong> the Palaeocene, several groups <strong>of</strong><br />

radically new kinds had arisen, <strong>and</strong> entered habitats<br />

not previously occupied by mammals during<br />

any <strong>of</strong> their previous 145 Ma <strong>of</strong> existence. <strong>The</strong>re<br />

(b)<br />

Buxolestes<br />

Figure 7.3 Two Early Eocene mammals from the Messel Oil<br />

Shales. (a) <strong>The</strong> leptictidan Leptictidium (Koenigswald et al. 1992).<br />

Presacral length approx. 30 cm. (b) <strong>The</strong> pantolestidan Buxolestes,<br />

with occlusal view <strong>of</strong> upper molars <strong>and</strong> lateral view <strong>of</strong><br />

dentition. Total length <strong>of</strong> body approx. 80 cm (Savage<br />

<strong>and</strong> Long 1986).<br />

LIVING AND FOSSIL PLACENTALS 231<br />

were middle to quite large-sized herbivorous groups,<br />

<strong>and</strong> dog-sized to small bear-sized carnivores.<br />

Anagalida<br />

<strong>The</strong> anagalidans were an Asian group, appearing in<br />

considerable diversity in the Early Palaeocene <strong>of</strong><br />

China. <strong>The</strong>y were small herbivores or omnivores,<br />

with somewhat enlarged, procumbent lower<br />

incisors (Fig. 7.4(a)). <strong>The</strong> premolar <strong>and</strong> molar teeth<br />

are broad, low-crowned, <strong>and</strong> hypsodont. <strong>The</strong>ir<br />

evolutionary affinities are not clear, but several<br />

authors accept the relationship with the Asian<br />

Cretaceous zalambdalestids that was proposed by<br />

Szalay <strong>and</strong> McKenna (1971).<br />

Mixodonta<br />

<strong>The</strong> mixodonts are also an exclusively Asian group <strong>of</strong><br />

small omnivorous mammals, also possibly related to<br />

zalambdalestids. <strong>The</strong>y are notable for their fully gliriform<br />

incisor teeth, <strong>and</strong> for this reason are widely<br />

regarded as related to the origin <strong>of</strong> the super-order<br />

Glires (rodents <strong>and</strong> lagomorphs). Li <strong>and</strong> Ting (1993)<br />

go so far as to claim that different families <strong>of</strong><br />

mixodonts are relatives <strong>of</strong> lagomorphs <strong>and</strong> rodents<br />

respectively. A mimotonid, such as the Early<br />

Palaeocene Mimotona (Fig. 7.4(b)), has the dental<br />

formula, 2/2: 0/0: 3/3: 3/3, along with several<br />

cranial characteristics <strong>of</strong> lagomorphs. In contrast, the<br />

early eurymylid Heomys (Fig. 7.4(c)) has a rodent-like<br />

dentition including a single upper incisor <strong>and</strong> only<br />

two lower premolars. <strong>The</strong>re are also characters <strong>of</strong> the<br />

postcranial skeleton that resemble those <strong>of</strong> rodents.<br />

Pantolestida<br />

<strong>The</strong> pantolestidans are almost, if not exclusively<br />

North American <strong>and</strong> European in occurrence,<br />

appearing in the Middle Palaeocene <strong>and</strong> surviving<br />

into the Oligocene (Koenigswald 1980). Otter-like in<br />

general size <strong>and</strong> proportions, the canines were well<br />

developed <strong>and</strong> while the molars <strong>of</strong> the earlier forms<br />

had sharp cutting edges, they tended to evolve a<br />

lower, broader form coupled with powerful adductor<br />

musculature suggestive <strong>of</strong> mollusc eating. <strong>The</strong><br />

Eocene Buxolestes (Fig. 7.3(b)) from Messel has fish<br />

remains preserved in its stomach (Koenigswald et al.<br />

1992). <strong>The</strong> relationships <strong>of</strong> pantolestidans are unclear,<br />

though it is generally assumed that they are a highly<br />

derived lineage related to palaeoryctidans.

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