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The Origin and Evolution of Mammals - Moodle

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252 THE ORIGIN AND EVOLUTION OF MAMMALS<br />

(a)<br />

Lepticis<br />

Macroscelidea (elephant shrews)<br />

Tenrecida (tenrecs <strong>and</strong> otter shrews)<br />

Chrysochlorida (golden moles)<br />

Tubulidentata (aardvark)<br />

Plesiorycteropus<br />

Hyopsodus (‘condylarth’)<br />

Meniscotherium (‘condylarth’)<br />

Phenacodus (‘condylarth’)<br />

Hyracoidea<br />

Desmostylia<br />

Sirenia (sea cows)<br />

Embrithopoda (arsinoitherians)<br />

Proboscidea (elephants) Pilgrimella<br />

Figure 7.15 (a) Asher et al.’s 2003 cladogram <strong>of</strong> Afrotheria based on an Adams consensus <strong>of</strong> molecular <strong>and</strong> morphological data. (b) Dentition<br />

<strong>of</strong> the anthacobunid Pilgrimella: lower teeth in occlusal <strong>and</strong> buccal views, upper teeth in buccal <strong>and</strong> occlusal views. Length <strong>of</strong> lower jaw<br />

fragement approx. 10 cm. (Wells <strong>and</strong> Gingerich 1983)<br />

are limited to a few low-yielding sites in North Africa<br />

(Gheerbrant 1995; Gheerbrant et al. 1996), <strong>and</strong><br />

the Early Eocene is barely any better represented. It is<br />

not until the great Qatrini Formation <strong>of</strong> the Fayum<br />

Depression in Egypt, which dates from Late Eocene<br />

through Early Oligocene (Rasmussen et al. 1992), that<br />

abundant collections <strong>of</strong> good quality fossil mammals<br />

have been made. <strong>The</strong> next oldest mammals come<br />

from the much sparser Late Oligocene fauna at<br />

Chilga in Ethiopia (Kappelman et al. 2003). What<br />

these <strong>and</strong> other localities indicate is that throughout<br />

the Oligocene afrotherians dominated the African<br />

mammalian fauna, with only a few <strong>of</strong> the northern<br />

groups, notably primates, present. <strong>The</strong> orders <strong>of</strong> large<br />

mammals that are so conspicuous in Africa today,<br />

artiodactyls, perissodactyls, <strong>and</strong> carnivorans, made<br />

little impact until the Miocene, from which time a<br />

good fossil record comprehensively illustrates the<br />

later Cenozoic history <strong>of</strong> mammals in Africa.<br />

(b)<br />

During the Late Eocene <strong>and</strong> Oligocene, the two<br />

main herbivorous mammalian taxa in Africa were<br />

the Hyracoidea <strong>and</strong> Proboscidea. Until recently<br />

opinion diverged about their mutual relationship<br />

(Prothero 1993). One group argued that the<br />

Hyracoidea were more closely related to<br />

Perissodactyla, citing a range <strong>of</strong> morphological<br />

details <strong>of</strong> the braincase, teeth, <strong>and</strong> foot structure in<br />

support (Fischer 1989; Prothero <strong>and</strong> Schoch 1989).<br />

Another group argued for a relationship between<br />

Hyracoidea <strong>and</strong> Proboscidea, along with the<br />

Sirenia, the three forming a taxon Paenungulata<br />

(Shoshani 1986; Novacek et al. 1988). Tassy <strong>and</strong><br />

Shoshani (1988) noted the incongruent evidence<br />

<strong>and</strong> were unable to decide between these possibilities.<br />

However, molecular evidence strongly supports<br />

the paenungulate group. <strong>The</strong> morphological<br />

characters that define Paenungulata, as listed by<br />

Archibald (1998), are an astragalus with flattened

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