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The Origin and Evolution of Mammals - Moodle

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have postcranial adaptations for arboreal leaping,<br />

such as an elongated calcaneum <strong>and</strong> opposable first<br />

digits. Several authors have argued that omomyoids<br />

are basal members <strong>of</strong> the Haplorhini, on the<br />

basis <strong>of</strong> the relatively large orbits, loss <strong>of</strong> the first<br />

premolars, <strong>and</strong> details <strong>of</strong> tooth structure <strong>and</strong> the<br />

ear region <strong>of</strong> the skull (Ross et al. 1998; Gunnell <strong>and</strong><br />

Rose 2002). Some go so far as to relate them to the<br />

tarsiers in particular; certainly the short snout,<br />

large, forwardly directed eyes, <strong>and</strong> highly agile<br />

skeleton give a very strong impression <strong>of</strong> a tarsier,<br />

as reviewed by Martin (1993).<br />

<strong>The</strong> modern groups <strong>of</strong> Strepsirhini have a poor fossil<br />

record, even for primates. <strong>The</strong>re is no certain<br />

lemuriform at all prior to the Late Pleistocene <strong>of</strong><br />

Madagascar, with the dubious <strong>and</strong> controversial<br />

exception <strong>of</strong> the Oligocene Bugtilemur from Pakistan<br />

(Marrivaux et al. 2001) <strong>The</strong> other strepsirhine branch,<br />

the lorisiformes, have been recorded in the late<br />

Middle Eocene <strong>of</strong> the Fayum region <strong>of</strong> Egypt, but<br />

only as isolated teeth: Karanisia is a possible loris, <strong>and</strong><br />

Saharagalago a possible galago (Seiffert et al. 2003).<br />

For the living haplorhines, the earliest member <strong>of</strong><br />

the modern tarsiiform family Tarsiidae has been<br />

identified from teeth attributed to the modern<br />

genus Tarsius, found in Middle Eocene deposits <strong>of</strong><br />

China (Beard et al. 1994).<br />

<strong>The</strong> diversity <strong>of</strong> the adapiforms, omomyoids,<br />

lorisiforms, <strong>and</strong> tarsiiforms show that the Eocene<br />

was a time <strong>of</strong> considerable radiation <strong>of</strong> primitive<br />

primates, producing a significant fauna <strong>of</strong> small,<br />

arboreal mammals. This observation might have<br />

been <strong>of</strong> only marginal interest had the radiation not<br />

also included basal members <strong>of</strong> the anthropoids, the<br />

monkeys, apes, <strong>and</strong> humans. What at the time was<br />

in effect another ‘prosimian’ grade <strong>of</strong> Eocene primates<br />

appears in the same Middle Eocene deposits <strong>of</strong><br />

China that yielded Tarsius (Beard et al. 1996; Dagosto<br />

et al. 1996). Eosimias is known only from dentitions<br />

<strong>and</strong> some probably associated limb bones, <strong>and</strong><br />

would indeed warrant little attention except for its<br />

probable position as the most basal member <strong>of</strong> the<br />

Anthropoidea (Ross et al. 1998; Ducrocq 2001). It<br />

was a small animal, the lower jaw length being<br />

about 2.6 cm. Characters that support its anthropoid<br />

affinity are mostly dental as no cranial <strong>and</strong> very little<br />

postcranial material has yet been found. <strong>The</strong> incisors<br />

are reduced in height, spatulate, <strong>and</strong> vertical<br />

LIVING AND FOSSIL PLACENTALS 271<br />

rather than slightly procumbent, <strong>and</strong> the talonid <strong>of</strong><br />

the last lower molar is much reduced. Isolated teeth<br />

<strong>of</strong> equally ancient basal anthropoids occur in the<br />

Glib region <strong>of</strong> Algeria (Godinot <strong>and</strong> Mahboubi<br />

1992), indicating that from the start anthropoids<br />

occurred in Africa as well as Asia.<br />

<strong>The</strong> exact relationship <strong>of</strong> the Anthropoidea to the<br />

other Eocene primates remains a matter <strong>of</strong> considerable<br />

debate, <strong>and</strong> virtually every possibility has<br />

been proposed by one author or another (Martin<br />

1993). Dagosto (2002) notes that hypotheses linking<br />

them to, respectively, adapiforms, omomyoids,<br />

tarsiids, omomyoids plus tarsiids, or to no known<br />

group have all been proposed by one author or<br />

another within the last decade or so.<br />

From the Late Eocene onwards, basal anthropoids<br />

radiated in Africa <strong>and</strong> Asia (Beard 2002a).<br />

Several genera are found in the Late Eocene <strong>and</strong><br />

early Oligocene <strong>of</strong> the Fayum deposits, notably<br />

Apidium, which is represented by partial skulls<br />

(Simons 1995). Siamopithecus from Thail<strong>and</strong><br />

(Ducrocq 1999) is an Asian representative although<br />

so far known only from the dentition. <strong>The</strong> interrelationships<br />

amongst these several lineages <strong>of</strong><br />

Eocene anthropoids are very unclear at present<br />

(Simons 1992; Ross et al. 1998; Ducrocq 2001), <strong>and</strong> it<br />

is not even established whether the respective taxa<br />

from the two continents are sister-groups, or part <strong>of</strong><br />

a contiguous fauna.<br />

This uncertainty about exact relationships is also<br />

true <strong>of</strong> the two modern groups <strong>of</strong> anthropoids. <strong>The</strong><br />

oldest <strong>of</strong> the New World monkeys, the Platyrrini, is<br />

Branisella, represented by Late Oligocene teeth <strong>and</strong><br />

fragmentary jaws from Bolivia. <strong>The</strong> oldest actual<br />

skull so far described is that <strong>of</strong> Chilicebus, which is<br />

Early Miocene in age <strong>and</strong> a member <strong>of</strong> the existing<br />

family Cebidae (Flynn et al. 1995). No African or<br />

Asian platyrrini have been found, although<br />

Simons (1997; Takei et al. 2000) noted some dental<br />

similarities between Branisella <strong>and</strong> the Fayum<br />

Proteopithecus. <strong>The</strong>y proposed a relationship<br />

between the two which, if true, <strong>of</strong>fers taxonomic<br />

support for the view that the origin <strong>of</strong> the New<br />

World monkeys was a dispersal event directly from<br />

Africa during the Oligocene. However, not all<br />

accept the proposed relationship, so the question is<br />

not closed, <strong>and</strong> an American or Asian origin for<br />

Platyrrini remains a possibility (Dagosto 2002).

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