07.12.2012 Views

The Origin and Evolution of Mammals - Moodle

The Origin and Evolution of Mammals - Moodle

The Origin and Evolution of Mammals - Moodle

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

<strong>The</strong> Late Cretaceous mammal record is extremely<br />

sparse, with the sole exceptions <strong>of</strong> North America<br />

<strong>and</strong> Mongolia, <strong>and</strong> the Garden <strong>of</strong> Eden hypothesis is<br />

that the origins <strong>and</strong> divergences <strong>of</strong> the modern placental<br />

orders occurred in a part <strong>of</strong> the world not at<br />

present adequately sampled for Cretaceous mammal<br />

fossils. <strong>The</strong> possibilities are the Gondwana continents<br />

<strong>of</strong> Australia, South America, Antarctica, <strong>and</strong><br />

Africa, in all <strong>of</strong> which, there is extremely little in the<br />

way <strong>of</strong> Cretaceous fossil mammals. On this view, the<br />

early Cenozoic explosive appearance <strong>of</strong> the modern<br />

orders was due to their immigration into the northern<br />

continents <strong>of</strong> Laurasia.<br />

<strong>The</strong> Garden <strong>of</strong> Eden hypothesis does have some<br />

arguments in its favour, principally that the molecular<br />

dates <strong>of</strong> the major divergences within placental<br />

mammals coincide approximately with the<br />

various phases <strong>of</strong> the break up <strong>of</strong> Gondwana.<br />

Murphy et al. (2001b) suggested that while the basal<br />

placentals (eutherian ancestors) occurred in the<br />

northern continents, as indicated by the Early<br />

Cretaceous fossil record, the Placentalia origininated<br />

in Gondwana (Fig. 7.26(a)). <strong>The</strong> Afrotheria<br />

diverged when Africa separated from the rest <strong>of</strong><br />

Gondwana about 105 Ma (Smith et al. 1994).<br />

Around 5–10 Ma later, according to the molecular<br />

dates, the Boreoeutheria differentiated, presumably<br />

as a consequence <strong>of</strong> their ancestor entering North<br />

America from South America. At this time, there<br />

may have been several small isl<strong>and</strong>s that would<br />

have <strong>of</strong>fered a route northwards.<br />

A second possibility is that the boreoeutherians<br />

originated in India, which separated from Africa at<br />

about the same time as did South America, becoming<br />

another isl<strong>and</strong> continent. It is not certain when<br />

India eventually collided with Asia (Hallam 1994).<br />

Most have believed this to be Late Palaeocene<br />

(Krause <strong>and</strong> Maas 1990; Prothero 1994), but<br />

other evidence, including geomorphological <strong>and</strong><br />

palaeontological, suggests that at least faunal interchange<br />

may have been possible a good deal earlier,<br />

around the end <strong>of</strong> the Cretaceous (Jaeger et al.<br />

1989). This hypothesis is consistent with the difficulty<br />

<strong>of</strong> resolving the base <strong>of</strong> the supraordinal tree,<br />

which is due to the very close dates <strong>of</strong> divergence<br />

between Afrotheria, Xenarthra, <strong>and</strong> Boreoeutheria.<br />

<strong>The</strong> only palaeontological support explicitly<br />

claimed for the Garden <strong>of</strong> Eden hypothesis comes<br />

LIVING AND FOSSIL PLACENTALS 277<br />

from Rich et al. (1999; Woodburne et al. 2003), who<br />

believe that the Early Cretaceous, 100 Ma Australian<br />

mammal jaw Ausktribosphenos is an erinaceid-like<br />

member <strong>of</strong> the Eulipotyphla, <strong>and</strong> therefore marks<br />

the area <strong>of</strong> origin <strong>of</strong> placentals. Only more material<br />

will confirm or refute this presently highly<br />

disputed identification.<br />

<strong>The</strong> Long Fuse hypothesis. <strong>The</strong> fourth possible explanation<br />

for the incongruent dates has been proposed<br />

most actively by Archibald <strong>and</strong> Deutschman (2001).<br />

<strong>The</strong> Long Fuse hypothesis is that most <strong>of</strong> the<br />

Cenozoic orders <strong>of</strong> placentals did diverge in the<br />

Late Cretaceous; that they are in fact represented by<br />

known fossils from that time; but that these have<br />

been unrecognised because they lack the diagnostic<br />

characters <strong>of</strong> the modern representatives. Rather,<br />

they remained small, unspecialised mammals with<br />

a basically insectivorous, or at most omnivorous<br />

dentition. On this view, the post-Cretaceous explosion<br />

<strong>of</strong> recognisable members <strong>of</strong> the modern orders<br />

was actually their evolution from existing, primitive,<br />

stem members. This, the hypothesis implies,<br />

occurred in many orders during a small window <strong>of</strong><br />

time, <strong>and</strong> was possibly in response to the extinction<br />

<strong>of</strong> the dinosaurs <strong>and</strong> therefore the release <strong>of</strong> the<br />

habitats <strong>and</strong> niches from which mammals had hitherto<br />

been competitively excluded. In this light, there<br />

are several c<strong>and</strong>idates for Cretaceous stemmembership<br />

<strong>of</strong> modern orders. <strong>The</strong> Leptictida have<br />

<strong>of</strong>ten been included with the Eulipotyphla<br />

(Novacek 1986a), <strong>and</strong> the Zalambdalestidae interpreted<br />

as stem Glires (Archibald et al. 2001). <strong>The</strong><br />

Zhelestidae possess incipiently ungulate-like molar<br />

teeth <strong>and</strong> could conceivably include stem members<br />

<strong>of</strong> any <strong>of</strong> the numerous Cenozoic ungulate groups<br />

(Archibald 1999). <strong>The</strong> Palaeoryctida have from time<br />

to time been allied to the Carnivora, <strong>and</strong> also some<br />

<strong>of</strong> the extinct Cenozoic orders such as Creodonta,<br />

Taeniodonta, Tillodonta, <strong>and</strong> Pantodonta. <strong>The</strong> evidence<br />

for all <strong>of</strong> these putative relationships is certainly<br />

not at all strong, <strong>of</strong>ten being based almost<br />

completely on minor dental similarities, although <strong>of</strong><br />

course this in itself is a prediction <strong>of</strong> the very<br />

hypothesis.<br />

Springer et al.’s (2003) molecular study already<br />

referred to (Fig. 7.25) is consistent with the Long<br />

Fuse hypothesis, ins<strong>of</strong>ar as it places most <strong>of</strong> the

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!