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The Origin and Evolution of Mammals - Moodle

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(d)<br />

Figure 6.6 (continued).<br />

Thylacosmilus<br />

Muizon et al. (1997) compared the primitive<br />

Tiupampan sparassodont Mayulestes with the<br />

didelphimorphs Pucadelphys <strong>and</strong> Andinodelphys<br />

from the same locality. <strong>The</strong>y found a number <strong>of</strong> similarities<br />

suggesting a relationship, such as a medial<br />

process <strong>of</strong> the squamosal, <strong>and</strong> an enlarged, slightly<br />

procumbent first upper incisor <strong>and</strong> staggered<br />

second lower incisor. Limited though this evidence<br />

is, it is rather more convincing because it does<br />

take cranial anatomy into account as well as dental.<br />

In fact, this is a reversion to another long-held<br />

view, that sparassodonts are simply specialised<br />

carnivores derived from a basal didelphimorph<br />

stock (Marshall 1978).<br />

Mayulestes (Fig. 6.6(a)) is the oldest known sparassodont<br />

<strong>and</strong> is represented by a skull <strong>and</strong> partial<br />

skeleton (Muizon 1998). It is placed in a separate<br />

family Mayulestidae, along with the contemporary<br />

Allqokirus that is only known from isolated teeth.<br />

Mayulestes has retained the primitive marsupial<br />

dental formula <strong>of</strong> I5/4 : C1/1 : P3/3 : M4/4, <strong>and</strong><br />

the structure <strong>of</strong> the molars is also <strong>of</strong> the basic<br />

marsupial form. However, the skull does show the<br />

diagnostic sparassodont characters <strong>of</strong> a contribution<br />

(e)<br />

LIVING AND FOSSIL MARSUPIALS 203<br />

Prothylacinus<br />

<strong>of</strong> the squamosal to the ro<strong>of</strong> <strong>of</strong> the alisphenoid<br />

sinus, <strong>and</strong> the loss <strong>of</strong> the prootic canal. <strong>The</strong> postcranial<br />

skeleton is indicative <strong>of</strong> a mammal with an<br />

agile, scansorial habit; the fore <strong>and</strong> hindlimbs had<br />

considerable three-dimensional mobility <strong>and</strong> the tail<br />

was probably prehensile. Muizon (1998) believed<br />

that the mode <strong>of</strong> life would have been comparable<br />

to that <strong>of</strong> a small pine marten, <strong>and</strong> suggested that<br />

this was the primitive habit, not only <strong>of</strong> sparassodonts,<br />

but <strong>of</strong> marsupials generally.<br />

Members <strong>of</strong> the more progressive, <strong>and</strong> far more<br />

abundant <strong>and</strong> diverse family Borhyaenidae appear<br />

in the slightly younger Itaboraí fauna, for example<br />

Patene. <strong>The</strong> Early Eocene Callistoe (Fig. 6.6(b)) is<br />

related to the sabre-toothed Thylacosmilus, as indicated<br />

by its already very enlarged canines. <strong>The</strong><br />

borhyaenids subsequently went on to radiate as<br />

middle-sized <strong>and</strong> larger carnivores <strong>of</strong> the South<br />

American Tertiary (Marshall 1978), a role shared<br />

amongst the mammals only with a few <strong>of</strong> the largest<br />

didelphids, <strong>and</strong> modest-sized procyonid placentals<br />

that entered the continent in the Late Miocene. In fact,<br />

the largest predators were amongst the flightless<br />

phorusrhacid birds, with some species approaching

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