The Origin and Evolution of Mammals - Moodle
The Origin and Evolution of Mammals - Moodle
The Origin and Evolution of Mammals - Moodle
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m<strong>and</strong>ibulae musculature. This, as in the dicynodonts,<br />
would have provided the necessary posteriorly<br />
directed force on the m<strong>and</strong>ible during occlusion<br />
(Rybczynski 2001). <strong>The</strong> form <strong>and</strong> evident action <strong>of</strong><br />
the teeth indicate a committed herbivorous animal.<br />
Dromasauroidea<br />
<strong>The</strong>re are three genera <strong>of</strong> small, primitive anomodonts<br />
from the Late Permian <strong>of</strong> South Africa generally<br />
included in a taxon Dromasauria. <strong>The</strong>y are<br />
represented by a total <strong>of</strong> four specimens, each one <strong>of</strong><br />
which is preserved as a natural s<strong>and</strong>stone cast <strong>of</strong> the<br />
skull <strong>and</strong> postcranial elements (Brinkman 1981).<br />
Galeops (Fig. 3.11(d)) shows features otherwise<br />
restricted to the dicynodonts, leading to the view<br />
that among the dromasaurs it is the dicynodont sistergroup<br />
(Modesto <strong>and</strong> Rybczynski 2000; Rybczynski<br />
2000). <strong>The</strong> characters in question include further<br />
shortening <strong>of</strong> the preorbital region <strong>of</strong> the skull,<br />
absence <strong>of</strong> anterior teeth on either premaxilla or dentary,<br />
<strong>and</strong> the remaining posterior dentition reduced to<br />
a short row <strong>of</strong> small, peg-like teeth. <strong>The</strong> arrangement<br />
implies that dicynodont-like horny tooth plates had<br />
evolved at the front <strong>of</strong> the jaws. Other features <strong>of</strong><br />
dicynodonts that are found are fusion <strong>of</strong> the paired<br />
dentaries at the symphysis, <strong>and</strong> a quadrate condyle<br />
divided by a longitudinal groove into lateral <strong>and</strong><br />
medial condyles, corresponding to elongated articular<br />
condyles. According to King (1994), the structure<br />
<strong>of</strong> the jaw articulation would have permitted propalinal<br />
movement <strong>of</strong> the lower jaw.<br />
<strong>The</strong> preserved specimens <strong>of</strong> neither <strong>of</strong> the other<br />
two genera <strong>of</strong> dromasaurs, Galepus <strong>and</strong> Galechirus<br />
(Fig. 3.11(c)) are as revealing. <strong>The</strong>y are much the same<br />
size as Galeops, <strong>and</strong> look superficially very similar.<br />
However, there are differences in the dentition which<br />
justifies at least the generic distinction between<br />
the three (Brinkman 1981). Galechirus possesses premaxillary<br />
teeth, which are decidedly procumbent,<br />
but the situation <strong>of</strong> the lower teeth is unclear. Galepus<br />
has at least six small lower teeth, but in this case the<br />
nature <strong>of</strong> the upper dentition is unclear.<br />
Dicynodontia<br />
<strong>The</strong> great majority <strong>of</strong> anomodonts belong to the very<br />
well-defined group Dicynodontia, which makes its<br />
appearance in the Eodicynodon Assemblage Zone<br />
EVOLUTION OF MAMMAL-LIKE REPTILES 43<br />
contemporary with the earliest <strong>of</strong> the primitive<br />
African anomodonts described above. Indeed, it is<br />
the eponymous Eodicynodon itself that first represents<br />
the group (Rubidge 1984, 1990a).<br />
Eodicynodon<br />
<strong>The</strong> trends in cranial evolution seen in the primitive<br />
anomodonts up to the stage represented by Galeops<br />
reach their culmination in the fully dicynodont form<br />
<strong>of</strong> Eodicynodon (Fig. 3.12(a)). All the incisor teeth are<br />
absent, there is an enlarged caniniform upper tusk,<br />
<strong>and</strong> its postcanine dentition is reduced to a small<br />
number <strong>of</strong> relatively minute teeth. (Rubidge, 1990b,<br />
described a specimen <strong>of</strong> Eodicynodon, which lacks the<br />
caniniform tooth; he attributed it to a different<br />
species). <strong>The</strong> triturating surface <strong>of</strong> the jaws consisted<br />
virtually exclusively <strong>of</strong> a horny beak. Cox (1998)<br />
concluded from the structure <strong>of</strong> the bony surfaces<br />
that there were three parts to the beak. A medial<br />
anterior dentary beak bit within the front margins<br />
<strong>of</strong> the palate, presumably with a cropping function.<br />
Further back, a pad on the dentary table acted against<br />
a palatal pad medial to the caniniform process. Third,<br />
a posterior horny blade on the dentary bit against a<br />
palatine pad behind the caniniform process. In some<br />
specimens the lower blade also carries a row <strong>of</strong> small<br />
teeth. In order to operate these biting structures, the<br />
external adductor musculature had a much enlarged<br />
lateral component, originating from the horizontally<br />
flattened <strong>and</strong> dorsally bowed zygomatic arch <strong>and</strong><br />
inserting at least partially on a lateral shelf on<br />
the dentary. <strong>The</strong> temporal fenestra as a whole is<br />
elongated, while the lateral pterygoid processes are<br />
reduced. <strong>The</strong>se point to the increased importance <strong>of</strong><br />
a posteriorly directed power stroke during the closing<br />
phase <strong>of</strong> the jaw action, accompanied by a reduction<br />
in the importance <strong>of</strong> the anterior pterygoideus<br />
musculature with its anteriorly directed component<br />
(Fig. 3.12(c)). <strong>The</strong> jaw articulation (Fig. 3.12(d)) has<br />
the uniquely dicynodont structure <strong>of</strong> more or less<br />
equal lateral <strong>and</strong> medial convex condyles on the<br />
quadrate <strong>and</strong> corresponding but elongated articular<br />
condyles on the lower jaw. King et al. (1989) believed<br />
that the feeding mechanism <strong>of</strong> Eodicynodon had<br />
achieved the essential dicynodont arrangement but<br />
that the degree <strong>of</strong> propaliny was still less than<br />
occurred in later dicynodonts, a view reiterated by<br />
King (1994) <strong>and</strong> Cox (1998).