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

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

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).

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

Saved successfully!

Ooh no, something went wrong!