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The Origin and Evolution of Mammals - Moodle

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ecognised four subgroups, based mainly on differences<br />

in jaw structure, although there are very few<br />

characters defining these taxa so that they are not at<br />

all well supported. A more recent <strong>and</strong> more detailed<br />

cladistic analysis is that <strong>of</strong> Angielczyk (2001), which<br />

includes 20 dicynodont taxa <strong>and</strong> 40 characters. It<br />

disagrees with many <strong>of</strong> King’s (1988) relationships,<br />

but again most <strong>of</strong> the clades are very weakly supported.<br />

Many <strong>of</strong> the nodes <strong>of</strong> his cladogram collapse<br />

with only a single additional step, <strong>and</strong> with as little<br />

as three additional steps virtually all the internal<br />

structure <strong>of</strong> the cladogram becomes unresolved.<br />

Thus convergent evolution <strong>and</strong> character reversal<br />

was extensive <strong>and</strong> it should be stressed again that the<br />

40–50 genera accepted nowadays, are all relatively<br />

conservative. Were they a modern herbivorous<br />

mammal group, dicynodonts might well have been<br />

incorporated into a single family with a disparity<br />

perhaps equivalent to that <strong>of</strong> the Bovidae. Given this<br />

lack <strong>of</strong> taxonomic resolution <strong>of</strong> Dicynodontia, the<br />

scheme <strong>of</strong> King (1988) based on evident differences in<br />

the feeding structures is followed, although more<br />

from convenience than conviction.<br />

Endothiodontoidea. Endothiodon has a highly atypical<br />

dentition. It is a large form found in the<br />

Tapinocephalus Assemblage Zone <strong>of</strong> South Africa<br />

<strong>and</strong> also in beds <strong>of</strong> equivalent age in Zambia, South<br />

America, <strong>and</strong> India (Ray 2000). <strong>The</strong> skull reaches<br />

over 50 cm in length with a distinctively high, very<br />

narrow sagittal crest. <strong>The</strong> lower jaw is distinguished<br />

by the very deep anterior part attached to the relatively<br />

slender postdentary part. <strong>The</strong> upturned front<br />

tip <strong>of</strong> the fused dentaries worked against the broad,<br />

deeply concave plate formed from the fused premaxillae.<br />

Canine tusks are absent, but the postcanine<br />

dentition is remarkably <strong>and</strong> presumably secondarily<br />

well developed. <strong>The</strong>re are about 10 long, slender<br />

upper teeth, fairly widely spaced <strong>and</strong> forming<br />

a single row that does not oppose the lower teeth,<br />

but must have bitten against a presumed horny pad<br />

lying in a longitudinal groove lateral to the lower<br />

teeth. <strong>The</strong> lower dentition consists <strong>of</strong> smaller, tightly<br />

packed teeth forming two or three irregular rows.<br />

Only the more lateral ones were functional, biting<br />

against a horny pad presumably present in life on<br />

the heavily rugose palatine bone, internal to the<br />

upper tooth row. <strong>The</strong> more medial <strong>of</strong> the lower teeth<br />

EVOLUTION OF MAMMAL-LIKE REPTILES 47<br />

are young teeth, just erupting <strong>and</strong> preparing to<br />

move laterally to replace existing functional teeth as<br />

they are discarded. An area <strong>of</strong> vascularised bone<br />

lying lateral to the upper tooth row has been interpreted<br />

by Cluver (1975) as the line <strong>of</strong> attachment <strong>of</strong><br />

a muscular, food-retaining cheek. Several authors<br />

have speculated on the diet <strong>of</strong> Endothiodon. King<br />

(1990) suggested that the teeth had a slicing action,<br />

perhaps by direct unilateral occlusion <strong>of</strong> uppers <strong>and</strong><br />

lowers. Latimer et al. (1995) proposed that they<br />

browsed on riverine vegetation using cheek pouches<br />

to collect the food. Cox (1998) rather ingeniously<br />

speculated that conifer cones were stripped by the<br />

action <strong>of</strong> the tip <strong>of</strong> the m<strong>and</strong>ibles acting against the<br />

premaxillae, followed by maceration <strong>of</strong> the seeds<br />

using the postcanine dentition.<br />

Kingorioidea. Kingoria (Cox 1959) lacks all signs <strong>of</strong><br />

postcanine teeth which, if its cladistic relationships<br />

are correctly inferred, were lost independently <strong>of</strong><br />

the loss in other lineages. Some specimens <strong>of</strong><br />

Kingoria retain but others have lost the pair <strong>of</strong> tusks.<br />

<strong>The</strong> horny, biting surfaces inferred from the structure<br />

<strong>of</strong> the bones <strong>of</strong> the upper <strong>and</strong> lower jaws differed<br />

considerably from those <strong>of</strong> other dicynodonts.<br />

<strong>The</strong> secondary palate formed from the premaxillae<br />

is smooth <strong>and</strong> flat, <strong>and</strong> the horn appears to have<br />

been restricted to the lateral margins <strong>of</strong> the adjacent<br />

maxillae. Instead <strong>of</strong> being sharp, these lateral edges<br />

are rounded, as are the corresponding anterior parts<br />

<strong>of</strong> the dentaries, implying a crushing rather than a<br />

shearing action. However, there are some sharp<br />

margins, a short one immediately in front <strong>of</strong> the<br />

caniniform process <strong>of</strong> the upper jaw, <strong>and</strong> another at<br />

the upturned anterior tip <strong>of</strong> the joined dentaries.<br />

Hotton (1986) attempted to explain this curious<br />

combination <strong>of</strong> features with the suggestion that<br />

Kingoria fed on a food source that was <strong>of</strong> small<br />

size, s<strong>of</strong>t, <strong>and</strong> abundant. King (1988) thought that<br />

perhaps its food consisted <strong>of</strong> invertebrates, grubbed<br />

up from mud <strong>and</strong> swallowed more or less without<br />

any chewing.<br />

Kingoria also has some distinct specialisations in<br />

its postcranial skeleton, reminiscent <strong>of</strong> the incipiently<br />

mammalian condition seen in cynodonts<br />

(Cox 1959; King 1985). <strong>The</strong> shoulder girdle has<br />

developed a particularly strongly everted spine<br />

extending from the acromion process up the front

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