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The Origin and Evolution of Mammals - Moodle

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(a)<br />

(b)<br />

(i)<br />

(ii)<br />

(iii)<br />

(iv)<br />

(v)<br />

a g<br />

canine<br />

2<br />

4<br />

3<br />

3<br />

1<br />

3<br />

Diademodon<br />

2<br />

2<br />

3<br />

<strong>The</strong> extreme version <strong>of</strong> this condition is seen in the<br />

tritylodontids, which do not replace any <strong>of</strong> their<br />

postcanines at all, but only discard them from the<br />

front <strong>and</strong> add to them at the back (Kühne 1956).<br />

Other cynodonts, including the basal form<br />

Thrinaxodon <strong>and</strong> the highly derived tritheledontid<br />

Pachygenelus (Crompton <strong>and</strong> Luo 1993), retained the<br />

primitive pattern <strong>of</strong> polyphyodonty, consisting <strong>of</strong><br />

waves <strong>of</strong> replacement from front to back affecting<br />

teeth alternately along the jaw.<br />

It is not until the basal mammal Morganucodon<br />

that the combination <strong>of</strong> determinate growth <strong>and</strong><br />

diphyodonty is known to have evolved, as was<br />

demonstrated by Parrington (1971), who found<br />

specimens amongst the hundreds <strong>of</strong> fragments that<br />

were either juvenile growth stages or, the great<br />

majority, identical-sized adults. <strong>The</strong> incisors, canines,<br />

s<br />

Postcanines<br />

1 2 3 4<br />

2 3 4<br />

2 3 4 5<br />

3 4 5<br />

3 4 5<br />

Sinoconodon<br />

Figure 4.12 Tooth replacement (a) Diademodon (Kemp 1982) <strong>and</strong><br />

(b) Sinoconodon. a-anterior teeth; g-gomphodont teeth. s-sectorial teeth<br />

(Crompton <strong>and</strong> Luo 1993).<br />

EVOLUTION OF MAMMALIAN BIOLOGY 121<br />

<strong>and</strong> anterior postcanines are replaced once, <strong>and</strong><br />

posterior postcanines are added sequentially at the<br />

back, not replaced, <strong>and</strong> therefore can properly be<br />

referred to as molar teeth. Given its correlation with<br />

growth pattern, it is assumed by this stage that lactation<br />

had evolved. However, the story is complicated<br />

by the situation in Sinoconodon, which is basal<br />

to Morganucodon. It still had indeterminate growth,<br />

for specimens are found that range in skull length<br />

from 2.2 to 6.2 cm, corresponding to an estimated<br />

body mass range <strong>of</strong> 13–517 g (Kielan-Jaworowska<br />

2004). <strong>The</strong> tooth replacement pattern is also more<br />

primitive in Sinoconodon, as indeed it had to be in<br />

order to allow for the very considerable growth in<br />

size <strong>of</strong> what must have been sub-adults not<br />

dependent on lactation for their growth. <strong>The</strong> incisors<br />

<strong>and</strong> canines still show alternate, multiple replacements.<br />

<strong>The</strong> postcanine teeth are not replaced, <strong>and</strong><br />

there is loss <strong>of</strong> anterior <strong>and</strong> addition <strong>of</strong> new posterior<br />

postcanines maintaining the relatively short<br />

postcanine tooth row <strong>of</strong> only three or four teeth<br />

(Fig. 4.12(b)). This condition in Sinoconodon is therefore<br />

intermediate between the primitive tritheledontid<br />

<strong>and</strong> the fully mammalian conditions. It may<br />

be speculated that the state <strong>of</strong> evolution <strong>of</strong> lactation<br />

was also intermediate, with maternal provision <strong>of</strong><br />

milk limited to an early neonate stage only, after<br />

which the juvenile was weaned <strong>and</strong> relied on its<br />

own foraging, or perhaps on a more limited conventional<br />

food supply provided by the mother.<br />

Temperature physiology<br />

Nothing is more fundamental to the life <strong>of</strong> mammals<br />

than their endothermic temperature physiology,<br />

if only because it entails a 10-fold increase in<br />

daily food requirements. Such a huge cost must be<br />

balanced by an equally large benefit for endothermy<br />

to have evolved <strong>and</strong> been maintained. Yet surprisingly<br />

there is no consensus about exactly how, why,<br />

or when endothermy evolved in the course <strong>of</strong> the<br />

evolution <strong>of</strong> the mammals. <strong>The</strong> fact that the birds<br />

share a virtually identical mode <strong>of</strong> endothermic<br />

temperature physiology with the mammals adds<br />

little elucidation: the same contentious issues apply<br />

to them. <strong>The</strong> problem arises because <strong>of</strong> the complex<br />

nature <strong>of</strong> endothermy. It has two distinct primary<br />

functions in modern mammals, <strong>and</strong> it also involves

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