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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(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