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The Origin and Evolution of Mammals - Moodle

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160 THE ORIGIN AND EVOLUTION OF MAMMALS<br />

(a)<br />

ol.b<br />

ol.b<br />

Pit<br />

Pit<br />

cer.hem<br />

cer.hem<br />

<strong>The</strong> brain <strong>of</strong> multituberculates (Fig. 5.12(a)) has<br />

been reconstructed from the endocranial casts <strong>of</strong> a<br />

number <strong>of</strong> genera, particularly Chulsanbaatar <strong>and</strong><br />

Nemegtbaatar (Kielan-Jaworowska et al. 1986) <strong>and</strong><br />

Ptilodus (Simpson 1937; modified by Krause<br />

<strong>and</strong> Kielan-Jaworowska 1993). All give the same<br />

general picture <strong>of</strong> relatively very large olfactory<br />

bulbs <strong>and</strong> no exposure <strong>of</strong> midbrain structures dorsally,<br />

which suggests that olfaction was a more<br />

important sense than vision, <strong>and</strong> therefore that they<br />

had a nocturnal habit. <strong>The</strong> relative size <strong>of</strong> the brain<br />

<strong>of</strong> Ptilodus, in terms <strong>of</strong> the Encephalisation Quotient<br />

(actual brain weight divided by expected brain<br />

weight for a mammal <strong>of</strong> that body weight) has been<br />

estimated by Krause <strong>and</strong> Kielan-Jaworowska (1993).<br />

It lies between a minimum <strong>of</strong> 0.37 <strong>and</strong> a maximum<br />

<strong>of</strong> 0.62, depending on which procedure is used, <strong>and</strong><br />

whether the olfactory bulbs are, or are not included<br />

in the estimates <strong>of</strong> brain mass. Despite the lack <strong>of</strong><br />

precision, it is clear that the multituberculate brain<br />

fl<br />

fl<br />

cerb<br />

Chulsanbatar<br />

(b)<br />

os tympanicum<br />

process anterior<br />

caput mallel<br />

manubrium<br />

crus breve incudis<br />

crus longum incudis<br />

stapes<br />

Didelphis<br />

Figure 5.12 Brain <strong>and</strong> ear ossicles <strong>of</strong> multituberculates. (a) Reconstructed brain <strong>of</strong> Nemegtbaatar in dorsal, ventral, <strong>and</strong> lateral views<br />

(Kielan-Jaworowska et al. 1986). (b) Ear ossicles <strong>of</strong> Chulsanbaatar (left) compared to Didelphis (right) in situ, <strong>and</strong> enlarged. (Hurum et al. 1996).<br />

cerb, cerebellum; cer.hem, cerebral hemisphere; fl, flocculus; ol.b, olfactory bulb; pit, pituitary.<br />

was significantly smaller than those <strong>of</strong> modern<br />

mammals.<br />

Multituberculates are one <strong>of</strong> the extremely few<br />

fossil mammal groups from any period where<br />

the presence <strong>of</strong> hair has been positively demonstrated.<br />

Skeletal remains <strong>of</strong> Lambdopsalis along with<br />

the exceptionally detailed impressions <strong>of</strong> hair have<br />

been found in fossilised coprolites from Late<br />

Palaeocene beds <strong>of</strong> Inner Mongolia (Meng <strong>and</strong><br />

Wyss 1997). Unfortunately, there are no organic<br />

remains <strong>of</strong> the material, so the attractive prospect <strong>of</strong><br />

finding out what colour the animal was cannot be<br />

satisfied.<br />

Kielan-Jaworowska (1979) speculated that multituberculates<br />

were viviparous. She based her argument<br />

on the very acute angle at which the two sides<br />

<strong>of</strong> the pelvic girdle meet ventrally, about 40 o instead<br />

<strong>of</strong> the 180 o or so typical <strong>of</strong> most mammals. This<br />

would allow an egg or neonate with a diameter<br />

no greater than about 3.4 mm in Kryptobaatar to

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