The Origin and Evolution of Mammals - Moodle

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(e) (a) Anomocephalus Ulemica (b) Suminia Patronomodon (f) Otsheria EVOLUTION OF MAMMAL-LIKE REPTILES 41 (g) (c) Galechirus (d) Galeops Figure 3.11 Basal anomodonts. (a) Anomocephalus: actual size 20 cm long (Modesto et al. 1999). (b) Patronomodon nyaphulii: approx. skull length 6 cm (Rubidge and Hopson 1996). (c) Galechirus scholzi: approx. skull length 6.8 cm. (d) Galeops whattsii: approx. skull length 4.5 cm (King 1988 after Brinkman 1981). (e) Ulemica invisa: skull length approx. 15 cm (Ivakhnenko 1996 and King 1994). (f) Otsheria netzvetajevi: skull length 10.5 cm (Ivakhnenko 1996). (g) Suminia getmanovi: skull length 5.8 cm (Rybszynski 2000).
42 THE ORIGIN AND EVOLUTION OF MAMMALS Copper sandstone lower jaw as the type and only safely attributed specimen of V. prima. Thus, although the family Venyukoviidae still stands, Venyukovia itself is scarcely known. Meanwhile, Tchudinov (1960, 1983) had described another primitive anomodont as Otsheria netzvetajevi, based on a skull without the lower jaw (Fig. 3.11(f)). This specimen is older than the Isheevo material, for it was found at the Ezhovo (Ocher) locality which is dated as late Kazanian. Most recently a fourth, quite different form, Suminia (Fig. 3.11(g)), has been described from the Late Tatarian Kotel’nich fauna, which is therefore the youngest of the Russian primitive anomodonts (Ivakhnenko 1994; Rybczynski 2000). A cladistic analysis by Modesto and Rybczynski (2000; Rybczynski 2000) concludes that the Russian genera are a monophyletic group characterised principally by an elongated postero-dorsal process of the premaxilla, a very broad parietal bone with the pineal foramen situated on a large boss at the front of it, and no preparietal bone. Otsheria (Fig. 3.11(f)) is thus the earliest of the at least reasonably well-known genera of venyukovioids (Tchudinov 1960, 1983). It has a small, 10-cm long skull that differs mainly from other venyukovioids in its dentition. Although not very well preserved, this can be seen to consist of probably four incisors in the premaxilla and nine maxillary teeth, all of which are short and stout, with laterally compressed points. In the absence of any information at all about the lower dentition, it is not clear whether there was tooth-totooth contact. So far as it may be inferred, the form of the individual upper teeth suggests a simple, foliagecutting action, although a generalised omnivorous diet might well have been the case. Ulemica (Fig. 3.11(e)) is based on two fairly complete skulls, several partial skulls and lower jaws, but so far no postcranial material, and all from the Early Tatarian Isheevo locality (Ivakhnenko 1996). The skull is 15–20 cm in length and heavily built. The dentition of Ulemica is very odd and difficult to understand functionally (King 1994; Ivakhnenko 1996). In the upper jaw, there is a large chisel-shaped first tooth, followed by three of similar size but with heellike internal extensions. These are followed by four small conical teeth and a much larger, swollen, caniniform tooth. Finally the tooth row is completed by a series of four or five very small pointed teeth. The corresponding lower dentition consists of a chisel-shaped first tooth, followed by three smaller teeth that are laterally flattened in younger specimens, but replaced by blunter, rounded ones in older ones, and apparently lost in fully grown specimens. The tooth row is completed by a series of small, medially displaced teeth. The biting action consists of direct tooth-to-tooth contact between the anterior teeth. The upper caniniform tooth does not meet a lower tooth, but bites into a pit on the bony surface of the jaw, to the side of the lower teeth. Behind this point, the teeth do not appear to meet either. The upper postcaniniform teeth bite towards the lateral side of the lower teeth. The latter, in turn, bite towards the bone of the palate, medial to the upper teeth. King (1994) suggested that horny biting surfaces were present in life on these bony surfaces. Judging both from the form of the dentition and the structure of the articular and quadrate, no propalinal movement of the lower jaw was possible. Instead, the anteriormost teeth would have provided a nipping, food-gathering action, and the posterior teeth acting against their respective opposing tooth plates would have provided a simple crushing function. The diet for which such a system was adapted is not clear, but presumably included various kinds of vegetation. Suminia (Fig. 3.11(g)), as well as being the youngest of the primitive Russian anomodonts is also the best known, on the basis of several skeletons and skulls (Ivakhnenko 1994; Rybczynski 2000). The skull is small, no more than about 6 cm in length, and unlike Ulemica it is lightly built. There are complete upper and lower marginal dentitions of 10 or 11 teeth, which decrease in size gradually from front to back. Each of the teeth has a broad base and a serrated, leaf-like crown, uppers with a concave front edge, lowers with a concave hind edge. Rybczynski and Reisz (2001) have analysed the jaw action, showing that the power stroke of the lower jaw was posteriorly directed and created a cutting action between lower and upper teeth that included direct tooth-to-tooth contact. Their interpretation is supported by the structure of the jaw hinge, which permitted propalinal movement of the jaw, and by the lateral flaring of the zygomatic arch, the squamosal component of which bears a lateral fossa for the origin of a lateral slip of the adductor
Page 2 and 3: The Origin and Evolution of Mammals
Page 4 and 5: The Origin and Evolution of Mammals
Page 6 and 7: Preface This book arose from twin a
Page 8 and 9: For Mal /gosia with love and thanks
Page 10 and 11: Contents 1 Introduction The definit
Page 12 and 13: CHAPTER 1 Introduction There are ab
Page 14 and 15: transversely occluding molar teeth
Page 16 and 17: the Mesozoic mammals, is to do with
Page 18 and 19: a hierarchy of committees under the
Page 20 and 21: it means that a 200 Ma igneous rock
Page 22 and 23: een a more fundamental impact than
Page 24 and 25: Table 2.3 Classification of marsupi
Page 26 and 27: (a) (b) (c) humerus radius aquatic
Page 28 and 29: (a) Westlothiana Limnoscelis PO Wes
Page 30 and 31: the ankle bones to form an astragal
Page 32 and 33: (b) (c) (a) Casea rutena Casea broi
Page 34 and 35: (Fig. 3.2(f)), is actually an ophia
Page 36 and 37: the first one is enlarged, often to
Page 38 and 39: Even at their initial appearance in
Page 40 and 41: (a) Nikkasaurus (b) (d) Reiszia (e)
Page 42 and 43: Nikkasauridae The family Nikkasauri
Page 44 and 45: has figured large in discussions of
Page 46 and 47: (c) Figure 3.9 (continued). Syodon
Page 48 and 49: a mixture of progressively more spe
Page 50 and 51: animal with interdigitating, but no
Page 54 and 55: mandibulae musculature. This, as in
Page 56 and 57: To date the postcranial skeleton of
Page 58 and 59: ecognised four subgroups, based mai
Page 60 and 61: the presence of a deep, longitudina
Page 62 and 63: collected from the Lystrosaurus Ass
Page 64 and 65: (a) (d) (c) (b) Leontocephalus V PA
Page 66 and 67: (a) (c) Lycosuchus Oliveria (d) EVO
Page 68 and 69: separate families. Lycosuchidae (Fi
Page 70 and 71: consist of a large labial cusp and
Page 72 and 73: Procynosuchus (d) Procynosuchus (a)
Page 74 and 75: They constitute the monophyletic gr
Page 76 and 77: Although there is no mammal-like co
Page 78 and 79: Cynognathidae Cynognathus (Fig. 3.2
Page 80 and 81: (e) (f) Figure 3.22 (continued). Ma
Page 82 and 83: (c) (d) (a) (b) Kayentatherium EVOL
Page 84 and 85: Pachygenelus (e) (a) (c) Therioherp
Page 86 and 87: palatal, and orbitosphenoid bones,
Page 88 and 89: Wible and Hopson 1993). The interor
Page 90 and 91: published a cladogram based on rece
Page 92 and 93: 310-320 Ma. By this time, the great
Page 94 and 95: are forms such as Casea itself, var
Page 96 and 97: lakes and swamps in the low-lying l
Page 98 and 99: urrowing and subsisting on a diet o
Page 100 and 101: Eothyris Haptodus sphenacodontine B
Page 102 and 103:
z (a) (c) a.pt.m. M B PO P P SQ P M
Page 104 and 105:
(b) a.pt.m. temp.m. a.pt.m. temp.m.
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the dentary bone above the level of
Page 108 and 109:
the therocephalian grade was not on
Page 110 and 111:
A balance was also achieved in the
Page 112 and 113:
Locomotion Ancestral amniote grade
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screw-shaped: the front part faces
Page 116 and 117:
For the recovery phase, the relativ
Page 118 and 119:
SC PRC p.i.f.i tr.min (c) dp.cr ect
Page 120 and 121:
the therocephalian pelvis and hindl
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spc s.sp s.sp SC PRC (d) delt T AST
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no significant novelties to what ha
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(a) (c) (e) (g) D D ex. au.m C tym
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(a) (c) (d) PMX (f) V n.turb mx.tur
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ol.b (a) (b) cer.hem gorgonopsian t
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(a) (b) (i) (ii) (iii) (iv) (v) a g
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cold stress, the part that usually
Page 136 and 137:
conducted the experiment of wrappin
Page 138 and 139:
mammals themselves that the enlarge
Page 140 and 141:
elevation of maximum aerobic activi
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a mammal. The difficulty with all s
Page 144 and 145:
collection, ingestion, and assimila
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were edaphosaurid and caseid pelyco
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CHAPTER 5 The Mesozoic mammals The
Page 150 and 151:
(a) (d) AL condylar foramina are se
Page 152 and 153:
evidence to relate the haramiyidans
Page 154 and 155:
postcranial skeleton, and Graybeal
Page 156 and 157:
side of the head can be in action a
Page 158 and 159:
the lower molars is relatively high
Page 160 and 161:
morganucodontan teeth. However, des
Page 162 and 163:
adjacent lower molars. A small exte
Page 164 and 165:
(a) (b) (d) P4 P4 Plagiaulax P4 Pau
Page 166 and 167:
American Morrison Formation genus C
Page 168 and 169:
a self-sharpening property. As the
Page 170 and 171:
near parasagittal gait (Fig. 5.11(e
Page 172 and 173:
pass through the birth canal. Relat
Page 174 and 175:
(a) 1 (b) (d) me (c) me.d 3 styl st
Page 176 and 177:
(a) (c) Henkelotherium Crusafontia
Page 178 and 179:
pubis is excluded from the acetabul
Page 180 and 181:
Cretaceous, (Aptian or Albian) of M
Page 182 and 183:
eautifully preserved placentals fro
Page 184 and 185:
subsequent specimens revealed that
Page 186 and 187:
Minimal divergence of tribosphenida
Page 188 and 189:
(c) (a) M 1 M 1 M 2 Steropodon M 2
Page 190 and 191:
(b) (a) (c) pa.d pr.d me.d Ambondro
Page 192 and 193:
crown-group therians) is accepted b
Page 194 and 195:
form of the tooth, must reflect sub
Page 196 and 197:
of feasibility that a taxon of endo
Page 198 and 199:
mammals, by creating environmental
Page 200 and 201:
the 11 species of marsupial, of whi
Page 202 and 203:
(d) (a) pa A Dasyuromorphia B C pr
Page 204 and 205:
earing two huge claws on digits thr
Page 206 and 207:
that contains the independent ances
Page 208 and 209:
(b) (d) (a) Glasbius Alphadon (c) D
Page 210 and 211:
elieved to be Late Cretaceous in ag
Page 212 and 213:
(Fig. 6.5(c)). The dentition exhibi
Page 214 and 215:
(d) Figure 6.6 (continued). Thylaco
Page 216 and 217:
(a) (c) Caroloameghina and Procarol
Page 218 and 219:
(a) (e) (d) Proargyrolagus Groeberi
Page 220 and 221:
(a) (b) Djarthia Thylacotinga (c) (
Page 222 and 223:
LIVING AND FOSSIL MARSUPIALS 211 M
Page 224 and 225:
Extinct Notoryctemorphia At present
Page 226 and 227:
(f) (g) Wakaleo Diprotodon optatum
Page 228 and 229:
fossil mammals, which might help cl
Page 230 and 231:
30 40 50 60 Figure 6.13 Southern Go
Page 232 and 233:
the Diprotodontia also included gen
Page 234 and 235:
1. Placentalia 2. Edentata 3. Epith
Page 236 and 237:
effectively absent. However, increa
Page 238 and 239:
Asioryctes (a) (b) (d) Cimolestes p
Page 240 and 241:
and Prokennalestes, although it doe
Page 242 and 243:
(a) Leptictidium Palaeoryctidans we
Page 244 and 245:
(a) Purgatorius (c) are part of the
Page 246 and 247:
(a) (d) (e) (g) Protoungulatum Meso
Page 248 and 249:
(a) (f) (e) Hyopsodus Phenacodus (d
Page 250 and 251:
Titanoides (pantodont) (a) (c) (b)
Page 252 and 253:
(a) (b) (d) Trogosus (tillodont) Es
Page 254 and 255:
Mioclaenus Paulacoutoia (didolodont
Page 256 and 257:
their presence. The five orders may
Page 258 and 259:
Xenungulata. Only two Late Palaeoce
Page 260 and 261:
(a) (b) (d) (c) Oxyaena Patriofelis
Page 262 and 263:
continuously growing, and form a gr
Page 264 and 265:
navicular facet, 19 or more thoraci
Page 266 and 267:
(a) (c) Phosphatherium Numidotheriu
Page 268 and 269:
coexist with other characters that
Page 270 and 271:
The salient feature of Widanelfasia
Page 272 and 273:
1 (a) 1. Perissodactyla 2. Titanoth
Page 274 and 275:
(a) (b) BUNODONTIA or SUIFORMES SUI
Page 276 and 277:
time, which suggests that it was on
Page 278 and 279:
condition. This particular combinat
Page 280 and 281:
etween Primates, Dermoptera (colugo
Page 282 and 283:
have postcranial adaptations for ar
Page 284 and 285:
undoubtedly related at a supraordin
Page 286 and 287:
indisputably to occur prior to the
Page 288 and 289:
The Late Cretaceous mammal record i
Page 290 and 291:
interordinal divergences in the Lat
Page 292 and 293:
diversity on Earth (Janis 1993). Fu
Page 294 and 295:
effect of the surrounding sea. Trop
Page 296 and 297:
was high. It lasted from 5 to 3 Ma
Page 298 and 299:
and thus remain in their preferred
Page 300 and 301:
agreement about exactly when within
Page 302 and 303:
References Abdala, F. Redescription
Page 304 and 305:
Ax, P. 1987. The phylogenetic syste
Page 306 and 307:
Bramble, D. M. 1978. Origin of the
Page 308 and 309:
Colbert, E.H. 1948. The mammal-like
Page 310 and 311:
Duvall, D. 1986. A new question of
Page 312 and 313:
Gow, C.E. 1980. The dentitions of t
Page 314 and 315:
Ivakhnenko, M.F. 1990. The late Pal
Page 316 and 317:
Kemp, T.S. 1988a. A note on the Mes
Page 318 and 319:
cladogenesis in dasyurid marsupials
Page 320 and 321:
(eds) Evolution of Tertiary mammals
Page 322 and 323:
McKenna, M.C. and Bell, S.K. 1997.
Page 324 and 325:
(ed) The phylogeny and classificati
Page 326 and 327:
Ray, S. 2000.Endothiodont dicynodon
Page 328 and 329:
Rubidge, B.S., Kitching, J.W., and
Page 330 and 331:
Simpson, G.G. 1970. The Argyrolagid
Page 332 and 333:
Thewissen, J.G.M. 1990. Evolution o
Page 334 and 335:
Novacek, M.J., and McKenna, M.C. (e
Page 336 and 337:
Index Note: Page numbers in italics
Page 338 and 339:
Equoidea 262 Equus 262 Erethizon 28
Page 340 and 341:
Overkill hypothesis 289, 290 Oxyaen
Page 342:
Tulerpeton 14, 18 Tylopoda 264 Ukha
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The Origin and Evolution of Mammals - Moodle

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