The Origin and Evolution of Mammals - Moodle

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(a) (b) (d) (c) Oxyaena Patriofelis Vulpavus Miacis Sinopa Pristinictis (e) (f) Protictis Figure 7.14 Creodonts and early carnivorores. (a) Skull of the oxyaenid creodont Oxyaena. Length approx. 20 cm (Romer 1966, after Wortman and Matthew). (b) Skeleton of the oxyaenid creodont Patriofelis (Gunnell 1998, after Osborn). (c) Skull and skeleton of the hyaenodont creodont Sinopa. Skull length approx. 15 cm (Romer 1966, after Wortman and Matthew, and Gunnell 1998 after Matthew). (d) Pristinictis. Upper molars in occlusal view and lingual view of an incomplete lower molar (Fox and Youzwyshn 1994). (e) Skull of the basal caniform miacid Vulpavus. Length approx. 10 cm (Savage and Long 1986). Upper dentition in occlusal view and lower dentition in labial view of Miacis. (Flynn 1998, figure 6.3). (f) Skull of the viverravid Protictis. Length approx. 9 cm (Carroll 1988 after MacIntyre 1996) and upper dentition in occlusal view and lower dentition in labial view of Protictis (Flynn 1998, figure 6.3).
250 THE ORIGIN AND EVOLUTION OF MAMMALS Hyaenodontids such as Sinopa (Fig. 7.14(c)) tended to be more agile, with slender jaws, and a range of body sizes comparable to the modern-day carnivorans. In feeding specialisations too, they radiated into a variety of highly active predators at one end and presumed bone-crushing scavengers at the other. The hyaenodontids also survived for far longer than the oxyaenids, not finally disappearing until the Miocene of Europe, Asia and Africa. As already mentioned, the Carnivora actually predated the creodonts, for the earliest members were Pappictidops, a viverravid from the Early Palaeocene Shanghuan Formation of China (Wang et al. 1998; Lucas 2001) and the Early Palaeocene Canadian genera Pristinictis and Ravenictis. At one time, all the early, primitive carnivorans were included in a group ‘Miacoidea’, which was assumed to be basal to the whole order. However, recent cladistic analysis indicates that different ‘miacoids’ are related to different lineages of the Carnivora (Flynn 1998; Janis et al. 1998b). The family Viverravidae (Fig. 7.14(f)), including the Early Palaeocene members mentioned, are related to the Feliformes on the basis of reduction of the number of molars to two, and an elongated skull. The family Miacidae (Fig. 7.14(e)), which do not appear until the very end of the Palaeocene, are primitive Caniformes as indicated, for example, by the relatively shorter skull, and the loss of contact between the calcaneum and fibula in the ankle. Surprisingly therefore, the two respective main lineages of modern Carnivora can be traced by fossils back to the Early Palaeocene. All the primitive, Palaeocene members were relatively small carnivores, varying only from the size of a weasel to that of a small cat, and it was not until the rise of the modern families such as the canids, felids, and ursids later in the Cenozoic that any members of the order achieved the large body size that certain mesonychids and creodonts had done far earlier. The origin and radiation of the modern orders The post-Palaeocene fossil record of placental mammals and its associated literature is enormous, and far beyond the scope of this work to deal with in more than the barest outline. The times of first occurrence, nature of the early members and their possible relationships, and palaeogeographic distribution are of the most immediate relevance for understanding the Cenozoic mammalian evolution. It is also, of course, irresistible to deal briefly with the more unfamiliar, often quite bizarre kinds of extinct mammals. Several modern orders appear in the Palaeocene fossil record, undoubtedly the Carnivora in the Early Palaeocene, and the Xenarthra, Rodentia, Eulipotyphla, and possibly Perissodactyla in the Late Palaeocene. However, even in these cases, their main radiation did not commence until the Eocene. The molecular evidence for four super-orders of modern placental mammals has been discussed earlier in the chapter, and is taken as the basis for the following account. As it happens, this radical new phylogenetic hypothesis is surprisingly little in conflict with conventional interpretations of the fossil record, apart from two aspects. One is the extensive taxonomic redistribution of the primitive ‘insectivorous’ orders; the other is the recognition that the ‘ungulate’ orders arose within two independent radiations. Otherwise, the molecular story contributes towards the formulation of new answers to several long-standing questions that the fossils alone have been unable to provide. Xenarthra Only the three living sub groups Cingulata (armadillos), Pilosa (sloths), and Vermilingua (anteaters) constitute the Xenarthra (Delsuc et al. 2001), now that a relationship with either the old world Pholidota, or the North America Palaeocene Palaeanodonta has been discarded (Rose and Emry 1993). The earliest fossil record consists of Late Palaeocene armadillo scutes of Prostegotherium in the Itaboraí fissure deposits of Brazil. Armadillos diverged throughout the Cenozoic of South America, and a few genera reached North America in the Late Pliocene faunal interchange. The Glyptodonta were the most spectacular cingulate xenarthrans, with their bony helmet, hugely exaggerated carapace of interlocking scutes, and tail massively protected by rings of bone. The cheek teeth, all that remain of the dentition, are tri-lobed,
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The Origin and Evolution of Mammals
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The Origin and Evolution of Mammals
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Preface This book arose from twin a
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For Mal /gosia with love and thanks
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Contents 1 Introduction The definit
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CHAPTER 1 Introduction There are ab
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transversely occluding molar teeth
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the Mesozoic mammals, is to do with
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a hierarchy of committees under the
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it means that a 200 Ma igneous rock
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een a more fundamental impact than
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Table 2.3 Classification of marsupi
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(a) (b) (c) humerus radius aquatic
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(a) Westlothiana Limnoscelis PO Wes
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the ankle bones to form an astragal
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(b) (c) (a) Casea rutena Casea broi
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(Fig. 3.2(f)), is actually an ophia
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the first one is enlarged, often to
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Even at their initial appearance in
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(a) Nikkasaurus (b) (d) Reiszia (e)
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Nikkasauridae The family Nikkasauri
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has figured large in discussions of
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(c) Figure 3.9 (continued). Syodon
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a mixture of progressively more spe
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animal with interdigitating, but no
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(e) (a) Anomocephalus Ulemica (b) S
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mandibulae musculature. This, as in
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To date the postcranial skeleton of
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ecognised four subgroups, based mai
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the presence of a deep, longitudina
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collected from the Lystrosaurus Ass
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(a) (d) (c) (b) Leontocephalus V PA
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(a) (c) Lycosuchus Oliveria (d) EVO
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separate families. Lycosuchidae (Fi
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consist of a large labial cusp and
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Procynosuchus (d) Procynosuchus (a)
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They constitute the monophyletic gr
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Although there is no mammal-like co
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Cynognathidae Cynognathus (Fig. 3.2
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(e) (f) Figure 3.22 (continued). Ma
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(c) (d) (a) (b) Kayentatherium EVOL
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Pachygenelus (e) (a) (c) Therioherp
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palatal, and orbitosphenoid bones,
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Wible and Hopson 1993). The interor
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published a cladogram based on rece
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310-320 Ma. By this time, the great
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are forms such as Casea itself, var
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lakes and swamps in the low-lying l
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urrowing and subsisting on a diet o
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Eothyris Haptodus sphenacodontine B
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z (a) (c) a.pt.m. M B PO P P SQ P M
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(b) a.pt.m. temp.m. a.pt.m. temp.m.
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the dentary bone above the level of
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the therocephalian grade was not on
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A balance was also achieved in the
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Locomotion Ancestral amniote grade
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screw-shaped: the front part faces
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For the recovery phase, the relativ
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SC PRC p.i.f.i tr.min (c) dp.cr ect
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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
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conducted the experiment of wrappin
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mammals themselves that the enlarge
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elevation of maximum aerobic activi
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a mammal. The difficulty with all s
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collection, ingestion, and assimila
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were edaphosaurid and caseid pelyco
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CHAPTER 5 The Mesozoic mammals The
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(a) (d) AL condylar foramina are se
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evidence to relate the haramiyidans
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postcranial skeleton, and Graybeal
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side of the head can be in action a
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the lower molars is relatively high
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morganucodontan teeth. However, des
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adjacent lower molars. A small exte
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(a) (b) (d) P4 P4 Plagiaulax P4 Pau
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American Morrison Formation genus C
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a self-sharpening property. As the
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near parasagittal gait (Fig. 5.11(e
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pass through the birth canal. Relat
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(a) 1 (b) (d) me (c) me.d 3 styl st
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(a) (c) Henkelotherium Crusafontia
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pubis is excluded from the acetabul
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Cretaceous, (Aptian or Albian) of M
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eautifully preserved placentals fro
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subsequent specimens revealed that
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Minimal divergence of tribosphenida
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(c) (a) M 1 M 1 M 2 Steropodon M 2
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(b) (a) (c) pa.d pr.d me.d Ambondro
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crown-group therians) is accepted b
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form of the tooth, must reflect sub
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of feasibility that a taxon of endo
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mammals, by creating environmental
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the 11 species of marsupial, of whi
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(d) (a) pa A Dasyuromorphia B C pr
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earing two huge claws on digits thr
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that contains the independent ances
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(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
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Page 236 and 237: effectively absent. However, increa
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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
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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
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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
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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
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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
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cladogenesis in dasyurid marsupials
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(eds) Evolution of Tertiary mammals
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McKenna, M.C. and Bell, S.K. 1997.
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(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
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Index Note: Page numbers in italics
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Equoidea 262 Equus 262 Erethizon 28
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Overkill hypothesis 289, 290 Oxyaen
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Tulerpeton 14, 18 Tylopoda 264 Ukha
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The Origin and Evolution of Mammals - Moodle

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