The Origin and Evolution of Mammals - Moodle

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coexist with other characters that are aquatic adaptations, including a dorsal position of the nostrils and a long row of very dense ribs. Desmostylia The desmostylians are a group of semiaquatic mammals as large as hippos. They were limited taxonomically to about half a dozen genera, temporally to the Late Oligocene and Miocene, and geographically to the east and west margins of the northern Pacific Ocean (Domning et al. 1986; Domning 2002a). Complete skeletons of Paleoparadoxia (Fig. 7.16(b)) demonstrate well-developed fore and hind limbs, with broad, presumably paddle-like hands and feet, suggesting an amphibious, seal-like existence. The incisors and canine teeth are well developed, and there is a long diastema. The cheek teeth resemble those of the early proboscideans, particularly Moeritherium. Clementz et al. (2003) analysed stable isotope contents of the bones of Desmostylia, and concluded that they could forage on sea grass, and also on land during terrestrial excursions. That desmostylians are paenungulates is not seriously disputed; a relationship with the tethythere grouping of proboscideans and sirenians is usually preferred (Fischer and Tassy 1993; Asher et al. 2003). Embrithopoda The embrithopods are best known as the celebrated Late Eocene Arsinoitherium (Fig. 7.16(d)) from the Fayum. It was a huge, rhinoceros-sized animal with the skull adorned with a pair of massive horns. Neither tusks nor enlarged canines are present, and the cheek teeth are bilophodont and hypsodont in form (Court 1992). Teeth of earlier and more primitive, hornless forms (Fig. 7.16(e)) occur in Romania and central Turkey (Maas et al. 1998). As with the desmostylians, there is more or less complete agreement that the Embrithopoda are part of the paenungulate radiation, but not about precisely where they fit. Tassy and Shoshani (1988) regarded them as the most basal paenungulate group. Court (1990), Fischer and Tassy (1993) and Asher et al. (2003) have them closer to the Proboscidea than to the Sirenia, within the Tethytheria. Macroscelidea Fossil elephant shrews occur only in Africa, the earliest record being teeth of Chambius from LIVING AND FOSSIL PLACENTALS 257 the Middle Eocene of Tunisia, followed by the Late Eocene Herodotius from the Fayum (Simons et al. 1991; Butler 1995). They have four-cusped, bunodont molars lacking lophs, similar to what is seen in later macroscelideans. Comparison of the dental characters of these primitive forms has been made with hyopsodontid ‘condylarths’ by a number of recent authors (Simons 1991 et al; Butler 1995). Most recently, Tabuce et al. (2001) have also supported the ‘condylarth’ origin of the Macroscelidea, and proposed that the dentition of the primitive members is close to the ancestral state for paenungulates. However, since the non-afrotherian Perissodactyla appear in their cladogram as relatives, convergent evolution of dental characters must have occurred at least at some level, which considerably blurs the picture. Tubulidentata Represented today by Orycteropus afer, the single species of aardvark, the fossil record of this group is scarcely any richer. The four or five known genera are all placed in the same family by McKenna and Bell (1997) and they date from the Oligocene of Europe and Early Miocene of Africa. There are subfossil bones of a very peculiar mammal in Madagascar, named Plesiorycteropus, preserved along with giant lemurs, elephant birds, etc. in Quaternary deposits. Representatives of most of the bones are present, except for the facial region, jaws, and dentition. It was a heavily built, digging animal, probably with a long narrow snout and lacking teeth. Most commentators have regarded Plesiorycteropus as a tubulidentate. However, McPhee (1994) failed to find any characters uniquely shared with Orycteropus, and his cladistic analysis did not group them exclusively together. He concluded that Plesiorycteropus is sufficiently distinct from any other placentals as to warrant its own order, Bibymalagasia. In the most recent analysis, Asher et al. (2003) found that Plesiorycteropus is at least a member of Afrotheria. Afrosoricida Seiffert and Simons (2000) have described lower jaw fragments of a Late Eocene form from the Fayum fauna, which they claim may be related to chrysochloridans and/or tenrecs, or possibly to the West Indian eulipotyphlan family Solenodontidae.
258 THE ORIGIN AND EVOLUTION OF MAMMALS (c) (d) Hoplophoneus (e) (a) Hesperocyon (b) Proailurus Enaliarctos Barbourofelis Figure 7.18 Carnivora. (a) Skull and skeleton of the Middle Eocene canid Hesperocyon. Presacral length approx. 45 cm (Munthe 1998 and Savage and Long 1986). (b) Skull and skeleton of the Early Miocene felid Proailurus (Carroll 1988 from Radinsky 1982, and Turner and Anton 1997). (c) The nimravid Hoplophoneus. Height at shoulder 48 cm (Turner and Anton 1997). (d) The nimravid Barbourofelis (Martin 1998). (e) The Oligoncene pinniped Enaliarctos. Presacral length approx. 1.25 m (Berta and Ray 1990).
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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
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elieved to be Late Cretaceous in ag
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(Fig. 6.5(c)). The dentition exhibi
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(d) Figure 6.6 (continued). Thylaco
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(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 240 and 241: and Prokennalestes, although it doe
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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
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 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 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
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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
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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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