The Origin and Evolution of Mammals - Moodle

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(a) (e) (d) Proargyrolagus Groeberia Klohnia immediately by high-crowned premolars, three upper and two lower. There are then four lowcrowned, squarish molars with flat occlusal surfaces covered in thick enamel. A powerful adductor (c) (b) LIVING AND FOSSIL MARSUPIALS 207 Argyrolagus Figure 6.8 Argyrolagoidea. (a) Partial skull of the Oligocene Proargyrolagus bolivianus in lateral view. Length approx. 3 cm (Sánchez-Villagra and Kay 1997). (b) Skull of Argyrolagus scaglii in ventral and lateral views. Skull length approx. 5 cm (Rich 1991 from Simpson 1970). (c) Postcranial skeleton of A. scaglii (Simpson 1970). (d) Jaw fragment of Klohnia in occlusal and lateral views (Flynn and Wyss 1999) (e) Anterior fragment of snout and lower jaw in lateral view and lower jaw in postero-lateral view of Groeberia minoprioi (redrawn after Pascual et al. 1994). musculature had evolved to operate the dentition, as indicated by a deep extension of the zygomatic arch below the orbit. Pascual et al. (1994), in describing new specimens, concluded that the diet was
208 THE ORIGIN AND EVOLUTION OF MAMMALS probably nuts and other hard food, and that in addition to the gnawing and breaking function of the incisors, a particularly well-developed tongue may have been involved, working in association with the symphyseal floor. Nothing is known of the rest of the skeleton. A second groeberiid, Klohnia (Fig. 6.8(d)), has been described from the late Eocene/Early Oligocene by Flynn and Wyss (1999), and it too is poorly known, with little more than partial dentitions preserved. A third possible member, Patagonia (Pascual and Carlini 1987), was originally described as a member of yet another family of specialised, rodent-like caenolestoids. It is based on a single partial lower jaw with a gliriform incisor. Flynn and Wyss (1999) made a preliminary analysis of the interrelationships and concluded that Groeberia, Klohnia, and Patagonia do indeed constitute a valid family, and that Groeberiidae is indeed the sister group of Argyrolagidae. Australidelphia: the Australian radiation As discussed earlier, the Australasian marsupials form a monophyletic group, provided that the living microbiotherian Dromiciops and its fossil relatives are admitted. Unlike the South American radiation, in Australia the marsupials included middle- to largesized herbivores as well as the carnivores. Indeed, apart from the minute number of monotremes, numerous bats, and a few dozen species of murid rodents that entered the continent in the later part of the Cenozoic from Asia, the entire pre-human mammalian fauna was marsupial. Also in contrast to South America, the fossil record of Tertiary Australian marsupials during the critical early stages is extremely poor (Vickers-Rich et al. 1991; Long et al. 2002). Nothing at all is known of the group during the Palaeocene, and the earliest record is the very modest Tingamarra fauna of Queensland, which is dated approximately 55 Ma, and so is latest Palaeocene or earliest Eocene. This is followed by another relatively huge gap; the next window on to the radiation is not until Late Oligocene times, around 25 Ma, with several fossil-yielding localities, including the start of the great tropical rain forest Riversleigh system, with its abundance of beautifully preserved fossils continuing through the Miocene (Archer et al. 1991). Other important, though less rich fossil-bearing areas from the Late Oligocene include the Etadunna Formation in the Lake Eyre Basin of South Australia, and the Namba Formation which is also in South Australia. With one exception, all the Oligocene and Miocene marsupials are accommodated in one or another of the four extant Australian orders. At a lower taxonomic level, 19 of the total of 23 families of australidelphians listed by McKenna and Bell (1997) occur in the Oligocene–Miocene, and of the 10 extant families only 4 are as yet unknown as fossils from that time. Thus, by Riversleigh times a taxonomically richer version of what is essentially the modern marsupial fauna existed; nevertheless there are a few surprises and mysteries. The Tingamarra marsupials The Early Eocene or possibly Late Palaeocene fossils from the Tingamarra Local Fauna, near Murgon in southeastern Queensland, are the oldest marsupial mammals from Australasia (Godthelp et al. 1992). Unfortunately, only a few very fragmentary jaws and isolated teeth are preserved, and none of them are clearly attributable to particular modern groups. Djarthia (Fig. 6.9(a)) was described by Godthelp et al. (1999) on the basis of several jaws and teeth. Its upper molars are dilambdodont, having a V-shaped centrocrista, and there are four well-developed stylar cusps. The proportions of the talonid and trigonid of the lower molars are of standard marsupial pattern. From these dental characters, it is impossible to tell whether Djarthia is a didelphid or a basal australidelphian, yet this is a fundamentally important issue for understanding Australian marsupial history. If Djarthia is a didelphid, then it means that both ameridelphians and australidelphians occurred together there. If it is a basal australidelphian, then the Australian radiation can be accounted for as a consequence of a single didelphid-like immigrant into the continent prior to the Eocene. A second Tingamarra form is equally tantalising. Archer et al. (1993) described isolated teeth of Thylacotinga (Fig. 6.9(b)) as a possible peramelemorph. However, this view has been revised, and Long et al. (2002) regard it as a possible member of the South American and Antarctic group Polydolopoidea. The molar teeth have swollen, bunodont cusps. The stylar shelf of the upper molar is
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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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Page 172 and 173: pass through the birth canal. Relat
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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
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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
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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 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
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Page 232 and 233: the Diprotodontia also included gen
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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
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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
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coexist with other characters that
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The salient feature of Widanelfasia
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1 (a) 1. Perissodactyla 2. Titanoth
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(a) (b) BUNODONTIA or SUIFORMES SUI
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time, which suggests that it was on
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condition. This particular combinat
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etween Primates, Dermoptera (colugo
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have postcranial adaptations for ar
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undoubtedly related at a supraordin
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indisputably to occur prior to the
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The Late Cretaceous mammal record i
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interordinal divergences in the Lat
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diversity on Earth (Janis 1993). Fu
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effect of the surrounding sea. Trop
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was high. It lasted from 5 to 3 Ma
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and thus remain in their preferred
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agreement about exactly when within
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References Abdala, F. Redescription
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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
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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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