The Origin and Evolution of Mammals - Moodle

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(b) (a) (c) pa.d pr.d me.d Ambondro Ausktribosphenos Asfaltomylos hy.d hyl.d en.d me.d pr.d pa.d Australosphenidans (d) Boreosphenidans Steropodon Ausktribosphenos Ambondro Wrapping cingulid Henkelotherium Peramus THE MESOZOIC MAMMALS 179 Twinned pa.d and me.d No cingulid 1mm Kielantherium Aegialodon 1mm Montanalestes No cingulid Wrapping cingulid Twinned pa.d and me.d Wrapping cingulid 1mm No cingulid No cingulid Figure 5.22 Australosphenidans. (a) Occlusal view of lower second molar of Ambondro mahabo. (Flynn et al. 1999). (b) External and internal views of lower jaw of Ausktribosphenos nyctos, with occlusal view of a generalized lower molar (Rich et al. 1997). (c) Lateral view of lower jaw of Asfaltomylos patagonicus. Length approx. 6.7 cm (Rauhut et al. 2002) (d) Characters of australosphenidans compared to tribosphenidans (boreosphenidans) (Luo et al. 2002). 1mm 1mm 1mm 1mm 1mm
180 THE ORIGIN AND EVOLUTION OF MAMMALS Since the first proposal of a taxon Australosphenida, additional Gondwana forms have been discovered, and attributed to the group, most notably Asfaltomylos (Fig. 5.21(c)), which extends the geographical range of the group to South America (Rauhut et al. 2002). It is from the Middle to Late Jurassic of Argentina, and the only specimen so far consists of a damaged but complete lower jaw showing a possible six premolar teeth and definitely three molars. Rich et al. (2001a) described another Early Cretaceous Australian jaw, Bishops, which resembles Ausktribosphenos and which Luo et al. (2002) therefore include in the australosphenidans. Several authors disagree strenuously with the Australosphenidan concept. Sigogneau-Russell et al. (2001) have commented that, while a relationship between Ausktribosphenos and Steropodon may be well supported, Ambondro certainly does not fit at all well, for its tooth is different in a number of respects. Rich et al. (2002) strongly argued against a relationship between on the one hand Teinolophos, Steropodon and the other monotremes, and on the other Ausktribosphenos and Bishops, disputing the characters supposedly uniting them. They continue to believe that the latter two are related to the true tribosphenidans. Pascual et al. (2002) go so far as to revert to an old view that monotreme molar teeth are not even basically tribosphenic, but that they were derived from a completely ‘non-therian’ ancestry. Woodburne et al. (2003) undertook a cladistic analysis, based on 51 characters, most dental but a few mandibular. They found that the monotremes, including Steropodon and Teinolophos as basal members, are the sister group of all the therian mammals. Furthermore, the disputed genera Ambondro, Ausktribosphenos, Asfaltomylos, and Bishops constitute a monophyletic group that nests within the stem placentals. So far only incomplete lower jaws and partial lower dentitions of putative australosphenidans have been found. It is therefore too early to be confident that it really is a well-founded taxon, let alone that it evolved independently such an apparently complex dental structure as the tribosphenic tooth, complete with neomorphic protocone and huge expansion of the talonid. More material will doubtless test this nevertheless exciting, and at least biogeographically satisfying concept. Meanwhile, the solution to the mystery of the monotremes continues to be elusive. An overview of the interrelationships and evolution Interrelationships Several cladistic analyses of the interrelationships of Mesozoic mammals (Fig. 5.23) have been published in recent years and there is good agreement on a number of issues, notably concerning the abandonment of any vestige of the old, simple dichotomy into ‘therian’ and ‘non-therian’ mammals. In its place, an increasingly crownward sequence consisting of Sinoconodon, morganucodontans, docodontans, eutriconodontans, and Trechnotheria (stem-group plus Haramiyida Adelobasileus Sinoconodon Morganucodonta Docodonta Hadrocodium Kuehneotherium Australosphenida Shuotherium Tinodon Eutriconodonta Multituberculata Spalacotheriidae Dryolestoidea Vincelestes Peramus Aegialodontidae Deltatherida Marsupialia Placentalia Figure 5.23 Simplified version of Luo et al.’s (2002) phylogeny of the major Mesozoic mammal groups.
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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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Page 142 and 143: a mammal. The difficulty with all s
Page 144 and 145: collection, ingestion, and assimila
Page 146 and 147: were edaphosaurid and caseid pelyco
Page 148 and 149: 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 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
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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 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
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Page 236 and 237: effectively absent. However, increa
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and Prokennalestes, although it doe
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(a) Leptictidium Palaeoryctidans we
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(a) Purgatorius (c) are part of the
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(a) (d) (e) (g) Protoungulatum Meso
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(a) (f) (e) Hyopsodus Phenacodus (d
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Titanoides (pantodont) (a) (c) (b)
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(a) (b) (d) Trogosus (tillodont) Es
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Mioclaenus Paulacoutoia (didolodont
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their presence. The five orders may
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Xenungulata. Only two Late Palaeoce
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(a) (b) (d) (c) Oxyaena Patriofelis
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continuously growing, and form a gr
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navicular facet, 19 or more thoraci
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(a) (c) Phosphatherium Numidotheriu
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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
Page 318 and 319:
cladogenesis in dasyurid marsupials
Page 320 and 321:
(eds) Evolution of Tertiary mammals
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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
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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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