The Origin and Evolution of Mammals - Moodle

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(a) (b) (d) P4 P4 Plagiaulax P4 Paulchoffatia (c) (e) THE MESOZOIC MAMMALS 153 Nemegtbaatar Ptilodus Sudamerica Figure 5.9 Multituberculate skulls and Gondwanatheria. (a) Paulchoffatia. (Kielan-Jaworowska and Nessov 1992 and Maio 1993). (b) Plagiaulax. (Kielan-Jaworowska and Nessov 1992) (c) Nemegtbaatar (Kielan-Jaworowska and Nessov 1992 and Maio 1993). (d) Ptilodus in lateral, ventral and dorsal views (Kielan-Jaworowska and Nessov 1992 and Maio 1993). (e) Sudamerica fragment of jaw and teeth (Pascual et al. 1999). P4 i i
154 THE ORIGIN AND EVOLUTION OF MAMMALS blade-like lower premolar tooth, it became clear that these are not multituberculates at all, but a quite separate group, Gondwanatheria. They have since been described from Madagascar and India as well (Krause et al. 1997), and there is a possible, but virtually indeterminate jaw fragment from Tanzania (Krause et al. 2003). Thus the group is restricted to the Gondwanan continents. The molar teeth of gondwanatheres are hypsodont, and have transverse enamel ridges that develop on the occlusal surface, once the softer dentine and enamel has worn down. They were presumably yet another group of Mesozoic mammals adopting the rodent-like habitat of small herbivore, feeding on tough, high-energy vegetable matter. In the absence of more material their relationships remain totally obscure. Diversity and evolution The skull of multituberculates (Fig. 5.9) is broad with a short, blunt snout, and laterally facing orbits. The jugal, long thought to be absent, is now known to consist of a very thin plate attached to the inner face of the otherwise robust zygomatic arch (Hopson et al. 1989). Other cranial characters in which they differ from other mammals are the absence of a component of the palatine bone in the orbital wall, although Hurum (1994) noted some exceptions, and the nature of the specialised jaw joint permitting antero-posterior movements of the lower jaw. Perhaps to free the jaw for this propalinal movement, multituberculates possessed ear ossicles free of any connection to the dentary bone (Fig. 5.12(b)), and which were therefore more or less identical to those of living mammals (Miao and Lillegraven 1986; Hurum et al. 1996). The dentition is the most strikingly unique character of the group. The upper incisors are reduced in number to a maximum of three, the second of which is enlarged, and worked against the single, large, procumbent lower incisor. The lower premolars are equally distinctive, with serrated, blade-like crowns that had a shearing or chopping action against the multicusped upper premolars. The molars are reduced to two in number. Both the uppers and the lowers have two rows (three in the upper molars of more advanced forms) of up to eight blunt cusps arranged longitudinally, and with no connecting ridges between them. When in occlusion, the rows of cusps of the upper molars fit into the valleys between cusp rows of the lowers, and vice versa, and the posterior shift of the lower jaw then caused a very effective grinding action, as discussed shortly. The traditional classification of the approximately 70 genera of multituberculates was into three superorders, the Plagiaulacoidea for the more primitive ones of the Upper Jurassic and European Early Cretaceous; the Ptilodontoidea for the advanced, predominantly North American forms; and the Taeniolabidoidea for the advanced, predominantly Asian forms (e.g. Clemens and Kielan- Jaworowska 1979; Hahn and Hahn 1983). However, subsequent attempts at formal cladistic analyses have amply confirmed the suspicion that this classification is a very poor reflection of the phylogeny of the group (Simmons 1993; Rougier et al. 1997; Kielan-Jaworowska and Hurum 1997). On the other hand, there is far from a consensus amongst these authors on what the true interrelationships are. In the most recent review, Kielan-Jaworowska and Hurum (2001) analysed 62 characters amongst 32 taxa and obtained 17,783 equally parsimonious trees. This extremely low resolution results from a combination of conservatism in many characters, extensive convergence and mosaic evolution particularly in the dentition, and the inevitable problem of missing characters in many of the taxa that are known from very incomplete material. At any event, they were unable to derive from the cladogram a satisfactory cladistic classification, and instead proposed to retain the admittedly paraphyletic group ‘Plagiaulacida’ for the generally primitive forms such as Plagiaulax itself, and a possible but by no means very strongly supported group Cimolodonta for many but not all the more advanced forms (Kielan-Jaworowska et al. 2004). Paulchoffia is the most plesiomorphic ‘plagiaulacid’ (Fig. 5.9(a)). It comes from the Upper Jurassic Guimarota Lignite Mine in Portugal and has retained what is presumed to be the primitive multituberculate dentition of I3/1: C0/0: PM5–4/4–3: M2/2. The lower premolars differ from those of all other multituberculates in their rectangular shape, and horizontal wear facets on the crowns that indicate a crushing rather than shearing function. The molar teeth have a relatively small number of cusps, there being only three or four in each row. The North
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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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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
Page 122 and 123: spc s.sp s.sp SC PRC (d) delt T AST
Page 124 and 125: no significant novelties to what ha
Page 126 and 127: (a) (c) (e) (g) D D ex. au.m C tym
Page 128 and 129: (a) (c) (d) PMX (f) V n.turb mx.tur
Page 130 and 131: ol.b (a) (b) cer.hem gorgonopsian t
Page 132 and 133: (a) (b) (i) (ii) (iii) (iv) (v) a g
Page 134 and 135: 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
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 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
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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
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(d) Figure 6.6 (continued). Thylaco
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(a) (c) Caroloameghina and Procarol
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(a) (e) (d) Proargyrolagus Groeberi
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(a) (b) Djarthia Thylacotinga (c) (
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LIVING AND FOSSIL MARSUPIALS 211 M
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Extinct Notoryctemorphia At present
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(f) (g) Wakaleo Diprotodon optatum
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fossil mammals, which might help cl
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30 40 50 60 Figure 6.13 Southern Go
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the Diprotodontia also included gen
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1. Placentalia 2. Edentata 3. Epith
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effectively absent. However, increa
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Asioryctes (a) (b) (d) Cimolestes p
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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
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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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The Origin and Evolution of Mammals - Moodle

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