The Origin and Evolution of Mammals - Moodle

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een a more fundamental impact than on mammalian systematics. It began with the now very primitivelooking immunological techniques generating such radical proposals as the close relationship of humans to the African apes (Sarich and Wilson 1967), and the monophyly of the Australian marsupials (Kirsch 1977). With the increased availability of comparative sequence, initially of proteins and subsequently nucleic acids, unfamiliar new hypotheses arrived, like the monophyly of Afrotheria, a morphologically very mixed bag of placental orders with no evident morphological characters in common at all, and the improbable-looking sister-group relationship of whales to hippos. However, both these are now widely, if not universally agreed upon by palaeontologists, who indeed have found support from fossils. In fairness of course, other early molecular proposals have disappeared without trace, such as guinea pigs not being rodents. In equal fairness, though, rejection of the latter was due primarily to further molecular evidence, rather than morphological evidence. The present position is that relatively enormous volumes of sequence data are already available across virtually the whole of living mammal diversity at the level of orders, and increasingly of families and lower taxa. This includes a mixture of sources. Many, and increasing numbers of complete mitochondrial genomes have been sequenced, along with a variety of nuclear genes. Databases for analysis are already ten to twenty thousand nucleotides long (10–20 kb), with presumably vastly increased potential resolving power. Nucleotide sequences have several advantages over morphological descriptions besides the more or less limitless amount of information available. It is easy in principle to recognise a single nucleotide as a unit character. It is also possible to have estimates of differential probabilities of different kinds of nucleotide substitutions, depending, for example, on whether it is a transition or a transversion, or first, second, or third placed position in the codon. Therefore, more realistic models of the evolutionary process are possible for inferring relationships from the data, with the help of suitably sophisticated methods such as maximum likelihood, and Bayesian inference. Again this subject is extensively discussed in several texts such as Salemi and Vandamme (2003). A few years ago, it was legitimate to question whether molecular phylogeny was more reliable than morphological, and consequently whether it should be allowed to overrule the latter in cases of conflicting hypotheses of relationships. A common reaction was that molecular and morphological data should be combined to produce a compromise phylogeny. This was always a conceptually difficult thing to achieve, and all too often simply resulted in a very poorly resolved cladogram instead. However, the increasing number of occasions on which newer palaeontological evidence, both morphological and biogeographical, has tended to support, or at least be compatible with the molecular evidence suggests that with a careful enough analysis, molecular data is perfectly capable of resolving the interrelationships of mammalian groups that have living members. Indeed, it is heralding an exciting new phase in mammalian palaeobiology. With much more strongly supported and widely agreed phylogenies to hand, the fossil record is open to a whole new set of improved interpretations about the complex interplay between phylogenesis, biogeography, and palaeoecology that constitutes the history of mammals. A classification of Synapsida TIME AND CLASSIFICATION 11 The classifications in Tables 2.1–2.4 are designed for general reference. All the formally expressed groups are monophyletic clades, but not all relationships between them are fully resolved to dichotomies. In the classifications of the Synapsida and the Mammalia, certain ancestral, or paraphyletic group names are included but in quotation marks, either because they are so familiar from the literature that they remain useful nomenclatural handles, or because the relationships of their constituent subgroups are inadequately known. Monophyletic groups whose cladistic position is relatively uncertain are indicated by a query. These classifications are designed for the relatively uninitiated in the field of synapsid systematics. They are therefore a compromise between a fully cladistic classification that would be overly name-bound, and would tend to convey overconfidence in the expressed relationships, and an ‘evolutionary’ classification with undeclared paraphyletic groups and the consequent loss of genealogical information. For more detail and discussion, the appropriate chapters should be consulted.
12 THE ORIGIN AND EVOLUTION OF MAMMALS Table 2.1 Classification of the Synapsida SYNAPSIDA Subclass “PELYCOSAURIA” Caseasauria Eothyrididae Caseidae “Eupelycosauria” Varanopseidae Ophiacodontidae Edaphosauridae Sphenacodontia Haptodontidae Sphenacodontidae Subclass THERAPSIDA Nikkasauridae (?) Niaftosuchus (?) Estemmenosuchidae (?) Biarmosuchia Biarmosuchidae Burnetiidae Dinocephalia Brithopia Titanosuchia Titanosuchidae Tapinocephalidae Styracocephalidae Anomodontia Anomocephalus Patronomodon Venyukovioidea Dromasauroidea Dicynodontia Eodicynodontidae Eudicynodontia Theriodontia Gorgonopsia (?) Eutheriodontia Therocephalia Cynodontia “Procynosuchia” Procynosuchidae Dviniidae Epicynodontia Galesauridae Thrinaxodontidae Eucynodontia For further details see Chapter 3. Modified after Reisz (1986), Kemp (1982, 1988c), Hopson (1991), and Hopson and Kitching (2001). Table 2.2 Classification of the Mesozoic mammals CLASS MAMMALIA Adelobasileus (?) Sinocondon Kuehneotheriidae (?) Morganucodontia Morganucodontidae Megazostrodontidae Docodonta Hadroconium “Eutriconodonta” Triconodontidae Gobiconodontidae Amphilestidae Subclass ALLOTHERIA Haramiyida (?) Multituberculata “Plagiaulacida” Cimolodonta TRECHNOTHERIA “Symmetrodonta” Amphidontidae Tinodontidae Spalacotheriidae “Eupantotheria” Dryolestidae Paurodontidae Vincelestidae Amphitheriidae Shuotheriidae (?) Subclass TRIBOSPHENIDA (�BOREOSPHENIDA) Aegialodontidae Metatheria Deltatheriidae (?) Marsupialia Placentalia (�Eutheria) Subclass AUSTRALOSPHENIDA (?) Ausktribosphenida Monotremata Simplified from Kielan-Jaworowska et al. (2004). The arrangement is cladistic except in the case of groups in quotation marks, which are paraphyletic, groups with query marks whose cladistic position is in doubt, and listed families which are left unresolved. Taxon names in parentheses are those used by Kielan-Jaworowska et al. (2004). For further details see Chapter 5.
Page 2 and 3: The Origin and Evolution of Mammals
Page 4 and 5: The Origin and Evolution of Mammals
Page 6 and 7: Preface This book arose from twin a
Page 8 and 9: For Mal /gosia with love and thanks
Page 10 and 11: Contents 1 Introduction The definit
Page 12 and 13: CHAPTER 1 Introduction There are ab
Page 14 and 15: transversely occluding molar teeth
Page 16 and 17: the Mesozoic mammals, is to do with
Page 18 and 19: a hierarchy of committees under the
Page 20 and 21: it means that a 200 Ma igneous rock
Page 24 and 25: Table 2.3 Classification of marsupi
Page 26 and 27: (a) (b) (c) humerus radius aquatic
Page 28 and 29: (a) Westlothiana Limnoscelis PO Wes
Page 30 and 31: the ankle bones to form an astragal
Page 32 and 33: (b) (c) (a) Casea rutena Casea broi
Page 34 and 35: (Fig. 3.2(f)), is actually an ophia
Page 36 and 37: the first one is enlarged, often to
Page 38 and 39: Even at their initial appearance in
Page 40 and 41: (a) Nikkasaurus (b) (d) Reiszia (e)
Page 42 and 43: Nikkasauridae The family Nikkasauri
Page 44 and 45: has figured large in discussions of
Page 46 and 47: (c) Figure 3.9 (continued). Syodon
Page 48 and 49: a mixture of progressively more spe
Page 50 and 51: animal with interdigitating, but no
Page 52 and 53: (e) (a) Anomocephalus Ulemica (b) S
Page 54 and 55: mandibulae musculature. This, as in
Page 56 and 57: To date the postcranial skeleton of
Page 58 and 59: ecognised four subgroups, based mai
Page 60 and 61: the presence of a deep, longitudina
Page 62 and 63: collected from the Lystrosaurus Ass
Page 64 and 65: (a) (d) (c) (b) Leontocephalus V PA
Page 66 and 67: (a) (c) Lycosuchus Oliveria (d) EVO
Page 68 and 69: separate families. Lycosuchidae (Fi
Page 70 and 71: consist of a large labial cusp and
Page 72 and 73:
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
Page 136 and 137:
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
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
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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
Page 194 and 195:
form of the tooth, must reflect sub
Page 196 and 197:
of feasibility that a taxon of endo
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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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that contains the independent ances
Page 208 and 209:
(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
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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
Page 234 and 235:
1. Placentalia 2. Edentata 3. Epith
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effectively absent. However, increa
Page 238 and 239:
Asioryctes (a) (b) (d) Cimolestes p
Page 240 and 241:
and Prokennalestes, although it doe
Page 242 and 243:
(a) Leptictidium Palaeoryctidans we
Page 244 and 245:
(a) Purgatorius (c) are part of the
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
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
Page 274 and 275:
(a) (b) BUNODONTIA or SUIFORMES SUI
Page 276 and 277:
time, which suggests that it was on
Page 278 and 279:
condition. This particular combinat
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etween Primates, Dermoptera (colugo
Page 282 and 283:
have postcranial adaptations for ar
Page 284 and 285:
undoubtedly related at a supraordin
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
Page 320 and 321:
(eds) Evolution of Tertiary mammals
Page 322 and 323:
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
Page 336 and 337:
Index Note: Page numbers in italics
Page 338 and 339:
Equoidea 262 Equus 262 Erethizon 28
Page 340 and 341:
Overkill hypothesis 289, 290 Oxyaen
Page 342:
Tulerpeton 14, 18 Tylopoda 264 Ukha
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The Origin and Evolution of Mammals - Moodle

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