The Origin and Evolution of Mammals - Moodle

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elieved to be Late Cretaceous in age, but this is disputed and it is more probably Early Palaeocene (Marshall et al. 1997). In any event, the fossils themselves are sparse and fragmentary, there being a possible tooth of the didelphid Peradectes and a possible member of the Pediomyidae (Sigé 1972). Taken at face value, these identifications are very significant because both taxa otherwise only occur in the latest Cretaceous, Maastrichtian, of North America. This suggests that a faunal interchange occurred, presumably from north to south, at a time very close to the end-Cretaceous, although the temporal resolution is not good enough to say whether just before, at, or just after that event. A richer and younger site in the same formation as Laguna Umayo has been described at Chulpas, and here there are reported to be as many as eight marsupial taxa, including a polydolopoid (Sillustania), a primitive caroloameghiniid (Chulpasia), caenolestids, and didelphids. All but the last of these groups are only known from South America (Crochet and Sigé 1996; Muizon and Cifelli 2001). The Chulpas fauna is tentatively dated close to the K/T boundary and presumably represents the beginning of the indigenous radiation that followed the initial immigration of marsupials from North America. Far the richest Palaeocene localities on the continent occur in the Santa Lucia Formation at Tiu Pampa in Bolivia, which is either Early (Muizon 1998) or possibly early Late (Marshall et al. 1997) Palaeocene in age, and at Itaboraí in Brazil, where the fissure-fill fauna is slightly younger. Both sites have yielded abundant and diverse marsupial faunas, including six or seven indigenous South American families between them. There are also representatives of the modern marsupial family Didelphidae, and the genus Khasia which is a member of the biogeographically challenging Microbiotheria to which the modern Dromiciops belongs. Other, less-rich Palaeocene mammal localities occur in the San Jorge Basin of Patagonia, some of which are Early Palaeocene (Marshall et al. 1997). There is inevitably a great deal of disagreement about the detailed phylogenetic interrelationships and thus taxonomy of ameridelphians, particularly as many of them are still represented only by teeth and jaws. However, the majority of authors accept three lineages (Aplin and Archer LIVING AND FOSSIL MARSUPIALS 199 1987; Marshall et al. 1990; McKenna and Bell 1997). Two of these have living members and have already been noted, although they both have a much wider range of fossil forms. Very generally speaking, the didelphimorphs retain primitive, sharp-cusped tribosphenic teeth; the sparassodonts have carnivorous, carnassial teeth; and the paucituberculates have broader, more bunodont teeth implying an omnivorous, somewhat rodent-like habit. Didelphimorphia Didelphimorphs consist of small- to medium-sized opossums. They are generally primitive in nature, mostly retaining the full marsupial dental formula, and possessing simple incisors and basic tribosphenic molars. The most controversial character is the dilambdodont condition, which is to say the V-shaped centrocrista, the crest on the upper molars that connects the metacone to the paracone (Fig. 6.1(b)). This has been taken by some authors to indicate that the didelphimorphs are a monophyletic group (Aplin and Archer 1987; Muizon and Cifelli 2001). However, a very similar dilambdodont condition is also found in primitive Australian dasyuromorphs and certain other primitive marsupials (Godthelp et al. 1999). As discussed later, this suggests the alternative view that dilambdodonty is primitive for both the ameridelphian and the australidelphian marsupials, and indeed that the didelphimorphs may be ancestral to both and therefore paraphyletic. There are actually certain possibly unique characters of the didelphimorph ankle joint (Szalay 1994), although this evidence is very limited because of the few adequately preserved postcranial skeletons. The Tiupampan form Szalinia (Fig. 6.5(a)) is the most primitive South American didelphimorph because its molar teeth are intermediate in structure between the basal North American marsupial Kokopellia, and more typical didelphimorphs from Tiu Pampa such as Andinodelphys and Pucadelphys (Muizon and Cifelli 2001). For instance, the centrocrista of Szalinia is only incipiently V-shaped. Pucadelphys is in fact much the best-known Tiupampan marsupial; several well-preserved skulls and skeletons have been found (Marshall et al. 1995). It is a typical didelphid in size and general appearance, with a skull length about 3 cm
200 THE ORIGIN AND EVOLUTION OF MAMMALS (a) (b) Szalinia (d) (c) Sparassocynus (e) Pucadelphys Figure 6.5 Didelphimorphia. (a) Upper molars of Szalinia gracilis in occlusal view (redrawn after Muizon and Cifelli 2001). (b) Upper second molar of Pucadelphys andinus in occlusal view (Marshall et al. 1995). (c) Skull of Pucadelphys andinus in dorsal, ventral, and lateral views. Skull length approx. 3 cm (Marshall et al. 1995). (d) Skull of Sparassocynus derivatus in dorsal and lateral views (Reig et al. 1987). (e) Sparassocynus sp. upper dentition in labial and occlusal views, and lower dentition in labial and occlusal views (Reig et al. 1987).
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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
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 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 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
Page 232 and 233: the Diprotodontia also included gen
Page 234 and 235: 1. Placentalia 2. Edentata 3. Epith
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
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
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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
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Simpson, G.G. 1970. The Argyrolagid
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Thewissen, J.G.M. 1990. Evolution o
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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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