The Origin and Evolution of Mammals - Moodle

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1. Placentalia 2. Edentata 3. Epitheria 4. Archonta 5. Glires 6. Paenungulata 7. Tethytheria 1 3 2 4 5 6 Monotremata Marsupialia Xenarthra Pholidota Carnivora Tubulidentata Lipotyphla Primates Scandentia Dermoptera Chiroptera Macroscelidea Lagomorpha Rodentia Artiodactyla Perissodactyla Cetacea Hyracoidea Sirenia 7 Proboscidea (a) (b) 100 100 99 76 100 100 A 100 100 100 94 B 98 59 100 100 100 100 * 100 100 100 100 100 100 * 74 42 100 * 100 100 100 100 100 77 100 79 100 * 100 91 100 75 100 * 100 100 100 86 100 * 99 70 100 99 100 * 100 * 100 100 100 * 100 * 100 * 100 85 100 * 100 100 100 81 100 * 100 100 76 100 40 * whale dolphin hippo Cetartiodactyla ruminant pig llama rhino tapir Perissodactyla horse cat Carnivora caniform pangolin Pholidota flying fox rousette fruit bat rhinolophoid bat Chiroptera phyllostomid microbat free tailed bat hedgehog shrew Eulipotyphla mole mouse rat Rodentia hystricid caviomorph sciurid rabbit Lagomorpha pika flying lemur Dermoptera tree shrew Scandentia strepsirrhine Primates human sloth Pilosa anteater armadillo Cingulata tenrec Afrosoricida golden mole s.e. elephant shrew Macroscelidea l.e. elephant shrew aardvark Tubulidentata sirenian Sirenia hyrax Hyracoidea elephant Proboscidea opossum diprotodontian Laurasiatheria Euarchontoglires Xenarthra Afrotheria Boreoeutheria Marsupialia Figure 7.1 Phylogeny of the living orders of placental mammals. (a) Novacek et al.’s (1988) morphological-based cladogram. (b) Murphy et al.’s (2001b) molecular-based cladogram. Numbers above the branches are percentage Bayesian posterior probabilities; numbers below the branches are maximum likelihood bootstrap support values.
224 THE ORIGIN AND EVOLUTION OF MAMMALS middle ear. Alternatively, the basal members of related orders may have shared significant derived characters, but these have subsequently been lost or transformed out of recognition as the individual orders evolved into their modern representatives. Further purely morphological analysis of living mammals is unlikely to lead to a recognition of which of these two phylogenetic patterns is true, let alone to resolve the interrelationships. Two centuries of study is surely approaching saturation. In principle the fossil record can improve the taxonomic resolution by providing character combinations not found amongst the living groups, and has indeed contributed much to the debates on supraordinal relationships, as discussed later. Unfortunately, palaeontological evidence will always be hampered by the bugbear of missing data: there are too many cases of incongruence amongst dental and skeletal characters for fossils alone to be regarded as reliable trackers of phylogeny. As with the marsupials discussed in Chapter 6, analysis of molecular sequence data is proving the likeliest source of information for solving the riddle of placental interrelationships, and some remarkable but by no means fanciful hypotheses have now emerged, almost to the extent of providing a tree that is fully resolved at the ordinal level. In one of the first contributions, de Jong et al. (1993) analysed the eye lens protein �A-crystallin and found that the macroscelidean Elephantulus rufescens had three amino acid replacements otherwise unique to the paenungulates (elephants, sirenians, and hyraxes) and the aardvark, suggesting a monophyletic group for these very different orders, whose sole nonmolecular common feature is that their history is mainly African. Within a few years, this relationship had become strongly supported by DNA sequences, both mitochondrial and nuclear, and even extended to include the Tenrecida (Springer et al. 1997b; Stanhope et al. 1998). These are the tenrecs of Madagascar and the related otter shrews of mainland southern Africa, which together had been regarded as a subgroup of lipotyphlan insectivores. One final taxon of African mammals has been added to what became named the Afrotheria. Chrysochlorids are the golden moles of southern Africa and, like the tenrecs, they too had been firmly believed to be lipotyphlans. The study of Stanhope et al. (1998) placed it firmly in the afrotherian clade. The Afrotheria concept goes a long way towards understanding the previously poorly understood relationships of the Tubulidentata and Macroscelidea. However, it contradicts all morphological based classifications of Lipotyphla, by removing the tenrecs and golden moles from their position nested deeply within the group, and related to the shrews and moles (Butler 1988). Meanwhile, another radical rearrangement was emerging. The relationships of the cetaceans had been the subject of a long dispute, even to the extent of whether they had evolved from a carnivorous, or an herbivorous ancestor. On the basis of the fossil evidence, the view prevailed that they had evolved from a secondarily carnivorous group of primitive, ‘condylarth’ ungulates known as mesonychids. This consensus was actually disturbed long ago, when, in a very early essay into molecular systematics, Boyden and Gemeroy’s (1950) immunological method suggested a relationship between whales and artiodactyls. Little notice was taken at the time. The recent era was marked by Graur and Higgins (1994), who analysed the molecular evidence then available, mainly protein sequences, and also concluded that cetaceans were most closely related to the Artiodactyla. Furthermore, their study pointed to the egregious conclusion that whales nested within the artiodactyls, as the sister-group of the hippos. Since then, a very large quantity of DNA sequence data bearing on the question has been analysed, with results overwhelming confirming the hippo-whale clade (Waddell et al. 1999; Gatesy and O’Leary 2001). At first, morphological evidence continued to fail to support even the overall group Cetartiodactyla, as it had come to be named (O’Leary 1999), although the similarities that do exist between whales and hippos in particular, such as the reduction of hair, development of subcutaneous fat, and ability to suckle the young underwater, were not lost on the proponents of the new grouping. Far from being the convergences hitherto supposed, they can be taken as evidence of a common ancestor between the two that was already adapted to a semiaquatic existence. Modern whales are so extremely derived that the key characters that would allow the recognition of their relationships, notably limb structure and dental structure, are
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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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a self-sharpening property. As the
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near parasagittal gait (Fig. 5.11(e
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pass through the birth canal. Relat
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(a) 1 (b) (d) me (c) me.d 3 styl st
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(a) (c) Henkelotherium Crusafontia
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pubis is excluded from the acetabul
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Cretaceous, (Aptian or Albian) of M
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eautifully preserved placentals fro
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Page 214 and 215: (d) Figure 6.6 (continued). Thylaco
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Page 218 and 219: (a) (e) (d) Proargyrolagus Groeberi
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Page 222 and 223: LIVING AND FOSSIL MARSUPIALS 211 M
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Page 256 and 257: their presence. The five orders may
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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
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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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