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Major Mammalian Clades: a Review Under Consideration of Molecular and Palaeontological Evidence

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Major Mammalian Clades: a Review Under Consideration of Molecular and Palaeontological Evidence Mamm. biol. 68 (2003) 1±15 Mammalian Biology ã Urban & Fischer Verlag http://www.urbanfischer.de/journals/mammbiol Zeitschrift fuÈr SaÈ ugetierkunde Review Major mammalian clades: a review under consideration of molecular and palaeontological evidence By K. M. HELGEN South Australian Museum, Adelaide, Australia and Department of Environmental Biology, University of Adelaide, Adelaide, Australia Receipt of Ms. 30. 10. 2001 Acceptance of Ms. 05. 06. 2002 Abstract Recent advances in mammalian higher-level molecular phylogenetics have generated support for the recognition of four fundamental superordinal clades of living placental mammals. These super- ordinal hypotheses are used as a framework for exploring patterns and trends that have occurred during mammalian evolution to give rise to the contemporary placental mammal fauna. Potential relationships of recent mammal groups not previously referred to one of these four clades are exam- ined here. Patterns of comparative disparity and diversity, historical and modern biogeography, and morphological and ecological convergence between and within placental superorders are discussed. Key words: Phylogenetics, eutherian, convergence, diversity, disparity Introduction ªThe stream of heredity makes phylogeny; exons) representing every eutherian order, in a sense, it is phylogeny. Complete genetic have generated well-resolved phylogenies analysis would provide the most priceless that suggest extant placental mammals can data for the mapping of this stream . .º be partitioned into four superordinal clades. One of these fundamental superordinal Simpson (1945) groups is Afrotheria, a clade of probable African origin comprising six orders of mostly African and Madagascan mammals: Comprehensive molecular studies of high- Hyracoidea, Proboscidea, Sirenia, Tubuli- er-level mammalian phylogenetics (Mad- dentata, Macroscelidea, and Afrosoricida sen et al. 2001; Scally et al. 2001; Murphy (tenrecs, otter shrews, and golden moles). et al. 2001 a, b) have recently infused fresh The superorder Xenarthra, in turn, is a tra- perspective into long-debated questions re- ditionally recognized assemblage compris- garding relationships and radiations among ing Folivora (sloths, see Delsuc et al. the living placental mammal lineages. These 2001), Vermilingua (vermilinguas), and Cin- studies, the first to employ very large mole- gulata (armadillos) (all of which are recog- cular datasets (dominated by nuclear nized here as full orders) with greatest di- 1616-5047/03/68/01-001 $ 15.00/0. 2 K. M. HELGEN versity and possible origin in South Ameri- Correct higher-level relationships among ca (Delsuc et al. 2001; or possibly Antarcti- placental orders, among marsupial orders ca, see Vizcaino et al. 1998). A third clade (Palma and Spotorno 1999; Phillips et was dubbed Euarchontoglires by Murphy al. 2001; Springer et al. 1998), and be- et al. (2001 b) to recognize that its member- tween placentals, marsupials, and mono- ship consists of the apparently monophy- tremes (Janke et al. 1997, 2002; Killian letic clades Glires (Rodentia and La- et al. 2001; Kirsch and Mayer 1998), are gomorpha) and Euarchonta (Primates, currently the subject of very active mole- Scandentia, and Dermoptera ± i. e. Archon- cular and morphological investigation. Re- ta sensu Gregory 1910, minus bats and ele- searchers employing varied data sets have phant shrews). The last of the four placental arrived at many conflicting hypotheses for superorders was designated Laurasiatheria placental relationships in particular (see by Madsen et al (2001 a) (to denote the ap- Liu et al. 2001). In this light, the potential parent origin of many of its member groups phylogenetic arrangements delineated by on northern continents); it includes the or- Murphy et al. (2001 b) should be consid- ders Lipotyphla (shrews, moles, and hedge- ered working hypotheses that will undergo hogs), Chiroptera, Carnivora, Pholidota, additional refinement (see Novacek 2001). Perissodactyla, and Artiodactyla (including Notably, superordinal relationships sup- cetaceans; n. b. the name Cetartiodactyla, ported by Murphy et al. (2001 b) (and dis- as currently employed by molecular sys- cussed here) strongly contradict the results tematists, should probably be considered a of many morphology-based phylogenetic term of convenience rather than a valid arrangements (e. g. Shoshani and McKen- superordinal name. Cetartiodactyla could na 1998; Fischer and Tassy 1993; Novacek be a valid name if cetaceans were consid- 1992; Novacek and Wyss 1986). However, ered the sister group of all other artiodac- molecular phylogenetic studies can em- tyls. However, cetaceans are properly ploy vastly larger datasets and potentially nested within the order Artiodactyla as tra- more powerful analytical methods, and ditionally defined, which is otherwise para- are thus of fundamental utility in resolving phyletic ± cf. Arnason et al. 2000; Lum et relationships between taxa when morpho- al. 2000; Shimamura et al. 1997 ± thus, the logical methods offer ambiguous results name Artiodactyla can be retained for this (cf. Murphy et al. 2001 b; Shoshani and assemblage, just as the order Carnivora, McKenna 1998). In addition to clarifying paraphyletic without pinnipeds, does not seemingly intractable phylogenetic pro- change its name upon their inclusion or ex- blems, molecular phylogenetics acts in tan- clusion). dem to enrich macroevolutionary studies This eutherian arrangement complements by pointing out anatomical convergences support previously generated from molecular that would be difficult to conclusively and morphological studies for the recognition identify otherwise (such as those used in of two marsupial superorders (Phillips et al. the past to unite paraphyletic superorders 2001; Szalay 1982, 1994; Temple-Smith 1987): such as Volitantia, Anagalida, Archonta, Australidelphia, including four Australasian Altungulata, and Edentata; cf. Shoshani orders as well as the enigmatic South Ameri- and McKenna 1998). This provides insight can marsupial Dromiciops (in the monotypic into the relationship between ecological order Microbiotheria); and Ameridelphia, similarity, homology, and homoplasy comprising the American opossums (order Di- through time. delphimorphia) and shrew-mice (order Pauci- Assuming that there are four fundamental tuberculata). Entertaining these superordinal placental clades provides a fresh framework hypotheses, a total of seven fundamental for exploring origins, comparative success, groupings can be recognized among living historical biogeography, and convergent mammals: four placental superorders, two adaptations of these major groups, and for marsupial superorders, and monotremes. addressing the unresolved relationships of Major mammalian clades 3 several poorly-known placental lineages The recent sundering of insectivore-grade within this framework. Some of these topics mammals to all branches of the placental have been discussed elsewhere (Eizirik et mammal tree, the suggestion that erinaceids al. 2001, Madsen et al. 2001, Murphy et al. closely resemble tribosphenic mammals 2001 a,b; Novacek 2001; Scally et al. 2001) from the Early Cretaceous of Australia but are explored in greater detail here. (Rich et al. 2001 b, 2002), and lingering ar- gument over hedgehog relationships (Cao et al. 2000; Mouchaty et al. 2000; Nikaido Unresolved clade membership et al. 2001) all emphasize the need to fur- ther clarify the phylogenetic position of Several poorly-known placental groups of both gymnures and hedgehogs. uncertain affinity were not included in the One last recent eutherian taxon omitted collective analyses of Murphy et al. from molecular investigations of higher-lev- (2001 b). While these are groups that ap- el placental relationships (due to lack of pear only peripherally in the Holocene, samples) is the endemic Madagascan order their phylogenetic relationships have signif- Bibymalagasia. MacPhee (1994) erected icant bearing on historical biogeography. this order to house Plesiorycteropus,anex- The first of these are the solenodons and tinct genus of medium-sized mammals that nesophonts, insectivore-grade mammals en- survived late into the Holocene, probably demic to the Greater Antilles. Nesophonts until about 1 000 ybp. The relationship be- have apparently become extinct in recent tween Bibymalagasia and other placental centuries and are known only by skeletal mammals is uncertain (MacPhee 1994; material, but solenodons survive as highly McKenna and Bell 1997). However, Ple- endangered species in Cuba and Hispanio- siorycteropus was earlier classified as a tu- la. The relationships of solenodons have bulidentate (Patterson 1975; Thewissen never been very clear (McDowell 1958; 1985), and MacPhee (1994) discussed mor- Van Valen 1967; Whidden and Asher phological resemblances between Plesioryc- 2001). They are probably referable to the teropus, aardvarks, and hyraxes. In light of laurasiatherian order Lipotyphla (in the re- current knowledge about major placental stricted sense, i. e. Eulipotyphla in recent groups, it seems highly possible that Plesior- literature) along with shrews, moles, and ycteropus is part of the radiation of hedgehogs (see Stanhope et al. 1998 b), afrotheres, all of which share a traditionally but further study is needed in the context enigmatic phylogenetic position and a dis- of the arrangement of Murphy et al. tribution restricted to Africa and/or Mada- (2001 b). The true affinities of nesophonts gascar. Interestingly, MacPhee (1994) remain almost completely
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