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41 EUTHERIA METATHERIA Tree of Mammals Chiroptera (bats) 1 1 Perissodactyla (horses, rhinos) Carnivora (lions, wolves) ) 2 Pholidota (pangolins) MYA s) Artiodactyla (deer, pigs) 2 c. 90 Laurasiatheria Cetacea (whales, dolphins) (laurasiatherians 3 Erinaceomorpha (hedgehogs) (boreoeutherian Sericomorpha (moles, shrews) Primates (monkeys, humans) 3 Boreoeutheria 4 Dermoptera (colugos, cobegos) (placentals) Scandentia (tree shrews) MYA Rodentia (rats, guinea pigs) 4 (supraprimates) MYA Euarchontoglires EUTHERIA 5 c. 120 Lagomorpha (rabbits, hares) 90–80 Xenarthra (armadillos, sloths, anteaters) 5 Proboscidea (elephants) 6 6 s) Sirenia (manatees, dugong) MYA 80–70 Hyracoidea (hyraxes) 7 (afrotherian MYA Tubulidentata (aardvark) 7 frotheria 180–170 Afrosoricida (tenrecs, golden moles) A Macroscelidea (elephant shrews) 8 Notoricteromorphia (marsupial moles) Dasyuromorphia (Tasmanian devil, numbat) 8 Peramelemorphia (bandicoots) 41 (marsupials) 9 Diprotodontia (kangaroos, koala) 9 MYA Microbiotheria (colocolo) c. 200 Paucituberculata (shrew opossums) METATHERIA 10 Didelphimorphia (opossums) Monotremata (equidnas, platypus) 10 Tree of Mammals showing the phylogenetic relationships among the most significant groups (28 orders). Colored boxes indicate high taxonomic ranks. Branches with thick lines indicate robust clades, and branches with thin lines less-supported clades. The number in the green circle indicates the chapter in which mammals are also included. Orange circles mark main nodes and their ages. Photographs illustrate principal clades; boxed numbers associate photographs with clades. ©2014 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 43_TOL1e_43.indd 502 5/8/14 3:50 PM Mammals Proliferation of Species after Dinosaurs’ Demise Pablo Vargas 43 SUMMARY Mammals are a natural group composed of three principal clades of extant species: monotremes (oviparous), metatherians (viviparous marsupials), and eutherians (viviparous placentals). From a systematics point of view, extant mammals are subdivided into 28 orders and 134 families, unequally distributed among monotremes (1 order/2 fami- lies), marsupials (7 orders/19 families), and placentals (20 orders/113 families). Placentals are in turn divided into 3 principal clades (afrotherians, xenarthrans, and boreoeutherians), with the relationships among the 3 difficult to resolve. A strong geographical component al- lows for a precise biogeographical reconstruction of the most ancient groups of mammals originated in the Late Triassic and distributed mainly in Australia (monotremes, marsupials), Africa (afrotherians), South America (marsupials, xenarthrans), North America (marsupials, boreoeutherians), and Eurasia (boreoeutherians). The phylogeny of mammals also makes it possible to point out five unique characters (dentary bone, no intermediate bones in the mandibular joint, three bones in the inner ear, development of hair, and production of milk) and a tendency toward giving birth to increasingly underdeveloped young, increasing separation between the digestive and urogenital orifices, and ribs limited to the thoracic region. Although morphological and physiological characters have historically provided the basis for a natural classification reflecting the evolution of mammals, there are other con- vergent characters in certain groups that have not facilitated systematics; notably among them, a derived epitheliochorial placenta, hooves (former “ungulates”), or the development of dentition related to the type of food (former “insectivores”). In this chapter the supraspe- cific systematics of the 5200 extant species is analyzed in the light of morphological and molecular data, and the most important groups of extinct tetrapods in the evolutionary line of mammals are discussed. The still-enigmatic origin of elephants and whales and the evo- lution of Homo sapiens are also addressed in detail in this chapter. U, - enough, the interesting fossil register left mainly by the organisms, in spite of their great organic complexity. bones of mammals makes a well-documented histori- Because of human beings’ interest in learning about cal reconstruction possible, although not without gaps. ourselves, and our intense study of species—either Mammals are thought to have been one of the domi- phylogenetically related (apes, chimpanzees), com- nant groups on Earth since the extinction of the dino- mercially useful (cows, goats, sheep), or medically saurs some 65 million years ago, not only because of the useful (mouse, rat)—we have been able to understand number of species (but see Chapter 34, Coleopterans), the evolution of mammals in detail. As if this were not but also because of their size and the adaptability of What is a mammal? Mammals (from the Latin mamma [teat]) are verte- primitive sinapsids (amniote tetrapods) that appeared brates with constant body temperature, hair, cere- in the Early Permian. Several radiations of mammals bral neocortex, and, in particular, the females that occurred in the last 65 million years, after the develop mammary glands that produce milk with mass extinction of dinosaurs, have been described. which they feed their young. All extant mammals Since then, they have diversified into multiple lin- descend from a common ancestor dating back to eages of species adapted to land, water, and air the Late Triassic, more than 200 million years ago. environments. Specifically, present-day mammals descend from ©2014 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured or disseminated in any form without express written permission from the publisher. 43_TOL1e_43.indd 503 5/8/14 3:50 PM 504 CHAPTER 43 • MAMMALS than extant species (Figure 43.1). Because the bones BOX 43.1 Morphological characters are so easily fossilized, there is a fairly ample mammal unique to mammals fossil record available. The five species of monotremes • Mandible consists of only the dentary bone. form a whole order of mammals, and are considered living fossils because they combine characteristics com- The dentary bone joins directly to the squamous • mon to the other mammals (hair, production of milk, bone of the cranium, making the mandibular joint. three bones in the ear, diaphragm, and heart with four • There are three bones in the middle ear (hammer, chambers) and to the sauropsids (laying eggs, feet with anvil, and stirrup), except in the monotremes spurs on hind legs of males, cranium tapering toward (reptilian ear). the beak-shaped snout, corneous beak). One platypus • All species develop hair to a greater or lesser degree. and four echidnas make up this ancient lineage, which • Mammary glands (derived from sebaceous glands) has survived in Australia and nearby islands in the face secrete milk, the necessary food for mammal young. of vigorous competition from placentals. Two hundred of the 275 species of marsupials are also Australian, which clearly shows that the island continent of Aus- many groups to nearly all the ecosystems on the planet. tralia has facilitated an independent and parallel evolu- Moreover, many mammal species are at the top of the tion of mammals with characteristics similar to those of trophic pyramid, making their ecological role one of su- placentals (in Australia there are few native placentals). preme interest. However, they had to wait for their evo- Standing out among the most notable paleontological lutionary moment, some 60 million years ago, for their findings is a marsupial (Sinodelphys) in a surprising impressive differentiation, even though the lineage that state of preservation, even with detectable hairs. Con- gave rise to them, starting with the therapsids, had comitant in space (China) and time (approximately appeared long before that. In fact, the extinction of the 125 Mya) with fossils, marsupials is the most ancient dinosaurs unleashed several adaptive radiations that placental found (Eomaia). Due to the wide diversity of launched a number of lineages and species of mammals placentals (some 5500 species) developed in the last 125 that colonized so many ecological environments. As with all living things, the number of extinct lin- eages of mammals is much greater than that of those BOX 43.2 Mammals by the numbers extant. Therefore, their evolutionary tree requires a combined knowledge of the fossil record and of ex- • Number of species: some 5500 placentals, 5 tant species. The first division of mammals in the strict monotremes, and around 275 marsupials. 70% sense separates the prototherians (eotherians, allote- of mammal species are rodents rians, and monotremes) and the therians (tritubercu- • Number of genera: about 1200 lata, pantotherians, metatherians or marsupials, and • Genera with the greatest number of species: eutherians or placentals), both with extant and extinct Crocidura (about 90 species), Myotis (about 90 groups. The term mammaliaformes is used to include a species), Rhinolophus (about 65 species) clade of mammals and related extinct forms. Mamma- • Number of families: 134 (although it varies liaforms derived from the therapsids (an extinct group, depending on the author) with the exception of the mammaliaforms), which be- • Number of orders: 28 (although it varies substantially long to the synapsids (sister group to the sauropsids; depending on the author) see Chapter 44, Sauropsids). Finally, the synapsids • Orders with the greatest number of species: are considered “mammal-like reptiles” because they Rodentia (rodents)
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