Mammal

For other uses, see Mammal (disambiguation).
"Mammalian" redirects here. For the 2010 documentary film, see Mammalian (film).
Mammal
Temporal range: 225–0 Ma (Kemp) or 167–0 Ma (Rowe) See discussion of dates in text
Common vampire bat Tasmanian devil Fox squirrel Platypus Humpback whale Armadillo Virginia opossum Human Tree pangolin Colugo Star nosed mole Plains zebra Eastern grey kangaroo Northern elephant seal African elephant Reindeer Giant panda Black and rufous elephant shrewMammal Diversity 2011.png
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Amniota
Clade: Synapsida
Clade: Mammaliaformes
Class: Mammalia
Linnaeus, 1758
Subgroups

Mammals (class Mammalia /məˈmeɪli.ə/ from Latin mamma "breast") are any members of a clade of endothermic amniotes distinguished from reptiles and birds by the possession of a neocortex (a region of the brain), hair,[lower-alpha 1] three middle ear bones, and mammary glands. The mammalian brain regulates body temperature and the circulatory system, including the four-chambered heart.

Mammals include the largest animals on the planet, the rorquals and other large whales, as well as some of the most intelligent, such as elephants, primates, including humans, and cetaceans. The basic body type is a four-legged land-borne animal, but some mammals are adapted for life at sea, in the air, in trees, or on two legs. The largest group of mammals, the placentals, have a placenta, which enables feeding the fetus during gestation. Mammals range in size from the 30–40 mm (1.2–1.6 in) bumblebee bat to the 33-meter (108 ft) blue whale.

The word "mammal" is modern, from the scientific name Mammalia coined by Carl Linnaeus in 1758, derived from the Latin mamma ("teat, pap"). All female mammals nurse their young with milk, which is secreted from special glands, the mammary glands. According to Mammal Species of the World, 5,416 species were known in 2006. These were grouped in 1,229 genera, 153 families and 29 orders.[1] In 2008 the IUCN completed a five-year, 1,700-scientist Global Mammal Assessment for its IUCN Red List, which counted 5,488 accepted species.[2]

In some classifications, extant mammals are divided into two subclasses: the Prototheria, that is, the order Monotremata; and the Theria, or the infraclasses Metatheria and Eutheria. The marsupials constitute the crown group of the Metatheria, and include all living metatherians as well as many extinct ones; the placentals are the crown group of the Eutheria.

Except for the five species of monotremes (egg-laying mammals), all modern mammals give birth to live young. Most mammals, including the six most species-rich orders, belong to the placental group. The three largest orders in numbers, are first Rodentia: mice, rats, porcupines, beavers, capybaras, and other gnawing mammals; then Chiroptera: bats; and then Soricomorpha: shrews, moles and solenodons. The next three orders, depending on the biological classification scheme used, are the Primates including the humans; the Cetartiodactyla including the whales and the even-toed hoofed mammals; and the Carnivora, that is, cats, dogs, weasels, bears, seals, and their relatives.[1]

While mammal classification at the 'family' level has been relatively stable, several contending classifications regarding the higher levels—subclass, infraclass, and order, especially of the marsupials—appear in contemporaneous literature. Much of the recent change reflects the advances of cladistic analysis and molecular genetics. Findings from molecular genetics, for example, have prompted adopting new groups, such as the Afrotheria, and abandoning traditional groups, such as the Insectivora.

The early synapsid mammalian ancestors were sphenacodont pelycosaurs, a group that produced the non-mammalian Dimetrodon. At the end of the Carboniferous period, this group diverged from the sauropsid line that led to today's reptiles and birds. The line following the stem group Sphenacodontia split-off several diverse groups of non-mammalian synapsids—sometimes referred to as mammal-like reptiles—before giving rise to the proto-mammals (Therapsida) in the early Mesozoic era. The modern mammalian orders arose in the Paleogene and Neogene periods of the Cenozoic era, after the extinction of the non-avian dinosaurs, and have been among the dominant terrestrial animal groups from 66 million years ago to the present.

Varying definitions, varying dates

In an influential 1988 paper, Timothy Rowe defined Mammalia phylogenetically as the crown group mammals, the clade consisting of the most recent common ancestor of living monotremes (echidnas and platypuses) and therian mammals (marsupials and placentals) and all descendants of that ancestor.[3] Since this ancestor lived in the Jurassic period, Rowe's definition excludes all animals from the earlier Triassic, despite the fact that Triassic fossils in the Haramiyida have been referred to the Mammalia since the mid-19th century.[4]

T. S. Kemp has provided a more traditional definition: "synapsids that possess a dentary–squamosal jaw articulation and occlusion between upper and lower molars with a transverse component to the movement" or, equivalently in Kemp's view, the clade originating with the last common ancestor of Sinoconodon and living mammals.[5]

If Mammalia is considered as the crown group, its origin can be roughly dated as the first known appearance of animals more closely related to some extant mammals than to others. Ambondro is more closely related to monotremes than to therian mammals while Amphilestes and Amphitherium are more closely related to the therians; as fossils of all three genera are dated about 167 million years ago in the Middle Jurassic, this is a reasonable estimate for the appearance of the crown group.[6] The earliest known synapsid satisfying Kemp's definitions is Tikitherium, dated 225 Ma, so the appearance of mammals in this broader sense can be given this Late Triassic date.[7][8] In any case, the temporal range of the group extends to the present day.

Distinguishing features

Living mammal species can be identified by the presence of sweat glands, including those that are specialized to produce milk to nourish their young. In classifying fossils, however, other features must be used, since soft tissue glands and many other features are not visible in fossils.

Many traits shared by all living mammals appeared among the earliest members of the group:

For the most part, these characteristics were not present in the Triassic ancestors of the mammals.

For palaeontologists who define Mammalia phylogenetically, no limit can be set on the features used to distinguish the group. Any feature may be relevant to a fossil's phylogenetic position. Palaeontologists defining Mammalia in terms of traits, on the other hand, need only consider those features that appear in the definition. The dentary-squamosal jaw joint is generally included.

Nearly all mammal groups possess an epipubic bone, the exception being modern placentals; while as a distinguishing feature it serves little purpose given the diversity of living placentals over other mammal groups, it is a useful tool among paleontologists.

Classification

Main article: Mammal classification
The orders Rodentia (blue), Chiroptera (red), and Soricomorpha (yellow) together comprise over 70% of mammal species.

George Gaylord Simpson's "Principles of Classification and a Classification of Mammals" (AMNH Bulletin v. 85, 1945) was the original source for the taxonomy listed here. Simpson laid out a systematics of mammal origins and relationships that was universally taught until the end of the 20th century. Since Simpson's classification, the paleontological record has been recalibrated, and the intervening years have seen much debate and progress concerning the theoretical underpinnings of systematization itself, partly through the new concept of cladistics. Though field work gradually made Simpson's classification outdated, it remained the closest thing to an official classification of mammals.

McKenna/Bell classification

In 1997, the mammals were comprehensively revised by Malcolm C. McKenna and Susan K. Bell, which has resulted in the McKenna/Bell classification. Their 1997 book, Classification of Mammals above the Species Level,[10] is the most comprehensive work to date on the systematics, relationships, and occurrences of all mammal taxa, living and extinct, down through the rank of genus, though recent molecular genetic data challenge several of the higher level groupings. The authors worked together as paleontologists at the American Museum of Natural History, New York. McKenna inherited the project from Simpson and, with Bell, constructed a completely updated hierarchical system, covering living and extinct taxa that reflects the historical genealogy of Mammalia.

The McKenna/Bell hierarchical listing of many terms used for mammal groups above the species includes extinct mammals, as well as modern groups, and introduces some fine distinctions, such as legions and sublegions (ranks that fall between classes and orders) that are likely to be glossed over by non-professionals.

Extinct groups are represented by a dagger (†).

Class Mammalia

Molecular classification of placentals

Molecular studies based on DNA analysis have suggested new relationships among mammal families over the last few years. Most of these findings have been independently validated by retrotransposon presence/absence data.[13] Classification systems based on molecular studies reveal three major groups or lineages of placental mammals- Afrotheria, Xenarthra, and Boreoeutheria- which diverged from early common ancestors in the Cretaceous. The relationships between these three lineages is contentious, and all three possible different hypotheses have been proposed with respect to which group is basal with respect to other placentals. These hypotheses are Atlantogenata (basal Boreoeutheria), Epitheria (basal Xenarthra), and Exafroplacentalia (basal Afrotheria).[14] Boreoeutheria in turn contains two major lineages- Euarchontoglires and Laurasiatheria.

Estimates for the divergence times between these three placental groups range from 105 to 120 million years ago, depending on type of DNA (e.g. nuclear or mitochondrial)[15] and varying interpretations of paleogeographic data.[14]

Mammalia

Monotremata


Theria

Marsupialia


Placentalia
Atlantogenata

Afrotheria

Xenarthra



Boreoeutheria
Euarchontoglires

Euarchonta

Glires



Laurasiatheria

Eulipotyphla


Scrotifera

Chiroptera

Cetartiodactyla

Perissodactyla


Ferae

Pholidota

Carnivora











Cladogram based on Tarver et al. (2016)[16]

Group I: Afrotheria

Group II: Xenarthra

Group III: Boreoeutheria

Evolutionary history

For more details on this topic, see Evolution of mammals.

Synapsida, the group which contains mammals and their extinct relatives, originated during the Pennsylvanian subperiod, when they split from the lineage that led to reptiles and birds. Crown group mammals evolved from earlier mammaliaforms during the Early Jurassic.

Cladogram following,[17] which takes Mammalia to be the crown group.

Mammaliaformes

Morganucodontidae

Docodonta

Haldanodon


Mammalia

Australosphenida (incl. Monotremata)

Fruitafossor


Haramiyavia

Multituberculata

Tinodon

Eutriconodonta (incl. Gobiconodonta)

Trechnotheria (incl. Theria)








A cladogram compiled by Mikko Haaramo and based on individual cladograms of After Rowe 1988; Luo, Crompton & Sun 2001; Luo, Cifelli & Kielan-Jaworowska 2001, Luo, Kielan-Jaworowska & Cifelli 2002, Kielan-Jaworowska, Cifelli & Luo 2004, and Luo & Wible 2005.[18]

Evolution from amniotes in the Paleozoic

The original synapsid skull structure contains one temporal opening behind the orbitals, in a fairly low position on the skull (lower right in this image). This opening might have assisted in containing the jaw muscles of these organisms which could have increased their biting strength.

The first fully terrestrial vertebrates were amniotes. Like their amphibious tetrapod predecessors, they have lungs and limbs. Amniotes' eggs, however, have internal membranes that allow the developing embryo to breathe but keep water in. Hence, amniotes can lay eggs on dry land, while amphibians generally need to lay their eggs in water.

The first amniotes apparently arose in the Late Carboniferous. They descended from earlier reptiliomorph amphibious tetrapods,[19] which lived on land that was already inhabited by insects and other invertebrates as well as by ferns, mosses and other plants. Within a few million years, two important amniote lineages became distinct: the synapsids, which would later include the common ancestor of the mammals; and the sauropsids, which would eventually come to include turtles, lizards, snakes, crocodilians, dinosaurs and birds.[20] Synapsids have a single hole (temporal fenestra) low on each side of the skull.

One synapsid group, the pelycosaurs, included the largest and fiercest animals of the early Permian.[21]

Therapsids descended from pelycosaurs in the Middle Permian, about 265 million years ago, and became the dominant land vertebrates.[22] They differ from basal eupelycosaurs in several features of the skull and jaws, including: larger temporal fenestrae and incisors which are equal in size.[23] The therapsid lineage leading to mammals went through a series of stages, beginning with animals that were very like their pelycosaur ancestors and ending with probainognathian cynodonts, some of which could easily be mistaken for mammals. Those stages were characterized by:

Nonmammalian synapsids are sometimes called "mammal-like reptiles".[22][25]

The mammals appear

The Permian–Triassic extinction event, which was a prolonged event due to the accumulation of several extinction pulses, ended the dominance of the carnivores among the therapsids. In the early Triassic, all the medium to large land carnivore niches were taken over by archosaurs which, over an extended period of time (35 million years), came to include the crocodylomorphs, the pterosaurs, and the dinosaurs. By the Jurassic, the dinosaurs had come to dominate the large terrestrial herbivore niches as well.

The first mammals (in Kemp's sense) appeared in the Late Triassic epoch (about 225 million years ago), 40 million years after the first therapsids. They expanded out of their nocturnal insectivore niche from the mid-Jurassic onwards; Castorocauda, for example, had adaptations for swimming, digging and catching fish.[26] Most, if not all, are thought to have remained nocturnal (the Nocturnal bottleneck), accounting for much of the typical mammalian traits.[27]

The majority of the mammal species that existed in the Mesozoic Era were multituberculates, eutriconodonts and spalacotheriids.[28]

The earliest known monotreme is Teinolophos, which lived about 123 million years ago in Australia. Monotremes have some features which may be inherited from the original amniotes:

Unlike other mammals, female monotremes do not have nipples and feed their young by "sweating" milk from patches on their bellies.

The earliest known metatherian is Sinodelphys, found in 125 million-year-old Early Cretaceous shale in China's northeastern Liaoning Province. The fossil is nearly complete and includes tufts of fur and imprints of soft tissues.[29]

The oldest known fossil among the Eutheria ("true beasts") is the small shrewlike Juramaia sinensis, or "Jurassic mother from China", dated to 160 million years ago in the Late Jurassic.[30] A later eutherian, Eomaia, dated to 125 million years ago in the Early Cretaceous, possessed some features in common with the marsupials but not with the placentals, evidence that these features were present in the last common ancestor of the two groups but were later lost in the placental lineage.[31] In particular:

Rise to dominance in the Cenozoic

Mammals took over the medium- to large-sized ecological niches in the Cenozoic, after the Cretaceous–Paleogene extinction event emptied ecological space once filled by non-avian dinosaurs and groups of reptiles that were now absent.[33] Then mammals diversified very quickly; both birds and mammals show an exponential rise in diversity.[33] For example, the earliest known bat dates from about 50 million years ago, only 16 million years after the extinction of the dinosaurs.[34]

Recent molecular phylogenetic studies suggest that most placental orders diverged about 100 to 85 million years ago and that modern families appeared in the period from the late Eocene through the Miocene.[35] But paleontologists object that no placental fossils have been found from before the end of the Cretaceous.[36] The earliest undisputed fossils of placentals come from the early Paleocene, after the extinction of the dinosaurs.[36] In particular, scientists have recently identified an early Paleocene animal named Protungulatum donnae as one of the first placental mammals.[37][38] The earliest known ancestor of primates is Archicebus achilles[39][40] from around 55 million years ago.[39][40] This tiny primate weighed 20–30 grams (0.7–1.1 ounce) and could fit within a human palm.[39][40]

During the Cenozoic, several groups of mammals appeared that were much larger than their nearest modern equivalents, but none was even close to the size of the largest dinosaurs with similar feeding habits.

Earliest appearances of features

Hadrocodium, whose fossils date from approximately 195 million years ago, in the Early Jurassic, provides the first clear evidence of a jaw joint formed solely by the squamosal and dentary bones; there is no space in the jaw for the articular, a bone involved in the jaws of all early synapsids.

It has been suggested that the original function of lactation (milk production) was to keep eggs moist. Much of the argument is based on monotremes, the egg-laying mammals.[41][42][43]

The earliest clear evidence of hair or fur is in fossils of Castorocauda, from 164 million years ago in the Middle Jurassic. In the 1950s, it was suggested that the foramina (passages) in the maxillae and premaxillae (bones in the front of the upper jaw) of cynodonts were channels which supplied blood vessels and nerves to vibrissae (whiskers) and so were evidence of hair or fur;[44][45] it was soon pointed out, however, that foramina do not necessarily show that an animal had vibrissae, as the modern lizard Tupinambis has foramina that are almost identical to those found in the nonmammalian cynodont Thrinaxodon.[25][46] Popular sources, nevertheless, continue to attribute whiskers to Thrinaxodon.[47]

The evolution of erect limbs in mammals is incomplete — living and fossil monotremes have sprawling limbs. The parasagittal (nonsprawling) limb posture appeared sometime in the Early Cretaceous or latest Jurassic; it is found in the eutherian Eomaia and the metatherian Sinodelphys, both dated 125 million years ago.[48]

When endothermy first appeared in the evolution of mammals is uncertain. Modern monotremes have lower body temperatures and more variable metabolic rates than marsupials and placentals,[49] but there is evidence that some of their ancestors, perhaps including ancestors of the therians, may have had body temperatures like those of modern therians.[50] Some of the evidence found so far suggests that Triassic cynodonts had fairly high metabolic rates, but it is not conclusive. For small animals, an insulative covering like fur is necessary for the maintenance of a high and stable body temperature.

Epipubic bones, a feature that strongly influenced the reproduction of most mammal clades, are first found in Tritylodontidae, suggesting that it is a synapomorphy between them and mammaliformes. They are omnipresent in non-placental mammaliformes, though Megazostrodon and Erythrotherium appear to have lacked them.[51]

Anatomy and morphology

Skeletal system

The majority of mammals have seven cervical vertebrae (bones in the neck), including bats, giraffes, whales, and humans. The exceptions are the manatee and the two-toed sloth, which have just six, and the three-toed sloth with nine cervical vertebrae.[52]

Respiratory system

The lungs of mammals have a spongy texture and are honeycombed with epithelium having a much larger surface area in total than the outer surface area of the lung itself. The lungs of humans are typical of this type of lung.

Breathing is largely driven by the muscular diaphragm, which divides the thorax from the abdominal cavity, forming a dome with its convexity towards the thorax. Contraction of the diaphragm flattens the dome, increasing the volume of the cavity in which the lung is enclosed. Air enters through the oral and nasal cavities; it flows through the larynx, trachea and bronchi and expands the alveoli. Relaxation of the diaphragm has the opposite effect, passively recoiling during normal breathing. During exercise, the abdominal wall contracts, increasing visceral pressure on the diaphragm, thus forcing the air out more quickly and forcefully. The rib cage itself also is able to expand and contract the thoracic cavity to some degree, through the action of other respiratory and accessory respiratory muscles. As a result, air is sucked into or expelled out of the lungs, always moving down its pressure gradient. This type of lung is known as a bellows lung as it resembles a blacksmith's bellows. Mammals take oxygen into their lungs, and discard carbon dioxide.

Nervous system

All mammalian brains possess a neocortex, a brain region unique to mammals. Placental mammals have a corpus callosum, unlike monotremes and marsupials. The size and number of cortical areas (Brodmann's areas) is least in monotremes (about 8-10) and most in placentals (up to 50).

Integumentary system

The integumentary system is made up of three layers: the outermost epidermis, the dermis, and the hypodermis.

The epidermis is typically 10 to 30 cells thick; its main function is to provide a waterproof layer. Its outermost cells are constantly lost; its bottommost cells are constantly dividing and pushing upward. The middle layer, the dermis, is 15 to 40 times thicker than the epidermis. The dermis is made up of many components, such as bony structures and blood vessels. The hypodermis is made up of adipose tissue. Its job is to store lipids, and to provide cushioning and insulation. The thickness of this layer varies widely from species to species.

Although other animals have features such as whiskers, feathers, setae, or cilia that superficially resemble it, no animals other than mammals have hair. It is a definitive characteristic of the class. Though some mammals have very little, careful examination reveals the characteristic, often in obscure parts of their bodies.

Color variation in mammals

Mammalian hair, also known as pelage, can vary in color between populations, organisms within a population, and even on the individual organism. Light-dark color variation is common in the mammalian taxa. Sometimes, this color variation is determined by age variation, however, in other cases, it is determined by other factors. Selective pressures, such as ecological interactions with other populations or environmental conditions, often lead to the variation in mammalian coloration.[53] These selective pressures favor certain colors in order to increase survival. Camouflage is thought to be a major selection pressure shaping coloration in mammals, although there is also evidence that sexual selection, communication, and physiological processes may influence the evolution of coloration as well.[54] Camouflage is the most predominant mechanism for color variation, as it aids in the concealment of the organisms from predators or from their prey. Coat color can also be for intraspecies communication such as warning members of their species about predators, indicating health for reproductive purposes, communicating between mother and young, and intimidating predators.[54] Studies have shown that in some cases, differences in female and male coat color could indicate information nutrition and hormone levels, which are important in the mate selection process.[53] One final mechanism for coat color variation is physiological response purposes, such as temperature regulation in tropical or arctic environments.[54] Although much has been observed about color variation, much of the genetic that link coat color to genes is still unknown. The genetic sites where pigmentation genes are found are known to affect phenotype by: 1) altering the spatial distribution of pigmentation of the hairs, and 2) altering the density and distribution of the hairs. Quantitative trait mapping is being used to better understand the distribution of loci responsible for pigmentation variation.[55] However, although the genetic sites are known, there is still much to learn about how these genes are expressed.[53]

Some primates and marsupials have shades of violet, green, or blue skin on parts of their bodies.[56] The two-toed sloth and the polar bear sometimes appear to have green fur, but this color is caused by algal growths.

Reproductive system

Goat kids will stay with their mother until they are weaned.

Most mammals are viviparous, giving birth to live young. However, the five species of monotreme, the platypuses and the echidnas, lay eggs. The monotremes have a sex determination system different from that of most other mammals.[57] In particular, the sex chromosomes of a platypus are more like those of a chicken than those of a therian mammal.[58] Like marsupials and most other mammals, monotreme young are larval and fetus-like, as the presence of epipubic bones prevents the expansion of the torso, forcing them to produce small young.

The mammary glands of mammals are specialized to produce milk, a liquid used by newborns as their primary source of nutrition. The monotremes branched early from other mammals and do not have the nipples seen in most mammals, but they do have mammary glands. The young lick the milk from a mammary patch on the mother's belly.

Viviparous mammals are in the subclass Theria; those living today are in the marsupial and placental infraclasses. A marsupial has a short gestation period, typically shorter than its estrous cycle, and gives birth to an undeveloped newborn that then undergoes further development; in many species, this takes place within a pouch-like sac, the marsupium, located in the front of the mother's abdomen. This is the plesyomorphic condition among viviparous mammals; the presence of epipubic bones in all non-placental mammals prevents the expansion of the torso needed for full pregnancy. Even non-placental eutherians probably reproduced this way.[59][60]

The placentals are unusual among mammals in giving birth to complete and fully developed young, usually after long gestation periods.

Physiology

Endothermy

Nearly all mammals are endothermic ("warm-blooded"). Most mammals also have hair to help keep them warm. Like birds, mammals can forage or hunt in weather and climates too cold for nonavian reptiles and large insects.

Endothermy requires plenty of food energy, so mammals eat more food per unit of body weight than most reptiles. Small insectivorous mammals eat prodigious amounts for their size.

A rare exception, the naked mole-rat, produces little metabolic heat, so it is considered an operational poikilotherm. Birds and tuna are also endothermic, so endothermy is not peculiar to mammals.

Intelligence

In intelligent mammals, such as primates, the cerebrum is larger relative to the rest of the brain. Intelligence itself is not easy to define, but indications of intelligence include the ability to learn, matched with behavioral flexibility. Rats, for example, are considered to be highly intelligent, as they can learn and perform new tasks, an ability that may be important when they first colonize a fresh habitat. In some mammals, food gathering appears to be related to intelligence: a deer feeding on plants has a brain smaller than a cat, which must think to outwit its prey.[61]

Social structure

Locomotion

Mammals evolved from four-legged ancestors. They use their limbs to walk, climb, swim, or fly. Some land mammals have toes that produce claws for climbing or hooves for running. Aquatic mammals like whales and dolphins have flippers which evolved from legs.

Terrestrial

Arboreal

Aquatic

Whales and dolphins propel themselves through the water by moving their tail flukes up and down, adjusting the angle of the flukes as needed. The more massive front of the body contributes stability.[62][63]

Aerial

Feeding

To maintain a high constant body temperature is energy expensive – mammals therefore need a nutritious and plentiful diet. While the earliest mammals were probably predators, different species have since adapted to meet their dietary requirements in a variety of ways. Some eat other animals – this is a carnivorous diet (and includes insectivorous diets). Other mammals, called herbivores, eat plants. A herbivorous diet includes subtypes such as fruit-eating and grass-eating. An omnivore eats both prey and plants. Carnivorous mammals have a simple digestive tract, because the proteins, lipids, and minerals found in meat require little in the way of specialized digestion. Plants, on the other hand, contain complex carbohydrates, such as cellulose. The digestive tract of an herbivore is therefore host to bacteria that ferment these substances, and make them available for digestion. The bacteria are either housed in the multichambered stomach or in a large cecum. The size of an animal is also a factor in determining diet type. Since small mammals have a high ratio of heat-losing surface area to heat-generating volume, they tend to have high energy requirements and a high metabolic rate. Mammals that weigh less than about 18 oz (500 g) are mostly insectivorous because they cannot tolerate the slow, complex digestive process of a herbivore. Larger animals, on the other hand, generate more heat and less of this heat is lost. They can therefore tolerate either a slower collection process (those that prey on larger vertebrates) or a slower digestive process (herbivores). Furthermore, mammals that weigh more than 18 oz (500 g) usually cannot collect enough insects during their waking hours to sustain themselves. The only large insectivorous mammals are those that feed on huge colonies of insects (ants or termites).[61]

Specializations in herbivory include: Granivory "seed eating", folivory "leaf eating", frugivory "fruit eating", nectivory "nectar eating", gummivory "gum eating", and mycophagy "fungus eating".

Hybrid mammals

Main article: Hybrid (biology)

The deliberate or accidental hybridising of two or more species of closely related animals through captive breeding is a human activity which has been in existence for millennia and has grown in recent times for economic purposes. The number of successful interspecific mammalian hybrids is relatively small, although it has come to be known that there is a significant number of naturally occurring hybrids between forms or regional varieties of a single species. These may form zones of gradation known as clines. Indeed, the distinction between some hitherto distinct species can become clouded once it can be shown that they may not only breed but produce fertile offspring. Some hybrid animals exhibit greater strength and resilience than either parent. This is known as hybrid vigor. The existence of the mule (donkey sire; horse dam) being used widely as a hardy draught animal throughout ancient and modern history is testament to this. Other well known examples are the lion/tiger hybrid, the liger, which is by far the largest big cat and sometimes used in circuses; and cattle hybrids such as between European and Indian domestic cattle or between domestic cattle and American bison, which are used in the meat industry and marketed as Beefalo. There is some speculation that the donkey itself may be the result of an ancient hybridisation between two wild ass species or sub-species. Hybrid animals are normally infertile partly because their parents usually have slightly different numbers of chromosomes, resulting in unpaired chromosomes in their cells, which prevents division of sex cells and the gonads from operating correctly, particularly in males. There are exceptions to this rule, especially if the speciation process was relatively recent or incomplete as is the case with many cattle and dog species. Normally behavior traits, natural hostility, natural ranges and breeding cycle differences maintain the separateness of closely related species and prevent natural hybridisation. However, the widespread disturbances to natural animal behaviours and range caused by human activity, cities, dumping grounds with food, agriculture, fencing, roads and so on do force animals together which would not normally breed. Clear examples exist between the various sub-species of grey wolf, coyote and domestic dog in North America. As many birds and mammals imprint on their mother and immediate family from infancy, a practice used by animal hybridizers is to foster a planned parent in a hybridization program with the same species as the one with which they are planned to mate.

See also

Note

  1. ↑ With a few exceptions, all of them cetaceans.

References

  1. 1 2 Wilson, D.E.; Reeder, D.M., eds. (2005). "Preface and introductory material". Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. xxvi. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ↑ "Initiatives". The IUCN Red List of Threatened Species. IUCN. April 2010.
  3. ↑ Rowe, T. (1988). "Definition, diagnosis, and origin of Mammalia" (PDF). Journal of Vertebrate Paleontology 8 (3): 241–264. doi:10.1080/02724634.1988.10011708.
  4. ↑ Lyell, Charles (1871). The Student's Elements of Geology. London: John Murray. p. 347. Retrieved August 12, 2013.
  5. ↑ Kemp, T. S. (2005). The Origin and Evolution of Mammals. Oxford University Press. p. 3. ISBN 0-19-850760-7.
  6. ↑ Cifelli, Richard L.; Davis, Brian M. (2003). "Marsupial origins". Science 302 (5652): 1899–1900. doi:10.1126/science.1092272. PMID 14671280.
  7. ↑ Datta, P. M. (2005). "Earliest mammal with transversely expanded upper molar from the Late Triassic (Carnian) Tiki Formation, South Rewa Gondwana Basin, India". Journal of Vertebrate Paleontology 25 (1): 200–207. doi:10.1671/0272-4634(2005)025[0200:EMWTEU]2.0.CO;2.
  8. ↑ Luo, Zhe-Xi; Martin, Thomas (2007). "Analysis of Molar Structure and Phylogeny of Docodont Genera" (PDF). Bulletin of Carnegie Museum of Natural History 39: 27–47. doi:10.2992/0145-9058(2007)39[27:AOMSAP]2.0.CO;2. Retrieved April 8, 2013.
  9. ↑ van Nievelt, Alexander F. H.; Smith, Kathleen K. (2005). "To replace or not to replace: the significance of reduced functional tooth replacement in marsupial and placental mammals". Paleobiology 31 (2): 324–346. doi:10.1666/0094-8373(2005)031[0324:trontr]2.0.co;2.
  10. ↑ McKenna, Malcolm C.; Bell, Susan Groag (1997). Classification of Mammals above the Species Level. Columbia University Press. ISBN 0-231-11013-8.
  11. ↑ Schiewe, Jessie (2010-07-28). "Australia's marsupials originated in what is now South America, study says". LATimes.Com. Los Angeles Times. Archived from the original on 1 August 2010. Retrieved 2010-08-01. External link in |work= (help)
  12. ↑ Nilsson, M. A.; Churakov, G.; , Sommer, M.; Van Tran, N.; Zemann, A.; Brosius, J.; Schmitz, J. (2010-07-27). "Tracking Marsupial Evolution Using Archaic Genomic Retroposon Insertions". PLoS Biology (Public Library of Science) 8 (7): e1000436. doi:10.1371/journal.pbio.1000436. PMC 2910653. PMID 20668664.
  13. ↑ Kriegs, Jan Ole; Churakov, Gennady; Kiefmann, Martin; Jordan, Ursula; Brosius, Jürgen; Schmitz, Jürgen (2006). "Retroposed Elements as Archives for the Evolutionary History of Placental Mammals". PLoS Biology 4 (4): e91. doi:10.1371/journal.pbio.0040091. PMC 1395351. PMID 16515367.
  14. 1 2 Nishihara, H.; Maruyama, S.; Okada, N. (2009). "Retroposon analysis and recent geological data suggest near-simultaneous divergence of the three superorders of mammals". Proceedings of the National Academy of Sciences 106 (13): 5235–5240. doi:10.1073/pnas.0809297106.
  15. ↑ Springer, Mark S.; Murphy, William J.; Eizirik, Eduardo; O'Brien, Stephen J. (2003). "Placental mammal diversification and the Cretaceous–Tertiary boundary". Proceedings of the National Academy of Sciences 100 (3): 1056–1061. doi:10.1073/pnas.0334222100. PMC 298725. PMID 12552136.
  16. ↑ Tarver, J.E. et al. (2016) The Interrelationships of Placental Mammals and the Limits of Phylogenetic Inference. Genome Biol Evol, (2016) 8 (2): 330-344. doi: 10.1093/gbe/evv261
  17. ↑ Jin Meng, Yuanqing Wang and Chuankui Li (2011). "Transitional mammalian middle ear from a new Cretaceous Jehol eutriconodont". Nature 472 (7342): 181–185. Bibcode:2011Natur.472..181M. doi:10.1038/nature09921. PMID 21490668.
  18. ↑ Haaramo, Mikko. "Mammaliaformes– mammals and near-mammals". Mikko's Phylogeny Archive.
  19. ↑ Ahlberg, P. E. and Milner, A. R. (April 1994). "The Origin and Early Diversification of Tetrapods". Nature 368 (6471): 507–514. Bibcode:1994Natur.368..507A. doi:10.1038/368507a0. Retrieved 2008-09-06.
  20. ↑ "Amniota – Palaeos". Archived from the original on 2010-12-20.
  21. ↑ "Synapsida overview – Palaeos". Archived from the original on 2010-12-20.
  22. 1 2 Kemp, T. S. (2006). "The origin and early radiation of the therapsid mammal-like reptiles: a palaeobiological hypothesis" (PDF). Journal of Evolutionary Biology 19 (4): 1231–47. doi:10.1111/j.1420-9101.2005.01076.x. PMID 16780524.
  23. ↑ "Therapsida – Palaeos".
  24. ↑ Kermack, D.M.; Kermack, K.A. (1984). The evolution of mammalian characters. Croom Helm. ISBN 0-7099-1534-9.
  25. 1 2 Bennett, A. F. and Ruben, J. A. (1986) "The metabolic and thermoregulatory status of therapsids"; pp. 207–218 in N. Hotton III, P. D. MacLean, J. J. Roth and E. C. Roth (eds), "The ecology and biology of mammal-like reptiles", Smithsonian Institution Press, Washington.
  26. ↑ "Jurassic "Beaver" Found; Rewrites History of Mammals".
  27. ↑ Hall, M. I.; Kamilar, J. M.; Kirk, E. C. (24 October 2012). "Eye shape and the nocturnal bottleneck of mammals". Proceedings of the Royal Society B: Biological Sciences 279 (1749): 4962–4968. doi:10.1098/rspb.2012.2258. PMID 23097513.
  28. ↑ Luo, Zhe-Xi (2007). "Transformation and diversification in early mammal evolution" (PDF). Nature 450 (7172): 1011–19. Bibcode:2007Natur.450.1011L. doi:10.1038/nature06277. PMID 18075580.
  29. ↑ "Oldest Marsupial Fossil Found in China". National Geographic News. December 15, 2003.
  30. ↑ Luo, Zhe-Xi; Yuan, Chong-Xi; Meng, Qing-Jin; Ji, Qiang (2011). "A Jurassic eutherian mammal and divergence of marsupials and placentals" (PDF). Nature 476 (7361): 442–445. Bibcode:2011Natur.476..442L. doi:10.1038/nature10291. PMID 21866158.
  31. ↑ "Eomaia scansoria: discovery of oldest known placental mammal".
  32. ↑ M. J. Novacek, G. W. Rougier, J. R. Wible, M. C. McKenna, D. Dashzeveg, and I. Horovitz (1997). "Epipubic bones in eutherian mammals from the Late Cretaceous of Mongolia". Nature 389 (6650): 483–486. Bibcode:1997Natur.389..483N. doi:10.1038/39020. PMID 9333234.
  33. 1 2 Sahney, S., Benton, M.J. and Ferry, P.A. (2010). "Links between global taxonomic diversity, ecological diversity and the expansion of vertebrates on land" (PDF). Biology Letters 6 (4): 544–547. doi:10.1098/rsbl.2009.1024. PMC 2936204. PMID 20106856.
  34. ↑ "Rogue finger gene got bats airborne". Newscientist.com. Retrieved 2009-03-08.
  35. ↑ Bininda-Emonds, O.R.P.; Cardillo, M.; Jones, K.E.; Beck, Robin M. D.; Grenyer, Richard; Price, Samantha A.; Vos, Rutger A.; et al. (2007). "The delayed rise of present-day mammals". Nature 446 (7135): 507–511. Bibcode:2007Natur.446..507B. doi:10.1038/nature05634. PMID 17392779.
  36. 1 2 Wible, J. R.; Rogier, G. W.; Novacek, M. J.; Asher, R. J. (2007). "Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary" (PDF). Nature 447 (7147): 1003–06. Bibcode:2007Natur.447.1003W. doi:10.1038/nature05854. PMID 17581585.
  37. ↑ O'Leary, Maureen A.; Bloch, Jonathan I.; Flynn, John J.; Gaudin, Timothy J.; Giallombardo, Andres; Giannini, Norberto P.; Goldberg, Suzann L.; Kraatz, Brian P.; Luo, Zhe-Xi; Meng, Jin; Novacek, Michael J.; Perini, Fernando A.; Randall, Zachary S.; Rougier, Guillermo; Sargis, Eric J.; Silcox, Mary T.; Simmons, Nancy b.; Spaulding, Micelle; Velazco, Paul M.; Weksler, Marcelo; Wible, John r.; Cirranello, Andrea L.; Cirranello, Andrea L. (8 February 2013). "The Placental Mammal Ancestor and the Post–K-Pg Radiation of Placentals". Science 339 (6120): 662–667. Bibcode:2013Sci...339..662O. doi:10.1126/science.1229237. PMID 23393258. Retrieved 9 February 2013.
  38. ↑ Wilford, John Noble (7 February 2013). "Rat-Size Ancestor Said to Link Man and Beast". New York Times. Retrieved 9 February 2013.
  39. 1 2 3 Wilford, John Noble (5 June 2013). "Palm-Size Fossil Resets Primates' Clock, Scientists Say". New York Times. Retrieved 5 June 2013.
  40. 1 2 3 Ni, Xijun; Gebo, Daniel L.; Dagosto, Marian; Meng, Jin; Tafforeau, Paul; Flynn, John J. Last7=Beard; Beard, K. Christopher (6 June 2013). "The oldest known primate skeleton and early haplorhine evolution". Nature 498 (7452): 60–64. Bibcode:2013Natur.498...60N. doi:10.1038/nature12200. PMID 23739424. Retrieved 5 June 2013.
  41. ↑ Oftedal, O.T. (2002). "The mammary gland and its origin during synapsid evolution". Journal of Mammary Gland Biology and Neoplasia 7 (3): 225–252. doi:10.1023/A:1022896515287. PMID 12751889.
  42. ↑ Oftedal, O.T. (2002). "The origin of lactation as a water source for parchment-shelled eggs". Journal of Mammary Gland Biology and Neoplasia 7 (3): 253–266. doi:10.1023/A:1022848632125. PMID 12751890.
  43. ↑ "Lactating on Eggs". Nationalzoo.si.edu. 2003-07-14. Retrieved 2009-03-08.
  44. ↑ Brink, A.S. (1955). "A study on the skeleton of Diademodon". Palaeontologia Africana 3: 3–39.
  45. ↑ Kemp, T.S. (1982). Mammal-like reptiles and the origin of mammals. London: Academic Press. p. 363. ISBN 0-12-404120-5.
  46. ↑ Estes, R. (1961). "Cranial anatomy of the cynodont reptile Thrinaxodon liorhinus". Bulletin of the Museum of Comparative Zoology (1253): 165–180.
  47. ↑ "Thrinaxodon: The Emerging Mammal". National Geographic Daily News. February 11, 2009. Retrieved August 26, 2012.
  48. ↑ Kielan−Jaworowska, Z.; Hurum, J.H.. (2006). "Limb posture in early mammals: Sprawling or parasagittal" (PDF). Acta Palaeontologica Polonica 51 (3): 10237–10239.
  49. ↑ Paul, G.S. (1988). Predatory Dinosaurs of the World. New York: Simon and Schuster. p. 464. ISBN 0-671-61946-2.
  50. ↑ J.M. Watson and J.A.M. Graves (1988). "Monotreme Cell-Cycles and the Evolution of Homeothermy". Australian Journal of Zoology (CSIRO) 36 (5): 573–584. doi:10.1071/ZO9880573.
  51. ↑ Jason A. Lillegraven, Zofia Kielan-Jaworowska, William A. Clemens, Mesozoic Mammals: The First Two-Thirds of Mammalian History, University of California Press, 17/12/1979 - 321
  52. ↑ CRC Handbook of Marine Mammal Medicine: Health, Disease, and Rehabilitation. Books.google.com. 2001-06-27. ISBN 978-1-4200-4163-7. Retrieved 2013-08-16.
  53. 1 2 3 Bradley et. al, Brenda year=2012. "Coat Color Variation and Pigmentation Gene Expression in Rhesus Macaques (Macaca Mulatta)" (PDF). Journal of Mammalian Evolution 20: 263–70. doi:10.1007/s10914-012-9212-3.
  54. 1 2 3 Caro, Tim (2005). "The Adaptive Significance of Coloration in Mammals" (PDF). BioScience 55 (2): 125–136. doi:10.1641/0006-3568(2005)055[0125:tasoci]2.0.co;2.
  55. ↑ Hoeskra, HE (2006). "genetics, development, and the evolution of adaptive pigmentation in vertebrates" (PDF). Heredity 97: 222–234. doi:10.1038/sj.hdy.6800861. PMID 16823403.
  56. ↑ Prum, Richard O.; Torres, Rodolfo H. (2004). "Structural colouration of mammalian skin: convergent evolution of coherently scattering dermal collagen arrays" (PDF). Journal of Experimental Biology 207 (12): 2157–72. doi:10.1242/jeb.00989.
  57. ↑ Wallis M.C., Waters P.D., Delbridge M.L., Kirby P.J., Pask A.J., Grützner F., Rens W., Ferguson-Smith M.A., Graves J.A.M.; Waters; Delbridge; Kirby; Pask; Grützner; Rens; Ferguson-Smith; Graves; et al. (2007). "Sex determination in platypus and echidna: autosomal location of SOX3 confirms the absence of SRY from monotremes". Chromosome Research 15 (8): 949–959. doi:10.1007/s10577-007-1185-3. PMID 18185981.
  58. ↑ Marshall Graves, Jennifer A. (2008). "Weird Animal Genomes and the Evolution of Vertebrate Sex and Sex Chromosomes" (PDF). Annual Review of Genetics 42: 568–586. doi:10.1146/annurev.genet.42.110807.091714. PMID 18983263.
  59. ↑ Giallombardo, Andres, 2009 New Cretaceous mammals from Mongolia and the early diversification of Eutheria Ph.D. dissertion, Columbia University, 2009402 pages; AAT 3373736 (abstract) The origin of Placental Mammals, Cimolestidae, Zalambdalestidae
  60. ↑ Michael L. Power,Jay Schulkin. The Evolution Of The Human Placenta. pp. 68–.
  61. 1 2 Don E. Wilson & David Burnie, ed. (2001). Animal: The Definitive Visual Guide to the World's Wildlife (1st ed.). DK Publishing. pp. 86–89. ISBN 978-0-7894-7764-4.
  62. ↑ Perry, D. A. (1949). "The anatomical basis of swimming in Whales". Journal of Zoology 119 (1): 49–60. doi:10.1111/j.1096-3642.1949.tb00866.x.
  63. ↑ Fish, F. E.; Hui, C. A. (1991). "Dolphin swimming — a review" (PDF). Mammal Review 21 (4): 181–195. doi:10.1111/j.1365-2907.1991.tb00292.x.

Further reading

External links

The Wikibook Dichotomous Key has a page on the topic of: Mammalia
External identifiers for Mammalia
Encyclopedia of Life 1642
ITIS 179913
NCBI 40674
Also found in: Wikispecies, Arctos


This article is issued from Wikipedia - version of the Saturday, April 23, 2016. The text is available under the Creative Commons Attribution/Share Alike but additional terms may apply for the media files.