哺乳动物或哺乳类是对脊椎动物下哺乳纲（学名：Mammalia，出自拉丁语mamma，即“胸部”）动物的统称。与同属羊膜动物、在石炭紀（約30億年前）分化的爬蟲類、鳥類不同的是，哺乳类具有新皮质、毛皮、三個听小骨和乳腺，雌性还會用乳腺分泌乳汁餵養幼崽（棕梠果蝠的雄性也可以餵乳）。目前，人们共发表了约 6400 个哺乳類物种，其中物种数量最多的目有囓齒目、翼手目和真盲缺目（刺猬、鼹鼠、鼩鼱等），多樣性略低於以上三目的有灵长目（包含人類在內的猿、猴等）、偶蹄目（鯨豚類和其他偶蹄目）和食肉目（猫、狗、海豹等）。
以支序學的角度來看，哺乳類是合弓綱中的唯一成員，而合弓綱和蜥形綱組合成了一個更大的演化支羊膜動物。合弓綱哺乳動物的先祖為楔齒龍類中的盤龍目，包含非哺乳類的異齒龍屬。在 30 億年前，石炭紀末期，合弓綱從蜥形綱中分化出來後，其中一個分支演化成了哺乳類，而蜥形綱則成為了現生的鳥類和爬蟲類。在二疊紀早期獸孔目出現之前，楔齒龍類的基群演化成了好幾支多樣性高的非哺乳類合弓綱，這些生物偶爾會被誤認為類似哺乳類的爬蟲類。現生的哺乳類起源於新生代的古近紀和新近紀，大約6千6百萬年前，也就是非鳥恐龍滅絕之後，且自出現以來便是陸生動物中最具優勢的類群。
自从卡尔·林奈首次将哺乳类分为一纲以来，哺乳动物的定义曾多次变动。目前，人们对哺乳动物的分类方式还没有明确的共识；最新有关方面的研究由麦肯纳与贝尔（1997年）和威尔森与里德尔（2005年）作出。喬治·蓋洛德·辛普森的“哺乳动物分类与分类原则”（AMNH Bulletin v. 85, 1945）从系統分類學上阐述了哺乳动物的起源、关系，他的理论直到20世纪末都还出现在各个教科书中。自辛普森发明了分类系统以来，化石记录已几经修正。分类系统理论本身也经多方讨论、发展壮大，在支序分類學出现后更是愈演愈烈。虽然实地考察中新发现的生物证据让辛普森的分类系统逐渐落伍，但他的分类方式依然是哺乳动物分类体系中最为权威的。
大多数哺乳动物，包括物种最多的六个目，属于胎盘动物。其中，物种最多的三个目是啮齿目（鼠、豪豬、河狸、水豚等啃咬哺乳动物）、翼手目（蝙蝠）和鼩形目（鼩鼱、鼹鼠、溝齒鼩等）。The next three biggest orders, depending on the biological classification scheme used, are the Primates including the apes, monkeys and lemurs; the Cetartiodactyla including whales and even-toed ungulates; and the Carnivora which includes cats, dogs, weasels, bears, seals and allies. According to Mammal Species of the World, 5,416 species were identified in 2006. These were grouped into 1,229 genera, 153 families and 29 orders. In 2008, the International Union for Conservation of Nature (IUCN) completed a five-year Global Mammal Assessment for its IUCN Red List, which counted 5,488 species. According to research published in the Journal of Mammalogy in 2018, the number of recognized mammal species is 6,495 including 96 recently extinct.
The word "mammal" is modern, from the scientific name Mammalia coined by Carl Linnaeus in 1758, derived from the Latin mamma ("teat, pap"). In an influential 1988 paper, Timothy Rowe defined Mammalia phylogenetically as the crown group of 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. 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. 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 in the Middle Jurassic, this is a reasonable estimate for the appearance of the crown group.
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. The earliest known synapsid satisfying Kemp's definitions is Tikitherium, dated , so the appearance of mammals in this broader sense can be given this Late Triassic date.
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, is a comprehensive work on the systematics, relationships and occurrences of all mammal taxa, living and extinct, down through the rank of genus, though 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.
- 原兽亚纲 Prototheria：
- 獸形亞綱 Theriiformes：現存的哺乳動物與其已滅絕的近親
- †异兽亚纲 Allotheria: multituberculates
- †真三尖齿兽目 Eutriconodonta: eutriconodonts
- 完獸亞綱 Holotheria：現存的哺乳動物與其已滅絕的近親
- 獸亞綱 Theria：現存的哺乳動物
- 有袋類 Marsupialia
- 胎盤動物 Placentalia
As of the early 21st century, molecular studies based on DNA analysis have suggested new relationships among mammal families. Most of these findings have been independently validated by retrotransposon presence/absence data. Classification systems based on molecular studies reveal three major groups or lineages of placental mammals—Afrotheria, Xenarthra and Boreoeutheria—which diverged in the Cretaceous. The relationships between these three lineages is contentious, and all three possible hypotheses have been proposed with respect to which group is basal. These hypotheses are Atlantogenata (basal Boreoeutheria), Epitheria (basal Xenarthra) and Exafroplacentalia (basal Afrotheria). 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 the type of DNA used (such as nuclear or mitochondrial) and varying interpretations of paleogeographic data.
- 非洲食蟲類 Afroinsectiphilia
- 近蹄類 Paenungulata
- 靈長總目 Euarchontoglires
- 勞亞獸總目 Laurasiatheria
- 顎关节 – The dentary (the lower jaw bone, which carries the teeth) and the squamosal (a small cranial bone) meet to form the joint. In most gnathostomes, including early therapsids, the joint consists of the articular (a small bone at the back of the lower jaw) and quadrate (a small bone at the back of the upper jaw).
- 中耳 – In crown-group mammals, sound is carried from the eardrum by a chain of three bones, the malleus, the incus and the stapes. Ancestrally, the malleus and the incus are derived from the articular and the quadrate bones that constituted the jaw joint of early therapsids.
- 换牙 – Teeth can be replaced once (diphyodonty) or (as in toothed whales and murid rodents) not at all (monophyodonty). Elephants, manatees, and kangaroos continually grow new teeth throughout their life (polyphyodonty).
- Prismatic enamel – The enamel coating on the surface of a tooth consists of prisms, solid, rod-like structures extending from the dentin to the tooth's surface.
- Occipital condyles – Two knobs at the base of the skull fit into the topmost neck vertebra; most other tetrapods, in contrast, have only one such knob.
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 which has nine cervical vertebrae. All mammalian brains possess a neocortex, a brain region unique to mammals. Placental mammals have a corpus callosum, unlike monotremes and marsupials.
The lungs of mammals are spongy and honeycombed. Breathing is mainly achieved with the diaphragm, which divides the thorax from the abdominal cavity, forming a dome convex to the thorax. Contraction of the diaphragm flattens the dome, increasing the volume of the lung cavity. Air enters through the oral and nasal cavities, and travels through the larynx, trachea and bronchi, and expands the alveoli. Relaxing the diaphragm has the opposite effect, decreasing the volume of the lung cavity, causing air to be pushed out of the lungs. During exercise, the abdominal wall contracts, increasing pressure on the diaphragm, which forces air out quicker and more forcefully. The rib cage is able to expand and contract the chest cavity through the action of other respiratory muscles. Consequently, 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 due to its resemblance to blacksmith bellows.
The mammalian heart has four chambers, two upper atria, the receiving chambers, and two lower ventricles, the discharging chambers. The heart has four valves, which separate its chambers and ensures blood flows in the correct direction through the heart (preventing backflow). After gas exchange in the pulmonary capillaries (blood vessels in the lungs), oxygen-rich blood returns to the left atrium via one of the four pulmonary veins. Blood flows nearly continuously back into the atrium, which acts as the receiving chamber, and from here through an opening into the left ventricle. Most blood flows passively into the heart while both the atria and ventricles are relaxed, but toward the end of the ventricular relaxation period, the left atrium will contract, pumping blood into the ventricle. The heart also requires nutrients and oxygen found in blood like other muscles, and is supplied via coronary arteries.
The integumentary system (skin) 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, which stores lipids and provides cushioning and insulation. The thickness of this layer varies widely from species to species;:97 marine mammals require a thick hypodermis (blubber) for insulation, and right whales have the thickest blubber at 20英寸（51厘米）. 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.:61
Herbivores have developed a diverse range of physical structures to facilitate the consumption of plant material. To break up intact plant tissues, mammals have developed teeth structures that reflect their feeding preferences. For instance, frugivores (animals that feed primarily on fruit) and herbivores that feed on soft foliage have low-crowned teeth specialized for grinding foliage and seeds. Grazing animals that tend to eat hard, silica-rich grasses, have high-crowned teeth, which are capable of grinding tough plant tissues and do not wear down as quickly as low-crowned teeth. Most carnivorous mammals have carnassialiforme teeth (of varying length depending on diet), long canines and similar tooth replacement patterns.
The stomach of Artiodactyls is divided into four sections: the rumen, the reticulum, the omasum and the abomasum (only ruminants have a rumen). After the plant material is consumed, it is mixed with saliva in the rumen and reticulum and separates into solid and liquid material. The solids lump together to form a bolus (or cud), and is regurgitated. When the bolus enters the mouth, the fluid is squeezed out with the tongue and swallowed again. Ingested food passes to the rumen and reticulum where cellulytic microbes (bacteria, protozoa and fungi) produce cellulase, which is needed to break down the cellulose in plants. Perissodactyls, in contrast to the ruminants, store digested food that has left the stomach in an enlarged cecum, where it is fermented by bacteria. Carnivora have a simple stomach adapted to digest primarily meat, as compared to the elaborate digestive systems of herbivorous animals, which are necessary to break down tough, complex plant fibers. The caecum is either absent or short and simple, and the large intestine is not sacculated or much wider than the small intestine.
The mammalian excretory system involves many components. Like most other land animals, mammals are ureotelic, and convert ammonia into urea, which is done by the liver as part of the urea cycle. Bilirubin, a waste product derived from blood cells, is passed through bile and urine with the help of enzymes excreted by the liver. The passing of bilirubin via bile through the intestinal tract gives mammalian feces a distinctive brown coloration. Distinctive features of the mammalian kidney include the presence of the renal pelvis and renal pyramids, and of a clearly distinguishable cortex and medulla, which is due to the presence of elongated loops of Henle. Only the mammalian kidney has a bean shape, although there are some exceptions, such as the multilobed reniculate kidneys of pinnipeds, cetaceans and bears. Most adult placental mammals have no remaining trace of the cloaca. In the embryo, the embryonic cloaca divides into a posterior region that becomes part of the anus, and an anterior region that has different fates depending on the sex of the individual: in females, it develops into the vestibule that receives the urethra and vagina, while in males it forms the entirety of the penile urethra. However, the tenrecs, golden moles, and some shrews retain a cloaca as adults. In marsupials, the genital tract is separate from the anus, but a trace of the original cloaca does remain externally. Monotremes, which translates from Greek into "single hole", have a true cloaca.
As in all other tetrapods, mammals have a larynx that can quickly open and close to produce sounds, and a supralaryngeal vocal tract which filters this sound. The lungs and surrounding musculature provide the air stream and pressure required to phonate. The larynx controls the pitch and volume of sound, but the strength the lungs exert to exhale also contributes to volume. More primitive mammals, such as the echidna, can only hiss, as sound is achieved solely through exhaling through a partially closed larynx. Other mammals phonate using vocal folds, as opposed to the vocal cords seen in birds and reptiles. The movement or tenseness of the vocal folds can result in many sounds such as purring and screaming. Mammals can change the position of the larynx, allowing them to breathe through the nose while swallowing through the mouth, and to form both oral and nasal sounds; nasal sounds, such as a dog whine, are generally soft sounds, and oral sounds, such as a dog bark, are generally loud.
Some mammals have a large larynx and thus a low-pitched voice, namely the hammer-headed bat (Hypsignathus monstrosus) where the larynx can take up the entirety of the thoracic cavity while pushing the lungs, heart, and trachea into the abdomen. Large vocal pads can also lower the pitch, as in the low-pitched roars of big cats. The production of infrasound is possible in some mammals such as the African elephant (Loxodonta spp.) and baleen whales. Small mammals with small larynxes have the ability to produce ultrasound, which can be detected by modifications to the middle ear and cochlea. Ultrasound is inaudible to birds and reptiles, which might have been important during the Mesozoic, when birds and reptiles were the dominant predators. This private channel is used by some rodents in, for example, mother-to-pup communication, and by bats when echolocating. Toothed whales also use echolocation, but, as opposed to the vocal membrane that extends upward from the vocal folds, they have a melon to manipulate sounds. Some mammals, namely the primates, have air sacs attached to the larynx, which may function to lower the resonances or increase the volume of sound.
The vocal production system is controlled by the cranial nerve nuclei in the brain, and supplied by the recurrent laryngeal nerve and the superior laryngeal nerve, branches of the vagus nerve. The vocal tract is supplied by the hypoglossal nerve and facial nerves. Electrical stimulation of the periaqueductal gray (PEG) region of the mammalian midbrain elicit vocalizations. The ability to learn new vocalizations is only exemplified in humans, seals, cetaceans, elephants and possibly bats; in humans, this is the result of a direct connection between the motor cortex, which controls movement, and the motor neurons in the spinal cord.
- 定期毛髮（Definitive）– 當達到一定長度後即會脫落。
- 感覺毛（Vibrissae）– 用於感測周遭環境。
- 毛皮（Pelage）– 包括護毛、絨毛與芒毛。
- 棘刺（Spine）– 堅硬的毛髮，用於防衛（例如豪豬背上的長刺）。
- 剛毛（Bristle）– 主要用於展示視覺訊號（例如獅子的鬃毛）。
- 柔毛（Velli）– 用於保暖，主要常見於新生的哺乳動物。
- 羊毛（Wool]）– 長而柔軟的卷曲毛髮。
毛髮長度與體溫調節無關：例如，部分棲息於熱帶地區的哺乳動物，如樹懶，具有與極區哺乳動物等長度的毛髮，然而並不具有相當保暖的能力；反而，部分僅具有短毛的熱帶哺乳動物，保暖能力卻與極區哺乳動物相當。毛髮生長的密度可以影響保暖的能力，尤其棲息於極區的哺乳動物都舉有相當高密度的毛髮，例如麝牛除了具有30 cm（12英寸）長的護毛外，還具有高密度的絨毛，能有效鎖住並減少體熱的散失，讓牠們能於−40 °C（−40 °F）的環境下生存:162–163。有些棲息於沙漠的哺乳動物，如駱駝，則透過高密度的毛來將外在環境的熱度隔絕於外，確保個體不會過熱；在夏天時，駱駝的毛髮溫度可達70 °C（158 °F），而皮膚溫度則能維持40 °C（104 °F）:188。水生哺乳動物也具有高密度的毛髮，透過將水隔絕於外保持皮膚乾燥的方式來減少體熱散失:162–163。
Mammalian coats are colored for a variety of reasons, the major selective pressures including camouflage, sexual selection, communication, and thermoregulation. Coloration in both the hair and skin of mammals is mainly determined by the type and amount of melanin; eumelanins for brown and black colors and pheomelanin for a range of yellow to reddish-brown colors, giving mammals an earth tone. Some mammals, like the mandrill, have more vibrant colors due to structural coloration. Many sloths appear green because their fur hosts green algae; this may be a symbiotic relation that affords camouflage to the sloths.
Camouflage is a powerful influence in a large number of mammals, as it helps to conceal individuals from predators or prey. In arctic and subarctic mammals such as the arctic fox (Alopex lagopus), collared lemming (Dicrostonyx groenlandicus), stoat (Mustela erminea), and snowshoe hare (Lepus americanus), seasonal color change between brown in summer and white in winter is driven largely by camouflage. Some arboreal mammals, notably primates and marsupials, have shades of violet, green, or blue skin on parts of their bodies, indicating some distinct advantage in their largely arboreal habitat due to convergent evolution.
Aposematism, warning off possible predators, is the most likely explanation of the black-and-white pelage of many mammals which are able to defend themselves, such as in the foul-smelling skunk and the powerful and aggressive honey badger. Coat color is sometimes sexually dimorphic, as in many primate species. Differences in female and male coat color may indicate nutrition and hormone levels, important in mate selection. Coat color may influence the ability to retain heat, depending on how much light is reflected. Mammals with a darker colored coat can absorb more heat from solar radiation, and stay warmer, and some smaller mammals, such as voles, have darker fur in the winter. The white, pigmentless fur of arctic mammals, such as the polar bear, may reflect more solar radiation directly onto the skin.:166–167 The dazzling black-and-white striping of zebras appear to provide some protection from biting flies.
In male placentals, the penis is used both for urination and copulation. Depending on the species, an erection may be fueled by blood flow into vascular, spongy tissue or by muscular action. A penis may be contained in a prepuce when not erect, and some placentals also have a penis bone (baculum). Marsupials typically have forked penises, while the echidna penis generally has four heads with only two functioning. The testes of most mammals descend into the scrotum which is typically posterior to the penis but is often anterior in marsupials. Female mammals generally have a clitoris, labia majora and labia minora on the outside, while the internal system contains paired oviducts, 1-2 uteri, 1-2 cervices and a vagina. Marsupials have two lateral vaginas and a medial vagina. The "vagina" of monotremes is better understood as a "urogenital sinus". The uterine systems of placental mammals can vary between a duplex, were there are two uteri and cervices which open into the vagina, a bipartite, were two uterine horns have a single cervix that connects to the vagina, a bicornuate, which consists where two uterine horns that are connected distally but separate medially creating a Y-shape, and a simplex, which has a single uterus.:220–221, 247
The ancestral condition for mammal reproduction is the birthing of relatively undeveloped, either through direct vivipary or a short period as soft-shelled eggs. This is likely due to the fact that the torso could not expand due to the presence of epipubic bones. The oldest demonstration of this reproductive style is with Kayentatherium, which produced undeveloped perinates, but at much higher litter sizes than any modern mammal, 38 specimens. Most modern mammals are viviparous, giving birth to live young. However, the five species of monotreme, the platypus and the four species of echidna, lay eggs. The monotremes have a sex determination system different from that of most other mammals. In particular, the sex chromosomes of a platypus are more like those of a chicken than those of a therian mammal.
Viviparous mammals are in the subclass Theria; those living today are in the marsupial and placental infraclasses. Marsupials have 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 plesiomorphic 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. The placentals give birth to relatively complete and developed young, usually after long gestation periods. They get their name from the placenta, which connects the developing fetus to the uterine wall to allow nutrient uptake. In placental mammals, the epipubic is either completely lost or converted into the baculum; allowing the torso to be able to expand and thus birth developed offspring.
The mammary glands of mammals are specialized to produce milk, the primary source of nutrition for newborns. 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. Compared to placental mammals, the milk of marsupials changes greatly in both production rate and in nutrient composition, due to the underdeveloped young. In addition, the mammary glands have more autonomy allowing them to supply separate milks to young at different development stages. Lactose is the main sugar in placental mammal milk while monotreme and marsupial milk is dominated by oligosaccharides. Weaning is the process in which a mammal becomes less dependent on their mother's milk and more on solid food.
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 ectothermic ("cold-blooded") reptiles and 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 are also endothermic, so endothermy is not unique to mammals.
Among mammals, species maximum lifespan varies significantly (for example the shrew has a lifespan of two years, whereas the oldest bowhead whale is recorded to be 211 years). Although the underlying basis for these lifespan differences is still uncertain, numerous studies indicate that the ability to repair DNA damage is an important determinant of mammalian lifespan. In a 1974 study by Hart and Setlow, it was found that DNA excision repair capability increased systematically with species lifespan among seven mammalian species. Species lifespan was observed to be robustly correlated with the capacity to recognize DNA double-strand breaks as well as the level of the DNA repair protein Ku80. In a study of the cells from sixteen mammalian species, genes employed in DNA repair were found to be up-regulated in the longer-lived species. The cellular level of the DNA repair enzyme poly ADP ribose polymerase was found to correlate with species lifespan in a study of 13 mammalian species. Three additional studies of a variety of mammalian species also reported a correlation between species lifespan and DNA repair capability.
Most vertebrates—the amphibians, the reptiles and some mammals such as humans and bears—are plantigrade, walking on the whole of the underside of the foot. Many mammals, such as cats and dogs, are digitigrade, walking on their toes, the greater stride length allowing more speed. Digitigrade mammals are also often adept at quiet movement. Some animals such as horses are unguligrade, walking on the tips of their toes. This even further increases their stride length and thus their speed. A few mammals, namely the great apes, are also known to walk on their knuckles, at least for their front legs. Giant anteaters and platypuses are also knuckle-walkers. Some mammals are bipeds, using only two limbs for locomotion, which can be seen in, for example, humans and the great apes. Bipedal species have a larger field of vision than quadrupeds, conserve more energy and have the ability to manipulate objects with their hands, which aids in foraging. Instead of walking, some bipeds hop, such as kangaroos and kangaroo rats.
Animals will use different gaits for different speeds, terrain and situations. For example, horses show four natural gaits, the slowest horse gait is the walk, then there are three faster gaits which, from slowest to fastest, are the trot, the canter and the gallop. Animals may also have unusual gaits that are used occasionally, such as for moving sideways or backwards. For example, the main human gaits are bipedal walking and running, but they employ many other gaits occasionally, including a four-legged crawl in tight spaces. Mammals show a vast range of gaits, the order that they place and lift their appendages in locomotion. Gaits can be grouped into categories according to their patterns of support sequence. For quadrupeds, there are three main categories: walking gaits, running gaits and leaping gaits. Walking is the most common gait, where some feet are on the ground at any given time, and found in almost all legged animals. Running is considered to occur when at some points in the stride all feet are off the ground in a moment of suspension.
Arboreal animals frequently have elongated limbs that help them cross gaps, reach fruit or other resources, test the firmness of support ahead and, in some cases, to brachiate (swing between trees). Many arboreal species, such as tree porcupines, silky anteaters, spider monkeys, and possums, use prehensile tails to grasp branches. In the spider monkey, the tip of the tail has either a bare patch or adhesive pad, which provides increased friction. Claws can be used to interact with rough substrates and reorient the direction of forces the animal applies. This is what allows squirrels to climb tree trunks that are so large to be essentially flat from the perspective of such a small animal. However, claws can interfere with an animal's ability to grasp very small branches, as they may wrap too far around and prick the animal's own paw. Frictional gripping is used by primates, relying upon hairless fingertips. Squeezing the branch between the fingertips generates frictional force that holds the animal's hand to the branch. However, this type of grip depends upon the angle of the frictional force, thus upon the diameter of the branch, with larger branches resulting in reduced gripping ability. To control descent, especially down large diameter branches, some arboreal animals such as squirrels have evolved highly mobile ankle joints that permit rotating the foot into a 'reversed' posture. This allows the claws to hook into the rough surface of the bark, opposing the force of gravity. Small size provides many advantages to arboreal species: such as increasing the relative size of branches to the animal, lower center of mass, increased stability, lower mass (allowing movement on smaller branches) and the ability to move through more cluttered habitat. Size relating to weight affects gliding animals such as the sugar glider. Some species of primate, bat and all species of sloth achieve passive stability by hanging beneath the branch. Both pitching and tipping become irrelevant, as the only method of failure would be losing their grip.
Bats are the only mammals that can truly fly. They fly through the air at a constant speed by moving their wings up and down (usually with some fore-aft movement as well). Because the animal is in motion, there is some airflow relative to its body which, combined with the velocity of the wings, generates a faster airflow moving over the wing. This generates a lift force vector pointing forwards and upwards, and a drag force vector pointing rearwards and upwards. The upwards components of these counteract gravity, keeping the body in the air, while the forward component provides thrust to counteract both the drag from the wing and from the body as a whole.
The wings of bats are much thinner and consist of more bones than those of birds, allowing bats to maneuver more accurately and fly with more lift and less drag. By folding the wings inwards towards their body on the upstroke, they use 35% less energy during flight than birds. The membranes are delicate, ripping easily; however, the tissue of the bat's membrane is able to regrow, such that small tears can heal quickly. The surface of their wings is equipped with touch-sensitive receptors on small bumps called Merkel cells, also found on human fingertips. These sensitive areas are different in bats, as each bump has a tiny hair in the center, making it even more sensitive and allowing the bat to detect and collect information about the air flowing over its wings, and to fly more efficiently by changing the shape of its wings in response.
A fossorial (from Latin fossor, meaning "digger") is an animal adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, and naked mole-rats. Many rodent species are also considered fossorial because they live in burrows for most but not all of the day. Species that live exclusively underground are subterranean, and those with limited adaptations to a fossorial lifestyle sub-fossorial. Some organisms are fossorial to aid in temperature regulation while others use the underground habitat for protection from predators or for food storage.
Fossorial mammals have a fusiform body, thickest at the shoulders and tapering off at the tail and nose. Unable to see in the dark burrows, most have degenerated eyes, but degeneration varies between species; pocket gophers, for example, are only semi-fossorial and have very small yet functional eyes, in the fully fossorial marsupial mole the eyes are degenerated and useless, talpa moles have vestigial eyes and the cape golden mole has a layer of skin covering the eyes. External ears flaps are also very small or absent. Truly fossorial mammals have short, stout legs as strength is more important than speed to a burrowing mammal, but semi-fossorial mammals have cursorial legs. The front paws are broad and have strong claws to help in loosening dirt while excavating burrows, and the back paws have webbing, as well as claws, which aids in throwing loosened dirt backwards. Most have large incisors to prevent dirt from flying into their mouth.
Fully aquatic mammals, the cetaceans and sirenians, have lost their legs and have a tail fin to propel themselves through the water. Flipper movement is continuous. Whales swim by moving their tail fin and lower body up and down, propelling themselves through vertical movement, while their flippers are mainly used for steering. Their skeletal anatomy allows them to be fast swimmers. Most species have a dorsal fin to prevent themselves from turning upside-down in the water. The flukes of sirenians are raised up and down in long strokes to move the animal forward, and can be twisted to turn. The forelimbs are paddle-like flippers which aid in turning and slowing.
Semi-aquatic mammals, like pinnipeds, have two pairs of flippers on the front and back, the fore-flippers and hind-flippers. The elbows and ankles are enclosed within the body. Pinnipeds have several adaptions for reducing drag. In addition to their streamlined bodies, they have smooth networks of muscle bundles in their skin that may increase laminar flow and make it easier for them to slip through water. They also lack arrector pili, so their fur can be streamlined as they swim. They rely on their fore-flippers for locomotion in a wing-like manner similar to penguins and sea turtles. Fore-flipper movement is not continuous, and the animal glides between each stroke. Compared to terrestrial carnivorans, the fore-limbs are reduced in length, which gives the locomotor muscles at the shoulder and elbow joints greater mechanical advantage; the hind-flippers serve as stabilizers. Other semi-aquatic mammals include beavers, hippopotamuses, otters and platypuses. Hippos are very large semi-aquatic mammals, and their barrel-shaped bodies have graviportal skeletal structures, adapted to carrying their enormous weight, and their specific gravity allows them to sink and move along the bottom of a river.
Many mammals communicate by vocalizing. Vocal communication serves many purposes, including in mating rituals, as warning calls, to indicate food sources, and for social purposes. Males often call during mating rituals to ward off other males and to attract females, as in the roaring of lions and red deer. The songs of the humpback whale may be signals to females; they have different dialects in different regions of the ocean. Social vocalizations include the territorial calls of gibbons, and the use of frequency in greater spear-nosed bats to distinguish between groups. The vervet monkey gives a distinct alarm call for each of at least four different predators, and the reactions of other monkeys vary according to the call. For example, if an alarm call signals a python, the monkeys climb into the trees, whereas the eagle alarm causes monkeys to seek a hiding place on the ground. Prairie dogs similarly have complex calls that signal the type, size, and speed of an approaching predator. Elephants communicate socially with a variety of sounds including snorting, screaming, trumpeting, roaring and rumbling. Some of the rumbling calls are infrasonic, below the hearing range of humans, and can be heard by other elephants up to 6英里（9.7公里） away at still times near sunrise and sunset.
Mammals signal by a variety of means. Many give visual anti-predator signals, as when deer and gazelle stot, honestly indicating their fit condition and their ability to escape, or when white-tailed deer and other prey mammals flag with conspicuous tail markings when alarmed, informing the predator that it has been detected. Many mammals make use of scent-marking, sometimes possibly to help defend territory, but probably with a range of functions both within and between species. Microbats and toothed whales including oceanic dolphins vocalize both socially and in echolocation.
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, which contain complex carbohydrates such as cellulose. An herbivorous diet includes subtypes such as granivory (seed eating), folivory (leaf eating), frugivory (fruit eating), nectarivory (nectar eating), gummivory (gum eating) and mycophagy (fungus eating). The digestive tract of an herbivore is host to bacteria that ferment these complex substances, and make them available for digestion, which are either housed in the multichambered stomach or in a large cecum. Some mammals are coprophagous, consuming feces to absorb the nutrients not digested when the food was first ingested.:131–137 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. Exceptions to this include baleen whales who also house gut flora in a multi-chambered stomach, like terrestrial herbivores.
The size of an animal is also a factor in determining diet type (Allen's rule). 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盎司（510克；1.1英磅） are mostly insectivorous because they cannot tolerate the slow, complex digestive process of an 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 (carnivores that feed on larger vertebrates) or a slower digestive process (herbivores). Furthermore, mammals that weigh more than 18盎司（510克；1.1英磅） 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).
Some mammals are omnivores and display varying degrees of carnivory and herbivory, generally leaning in favor of one more than the other. Since plants and meat are digested differently, there is a preference for one over the other, as in bears where some species may be mostly carnivorous and others mostly herbivorous. They are grouped into three categories: mesocarnivory (50–70% meat), hypercarnivory (70% and greater of meat), and hypocarnivory (50% or less of meat). The dentition of hypocarnivores consists of dull, triangular carnassial teeth meant for grinding food. Hypercarnivores, however, have conical teeth and sharp carnassials meant for slashing, and in some cases strong jaws for bone-crushing, as in the case of hyenas, allowing them to consume bones; some extinct groups, notably the Machairodontinae, had saber-shaped canines.
Some physiological carnivores consume plant matter and some physiological herbivores consume meat. From a behavioral aspect, this would make them omnivores, but from the physiological standpoint, this may be due to zoopharmacognosy. Physiologically, animals must be able to obtain both energy and nutrients from plant and animal materials to be considered omnivorous. Thus, such animals are still able to be classified as carnivores and herbivores when they are just obtaining nutrients from materials originating from sources that do not seemingly complement their classification. For example, it is well documented that some ungulates such as giraffes, camels, and cattle, will gnaw on bones to consume particular minerals and nutrients. Also, cats, which are generally regarded as obligate carnivores, occasionally eat grass to regurgitate indigestible material (such as hairballs), aid with hemoglobin production, and as a laxative.
Many mammals, in the absence of sufficient food requirements in an environment, suppress their metabolism and conserve energy in a process known as hibernation. In the period preceding hibernation, larger mammals, such as bears, become polyphagic to increase fat stores, whereas smaller mammals prefer to collect and stash food. The slowing of the metabolism is accompanied by a decreased heart and respiratory rate, as well as a drop in internal temperatures, which can be around ambient temperature in some cases. For example, the internal temperatures of hibernating arctic ground squirrels can drop to −2.9 °C（26.8 °F）, however the head and neck always stay above 0 °C（32 °F）. A few mammals in hot environments aestivate in times of drought or extreme heat, namely the fat-tailed dwarf lemur (Cheirogaleus medius).
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.
Tool use by animals may indicate different levels of learning and cognition. The sea otter uses rocks as essential and regular parts of its foraging behaviour (smashing abalone from rocks or breaking open shells), with some populations spending 21% of their time making tools. Other tool use, such as chimpanzees using twigs to "fish" for termites, may be developed by watching others use tools and may even be a true example of animal teaching. Tools may even be used in solving puzzles in which the animal appears to experience a "Eureka moment". Other mammals that do not use tools, such as dogs, can also experience a Eureka moment.
Brain size was previously considered a major indicator of the intelligence of an animal. Since most of the brain is used for maintaining bodily functions, greater ratios of brain to body mass may increase the amount of brain mass available for more complex cognitive tasks. Allometric analysis indicates that mammalian brain size scales at approximately the 2⁄3 or 3⁄4 exponent of the body mass. Comparison of a particular animal's brain size with the expected brain size based on such allometric analysis provides an encephalisation quotient that can be used as another indication of animal intelligence. Sperm whales have the largest brain mass of any animal on earth, averaging 8,000立方厘米（490立方英寸） and 7.8公斤（17英磅） in mature males.
Self-awareness appears to be a sign of abstract thinking. Self-awareness, although not well-defined, is believed to be a precursor to more advanced processes such as metacognitive reasoning. The traditional method for measuring this is the mirror test, which determines if an animal possesses the ability of self-recognition. Mammals that have passed the mirror test include Asian elephants (some pass, some do not); chimpanzees; bonobos; orangutans; humans, from 18 months (mirror stage); bottlenose dolphins[a] killer whales; and false killer whales.
Eusociality is the highest level of social organization. These societies have an overlap of adult generations, the division of reproductive labor and cooperative caring of young. Usually insects, such as bees, ants and termites, have eusocial behavior, but it is demonstrated in two rodent species: the naked mole-rat and the Damaraland mole-rat.
Presociality is when animals exhibit more than just sexual interactions with members of the same species, but fall short of qualifying as eusocial. That is, presocial animals can display communal living, cooperative care of young, or primitive division of reproductive labor, but they do not display all of the three essential traits of eusocial animals. Humans and some species of Callitrichidae (marmosets and tamarins) are unique among primates in their degree of cooperative care of young. Harry Harlow set up an experiment with rhesus monkeys, presocial primates, in 1958; the results from this study showed that social encounters are necessary in order for the young monkeys to develop both mentally and sexually.
A fission-fusion society is a society that changes frequently in its size and composition, making up a permanent social group called the "parent group". Permanent social networks consist of all individual members of a community and often varies to track changes in their environment. In a fission–fusion society, the main parent group can fracture (fission) into smaller stable subgroups or individuals to adapt to environmental or social circumstances. For example, a number of males may break off from the main group in order to hunt or forage for food during the day, but at night they may return to join (fusion) the primary group to share food and partake in other activities. Many mammals exhibit this, such as primates (for example orangutans and spider monkeys), elephants, spotted hyenas, lions, and dolphins.
Solitary animals defend a territory and avoid social interactions with the members of its species, except during breeding season. This is to avoid resource competition, as two individuals of the same species would occupy the same niche, and to prevent depletion of food. A solitary animal, while foraging, can also be less conspicuous to predators or prey.
In a hierarchy, individuals are either dominant or submissive. A despotic hierarchy is where one individual is dominant while the others are submissive, as in wolves and lemurs, and a pecking order is a linear ranking of individuals where there is a top individual and a bottom individual. Pecking orders may also be ranked by sex, where the lowest individual of a sex has a higher ranking than the top individual of the other sex, as in hyenas. Dominant individuals, or alphas, have a high chance of reproductive success, especially in harems where one or a few males (resident males) have exclusive breeding rights to females in a group. Non-resident males can also be accepted in harems, but some species, such as the common vampire bat (Desmodus rotundus), may be more strict.
Some mammals are perfectly monogamous, meaning that they mate for life and take no other partners (even after the original mate's death), as with wolves, Eurasian beavers, and otters. There are three types of polygamy: either one or multiple dominant males have breeding rights (polygyny), multiple males that females mate with (polyandry), or multiple males have exclusive relations with multiple females (polygynandry). It is much more common for polygynous mating to happen, which, excluding leks, are estimated to occur in up to 90% of mammals. Lek mating occurs when males congregate around females and try to attract them with various courtship displays and vocalizations, as in harbor seals.
All higher mammals (excluding monotremes) share two major adaptations for care of the young: live birth and lactation. These imply a group-wide choice of a degree of parental care. They may build nests and dig burrows to raise their young in, or feed and guard them often for a prolonged period of time. Many mammals are K-selected, and invest more time and energy into their young than do r-selected animals. When two animals mate, they both share an interest in the success of the offspring, though often to different extremes. Mammalian females exhibit some degree of maternal aggression, another example of parental care, which may be targeted against other females of the species or the young of other females; however, some mammals may "aunt" the infants of other females, and care for them. Mammalian males may play a role in child rearing, as with tenrecs, however this varies species to species, even within the same genus. For example, the males of the southern pig-tailed macaque (Macaca nemestrina) do not participate in child care, whereas the males of the Japanese macaque (M. fuscata) do.
Non-human mammals play a wide variety of roles in human culture. They are the most popular of pets, with tens of millions of dogs, cats and other animals including rabbits and mice kept by families around the world. Mammals such as mammoths, horses and deer are among the earliest subjects of art, being found in Upper Paleolithic cave paintings such as at Lascaux. Major artists such as Albrecht Dürer, George Stubbs and Edwin Landseer are known for their portraits of mammals. Many species of mammals have been hunted for sport and for food; deer and wild boar are especially popular as game animals. Mammals such as horses and dogs are widely raced for sport, often combined with betting on the outcome. There is a tension between the role of animals as companions to humans, and their existence as individuals with rights of their own. Mammals further play a wide variety of roles in literature, film, mythology, and religion.
Domestic mammals form a large part of the livestock raised for meat across the world. They include (2009) around 1.4 billion cattle, 1 billion sheep, 1 billion domestic pigs, and (1985) over 700 million rabbits. Working domestic animals including cattle and horses have been used for work and transport from the origins of agriculture, their numbers declining with the arrival of mechanised transport and agricultural machinery. In 2004 they still provided some 80% of the power for the mainly small farms in the third world, and some 20% of the world's transport, again mainly in rural areas. In mountainous regions unsuitable for wheeled vehicles, pack animals continue to transport goods. Mammal skins provide leather for shoes, clothing and upholstery. Wool from mammals including sheep, goats and alpacas has been used for centuries for clothing. Mammals serve a major role in science as experimental animals, both in fundamental biological research, such as in genetics, and in the development of new medicines, which must be tested exhaustively to demonstrate their safety. Millions of mammals, especially mice and rats, are used in experiments each year. A knockout mouse is a genetically modified mouse with an inactivated gene, replaced or disrupted with an artificial piece of DNA. They enable the study of sequenced genes whose functions are unknown. A small percentage of the mammals are non-human primates, used in research for their similarity to humans.
Charles Darwin, Jared Diamond and others have noted the importance of domesticated mammals in the Neolithic development of agriculture and of civilization, causing farmers to replace hunter-gatherers around the world.[b] This transition from hunting and gathering to herding flocks and growing crops was a major step in human history. The new agricultural economies, based on domesticated mammals, caused "radical restructuring of human societies, worldwide alterations in biodiversity, and significant changes in the Earth's landforms and its atmosphere... momentous outcomes".
Hybrids are offspring resulting from the breeding of two genetically distinct individuals, which usually will result in a high degree of heterozygosity, though hybrid and heterozygous are not synonymous. The deliberate or accidental hybridizing 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 for economic purposes. Hybrids between different subspecies within a species (such as between the Bengal tiger and Siberian tiger) are known as intra-specific hybrids. Hybrids between different species within the same genus (such as between lions and tigers) are known as interspecific hybrids or crosses. Hybrids between different genera (such as between sheep and goats) are known as intergeneric hybrids. Natural hybrids will occur in hybrid zones, where two populations of species within the same genera or species living in the same or adjacent areas will interbreed with each other. Some hybrids have been recognized as species, such as the red wolf (though this is controversial).
Artificial selection, the deliberate selective breeding of domestic animals, is being used to breed back recently extinct animals in an attempt to achieve an animal breed with a phenotype that resembles that extinct wildtype ancestor. A breeding-back (intraspecific) hybrid may be very similar to the extinct wildtype in appearance, ecological niche and to some extent genetics, but the initial gene pool of that wild type is lost forever with its extinction. As a result, bred-back breeds are at best vague look-alikes of extinct wildtypes, as Heck cattle are of the aurochs.
Purebred wild species evolved to a specific ecology can be threatened with extinction through the process of genetic pollution, the uncontrolled hybridization, introgression genetic swamping which leads to homogenization or out-competition from the heterosic hybrid species. When new populations are imported or selectively bred by people, or when habitat modification brings previously isolated species into contact, extinction in some species, especially rare varieties, is possible. Interbreeding can swamp the rarer gene pool and create hybrids, depleting the purebred gene pool. For example, the endangered wild water buffalo is most threatened with extinction by genetic pollution from the domestic water buffalo. Such extinctions are not always apparent from a morphological standpoint. Some degree of gene flow is a normal evolutionary process, nevertheless, hybridization threatens the existence of rare species.
The loss of species from ecological communities, defaunation, is primarily driven by human activity. This has resulted in empty forests, ecological communities depleted of large vertebrates. In the Quaternary extinction event, the mass die-off of megafaunal variety coincided with the appearance of humans, suggesting a human influence. One hypothesis is that humans hunted large mammals, such as the woolly mammoth, into extinction. The 2019 Global Assessment Report on Biodiversity and Ecosystem Services by IPBES states that the total biomass of wild mammals has declined by 82 percent since the beginning of human civilization.
Various species are predicted to become extinct in the near future, among them the rhinoceros, primates, pangolins, and giraffes. Hunting alone threatens hundreds of mammalian species around the world. Scientists claim that the growing demand for meat is contributing to biodiversity loss as this is a significant driver of deforestation and habitat destruction; species-rich habitats, such as significant portions of the Amazon rainforest, are being converted to agricultural land for meat production. According to the World Wildlife Fund's 2016 Living Planet Index, global wildlife populations have declined 58% since 1970, primarily due to habitat destruction, over-hunting and pollution. They project that if current trends continue, 67% of wildlife could disappear by 2020. Another influence is over-hunting and poaching, which can reduce the overall population of game animals, especially those located near villages, as in the case of peccaries. The effects of poaching can especially be seen in the ivory trade with African elephants.[來源請求] Marine mammals are at risk from entanglement from fishing gear, notably cetaceans, with discard mortalities ranging from 65,000 to 86,000 individuals annually.
Attention is being given to endangered species globally, notably through the Convention on Biological Diversity, otherwise known as the Rio Accord, which includes 189 signatory countries that are focused on identifying endangered species and habitats. Another notable conservation organization is the IUCN, which has a membership of over 1,200 governmental and non-governmental organizations.
Recent extinctions can be directly attributed to human influences. The IUCN characterizes 'recent' extinction as those that have occurred past the cut-off point of 1500, and around 80 mammal species have gone extinct since that time and 2015. Some species, such as the Père David's deer are extinct in the wild, and survive solely in captive populations. Other species, such as the Florida panther, are ecologically extinct, surviving in such low numbers that they essentially have no impact on the ecosystem.:318 Other populations are only locally extinct (extirpated), still existing elsewhere, but reduced in distribution,:75–77 as with the extinction of gray whales in the Atlantic.
- List of mammal genera – living mammals
- List of mammalogists
- List of monotremes and marsupials
- List of placental mammals
- List of prehistoric mammals
- List of threatened mammals of the United States
- Lists of mammals by population size
- Lists of mammals by region
- Mammals described in the 2000s
- Mammals in culture
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[[Category:哺乳類| ]] [[Category:Bathonian first appearances]] [[Category:Extant Middle Jurassic first appearances]] [[Category:卡尔·林奈命名的生物分类]]