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77-404: Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The process, which takes place in the front part of the digestive system and therefore is called foregut fermentation , typically requires

154-581: A clade sister to Cervidae . According to the study, Cervidae diverged from the Bovidae-Moschidae clade 27 to 28 million years ago. The following cladogram is based on the 2003 study. Tragulidae [REDACTED] Antilocapridae [REDACTED] Giraffidae [REDACTED] Cervidae [REDACTED] Bovidae [REDACTED] Moschidae [REDACTED] Infraorder Pecora ("horned ruminants", "higher ruminants") Pecorans share characteristics with other artiodactyls, including

231-615: A clade sister to Cervidae . According to the study, Cervidae diverged from the Bovidae-Moschidae clade 27 to 28 million years ago. The following cladogram is based on a large-scale genome ruminant genome sequence study from 2019: Tragulidae [REDACTED] Antilocapridae [REDACTED] Giraffidae [REDACTED] Cervidae [REDACTED] Bovidae [REDACTED] Moschidae [REDACTED] Hofmann and Stewart divided ruminants into three major categories based on their feed type and feeding habits: concentrate selectors, intermediate types, and grass/roughage eaters, with

308-445: A global warming potential of 86 compared to CO 2 over a 20-year period. As a by-product of consuming cellulose, cattle belch out methane, there-by returning that carbon sequestered by plants back into the atmosphere. After about 10 to 12 years, that methane is broken down and converted back to CO 2 . Once converted to CO 2 , plants can again perform photosynthesis and fix that carbon back into cellulose. From here, cattle can eat

385-499: A common ancestor, but instead evolved independently on at least two occasions. Likewise, while Pecora as a group is supported by both molecular and morphological studies, morphological support for interrelationships between pecoran families is disputed. The first fossil ruminants appeared in the Early Eocene and were small, likely omnivorous , forest-dwellers. Molecular dating studies estimate that Ruminantia split into

462-484: A cow. The role of saliva is to provide ample fluid for rumen fermentation and to act as a buffering agent. Rumen fermentation produces large amounts of organic acids, thus maintaining the appropriate pH of rumen fluids is a critical factor in rumen fermentation. After digesta passes through the rumen, the omasum absorbs excess fluid so that digestive enzymes and acid in the abomasum are not diluted. Tannins are phenolic compounds that are commonly found in plants. Found in

539-653: A few bumps. In contrast, camels and ruminants have bumps that are crescent-shaped cusps ( selenodont ). Pecora Pecora is an infraorder of even-toed hoofed mammals with ruminant digestion. Most members of Pecora have cranial appendages projecting from their frontal bones ; only two extant genera lack them, Hydropotes and Moschus . The name "Pecora" comes from the Latin word pecus , which means "cattle". Although most pecorans have cranial appendages, only some of these are properly called "horns", and many scientists agree that these appendages did not arise from

616-428: A four-chambered stomach, and a paraxonic foot, meaning that it supports weight on the third and fourth digits. Several characteristics distinguish Pecora from its sister taxon, Tragulina : an astragalus with parallel sides, a loss of the trapezium , and differences in parts of the skull such as the petrosal bone . The distinguishing features of most pecoran families are cranial appendages. Most modern pecorans (with

693-576: A similar aquatic lifestyle. Hippopotamuses appeared in the late Miocene and occupied Africa and Asia—they never got to the Americas. The camels ( Tylopoda ) were, during large parts of the Cenozoic , limited to North America; early forms like Cainotheriidae occupied Europe. Among the North American camels were groups like the stocky, short-legged Merycoidodontidae . They first appeared in

770-415: A simple single-chambered stomach. Being hindgut fermenters , these animals ferment cellulose in an enlarged cecum . In smaller hindgut fermenters of the order Lagomorpha (rabbits, hares, and pikas), and Caviomorph rodents ( Guinea pigs , capybaras , etc.), material from the cecum is formed into cecotropes , passed through the large intestine, expelled and subsequently reingested to absorb nutrients in

847-477: A simple stomach that digests food. Thus, they were grouped together as non-ruminants (Porcine). All other even-toed ungulates have molars with a selenodont construction (crescent-shaped cusps) and have the ability to ruminate , which requires regurgitating food and re-chewing it. Differences in stomach construction indicated that rumination evolved independently between tylopods and ruminants ; therefore, tylopods were excluded from Ruminantia . The taxonomy that

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924-474: A slim build, lanky legs, and a long tail. Their hind legs were much longer than their front legs. The early to middle Eocene saw the emergence of the ancestors of most of today's mammals. Two formerly widespread, but now extinct, families of even-toed ungulates were Entelodontidae and Anthracotheriidae . Entelodonts existed from the middle Eocene to the early Miocene in Eurasia and North America. They had

1001-473: A stocky body with short legs and a massive head, which was characterized by two humps on the lower jaw bone. Anthracotheres had a large, porcine ( pig -like) build, with short legs and an elongated muzzle . This group appeared in the middle Eocene up until the Pliocene , and spread throughout Eurasia, Africa, and North America. Anthracotheres are thought to be the ancestors of hippos, and, likewise, probably led

1078-469: A three-compartment stomach instead of four like ruminants. The Hippopotamidae (comprising hippopotamuses ) are well-known examples. Pseudoruminants, like traditional ruminants, are foregut fermentors and most ruminate or chew cud . However, their anatomy and method of digestion differs significantly from that of a four-chambered ruminant. Monogastric herbivores , such as rhinoceroses , horses , guinea pigs , and rabbits , are not ruminants, as they have

1155-424: A very flexible body, contributing to their speed by increasing their stride length. Many even-toed ungulates have a relatively large head. The skull is elongated and rather narrow; the frontal bone is enlarged near the back and displaces the parietal bone , which forms only part of the side of the cranium (especially in ruminants). Four families of even-toed ungulates have cranial appendages. These Pecora (with

1232-459: A warm, moist, anaerobic environment with a temperature range of 37.7 to 42.2 °C (99.9 to 108.0 °F) and a pH between 6.0 and 6.4. Without the help of microbes, ruminants would not be able to use nutrients from forages. The food is mixed with saliva and separates into layers of solid and liquid material. Solids clump together to form the cud or bolus . The cud is then regurgitated and chewed to completely mix it with saliva and to break down

1309-626: A wide range of climates (from tropic to arctic) and habitats (from open plains to forests). The population of domestic ruminants is greater than 3.5 billion, with cattle, sheep, and goats accounting for about 95% of the total population. Goats were domesticated in the Near East circa 8000 BC. Most other species were domesticated by 2500 BC., either in the Near East or southern Asia. Ruminating animals have various physiological features that enable them to survive in nature. One feature of ruminants

1386-570: Is a stem-based definition for Ruminantiamorpha, and is more inclusive than the crown group Ruminantia. As a crown group, Ruminantia only includes the last common ancestor of all extant (living) ruminants and their descendants (living or extinct ), whereas Ruminantiamorpha, as a stem group, also includes more basal extinct ruminant ancestors that are more closely related to living ruminants than to other members of Artiodactyla. When considering only living taxa ( neontology ), this makes Ruminantiamorpha and Ruminantia synonymous , and only Ruminantia

1463-643: Is an infraorder within the larger suborder Ruminantia , and is the sister clade to the infraorder Tragulina (of which Tragulidae is the only surviving family ). Pecora's placement within Artiodactyla can be represented in the following cladogram : Tylopoda (camels) [REDACTED]   Suina (pigs) [REDACTED]   Tragulidae (mouse deer) [REDACTED]   Pecora (horn bearers) [REDACTED]   Hippopotamidae (hippopotamuses) [REDACTED]   Cetacea (whales) [REDACTED] Current attempts to determine

1540-530: Is estimated to contain 10–50 billion bacteria and 1 million protozoa, as well as several yeasts and fungi. Since the environment inside a rumen is anaerobic , most of these microbial species are obligate or facultative anaerobes that can decompose complex plant material, such as cellulose , hemicellulose , starch , and proteins . The hydrolysis of cellulose results in sugars, which are further fermented to acetate, lactate, propionate, butyrate, carbon dioxide, and methane . As bacteria conduct fermentation in

1617-475: Is more adapted to a squeezing mastication , which is characteristic of omnivores . Camels and ruminants have fewer teeth; there is often a yawning diastema , a designated gap in the teeth where the molars are aligned for crushing plant matter. The incisors are often reduced in ruminants, and are completely absent in the upper jaw . The canines are enlarged and tusk-like in the Suina, and are used for digging in

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1694-500: Is shown to be closely related to their environment and anatomy, where light Pecorian species use both flexed and extended suspensions in their fast gallops. The white-tail and mule-deer have been observed to primarily use the extended suspension, since in this phase of their gallop they leap over bushes and logs that are present in their brush environment. However, heavy Pecorian species do not use extended suspensions as most have backs that slope downward with shorter hind legs. Pecora

1771-407: Is the large ruminal storage capacity that gives them the ability to consume feed rapidly and complete the chewing process later. This is known as rumination, which consists of the regurgitation of feed, rechewing, resalivation, and reswallowing. Rumination reduces particle size, which enhances microbial function and allows the digesta to pass more easily through the digestive tract. Vertebrates lack

1848-438: Is their continuously growing teeth. During grazing, the silica content in forage causes abrasion of the teeth. This is compensated for by continuous tooth growth throughout the ruminant's life, as opposed to humans or other nonruminants, whose teeth stop growing after a particular age. Most ruminants do not have upper incisors; instead, they have a thick dental pad to thoroughly chew plant-based food. Another feature of ruminants

1925-708: Is therefore focused on anthracotheres (family Anthracotheriidae); one dating from the Eocene to Miocene was declared to be "hippo-like" upon discovery in the 19th century. A study from 2005 showed that the anthracotheres and hippopotamuses had very similar skulls , but differed in the adaptations of their teeth. It was nevertheless believed that cetaceans and anthracothereres descended from a common ancestor, and that hippopotamuses developed from anthracotheres. A study published in 2015 confirmed this, but also revealed that hippopotamuses were derived from older anthracotherians. The newly introduced genus Epirigenys from Eastern Africa

2002-525: Is thus the sister group of hippos. Linnaeus postulated a close relationship between camels and ruminants as early as the mid-1700s. Henri de Blainville recognized the similar anatomy of the limbs of pigs and hippos, and British zoologist Richard Owen coined the term "even-toed ungulates" and the scientific name "Artiodactyla" in 1848. Internal morphology (mainly the stomach and the molars) were used for classification. Suines (including pigs ) and hippopotamuses have molars with well-developed roots and

2079-872: Is used. Thus, Ruminantiamorpha is only used in the context of paleontology . Accordingly, Spaulding grouped some genera of the extinct family Anthracotheriidae within Ruminantiamorpha (but not in Ruminantia), but placed others within Ruminantiamorpha's sister clade, Cetancodontamorpha . Ruminantia's placement within Artiodactyla can be represented in the following cladogram : Tylopoda (camels) [REDACTED]   Suina (pigs) [REDACTED]   Tragulidae (mouse deer) [REDACTED]   Pecora (horn bearers) [REDACTED]   Hippopotamidae (hippopotamuses) [REDACTED]   Cetacea (whales) [REDACTED] Within Ruminantia,

2156-455: Is very important because it provides liquid for the microbial population, recirculates nitrogen and minerals, and acts as a buffer for the rumen pH. The type of feed the animal consumes affects the amount of saliva that is produced. Though the rumen and reticulum have different names, they have very similar tissue layers and textures, making it difficult to visually separate them. They also perform similar tasks. Together, these chambers are called

2233-563: The Miocene suggests that its rapid diversification may correspond to the climate change events of that epoch, as this time period was marked by much of Earth's forest habitats being replaced by grasslands due to widespread cooling and drying. It is likely that the antelopes, giraffids, and pronghorns evolved in an open environment while the cervids, including the caribou, evolved in a woodland habitat. The type of gallop in Pecorian species

2310-694: The Old World , exist today only in the Americas . South America was settled by even-toed ungulates only in the Pliocene, after the land bridge at the Isthmus of Panama formed some three million years ago. With only the peccaries, lamoids (or llamas ), and various species of capreoline deer , South America has comparatively fewer artiodactyl families than other continents, except Australia, which has no native species. The classification of artiodactyls

2387-555: The Tragulidae (mouse deer) are considered the most basal family, with the remaining ruminants classified as belonging to the infraorder Pecora . Until the beginning of the 21st century it was understood that the family Moschidae (musk deer) was sister to Cervidae . However, a 2003 phylogenetic study by Alexandre Hassanin (of National Museum of Natural History, France ) and colleagues, based on mitochondrial and nuclear analyses, revealed that Moschidae and Bovidae form

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2464-507: The hind legs have a reduction in the number of toes to three. The central axis of the leg is between the third and fourth toe. The first toe is missing in modern artiodactyls, and can only be found in now-extinct genera. The second and fifth toes are adapted differently between species: When camels have only two toes present, the claws are transformed into nails (while both are made of keratin , claws are curved and pointed while nails are flat and dull). These claws consist of three parts:

2541-537: The order Artiodactyla ( / ˌ ɑːr t i oʊ ˈ d æ k t ɪ l ə / AR -tee-oh- DAK -tih-lə , from Ancient Greek ἄρτιος , ártios  'even' and δάκτυλος , dáktylos  'finger, toe'). Typically, they are ungulates which bear weight equally on two (an even number) of their five toes (the third and fourth, often in the form of a hoof ). The other three toes are either present, absent, vestigial , or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of

2618-412: The ruminants as the sister group of cetaceans. Subsequent studies established the close relationship between hippopotamuses and cetaceans; these studies were based on casein genes , SINEs , fibrinogen sequences, cytochrome and rRNA sequences, IRBP (and vWF ) gene sequences, adrenergic receptors , and apolipoproteins . In 2001, the fossil limbs of a Pakicetus (amphibioid cetacean

2695-409: The (one's) cud' is to reflect or meditate. In psychology, "rumination" refers to a pattern of thinking, and is unrelated to digestive physiology. Methane is produced by a type of archaea , called methanogens , as described above within the rumen, and this methane is released to the atmosphere. The rumen is the major site of methane production in ruminants. Methane is a strong greenhouse gas with

2772-399: The U.S., and 22% of the total U.S. methane emissions . The meat from domestically raised ruminants has a higher carbon equivalent footprint than other meats or vegetarian sources of protein based on a global meta-analysis of lifecycle assessment studies. Methane production by meat animals, principally ruminants, is estimated 15–20% global production of methane, unless the animals were hunted in

2849-442: The ability to hydrolyse the beta [1–4] glycosidic bond of plant cellulose due to the lack of the enzyme cellulase . Thus, ruminants completely depend on the microbial flora, present in the rumen or hindgut, to digest cellulose. Digestion of food in the rumen is primarily carried out by the rumen microflora, which contains dense populations of several species of bacteria , protozoa , sometimes yeasts and other fungi – 1 ml of rumen

2926-418: The abomasum. This is the gastric compartment of the ruminant stomach. The abomasum is the direct equivalent of the monogastric stomach, and digesta is digested here in much the same way. This compartment releases acids and enzymes that further digest the material passing through. This is also where the ruminant digests the microbes produced in the rumen. Digesta is finally moved into the small intestine , where

3003-512: The absorption of nutrients by reducing the populations of proteolytic rumen bacteria. Very high levels of tannin intake can produce toxicity that can even cause death. Animals that normally consume tannin-rich plants can develop defensive mechanisms against tannins, such as the strategic deployment of lipids and extracellular polysaccharides that have a high affinity to binding to tannins. Some ruminants (goats, deer, elk, moose) are able to consume food high in tannins (leaves, twigs, bark) due to

3080-460: The assumption that feeding habits in ruminants cause morphological differences in their digestive systems, including salivary glands, rumen size, and rumen papillae. However, Woodall found that there is little correlation between the fiber content of a ruminant's diet and morphological characteristics, meaning that the categorical divisions of ruminants by Hofmann and Stewart warrant further research. Also, some mammals are pseudoruminants , which have

3157-447: The cecotropes. The primary difference between ruminants and nonruminants is that ruminants' stomachs have four compartments: The first two chambers are the rumen and the reticulum. These two compartments make up the fermentation vat and are the major site of microbial activity. Fermentation is crucial to digestion because it breaks down complex carbohydrates, such as cellulose, and enables the animal to use them. Microbes function best in

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3234-400: The digestion and absorption of nutrients occurs. The small intestine is the main site of nutrient absorption. The surface area of the digesta is greatly increased here because of the villi that are in the small intestine. This increased surface area allows for greater nutrient absorption. Microbes produced in the reticulorumen are also digested in the small intestine. After the small intestine is

3311-462: The early Eocene (about 53 million years ago). Since these findings almost simultaneously appeared in Europe , Asia , and North America , it is very difficult to accurately determine the origin of artiodactyls. The fossils are classified as belonging to the family Diacodexeidae ; their best-known and best-preserved member is Diacodexis . These were small animals, some as small as a hare , with

3388-482: The early Miocene . Ruminantia is a crown group of ruminants within the order Artiodactyla , cladistically defined by Spaulding et al. as "the least inclusive clade that includes Bos taurus (cow) and Tragulus napu (mouse deer)". Ruminantiamorpha is a higher-level clade of artiodactyls, cladistically defined by Spaulding et al. as "Ruminantia plus all extinct taxa more closely related to extant members of Ruminantia than to any other living species." This

3465-609: The early Cenozoic ( Paleocene and Eocene ), which had hooves instead of claws on their feet. Their molars were adapted to a carnivorous diet, resembling the teeth in modern toothed whales , and, unlike other mammals, had a uniform construction. The suspected relations can be shown as follows:   Artiodactyla [REDACTED]   Mesonychia † [REDACTED]   Cetacea [REDACTED] Molecular findings and morphological indications suggest that artiodactyls, as traditionally defined, are paraphyletic with respect to cetaceans. Cetaceans are deeply nested within

3542-419: The exception of the musk deer ), have one of four types of cranial appendages: true horns, antlers , ossicones , or pronghorns . True horns have a bone core that is covered in a permanent sheath of keratin, and are found only in the bovids . Antlers are bony structures that are shed and replaced each year; they are found in deer (members of the family Cervidae ). They grow from a permanent outgrowth of

3619-624: The fermented ingesta (known as cud ) to be regurgitated and chewed again. The process of rechewing the cud to further break down plant matter and stimulate digestion is called rumination . The word "ruminant" comes from the Latin ruminare , which means "to chew over again". The roughly 200 species of ruminants include both domestic and wild species. Ruminating mammals include cattle , all domesticated and wild bovines , goats , sheep , giraffes , deer , gazelles , and antelopes . It has also been suggested that notoungulates also relied on rumination, as opposed to other atlantogenatans that rely on

3696-456: The five toes. Another difference between the two orders is that many artiodactyls (except for Suina ) digest plant cellulose in one or more stomach chambers rather than in their intestine (as perissodactyls do). Molecular biology, along with new fossil discoveries, has found that cetaceans ( whales , dolphins , and porpoises ) fall within this taxonomic branch, being most closely related to hippopotamuses . Some modern taxonomists thus apply

3773-1021: The former; the two groups together form a monophyletic taxon, for which the name Cetartiodactyla is sometimes used. Modern nomenclature divides Artiodactyla (or Cetartiodactyla) in four subordinate taxa: camelids (Tylopoda), pigs and peccaries (Suina), ruminants (Ruminantia), and hippos plus cetaceans (Whippomorpha). The presumed lineages within Artiodactyla can be represented in the following cladogram :   Tylopoda (camels) [REDACTED]   Suina (pigs) [REDACTED]   Tragulidae (mouse deer) [REDACTED]   Pecora (horn bearers) [REDACTED]   Hippopotamidae (hippopotamuses) [REDACTED]   Cetacea (whales) [REDACTED] The four summarized Artiodactyla taxa are divided into ten extant families: Although deer, musk deer, and pronghorns have traditionally been summarized as cervids (Cervioidea), molecular studies provide different—and inconsistent—results, so

3850-674: The frontal bone called the pedicle and can be branched, as in the white-tailed deer ( Odocoileus virginianus ), or palmate , as in the moose ( Alces alces ). Ossicones are permanent bone structures that fuse to the frontal or parietal bones during an animal's life and are found only in the Giraffidae . Pronghorns , while similar to horns in that they have keratinous sheaths covering permanent bone cores, are deciduous. All these cranial appendages can serve for posturing, battling for mating privilege, and for defense. In almost all cases, they are sexually dimorphic, and are often found only on

3927-401: The ground and for defense. In ruminants, the males' upper canines are enlarged and used as a weapon in certain species (mouse deer, musk deer, water deer ); species with frontal weapons are usually missing the upper canines. The lower canines of ruminants resemble the incisors, so that these animals have eight uniform teeth in the frontal part of the lower jaw . The molars of porcine have only

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4004-417: The large intestine. The major roles here are breaking down mainly fiber by fermentation with microbes, absorption of water (ions and minerals) and other fermented products, and also expelling waste. Fermentation continues in the large intestine in the same way as in the reticulorumen. Only small amounts of glucose are absorbed from dietary carbohydrates. Most dietary carbohydrates are fermented into VFAs in

4081-601: The late Eocene and developed a great diversity of species in North America. Only in the late Miocene or early Pliocene did they migrate from North America into Eurasia. The North American varieties became extinct around 10,000 years ago. Suina (including pigs ) have been around since the Eocene. In the late Eocene or the Oligocene , two families stayed in Eurasia and Africa; the peccaries , which became extinct in

4158-622: The leaf, bud, seed, root, and stem tissues, tannins are widely distributed in many different species of plants. Tannins are separated into two classes: hydrolysable tannins and condensed tannins . Depending on their concentration and nature, either class can have adverse or beneficial effects. Tannins can be beneficial, having been shown to increase milk production, wool growth, ovulation rate, and lambing percentage, as well as reducing bloat risk and reducing internal parasite burdens. Tannins can be toxic to ruminants, in that they precipitate proteins, making them unavailable for digestion, and they inhibit

4235-407: The limbs are predominantly localized, which ensures that artiodactyls often have very slender legs. A clavicle is never present, and the scapula is very agile and swings back and forth for added mobility when running. The special construction of the legs causes the legs to be unable to rotate, which allows for greater stability when running at high speeds. In addition, many smaller artiodactyls have

4312-634: The males are consistently larger and heavier than the females. In deer, only the males boast antlers, and the horns of bovines are usually small or not present in females. Male Indian antelopes have a much darker coat than females. Almost all even-toed ungulates have fur, with the exception being the nearly hairless hippopotamus. Fur varies in length and coloration depending on the habitat. Species in cooler regions can shed their coat. Camouflaged coats come in colors of yellow, gray, brown, or black tones. Even-toed ungulates bear their name because they have an even number of toes (two or four)—in some peccaries,

4389-463: The males. One exception is the species Rangifer tarandus , known as reindeer in Europe or caribou in North America, where both sexes can grow antlers yearly, though the females' antlers are typically smaller and not always present. There are two trends in terms of teeth within Artiodactyla. The Suina and hippopotamuses have a relatively large number of teeth (with some pigs having 44); their dentition

4466-452: The more typical hindgut fermentation , though this is not entirely certain. Ruminants represent the most diverse group of living ungulates . The suborder Ruminantia includes six different families: Tragulidae , Giraffidae , Antilocapridae , Cervidae , Moschidae , and Bovidae . The first fossil ruminants appeared in the Early Eocene and were small, likely omnivorous, forest-dwellers. Artiodactyls with cranial appendages first occur in

4543-421: The mouse deer, often reaches a body length of only 45 centimeters (18 in) and a weight of 1.5 kilograms (3.3 lb). The largest member, the hippopotamus, can grow up to 5 meters (16 ft) in length and weigh 4.5 metric tons (5 short tons), and the giraffe can grow to be 5.5 meters (18 ft) tall and 4.7 meters (15 ft) in body length. All even-toed ungulates display some form of sexual dimorphism :

4620-948: The name Cetartiodactyla ( / s ɪ ˌ t ɑːr t i oʊ ˈ d æ k t ɪ l ə / ) to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use " even-toed ungulates " to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature. The roughly 270 land-based even-toed ungulate species include pigs , peccaries , hippopotamuses , antelopes , deer , giraffes , camels , llamas , alpacas , sheep , goats and cattle . Many are herbivores, but suids are omnivorous, whereas cetaceans are entirely carnivorous. Artiodactyls are also known by many extinct groups such as anoplotheres , cainotheriids , merycoidodonts , entelodonts , anthracotheres , basilosaurids , and palaeomerycids . Many artiodactyls are of great dietary, economic, and cultural importance to humans. The oldest fossils of even-toed ungulates date back to

4697-460: The particle size. Smaller particle size allows for increased nutrient absorption. Fiber, especially cellulose and hemicellulose , is primarily broken down in these chambers by microbes (mostly bacteria , as well as some protozoa , fungi , and yeast ) into the three volatile fatty acids (VFAs): acetic acid , propionic acid , and butyric acid . Protein and nonstructural carbohydrate ( pectin , sugars , and starches ) are also fermented. Saliva

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4774-428: The plants and the cycle begins once again. In essence, the methane belched from cattle is not adding new carbon to the atmosphere. Rather it is part of the natural cycling of carbon through the biogenic carbon cycle . In 2010, enteric fermentation accounted for 43% of the total greenhouse gas emissions from all agricultural activity in the world, 26% of the total greenhouse gas emissions from agricultural activity in

4851-442: The plate (top and sides), the sole (bottom), and the bale (rear). In general, the claws of the forelegs are wider and blunter than those of the hind legs, and they are farther apart. Aside from camels, all even-toed ungulates put just the tip of the foremost phalanx on the ground. In even-toed ungulates, the bones of the stylopodium (upper arm or thigh bone) and zygopodiums (tibia and fibula) are usually elongated. The muscles of

4928-566: The presence in their saliva of tannin-binding proteins. The Law of Moses in the Bible allowed the eating of some mammals that had cloven hooves (i.e. members of the order Artiodactyla ) and "that chew the cud", a stipulation preserved to this day in Jewish dietary laws . The verb 'to ruminate' has been extended metaphorically to mean to ponder thoughtfully or to meditate on some topic. Similarly, ideas may be 'chewed on' or 'digested'. 'Chew

5005-413: The question of phylogenetic systematics of infraorder Pecora (the horned ruminants) for the time being, cannot be answered. Artiodactyls are generally quadrupeds . Two major body types are known: suinids and hippopotamuses are characterized by a stocky body, short legs, and a large head; camels and ruminants, though, have a more slender build and lanky legs. Size varies considerably; the smallest member,

5082-645: The relationships among pecoran families (as well as all artiodactyls) rely on molecular studies, as little consensus exists in morphological studies. Different families within Pecora are recognized as valid by different groups of scientists. Until the beginning of the 21st century it was understood that the family Moschidae (musk deer) was sister to Cervidae . However, a 2003 phylogenetic study by Alexandre Hassanin (of National Museum of Natural History, France ) and colleagues, based on mitochondrial and nuclear analyses, revealed that Moschidae and Bovidae form

5159-407: The reticulorumen. The degraded digesta, which is now in the lower liquid part of the reticulorumen, then passes into the next chamber, the omasum. This chamber controls what is able to pass into the abomasum. It keeps the particle size as small as possible in order to pass into the abomasum. The omasum also absorbs volatile fatty acids and ammonia. After this, the digesta is moved to the true stomach,

5236-538: The rumen, they consume about 10% of the carbon, 60% of the phosphorus, and 80% of the nitrogen that the ruminant ingests. To reclaim these nutrients, the ruminant then digests the bacteria in the abomasum . The enzyme lysozyme has adapted to facilitate digestion of bacteria in the ruminant abomasum. Pancreatic ribonuclease also degrades bacterial RNA in the ruminant small intestine as a source of nitrogen. During grazing, ruminants produce large amounts of saliva – estimates range from 100 to 150 litres of saliva per day for

5313-594: The rumen. The glucose needed as energy for the brain and for lactose and milk fat in milk production, as well as other uses, comes from nonsugar sources, such as the VFA propionate, glycerol, lactate, and protein. The VFA propionate is used for around 70% of the glucose and glycogen produced and protein for another 20% (50% under starvation conditions). Wild ruminants number at least 75 million and are native to all continents except Antarctica and Australia. Nearly 90% of all species are found in Eurasia and Africa. Species inhabit

5390-472: The sequence with that of other living beings—the more similar they are, the more closely they are related. Comparison of even-toed ungulate and cetaceans genetic material has shown that the closest living relatives of whales and hippopotamuses is the paraphyletic group Artiodactyla. Dan Graur and Desmond Higgins were among the first to come to this conclusion, and included a paper published in 1994. However, they did not recognize hippopotamuses and classified

5467-668: The size of a wolf) and Ichthyolestes (an early whale the size of a fox) were found in Pakistan. They were both archaeocetes ("ancient whales") from about 48 million years ago (in the Eocene). These findings showed that archaeocetes were more terrestrial than previously thought, and that the special construction of the talus (ankle bone) with a double-rolled joint surface, previously thought to be unique to even-toed ungulates, were also in early cetaceans. The mesonychians , another type of ungulate, did not show this special construction of

5544-521: The talus, and thus was concluded to not have the same ancestors as cetaceans. The oldest cetaceans date back to the early Eocene (53 million years ago), whereas the oldest known hippopotamus dates back only to the Miocene (15 million years ago). The hippopotamids are descended from the anthracotheres, a family of semiaquatic and terrestrial artiodactyls that appeared in the late Eocene, and are thought to have resembled small- or narrow-headed hippos. Research

5621-479: The traditional order Artiodactyla and infraorder Cetacea are sometimes subsumed into the more inclusive Cetartiodactyla taxon. An alternative approach is to include both land-dwelling even-toed ungulates and ocean-dwelling cetaceans in a revised Artiodactyla taxon. In the 1990s, biological systematics used not only morphology and fossils to classify organisms, but also molecular biology . Molecular biology involves sequencing an organism's DNA and RNA and comparing

5698-460: The two sister clades Pecora and Tragulina around 45 million years ago, during the Eocene . However, it was not until 15 million years later, at around 30 million years ago during the Oligocene , that the evolutionary radiation of Pecora began and the five families appeared ( Bovidae , Cervidae , Moschidae , Giraffidae , and Antilocapridae ). The appearance of many Pecoran fossils during

5775-478: The wild. The current U.S. domestic beef and dairy cattle population is around 90 million head, approximately 50% higher than the peak wild population of American bison of 60 million head in the 1700s, which primarily roamed the part of North America that now makes up the United States. Artiodactyl Cetartiodactyla Montgelard et al. 1997 Artiodactyls are placental mammals belonging to

5852-426: Was hotly debated because ocean-dwelling cetaceans evolved from land-dwelling even-toed ungulates. Some semiaquatic even-toed ungulates ( hippopotamuses ) are more closely related to ocean-dwelling cetaceans than to other even-toed ungulates. Phylogenetic classification only recognizes monophyletic taxa; that is, groups that descend from a common ancestor and include all of its descendants. To address this problem,

5929-621: Was widely accepted by the end of the 20th century was:   Suidae [REDACTED]   Hippopotamidae [REDACTED]   Tylopoda [REDACTED]   Tragulidae [REDACTED]   Pecora [REDACTED] Modern cetaceans are highly adapted sea creatures which, morphologically, have little in common with land mammals; they are similar to other marine mammals , such as seals and sea cows , due to convergent evolution . However, they evolved from originally terrestrial mammals. The most likely ancestors were long thought to be mesonychians—large, carnivorous animals from

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