Tylopoda (meaning "calloused foot") is a

suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...

of terrestrial

herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...

even-toed ungulate Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). 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 t ...

s belonging to the

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

Artiodactyla Artiodactyls are placental mammals belonging to the order (biology), order Artiodactyla ( , ). 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 ...

. They are found in the wild in their

native range Species distribution, or species dispersion, is the manner in which a biological taxon is spatially arranged. The geographic limits of a particular taxon's distribution is its range, often represented as shaded areas on a map. Patterns of distr ...

s of

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

and

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

, while Australian feral camels are introduced. The group has a long fossil history in

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

and

Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...

. Tylopoda appeared during the

Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...

around 50 million years ago. Tylopoda has only one

extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...

family,

Camelidae Camelids are members of the biological family Camelidae, the only currently living family in the suborder Tylopoda. The seven extant members of this group are: dromedary camels, Bactrian camels, wild Bactrian camels, llamas, alpacas, vicuñas ...

, which includes

camel A camel (from and () from Ancient Semitic: ''gāmāl'') is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. Camels have long been domesticated and, as livestock, they provid ...

llama The llama (; or ) (''Lama glama'') is a domesticated South American camelid, widely used as a List of meat animals, meat and pack animal by Inca empire, Andean cultures since the pre-Columbian era. Llamas are social animals and live with ...

guanaco The guanaco ( ; ''Lama guanicoe'') is a camelid native to South America, closely related to the llama. Guanacos are one of two wild South American camelids; the other species is the vicuña, which lives at higher elevations. Etymology The gua ...

alpaca The alpaca (''Lama pacos'') is a species of South American camelid mammal. Traditionally, alpacas were kept in herds that grazed on the level heights of the Andes of Southern Peru, Western Bolivia, Ecuador, and Northern Chile. More recentl ...

s and

vicuña The vicuña (''Lama vicugna'') or vicuna (both , very rarely spelled ''vicugna'', Vicugna, its former genus name) is one of the two wild South American camelids, which live in the high alpine tundra, alpine areas of the Andes; the other cameli ...

s. This group was much more diverse in the past, containing a number of extinct

families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...

in addition to the ancestors of living camelids (see below). Tylopods are not

ruminants Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...

Taxonomy and systematics

Tylopoda was named by Illiger (1811) and considered

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

by Matthew (1908). It was treated as an unranked clade by Matthew (1908) and as a suborder by Carroll (1988), Ursing et al. (2000) and Whistler and Webb (2005). It was assigned to

Ruminantia Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...

by Matthew (1908); to

by Flower (1883) and Carroll (1988); to Neoselenodontia by Whistler and Webb (2005); and to

Cetartiodactyla Artiodactyls are placental mammals belonging to the order (biology), order Artiodactyla ( , ). 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 ...

by Ursing et al. (2000) and by Agnarsson and May-Collado (2008). The main problem with circumscription of Tylopoda is that the extensive fossil record of camel-like mammals has not yet been thoroughly examined from a

cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...

standpoint. Tylopoda is a highly distinctive lineage among the

artiodactyl Artiodactyls are placental mammals belonging to the order Artiodactyla ( , ). 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 t ...

s, but its exact relationships are somewhat elusive because the six living species are all closely related and can be considered "

living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...

s", the sole surviving lineage of a prehistorically wildly successful

radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...

. More recent studies suggest that tylopods are not as closely related to

ruminant 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 microb ...

s as traditionally believed, expressed in

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

form as:Spaulding, M., O'Leary, M.A. & Gatesy, J. (2009): Relationships of Cetacea (Artiodactyla) Among Mammals: Increased Taxon Sampling Alters Interpretations of Key Fossils and Character Evolution. '' PLoS ONE'' no 4(9): e7062.
article
/ref>(see e.g. Fig S10) Tylopoda are extremely conservative in their lifestyle and (like ruminants) seem to have occupied the same

ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...

since their origin over 40 million years ago. Thus, it seems that the previous assumption of a close relationship between Tylopoda and ruminants is simply because all other close relatives (whales,

pig The pig (''Sus domesticus''), also called swine (: swine) or hog, is an omnivorous, domesticated, even-toed, hoofed mammal. It is named the domestic pig when distinguishing it from other members of the genus '' Sus''. Some authorities cons ...

s etc.) are so divergent in their adaptations as to have obscured most indications of relationship, or at least those visible to

phenetic In biology, phenetics (; ), also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually with respect to Morphology (biology), morphology or other observable traits, regardless of their phylogeny or evoluti ...

analyses. However, the rather basal position that Tylopoda appears to have among the

s and relatives means that the oldest members of this lineage are still morphologically very primitive and hard to distinguish from the ancestors of related lineages. The first major modern and comprehensive analysis of the problem (in 2009) supported this; while some taxa traditionally considered Tylopoda could be confirmed to belong to this suborder (and a few refuted), the delimitation of this group is still very much disputed despite (or because of) an extensive fossil record. Agriochoerus

The taxa currently assigned (with some reliability) to Tylopoda are: Basal and ''

incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...

'' * Genus †'' Gobiohyus''? * Family † Homacodontidae Superfamily Cameloidea * Family † Oromerycidae * Family

Superfamily †

Merycoidodontoidea Merycoidodontoidea, previously known as "oreodonts" or "ruminating hogs," are an extinct Superfamily (taxonomy), superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the b ...

(=Oreodontoidea) * Family †" Agriochoeridae" (

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

) * Family † Merycoidodontidae Diacodexis pakistanensis e

Disputed Tylopoda

Several additional prehistoric (cet)artiodactyl taxa are sometimes assigned to the Tylopoda, but other authors consider them ''incertae sedis'' or basal lineages among the (Cet)artiodactyla or as more closely related to other artiodactyl groups like

s: * Family † Antiacodontidae * Family † Choeropotamidae (= Haplobunodontidae) * Family †" Diacodexeidae" (

) * Family † Leptochoeridae * Family † Anoplotheriidae * Family † Cainotheriidae * Family † Xiphodontidae Some studies have considered Protoceratidae closely related to Tylopoda, but others have considered them more closely related to the

References

External links

* * {{Taxonbar, from=Q1077169 Artiodactyls Extant Eocene first appearances de:Kamele