Tylopoda (meaning "calloused foot") is a
suborder
Order ( la, ordo) 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 ...
of
terrestrial
Terrestrial refers to things related to land or the planet Earth.
Terrestrial may also refer to:
* Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
herbivorous
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
even-toed ungulate
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
s belonging to the
order Artiodactyla
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
. 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 ...
s of
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
and
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, while
Australian feral camel
Australian feral camels are feral populations of dromedaries (''Camelus dromedarius'') in Australia. Imported from British India and Afghanistan during the 19th century for transport and construction during the colonisation of the central and ...
s are
introduced. The group has a long fossil history in
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
and
Eurasia
Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
. Tylopoda appeared during the
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
around 50 million years ago.
Tylopoda has only one
extant
Extant 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
* Extant taxon, a taxon which is not extinct, ...
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: la, camelus and grc-gre, κάμηλος (''kamēlos'') from Hebrew or Phoenician: גָמָל ''gāmāl''.) is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. C ...
s,
llama
The llama (; ) (''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 othe ...
s,
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 being the vicuña, which lives at higher elevations.
Etymology
The guanaco g ...
s,
alpaca
The alpaca (''Lama pacos'') is a species of South American camelid mammal. It is similar to, and often confused with, the llama. However, alpacas are often noticeably smaller than llamas. The two animals are closely related and can successfu ...
s and
vicuña
The vicuña (''Lama vicugna'') or vicuna (both , very rarely spelled ''vicugna'', its former genus name) is one of the two wild South American camelids, which live in the high alpine areas of the Andes, the other being the guanaco, which live ...
s. This group was much more diverse in the past, containing a number of extinct
families
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
in addition to the ancestors of living camelids (see below).
Tylopods are not
ruminants
Ruminants ( suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals 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 ...
.
Taxonomy and systematics
Tylopoda was named by Illiger (1811) and considered
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
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 ( suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals 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 ...
by Matthew (1908); to
Artiodactyla
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
by Flower (1883) and Carroll (1988); to
Neoselenodontia by Whistler and Webb (2005); and to
Cetartiodactyla
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
by Ursing et al. (2000) and by Agnarsson and May-Collado (2008).
[R. L. Carroll. 1988. Vertebrate Paleontology and Evolution. W. H. Freeman and Company, New York 1-698]
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 (; ) 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 typically shared derived char ...
standpoint. Tylopoda is a highly distinctive lineage among the
artiodactyl
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
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 an extant taxon that cosmetically 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 origin of the extant clade. Living fossi ...
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 through a material medium. This includes:
* ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...
. More recent studies suggest that tylopods are not as closely related to
ruminant
Ruminants (suborder Ruminantia) are ungulate, hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principally ...
s as traditionally believed, expressed in
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
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 resources and competitors (for ...
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''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
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 ( el, phainein – to appear) , also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary rel ...
analyses. However, the rather basal position that Tylopoda appears to have among the even-toed ungulate
The even-toed ungulates (Artiodactyla , ) are ungulates—hoofed animals—which bear weight equally on two (an even number) of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing poster ...
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.
The taxa currently assigned (with some reliability) to Tylopoda are:
Basal and ''incertae sedis
' () or ''problematica'' is a term used for a 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
Homacodontidae is an extinct family (biology), family of Basal (phylogenetics), basal Artiodactyla, artiodactyl mammals from the early Eocene to late Oligocene of North America, Europe, and Asia.
Description
They were small animals, averaging a ...
Superfamily † Anoplotherioidea
* Family †Anoplotheriidae
Anoplotheriidae is an extinct family of even-toed ungulates ( order Artiodactyla). They were endemic to Western Europe during the Eocene and Oligocene epochs about 48—23 million years ago (Mya), existing for about 25 million years. They disa ...
* Family †Cainotheriidae
Cainotheriidae is an extinct family of artiodactyls known from the Late Eocene to Middle Miocene of Europe. They are mostly found preserved in karstic deposits.
These animals were small in size, and generally did not exceed in height at the ...
* Family † Dacrytheriidae
Superfamily Cameloidea
* Family †Oromerycidae
Oromerycidae is a small (both in size and diversity), extinct family (biology), family of artiodactyls (even-toed hoofed mammals) closely related to living camels, known from the early to late Eocene of western North America.
Oromerycids are pl ...
* 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, ...
Superfamily †Merycoidodontoidea
Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the better known forms were gen ...
(=Oreodontoidea)
* Family †" Agriochoeridae" (paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
)
* Family †Merycoidodontidae
Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the better known forms were gen ...
Superfamily † Xiphodontoidea
* Family †Xiphodontidae
Xiphodontidae is an extinct family of even-toed ungulates (order Artiodactyla), endemic to Europe during the Eocene 40.4—33.9 million years ago, existing for about 7.5 million years. They were, most likely, all terrestrial herbivore ...
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:
* Family † Antiacodontidae
* Family †Choeropotamidae
Choeropotamidae, also known as Haplobunodontidae, are a family of extinct mammals, extinct herbivores, belonging to artiodactyls. They lived between the lower/middle Eocene and lower Oligocene (about 48 - 30 million years ago) and their remai ...
(= Haplobunodontidae)
* Family †" Diacodexeidae" (paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
)
* Family † Leptochoeridae
The Cebochoeridae and Protoceratidae
Protoceratidae is an extinct family of herbivorous North American artiodactyls (even-toed ungulates) that lived during the Eocene through Pliocene at around 46.2—4.9 Mya, existing for about 41 million years.
Classification
Protoceratidae was ...
, on the other hand, are prehistoric families that were occasionally placed within Tylopoda in the past, but are now considered more closely related to cetaceans and ruminant
Ruminants (suborder Ruminantia) are ungulate, hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principally ...
s respectively than to camels.
References
External links
*
*
{{Taxonbar, from=Q1077169
Even-toed ungulates
Extant Eocene first appearances
de:Kamele