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Ungulata

In traditional classification, "Ungulata" is a general term used to refer to mammals with hooves, including Perissodactyla and Artiodactyla. However, with the progress of molecular phylogenetic research, the traditional "ungulates" or "Ungulata" is not a strict monophyletic concept. In modern classification, whales (Cetacea) are included in even-toed ungulates and merged with even-toed ungulates (Artiodactyla) into "Cetartiodactyla". Therefore, when discussing ungulates today, they are usually broken down into two independent major living groups:

  1. Perissodactyla: including horses, rhinos, tapirs, etc., with an odd number of toes (the middle toe bears weight).

  2. Cetartiodactyla: integrates traditional even-toed ungulates (such as pigs, cattle, deer, camels) and cetaceans (whales, dolphins, porpoises) into a natural monophyletic group. Even-toed ungulates are characterized by an even number of toes on each foot (usually 2 or 4), while the evolution of whales shows that they are closely related to hippos.

Historically, "Ungulates" also included some extinct primitive groups (such as Callostomidae and "Obtsodontia" in a broad sense), but here we mainly focus on the two major living groups: Perissodactyla and Cetiformes.

I. Introduction to Hoofed Mammals

1. Traditional vs. Modern Classification:
Traditionally, “Ungulata” was used as a broad term to include all hoofed mammals. This encompassed horses, rhinos, tapirs (odd-toed) as well as pigs, camels, deer, cattle, and others (even-toed). However, with advances in molecular phylogenetics, it became clear that whales (Cetacea) share a close evolutionary relationship with hippopotamuses (Hippopotamidae), which are even-toed ungulates. As a result, whales have been placed within the same clade as even-toed ungulates, creating the order Cetartiodactyla. This modern grouping better reflects their evolutionary origins.

2. The Two Main Orders of “Ungulates”:

  • Perissodactyla (Odd-toed Ungulates): Horses, rhinos, and tapirs—characterized by having an odd number of toes (usually one or three) and bearing weight mainly on the middle toe.

  • Cetartiodactyla (Even-toed Ungulates + Whales): This group combines traditional Artiodactyla (even-toed ungulates like pigs, camels, deer, bovids) with Cetacea (whales, dolphins, porpoises). Even-toed ungulates typically have two or four toes, and whales are highly specialized aquatic descendants of ancient terrestrial ungulate ancestors.


II. Evolutionary History

  • In the Paleogene and Neogene periods (roughly 20–50 million years ago), hoofed mammals underwent extensive adaptive radiations, spreading across the globe.

  • Perissodactyls once included a much greater diversity, but only three families survive today. They dominated certain habitats in the past but declined due to changing climates and the rise of grasslands.

  • Artiodactyls diversified more extensively, giving rise to a broad range of large herbivores on land. Within the even-toed lineage, one group returned to the water, evolving into modern whales and dolphins. Molecular data confirm that whales are nested within the even-toed ungulates, closely related to hippos, leading to the modern concept of Cetartiodactyla.


III. Classification Tables

A. Perissodactyla (Odd-Toed Ungulates)

FamilyExample Genera/SpeciesNotes
Equidae (Horses, Zebras, Donkeys)Equus (horses, zebras, donkeys)Weight-bearing on a single central toe; adapted for running in open habitats.
Rhinocerotidae (Rhinoceroses)Rhinoceros (Indian rhino), Ceratotherium (white rhino)Thick-skinned herbivores with one or two horns, found in Africa and Asia.
Tapiridae (Tapirs)Tapirus (tapirs of Central/South America, Southeast Asia)Short, flexible proboscis; forest dwellers that feed on leaves and fruits.

B. Cetartiodactyla (Even-Toed Ungulates + Whales)

Even-Toed Ungulates (Artiodactyla)

FamilyExample Genera/SpeciesNotes
Suidae (Pigs)Sus (wild boar, domestic pig), Phacochoerus (warthog)Omnivorous, often with prominent canines; Old World distribution.
Hippopotamidae (Hippos)Hippopotamus amphibiusSemi-aquatic, closely related to whales at the molecular level; African.
Camelidae (Camels, Llamas)Camelus (camels), Lama (llamas), Vicugna (vicuñas)Adapted to arid and high-altitude environments in Africa, Asia, and South America.
Cervidae (Deer)Cervus (red deer), Rangifer (reindeer), Muntiacus (muntjacs)Males typically bear antlers (bone structures shed annually); widespread across Eurasia and the Americas.
Giraffidae (Giraffe, Okapi)Giraffa (giraffe), Okapia (okapi)African endemics, the giraffe is adapted to browsing high foliage.
Bovidae (Cattle, Sheep, Goats, Antelopes)Bos (cattle), Ovis (sheep), Capra (goats), various antelopesExtremely successful and diverse herbivores in Africa, Eurasia, and North America.

Whales (Cetacea, included in Cetartiodactyla)

FamilyExample Genera/SpeciesNotes
Balaenopteridae (Rorqual Whales)Balaenoptera (blue whale), Megaptera (humpback whale)Baleen whales filter-feeding on krill and small fish.
Balaenidae (Right Whales)Eubalaena (right whales)Slow-moving baleen whales that feed on zooplankton.
Physeteridae (Sperm Whales)Physeter (sperm whale)Deep-diving predators of squid; have a large, oil-filled head.
Delphinidae (Dolphins, Orcas)Tursiops (bottlenose dolphin), Orcinus (killer whale)Tooth-bearing whales, highly social and widespread.
Phocoenidae (Porpoises)Phocoena (harbor porpoise)Small toothed whales with shorter beaks, found in coastal waters.

(There are many other whale families; the above are representative examples.)


IV. Summary

The concept of “hoofed mammals” as a single order is outdated. Modern evolutionary and molecular studies have clarified their relationships, placing whales within the lineage of even-toed ungulates. Thus, today we speak of:

  • Perissodactyla (Odd-Toed Ungulates): Horses, rhinos, tapirs.

  • Cetartiodactyla (Even-Toed Ungulates + Cetaceans): Pigs, hippos, camels, deer, cattle, antelopes, and whales/dolphins.

This modern classification highlights the incredible evolutionary journey of some terrestrial herbivores that eventually took to the oceans and became fully aquatic cetaceans. By understanding these relationships, users can appreciate how modern taxonomy accurately reflects the deep evolutionary history and adaptation of hoofed mammals.

Carnivora Primates Rodents Chiroptera Eulipotyphla Ungulata Cetacea Lagoiformes Pholidota Proboscidea Monotremata
Capra walie

Capra walie

Capra walie

Features:Both sexes have horns, and the front surface of the male horns has very obvious ridges.

The Walia Ibex (Capra walie) has no subspecies. Walia Ibex are active or feeding only in the early morning and late evening. They usually become lethargic around midday and tend to hide under dense bushes, away from the strong midday sun and any nearby predators.Unlike most species of the genus Capr...

Capra pyrenaica

Capra pyrenaica

Iberian ibex, Iberian wild goat, Spanish goat, Iberian wild goat, Alpine wild goat

Features:The male's horns are farther apart than those of other species.

Spanish ibex (scientific name: Capra pyrenaica) is called Iberian Wild Goat, Iberian Ibex in English, Bouquetin Ibérique in French, Cabra Montesa, Cabra Montés in Spanish, Iberiensteinbock, Spanischer Steinbock in German, and Cabra-montês in Portuguese. There are 4 subspecies.Spanish ibexes exhib...

Capra nubiana

Capra nubiana

Capra nubiana

Features:There are huge and majestic horns on the head, the longest record is 147.3 cm

Nubian ibex (scientific name: Capra nubiana) English Nubian Ibex, Spanish Íbice Núbico, Arabic Wa'al, German Nubischer Steinbock, Syrischer Steinbock, no subspecies. Nubian ibex was first described as a subspecies of ibex by F. Cuvier in 1825. Synonyms of the species include Capra arabica, Cap...

Capra ibex

Capra ibex

Ibex, Alpine Ibex, Capra Ibex, Alpine Ibex

Features:They use their horns to protect themselves from predators.

Ibex (scientific name: Capra ibex) English Alpine ibex, French Bouquetin des Alpes, Spanish Íbice de los Alpes, German Alpensteinbock, Italian Stambecco alpino, no subspecies.Ibex are herbivores and feed on grass, moss, flowers, leaves and twigs. If leaves and twigs are beyond their reach, ibexes w...

Capra falconeri

Capra falconeri

Capra falconeri,spiral-horned goat

Features:The largest goat

Markhor (scientific name: Capra falconeri) is called Markhor in English, Markhor and Schraubenziege in German, and has three subspecies.Markhor ewes live in groups of up to 9, while rams generally live alone. During the breeding season, rams fight to attract the attention of ewes. The rams will lock...

Capra cylindricornis

Capra cylindricornis

Capra cylindricornis

Features:The body is stout, the legs are short, and the beard is most noticeable in winter.

Eastern Caucasian ibex (scientific name: Capra cylindricornis) English Eastern Tur, East Caucasian Tur, Spanish Tur oriental, Russian Bосточнокавказский тур, Дагестанский тур, no subspecies.The coat of male East Caucasian ibex is brown in winter, which helps to dis...

Capra caucasica

Capra caucasica

Capra caucasica

Features:The fur is chestnut, the lower abdomen is yellow, the legs are darker, and the horns are scimitar-shaped.

West Caucasian ibex (scientific name: Capra caucasica) English Western Tur, Tur, West Caucasian Tur, Spanish Tur occidental, German Westkaukasischer Steinbock, Russian Западнокавказский тур, Кубанский тур, no subspecies.The herd composition of the West Caucasian ibex...

Capra hircus

Capra hircus

Summer sheep, black sheep, domestic goat

Features:The most geographically widespread of all domestic animals

Goats (scientific name: Capra hircus) are one of the earliest domesticated livestock by humans.Modern goats are domesticated from wild goats. Archaeological excavations have shown that goats in Central and West Asia were domesticated as livestock as early as 5700 BC. Since goats belong to one of the...

Capra aegagrus

Capra aegagrus

Capra aegagrus,Persian Ibex

Features:Both sexes have distinct curved horns with notches, and males have long whiskers under their chins.

Wild goats (scientific name: Capra aegagrus) are also known as Wild Goat and Bezoar in foreign languages. There are 7 subspecies.In summer, wild goats stay high in the mountains and feed on grass and moss; in winter, they move to lower places to avoid the thick snow. They feed on herbs, shrubs and o...

Ammotragus lervia

Ammotragus lervia

Ammotragus lervia,Bearded sheep, Barbary sheep, Berber sheep

Features:Males have a sparse mane from throat to chest

The scientific name of the Barbary Sheep is Ammotragus lervia, which is also known as Aoudad, Barbary Sheep, Uaddan in English, Mouflon À Manchettes in French, Aoudad, Berberschaf, Mähnenschaf, Mähnenspringer in German. It is the only species in the genus Ammotragus and has 6 subspecies.The ram u...

Addax nasomaculatus

Addax nasomaculatus

Features:It is large in size and is named after its twisted horns.

Addax nasomaculatus (scientific name: Addax nasomaculatus) has two subspecies.Addax has a strong gregarious nature. It lives in small groups in the wild of Africa and leads a wandering life. The population includes both males and females, ranging from 5 to 20, led by older male addacs, and forages c...

Oryx leucoryx

Oryx leucoryx

Oryx leucoryx,Arabian Oryx, Arabian Oryx, Arabian Oryx

Features:It is the smallest member of the genus Oryx.

Arabian Oryx (scientific name: Oryx leucoryx) is called Arabian Oryx in foreign language. It has no subspecies and is the smallest member of the genus Oryx.Arabian Oryx live in groups, with groups of different sexes, 5-30 in each group, and groups of more than 100 have also appeared. They are genera...

Oryx gazella

Oryx gazella

Oryx gazella,Gemsbok,Gemsbok

Features:It is the largest species in the genus Oryx.

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Oryx dammah

Oryx dammah

Oryx dammah,Scimitar-horned oryx,Scimitar-horned oryx, white oryx, white oryx, scimitar-horned oryx, scimitar-horned oryx

Features:The horns are long and curve backwards like a scimitar.

Scimitar-horned oryx (scientific name: Oryx dammah) is called Scimitar-horned oryx in English. It has no subspecies. It was discovered in 1816 and named by biologist Philipp Jakob Cretzschmar in 1827.Scimitar-horned oryx are active in the early morning and evening, usually gathering in groups of mor...

Oryx beisa

Oryx beisa

East African Gemsbok, Oryx oryx, East African Gemsbok

Features:There is no black area on the hind legs, and the black markings only extend to the abdomen

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Hippotragus niger

Hippotragus niger

Hippotragus niger,Sable Antelope

Features:It is the national animal of Zimbabwe, and its subspecies, the blackbuck, is the national animal of Angola.

Sable Antelope (scientific name: Hippotragus niger) is also known as Sable Antelope in English. There are 5 subspecies.Sable Antelope populations usually have 30-75 females and young antelopes in good habitats. They are led by 5 male antelopes. The herd will be divided into several small groups duri...

Hippotragus leucophaeus

Hippotragus leucophaeus

Hippotragus leucophaeus,Bluebuck

Features:Extinct, the first large mammal to disappear in Africa

The scientific name of the bluebuck is Hippotragus leucophaeus, and its foreign name is Bluebuck. It is an extinct antelope and the first large mammal to disappear in Africa. They are close relatives of the black antelope and the black antelope, but are slightly smaller.Like the antelope and black a...

Hippotragus equinus

Hippotragus equinus

Hippotragus equinus

Features:Black markings on face

The scientific name of the Hippotragus equinus is small groups living in plains and shrub areas. It is closely related to the sable antelope (H. niger) and the extinct blue scimitar-horned antelope (H. leucophoeus). It feeds on the fresh and tender parts or leaves of plants. The favorite grasses of...

Sylvicapra grimmia

Sylvicapra grimmia

Gray duiker, Gray duiker, Gray duiker, Gray duiker

Features:Light red to dark grey, white underparts, black muzzle, nose and front legs

Common Duiker (scientific name: Sylvicapra grimmia) English Common Duiker, Bush Duiker, Grey Duiker, French Céphalophe de Grimm, Céphalophe du Cap, South African Gewone Duiker, German Kronenducker, there are 13 subspecies.Common duikers are active in the early morning, evening, and night. During t...

Cephalophus zebra

Cephalophus zebra

Cephalophus zebra

Features:Possessing one of the most unique fur coats of any mammal

The spotted duiker (scientific name: Cephalophus zebra) is called Zebra Duiker, Banded Duiker, Zebra Antelope in English, Céphalophe Zébré, Céphalophe rayé in French, Duiquero Cebrado in Spanish, and Zebraducker in German. It has no subspecies and is a species endemic to the Upper Guinea Forest...

Cephalophus weynsi

Cephalophus weynsi

Cephalophus weynsi

Features:Overall color is chestnut brown or dark red with olive tones, pale underparts, and black legs

Weyns's Duiker (scientific name: Cephalophus weynsi) English Weyns's Duiker, French Céphalpohe de Weyns, German Weynsducker, has 3 subspecies. Weyns's Duiker can be distinguished from the bay duiker and white-bellied duiker distributed in the same area because they do not have obvious d...