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Pachydermata
Pachydermata
Pachydermata
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Animals described as pachyderms, clockwise from top left: a hippopotamus, an Indian rhinoceros, a Malayan tapir, and an African bush elephant

Pachydermata (meaning 'thick skin', from the Greek παχύς, pachys, 'thick', and δέρμα, derma, 'skin') is an obsolete order of mammals described by Gottlieb Storr, Georges Cuvier, and others, at one time recognized by many systematists. The term pachyderm is commonly used to describe elephants, rhinoceroses, hippopotamuses and tapirs. The grouping was determined to be artificial as a biological classification due to genetic studies.

Description

[edit]

Pachydermata is an obsolete order of mammals described by Gottlieb Storr, Georges Cuvier, and others, at one time recognized by many systematists. The grouping is polyphyletic, so the order is no longer in use as a biological classification.[when?] Outside strict biological classification, the related term pachyderm is commonly used to describe elephants, rhinoceroses, hippopotamuses and tapirs.

Cuvier himself defined Pachydermata as "animals with hoofs, non ruminants", whereas Storr had described it as "mammals with hoofs with more than two toes". The classification Pachydermata included the three families of mammals he called Proboscidiana, Pachydermata Ordinaria, and Solipedes, which are all herbivorous.[1] The members of the grouping are now divided into the Proboscidea (represented among living species only by three species of elephants), the Perissodactyla (odd-toed ungulates, including horses, tapirs, and rhinoceroses), the Suina (pigs and peccaries), the Hippopotamidae, Hyracoidea (hyraxes) and Sirenians (Manatee and dugong).

The term is important in the history of systematics. Although the former order of Pachydermata is often described as an artificial grouping of unrelated mammals, it was recognised by notable zoologists, including Charles Darwin, as a grade of hoofed mammals to the exception of other ungulates; and anatomical characters support the affinities of "pachyderm" mammals to each other and to other ungulates.

History

[edit]

In a series of research papers published in 1796, Georges Cuvier created the classification Pachydermata.[1] Cuvier added horses to the order.[2] One naturalist, Delabere Pritchett Blaine, has speculated that:

Baron Cuvier, it is probable, was led to arrange the horse genus among the Pachydermata, less on account of the thickness and tenacity of the skin, than on the slight departure from a true monodactylous character, which every member of this family exhibits in having vestiges of two additional toes under the skin.

— Delabere Pritchett Blaine, An Encyclopaedia of Rural Sports, Vol. 1, page 240

According to genetic studies, elephants, rhinoceroses, tapirs and hippopotamuses are classified as separate clades altogether. Rhinos, hippos, pigs, peccaries, horses, zebras, donkeys and tapirs are classified in clade Laurasiatheria, while elephants, hyraxes, manatees and dugongs are classified in clade Afrotheria.

References

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Pachydermata

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Pachydermata is an obsolete taxonomic order within the class Mammalia, first proposed by Gottlieb Storr in 1780 and formalized in 1796 by to group large, thick-skinned mammals characterized by hoofed feet and non-ruminant digestion, including elephants, rhinoceroses, hippopotamuses, tapirs, pigs, peccaries, and . The term derives from the Greek words pachys (thick) and derma (skin), reflecting the superficial morphological trait of thickened hide that unified these animals in early classifications. Historically, the concept traces back to Carl Linnaeus's (1758), where similar ungulates were loosely categorized, but Cuvier formalized Pachydermata in a series of 1796 papers as "hoofed, non-ruminant" mammals, later expanding it to encompass and even marine forms like walruses and dugongs. This grouping persisted into the 19th century, with naturalists like refining ungulate taxonomy by establishing orders such as (odd-toed ungulates) in 1848, which separated rhinos, tapirs, and from even-toed like hippos and pigs. By the early , works like Richard Lydekker's surveys (1913–1915) still referenced Pachydermata as a broad category based on traits like enlarged toenails, but its artificial nature became evident. Pachydermata fell out of use due to its polyphyletic composition, meaning the included taxa do not share a common ancestor exclusive to the group, as revealed by advances in comparative anatomy, paleontology, and molecular phylogenetics. Modern classifications reassign its members: elephants and hyraxes to the clade Afrotheria (Proboscidea and Hyracoidea, respectively), rhinos, tapirs, and horses to Perissodactyla, and hippos, pigs, and peccaries to Artiodactyla (now Cetartiodactyla, incorporating whales). This shift, detailed in comprehensive reviews like Groves and Grubb's Ungulate Taxonomy (2011), emphasizes monophyletic groups based on evolutionary relationships rather than convergent traits like skin thickness. Today, "pachyderm" persists colloquially for elephants and rhinos but holds no systematic validity in zoology.

Etymology and Definition

Origin of the Term

The term "Pachydermata" originates from the Greek roots pachys (παχύς), meaning "thick," and derma (δέρμα), meaning "skin," literally translating to "thick-skinned." French naturalist Georges Cuvier introduced the singular form "pachyderme" in 1797 as a biological descriptor for certain non-ruminant hoofed mammals, emphasizing their shared anatomical trait of dense, often hairless skin. This initial usage appeared in Cuvier's Tableau élémentaire de l'histoire naturelle des animaux (1797–1798), where the term served to categorize animals based on this prominent physical characteristic in comparative anatomy. Subsequently, "pachyderme" transitioned into the plural Latinized form "Pachydermata" as a formal taxonomic order within the developing framework of zoological classification in the early 19th century.

Original Characteristics

The order Pachydermata, as originally conceived by , grouped non-ruminant ungulates—hoofed mammals with simple digestive systems lacking the multi-chambered stomachs of ruminants—based on shared traits including thick, leathery skin as a protective adaptation, quadrupedal locomotion with pillar-like limbs and hoofed feet for weight support, and herbivorous diets processed via or similar mechanisms. Body sizes varied widely across members, from small forms like to massive species such as and rhinoceroses. was generally adapted for grinding fibrous , with large molars, though specifics differed (e.g., prominent tusks in ). Some exhibited semi-aquatic adaptations, like webbed toes in hippopotamuses.

Historical Classification

Cuvier's Proposal

and introduced the term Pachydermata in 1795, which Cuvier formalized in a series of papers and detailed as a taxonomic order in 1798 as part of his foundational classification system for mammals, in Tableau élémentaire de l'histoire naturelle des animaux. This served as preparatory groundwork for his later comprehensive work, Le Règne Animal (), and proposed a quadripartite division of mammals into four orders: (encompassing carnivorous forms), (including and related gnawing animals), Cete (cetaceans), and Pachydermata. Cuvier's system marked a shift toward a more anatomically grounded , integrating evidence with living to establish natural groupings based on organizational principles rather than arbitrary divisions. At the core of Cuvier's proposal, Pachydermata was defined as a cohesive order of thick-skinned, hoofed mammals characterized by robust skeletal structures and non-ruminant digestive systems, distinguishing them from other s. He initially highlighted (Elephas), (Rhinoceros), and (Hippopotamus) as archetypal members, emphasizing their shared integumentary features—such as tough, nearly hairless skin—and skeletal adaptations for weight-bearing and terrestrial locomotion. These animals were grouped together due to their uniform possession of thick cuticles and heavy, columnar limb bones, which Cuvier viewed as correlated traits supporting a semi-aquatic or heavily armored lifestyle in and , and amphibious habits in . By excluding ruminants, despite superficial similarities in ungulate foot structure, Cuvier underscored that Pachydermata lacked the multi-chambered stomachs and cud-chewing behavior typical of ruminants, thereby rejecting any close affinity based on those traits. Cuvier's formulation of Pachydermata drew heavily from his pioneering work in , prioritizing functional correlations among organ systems—such as the integration of skin, skeleton, and dentition for survival—over the external morphological criteria emphasized in . This approach allowed him to infer whole-animal adaptations from partial remains, including fossils, and positioned Pachydermata as a functionally coherent assemblage adapted to environments demanding physical durability over agility or predation. His emphasis on these adaptive interdependencies influenced subsequent naturalists, establishing as a cornerstone of classification.

19th-Century Revisions

In the early decades of the , naturalists revised Georges Cuvier's original Pachydermata order to accommodate new anatomical insights and discoveries, shifting emphasis from skin thickness to skeletal and dental features. , in his 1821 classification of vertebrose animals, subdivided Pachydermata into two primary groups: Bruta, which included and their allies characterized by trunk-like appendages and reduced , and Pachydermes proper, encompassing rhinoceroses, hippopotamuses, and swine-like forms with more robust, hoofed limbs. This subdivision aimed to highlight distinctions in locomotion and feeding adaptations within the thick-skinned mammals. By the mid-19th century, refined the classification in his 1848 work on mammalian classification by establishing the order for odd-toed ungulates, including (Equidae), tapirs (Tapiridae), and rhinoceroses based on their shared odd-toed (perissodactylous) foot structure and grinding , thus separating them from even-toed like hippos and pigs. These changes sparked debates among systematists over boundary definitions, with some proposals including armadillos () due to armored skin or (Hyracoidea) for superficial resemblances to small pachyderms, though such inclusions were contested for lacking consistent anatomical ties. The evolving focus on hoof morphology over cutaneous traits marked a conceptual shift toward functional adaptations for terrestrial locomotion, influencing subsequent classifications. The influence of Charles Darwin's evolutionary theory, published in 1859, intensified scrutiny of Pachydermata's coherence, as naturalists questioned whether the order represented true phylogenetic relationships or merely convergent forms adapted to similar environments. Expansions occurred primarily between the 1820s and 1850s, but by the 1860s, the group fragmented into specialized suborders, such as for proboscideans and Solidungula for solid-hoofed forms, reflecting growing specialization in .

Taxonomic Composition

Included Orders and Families

The primary groups included within Pachydermata across historical classifications encompassed the order , the family Rhinocerotidae, the family , the family Tapiridae, the families and Tayassuidae, and the family Procaviidae (), unified by their thick skins, hoofed feet, and non-ruminant digestion. encompassed modern elephants (family Elephantidae) along with numerous extinct relatives, such as mammoths (Mammuthus), mastodons (family ), and deinotheres (family ), characterized by elongated trunks for manipulation and ivory tusks for foraging and combat. These forms ranged from the to the Pleistocene, with mastodons featuring low-crowned molars suited to in forested environments, while deinotheres possessed downward-curving lower tusks adapted for stripping bark. Rhinocerotidae included the various of rhinoceroses, distinguished by one or two prominent nasal horns formed from compacted and odd-toed (perissodactyl) hooves that supported their massive, tank-like bodies weighing up to 2,300 kg in some . Historical taxonomies placed both extant African and Asian rhinoceroses within this family as quintessential pachyderms due to their armored skin and solitary habits. Hippopotamidae comprised the hippopotamuses, semi-aquatic mammals with barrel-shaped torsos, wide mouths featuring large canines, and skin up to 5 cm thick that secretes a oil for protection in aquatic habitats; the common hippopotamus (Hippopotamus amphibius) and (Choeropsis liberiensis) were the principal members. Tapiridae included tapirs, odd-toed ungulates with elongated, flexible snouts for foraging in dense forests and wetlands, featuring short, thick legs and skin folds that provided some protection; species like the (Tapirus indicus) were viewed as linking rhinos and more primitive forms. Suidae and Tayassuidae encompassed pigs and peccaries, even-toed ungulates with bristly coats, mobile snouts, and omnivorous to herbivorous diets; pigs (e.g., Sus scrofa) and peccaries (e.g., Pecari tajacu) were included for their robust builds and non-ruminant digestion, though later separated due to foot structure differences. Procaviidae included hyraxes, small, rodent-like mammals with rubbery feet, short tails, and leathery skin resembling that of elephants, leading to their early placement as primitive pachyderms despite closer relations to elephants in modern phylogeny. Extinct inclusions extended to palaeotheres (family Palaeotheriidae), primitive tapir-like perissodactyls from the Eocene epoch, which were grouped under the Solipedes division of Pachydermata for their single-toed feet and inferred thick-skinned anatomy, bridging early horse and rhino lineages.

Variations Across Classifications

Throughout the 19th century, the taxonomic group Pachydermata exhibited significant variations in composition as naturalists refined classifications based on anatomical, ecological, and evidence. Originally proposed by in the late as a broad category for non-ruminant ungulates with thick skin, Pachydermata encompassed elephants (), rhinoceroses (Rhinocerotidae), hippopotamuses (), tapirs (Tapiridae), pigs and peccaries ( and Tayassuidae), and (Procaviidae). Subsequent systems built on this foundation, with some classifications emphasizing dental and skeletal similarities. Further variations arose with the occasional addition of (horses) by figures like , who briefly aligned them with Pachydermata based on foot structure and resemblances to early perissodactyls, despite their adaptations. (Procaviidae) were included in core and peripheral classifications owing to their tough, leathery hides akin to elephants, though these placements were debated and not always central. Meanwhile, exclusions intensified for (pigs), which were progressively removed from core Pachydermata in mid-century revisions due to their even-toed feet and omnivorous digestion contrasting with the odd-toed, herbivorous traits of rhinos and tapirs; this shift reflected emerging distinctions between perissodactyls and . Debates also surrounded sirenians (manatees and dugongs, order ), with some 19th-century naturalists proposing affinities to Pachydermata based on aquatic lifestyles and blunt, herbivorous forms, interpreting them as "pachydermous" sea mammals derived from terrestrial ungulates. Specific schemes underscored these inconsistencies: Henri-Marie Ducrotay de Blainville's 1839 classification expanded Pachydermata to include hippopotamuses, pigs, tapirs, , rhinoceroses, and , emphasizing integumentary and locomotor traits over digestive uniformity. In contrast, some works broadened it further to encompass all non- ungulates, incorporating equids and sirenians while questioning ruminant exclusions, reflecting a trend toward inclusive, phenotype-driven amid discoveries. These fluctuations illustrate how Pachydermata served as a flexible construct in early mammalian , prone to revision as evolved.

Modern Perspectives

Reasons for Obsolescence

The obsolescence of Pachydermata as a taxonomic order stemmed primarily from its nature, wherein the included groups—such as , rhinoceroses, and hippopotamuses—did not share a recent common ancestor exclusive to them, rendering the grouping artificial and non-reflective of evolutionary history. This arose because the classification relied on superficial morphological traits, notably the thickness and wrinkling of the skin, which represented convergent adaptations to similar ecological niches rather than shared phylogenetic origins. The rise of in the mid-20th century further underscored this flaw, as it prioritized monophyletic clades defined by shared derived characteristics and evolutionary branching patterns over Linnaean hierarchies based on overall similarity. A pivotal development in abandoning Pachydermata was George Gaylord Simpson's 1945 reclassification of mammals, which fragmented ungulates into distinct orders like (elephants), (odd-toed ungulates including rhinoceroses), and Artiodactyla (even-toed ungulates including hippopotamuses), emphasizing phylogenetic relationships derived from fossil evidence over Cuvierian morphology. Simpson explicitly critiqued artificial groupings like Pachydermata for failing to align with ary principles, advocating instead for a system grounded in the modern synthesis of Darwinian and . The shift from descriptive morphology to evolutionary and phylogenetic , beginning in the and accelerating with the modern synthesis in –, ultimately rendered Pachydermata untenable, with molecular evidence in the late providing further confirmation. Molecular evidence from the 1990s and 2000s provided definitive confirmation of Pachydermata's , revealing distant relationships among its former members; for instance, hippopotamuses were shown to be more closely related to cetaceans (whales and dolphins) than to or rhinoceroses, forming the monophyletic Whippomorpha within Artiodactyla. Concurrently, genomic analyses established and within the superordinal , a grouping evolutionarily remote from artiodactyls like hippopotamuses, thus highlighting the of thick skin as a misleading classificatory trait. These findings, building on cladistic principles, solidified the rejection of Pachydermata by demonstrating that its constituents diverged phylogenetically over 60 million years ago.

Current Taxonomic Placement

In contemporary mammalian taxonomy, the taxa historically united under Pachydermata are dispersed across distinct phylogenetic lineages within the subclass of the class Mammalia. The order , encompassing elephants, stands as an independent order within the superorder , a primarily originating from African lineages and supported by molecular evidence. The sole extant family, , includes two genera: Loxodonta (African elephants, with savanna and forest species) and Elephas (). Also within , the order Hyracoidea includes in the family Procaviidae, comprising three genera (, Heterohyrax, Dendrohyrax) and 11 species distributed in and the . The odd-toed ungulates formerly included in Pachydermata are classified in the order , which falls under the superorder —a diverse group encompassing many Eurasian-derived mammals. This order features the family Rhinocerotidae, comprising five extant species in four genera (two African and three Asian), all characterized by their single or double nasal horns and critically endangered status due to and habitat loss. The family Tapiridae includes four species of the genus , distributed across Central and as well as , noted for their prehensile snouts and semi-aquatic habits. The family Equidae includes horses and relatives, with seven extant species in the single genus Equus. Even-toed ungulates from the historical Pachydermata, such as hippopotamuses, are placed in the order Artiodactyla, also within . The family contains two extant species: the common (Hippopotamus amphibius), widespread in sub-Saharan African rivers, and the (Choeropsis liberiensis), restricted to West African forests. These hippopotamids form a close to cetaceans (whales and dolphins), together comprising the , a relationship established through molecular and morphological analyses. Other even-toed ungulates formerly included, such as pigs and peccaries, are classified in the infraorder within Artiodactyla. The family (pigs) includes about 18 species across six genera, primarily in and , while the family Tayassuidae (peccaries) includes three species in three genera, native to the . Later expansions of Pachydermata included marine forms like walruses (family , order , superorder ) and dugongs (family , order , superorder ). Collectively, these lineages—spanning and —lack a shared monophyletic beyond , underscoring the polyphyletic composition of the obsolete Pachydermata grouping in modern .

References

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  23. https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=91561
  24. https://animaldiversity.org/accounts/Rhinocerotidae/
  25. https://animaldiversity.org/accounts/Tapiridae/
  26. https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=9793
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  29. https://ufdcimages.uflib.ufl.edu/AA/00/03/75/36/00001/toxodontiaofsall00shoc.pdf
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