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Hipparion
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Hipparion
Temporal range: Late Miocene 9.9–5.3 Ma
↓
Skeleton on display at the National Natural History Museum of China
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Perissodactyla
Family: Equidae
Subfamily: Equinae
Tribe: †Hipparionini
Genus: †Hipparion
De Christol, 1832
Species

See text

Synonyms
  • Hemihipparion

Hipparion is an extinct genus of three-toed, medium-sized equine belonging to the extinct tribe Hipparionini, which lived about 10-5 million years ago.[1][2] While the genus formerly included most hipparionines, the genus is now more narrowly defined as hipparionines from Eurasia spanning the Late Miocene.[2] Hipparion was a mixed-feeder who ate mostly grass, and lived in the savannah biome.[2][3] Hipparion evolved from Cormohipparion,[2] and went extinct due to environmental changes like cooling climates and decreasing atmospheric carbon dioxide levels.[4]

Taxonomy

[edit]

"Hipparion" in sensu lato

[edit]

The genus "Hipparion" was used for over a century as a form classification to describe over a hundred species of Holartic hipparionines from the Pliocene and Miocene eras that had three toes and isolated protocones. Since then, groups such as the genera Cormohipparion and Neohipparion were proposed to further sort these species, typically based on differences in skull morphology. These species are now known as "Hipparion" in sensu lato (s.l.), or a broad sense.[5]

Hipparion in sensu stricto

[edit]

Hipparion in sensu stricto (s.s.), or a strict sense, describes the genus of Old World hipparionines from remains found in Eurasia (France, Greece, Turkey, Iran, and China) from the Late Miocene era (~10-5 Ma, or million years ago). The assignment of remains from elsewhere to the genus, such as North America and Africa, is uncertain.[2]

Morphology

[edit]
Life reconstruction of H. forcei
Skull of H. gracile against a black background
H. gracile skull
Fossil of three-toed Hipparion lower hind leg, held up on a stand and against a brown cloth background
Three-toed Hipparion hind leg fossil

Hipparion generally resembled a smaller version of the modern horse, but was tridactyl, or three-toed. It had two vestigial outer toes on each limb in addition to its hoof.[2] In some species, these outer toes were functional.[6] Hipparion was typically medium in size, at about 1.4 m (4.6 ft) tall at the shoulder.[7][8] The estimated body mass of Hipparion depends on the species, but ranges from about 135 to 200 kg (about 298 to 441 lbs).[2] Hipparion had hypsodont dentition (high-crowned teeth) for its premolars and molars, with a crown height of about 60 mm (2.36 in). Hipparion had isolated protocones in the upper molars, meaning a cusp of the teeth called a protocone was not connected to a tooth crest called a protoloph.[2] Hipparion is also characterized by its facial fossa, or deep depression in the skull, located high on the head in front of the orbit.[9][10]

Paleobiology and paleoecology

[edit]
Mauricio Antón's illustration of Hipparion cornelianum that resembles a modern-day zebra

Hipparion lived in the Old World Savannah Biome, or OWSB, consisting of woodlands to grasslands.[2] Hipparion ate a mixed-feed diet, mostly consisting of grass. This diet is indicated by fossil evidence of microscopic wear patterns of scratches and pits on the enamel of Hipparion's teeth, observed using scanning electron microscopy (SEM).[3]

Hipparion achieved skeletal maturity and possibly sexual maturity at about 3 years old. Fossils of Hipparion individuals are up to 10 years old at death.[8]

Isotopic analysis indicates that in the late Miocene Batallones 3 fossil site in Spain, the sabertooth cats Promegantereon and Machairodus, the amphicyonids (bear-dogs) Magericyon and Thaumastocyon, the large mustelid Eomellivora and possibly the early omnivorous giant panda relativa Indarctos likely considerably predated upon Hipparion.[11]

Evolution and extinction

[edit]

Hipparion likely evolved from a species of Cormohipparion during the Late Miocene, about 11.4–11.0 Ma. This species, C. occidentale, came to Eurasia and Africa from North America.[2] The last common ancestor of Hipparion and the modern horse was Merychippus.[12]

In the Old World, Hipparion experienced population decline and extinction down a North to South gradient, as did many other Miocene vertebrates. This trend is believed to be due to environmental changes caused by global cooling and decreasing carbon dioxide levels in the atmosphere.[4]

Species

[edit]
  • H. chiai Liu et al., 1978
  • H. concudense Pirlot, 1956
  • H. condoni Merrian, 1915
  • H. crassum Gervais, 1859
  • H. dietrichi Wehrli, 1941
  • H. fissurae Crusafont and Sondaar, 1971
  • H. forcei Richey, 1948
  • H. gromovae Villalta and Crusafont, 1957
  • H. laromae Pesquero et al., 2006
  • H. longipes Gromova, 1952
  • H. lufengense Sun, 2013
  • H. macedonicum Koufos, 1984
  • H. matthewi Abel, 1926
  • H. mediterraneum Roth and Wagner, 1855
  • H. molayanense Zouhri, 1992
  • H. minus Pavlow, 1890
  • H. periafricanum Villalta and Crusafont, 1957
  • H. philippus Koufos & Vlachou, 2016
  • H. phlegrae Lazaridis and Tsoukala, 2014
  • H. prostylum Gervais, 1849 (type)
  • H. rocinantis Pacheco, 1921
  • H. sellardsi Matthew and Stirton, 1930
  • H. shirleyae MacFadden, 1984
  • H. sithonis Koufos & Vlachou, 2016
  • H. sitifense Pomel, 1897
  • H. tchicoicum Ivanjev, 1966[13]
  • H. tehonense (Merriam, 1916)
  • H. theniusi Melentis, 1969
  • H. venustum Leidy, 1860

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Hipparion

View on Grokipedia
from Grokipedia
Hipparion is an extinct genus of three-toed horses belonging to the family Equidae, subfamily Equinae, and tribe Hipparionini, which flourished during the late Miocene through the Pliocene and into the early Pleistocene epochs approximately 11 to 1 million years ago. The genus is characterized by medium body sizes, hypsodont (high-crowned) teeth adapted for grazing on abrasive vegetation, and a tridactyl (three-toed) foot with reduced lateral toes, representing an intermediate stage in equid evolution between earlier browsers and later single-toed grazers. Originating in North America, early hipparionins like Cormohipparion migrated to the Old World via the Bering land bridge around 11.4 million years ago, establishing the "Hipparion Datum" that marks a significant faunal turnover in Eurasian mammal communities. Following their dispersal, Hipparion and related genera rapidly diversified across , , and the , peaking in during the Turolian stage (8.9–5.3 million years ago) with lineages adapting to varied environments through differences in body mass (from under 100 kg to over 300 kg) and tooth morphology. Notable species include Hippotherium primigenium, considered one of the earliest and most primitive in , and Hipparion concudense, known from Spanish sites where bone histology reveals slow growth rates and growth patterns similar to modern equids but with a shorter lifespan (up to about 10 years). This diversification reflected adaptations to increasingly seasonal grasslands, but hipparionins failed to evolve traits for cold Pleistocene climates, leading to their decline at the Miocene-Pliocene boundary around 5.3 million years ago and eventual by about 1 million years ago in surviving pockets in and . Paleontologically, Hipparion fossils are crucial biostratigraphic markers, defining the "Hipparion faunas" of late deposits and providing insights into global climatic shifts, such as the expansion of C4 grasslands and mammalian dispersals. Although polyphyletic in modern —with genera like Neohipparion, Nannippus, and Cormohipparion distinguished from Hipparion sensu stricto—the group exemplifies the of equids before the dominance of the single-toed Equus lineage.

Taxonomy

Historical classification

The genus Hipparion was originally established by French paleontologist in 1832, based on dental remains of the type species H. prostylum from the deposits of Mont Léberon in , where it was recognized as a three-toed equid with cheek teeth and an isolated protocone, distinguishing it from contemporary relatives. Early studies emphasized its tridactyl foot structure, positioning it as an intermediate form between earlier three-toed horses and later monodactyl equines. The discovery of abundant Hipparion fossils at Pikermi, , starting in 1836, rapidly expanded knowledge of the , with material from this site providing the bulk of early European specimens and reinforcing its status as a widespread grazer. In the early , American paleontologists William Diller Matthew and significantly broadened the scope of Hipparion, incorporating numerous Eurasian under a sensu lato concept that treated it as a catch-all for three-toed equids with similar dental features across Holarctic faunas. 's comprehensive assembly of collections at the , detailed in his , highlighted Hipparion's morphological diversity and proposed it as a key evolutionary link in equid phylogeny, while Matthew's analysis outlined a linear progression of that lumped many forms under the . This expansive approach included from , , and , reflecting limited understanding of intercontinental dispersal and convergence at the time. Key debates in historical classification centered on the inclusion of North American forms within Hipparion, initially viewed as conspecific with Eurasian ones due to shared traits like the preorbital fossa and tridactyl limbs, but later questioned as evidence mounted for separate evolutionary origins. For instance, early workers like Gidley in 1907 distinguished Hipparion from related genera such as Protohippus, yet North American specimens were routinely assigned to Hipparion until the mid-20th century recognition of distinct genera like Neohipparion by Stirton in 1940, which highlighted regional endemism and migration patterns. Over time, the genus concept evolved from this broad "form genus" encompassing polyphyletic assemblages—used for over a century to classify Mio-Pliocene three-toed horses—to more restricted definitions emphasizing clade-specific synapomorphies, as pioneered in revisions by Woodburne and Bernor in 1980 that segregated Old World lineages.

Modern taxonomy

In modern taxonomy, Hipparion sensu stricto is narrowly defined as a genus restricted to a small number of Eurasian species from the late Miocene, primarily characterized by a shallow, poorly defined anterior dorsal preorbital fossa and specific dental features such as moderately hypsodont cheek teeth with isolated protocones. This concept, originating from revisions in the late 20th century, limits the genus to about five species, including the type species H. prostylum from the Turolian (late Miocene) locality of Mont-Léberon in southern France. In contrast, Hipparion sensu lato represents a historically broader, polyphyletic assemblage of three-toed equids that dispersed widely during the Miocene, encompassing diverse forms now reclassified into separate genera based on cranial, dental, and postcranial distinctions; examples include Cormohipparion (ancestral immigrant forms with robust builds), Neohipparion (North American-derived species with variable preorbital fossae), and Pseudhipparion (characterized by slender limbs and mixed browsing-grazing adaptations). Phylogenetically, Hipparion sensu stricto belongs to the tribe Hipparionini within the subfamily of , forming part of the hipparionine radiation that originated in and migrated to the around 11.4 million years ago. Cladistic analyses from the onward position Hipparionini as a sister to the tribe , which gave rise to modern monodactyl horses in the genus Equus, with shared synapomorphies including dentition and tridactyl feet in the former. This placement is supported by integrative studies combining cranial morphology, dental metrics, and stratigraphic data, highlighting a single dispersal event via an early Cormohipparion as the stem for Old World diversification. Recent taxonomic revisions, particularly from 2021 to 2023, have further refined these boundaries using advanced morphometric techniques such as principal component analysis on dental and cranial measurements, alongside cladistic parsimony methods, to address the longstanding polyphyly of Hipparion sensu lato. For instance, a comprehensive 2021 synthesis recognized up to 10 Old World hipparionine genera and 63 species, elevating forms like Cremohipparion and Plesiohipparion based on quantitative traits such as metaconule size and preorbital fossa depth, while confirming H. prostylum as a valid Eurasian endemic. A 2023 review of Plio-Pleistocene western Eurasian hipparions reclassified late-surviving species—such as H. crassum into Proboscidipparion and H. rocinantis into Plesiohipparion—emphasizing regional endemism and evolutionary transitions toward extinction, without invoking molecular data due to the fossil nature of the record. As of 2025, further revisions include the description of a new species, Cormohipparion sofularensis, from early Turolian deposits in Turkey, and reassessments assigning certain Eurasian hipparion species to Hippotherium. These updates underscore a consensus on Hipparionini's monophyletic origins while resolving paraphyletic groupings through rigorous quantitative evidence.

Anatomy

Body size and build

Hipparion species exhibited a medium-sized , with shoulder heights typically ranging from 1.0 to 1.4 meters, making them pony-sized compared to larger modern equids like Equus caballus, which often exceed 1.5 meters at the . Body lengths measured approximately 2 to 2.5 meters, and weight estimates varied by and region but generally fell between 135 and 200 kilograms for the genus Hipparion sensu stricto, placing them larger than earlier equids such as Mesohippus (around 20-50 kg) but substantially smaller than Pleistocene Equus (400-600 kg or more). The overall build of Hipparion was slender and gracile, featuring a lightweight frame suited for agile movement and speed across open savanna terrains, in contrast to the more robust structure of later grazing horses. This adaptation included a relatively short neck and elongated trunk, which contributed to a low center of gravity and enhanced cursorial capabilities when compared to the proportionally longer-necked Equus. Hipparions retained three functional toes on each foot, supporting their lightweight morphology. Across their temporal range, Hipparion body sizes showed variation, with early forms (MN9-MN10, around 11-9 Ma) tending to be smaller (under 110 kg in some lineages) and more browsing-oriented, while species (MN14-MN16, 5.3-2.5 Ma) averaged up to 20% larger (often exceeding 210 kg) in response to expanding open grasslands and increased . This trend toward larger sizes in later forms reflected adaptations to more arid, grass-dominated environments.

Cranial features

The skull of Hipparion exhibits an elongated facial structure, a hallmark of the genus, with a prominent preorbital fossa serving as a primary diagnostic feature that distinguishes it from other equids. This fossa is typically subtriangular in shape, well-developed, and positioned close to the , often with a distinct anterior margin and posterior pocket in certain species. The braincase is comparable in overall size to that of modern horses when scaled to body proportions but features fewer and less intricate convolutions, reflecting a lower degree of encephalization. Dentition in Hipparion is characterized by molars displaying complex enamel folding patterns, adapted for processing grasses through increased wear resistance and grinding efficiency. The premolars show greater molarization compared to those of earlier equids like , with heightened crowns and more elaborate occlusal patterns that facilitated a shift toward a more , grass-dominated diet. The nasal notch is retracted, typically positioned above the anterior margin of P² or the middle of the C-P² , indicating a mobile upper capable of both and behaviors. Orbital placement is posterior to the preorbital fossa, with the anterior rim often aligned near the fossa's caudal extent. Morphological variations exist across Hipparion taxa, particularly in preorbital fossa depth; forms classified as Hipparion sensu stricto possess deeper, more prominent fossae compared to some sensu lato relatives, such as Cormohipparion, which feature shallower or more rimmed structures.

Postcranial skeleton

The postcranial skeleton of Hipparion exhibits adaptations for cursorial locomotion, characterized by elongated limbs suited to open terrains. The limb bones feature long, slender metapodials, with metacarpals and metatarsals that are notably narrow and extended, enhancing stride length and speed in Miocene forms. These metapodials support a tridactyl foot configuration, where the central digit (III) is dominant and bears the majority of the body weight, while the lateral digits (II and IV) are reduced but functional, measuring approximately 75-85% the length of the central digit and providing auxiliary support. The forelimbs are slightly longer than the hindlimbs overall, contributing to a balanced gait for agile movement across varied substrates. The vertebral column of Hipparion demonstrates a flexible optimized for dynamic locomotion. It typically includes 18 , with elongated and craniocaudally thin spinous processes on the anterior (T1-T13) that facilitate strong dorsal musculature and mediolateral flexibility, unlike the more rigid structure in modern Equus. This configuration, combined with 6 , supports agile behaviors by allowing greater spinal rotation and spring-like motion, particularly in forested or mixed habitats. Foot morphology in Hipparion retains three functional toes per foot, contrasting with the monodactyl condition of later Equus species, which enables enhanced stability and traction on uneven or soft terrains. The central toe features a keratinous hoof with a spring mechanism for shock absorption, while the lateral toes contact the ground during weight transfer, as evidenced by fossil trackways like those from Laetoli showing side toe impressions. Regional variations in postcranial robustness are also noted, with African forms, such as those from Ethiopian sites, displaying more robust metapodials and limb elements compared to Eurasian counterparts, reflecting adaptations to denser vegetation or differing locomotor demands.

Biology

Habitat and diet

Hipparion species inhabited a range of environments from woodlands to open grasslands across Miocene-Pliocene Eurasia and Africa, as inferred from fossil associations, pollen records, and sedimentary analyses at key sites such as Pikermi in Greece and Laetoli in Tanzania. At Pikermi, during the late Miocene Turolian stage (approximately 8-5 Ma), pollen and paleosol carbonates indicate a dry woodland or forest-dominated landscape with C3 vegetation, rather than expansive C4 savannas, supporting a mosaic habitat suitable for mixed feeders. In contrast, Laetoli's Pliocene deposits (3.8-3.5 Ma) reveal a heterogeneous wooded savanna with grassy patches, evidenced by volcanic sediments and associated faunal diversity, highlighting Hipparion's adaptability to transitional ecosystems. Dietary reconstructions from stable carbon isotopes in demonstrate that Hipparion consumed a mixed diet of C3 browse (trees, shrubs) and C4 grasses, with grass comprising 40-60% of intake in many populations, reflecting opportunistic feeding strategies. Increasing hypsodonty (high-crowned teeth) in later forms facilitated processing tougher, more abrasive vegetation, including silica-rich grasses, as seen in dental adaptations that reduced wear from grit. At sites like the Xunhua Basin in (~9 Ma), enamel δ¹³C values averaging -8.9‰ indicate predominantly C3 diets with minor C4 input (5-10%), while East African records from Nakali (9.9 Ma) show early shifts to >50% C4 grass consumption. Hipparion exhibited environmental tolerances to seasonal , as indicated by oxygen data (δ¹⁸O) in enamel suggesting wetter conditions than modern analogs but with dry spells, and dental microwear textures revealing and grit ingestion from open, windy habitats. In circum-Mediterranean regions, microwear increased from the Vallesian (11.2-8.9 Ma) to Turolian, pointing to greater exposure to abrasive soils during seasonal droughts. Regional differences show early forms favoring more browsing in wooded areas, while late populations, particularly in and eastern , shifted toward dominance amid expanding grasslands.

Behavior and lifespan

Hipparion and related hipparionins likely exhibited social behaviors akin to modern equids, forming herds that facilitated predator avoidance through collective vigilance and rapid flight responses. beds, such as the Love Bone Bed in containing remains of over Neohipparion individuals, suggest accumulation from group migrations rather than solitary deaths, implying as a strategy for traversing open landscapes. Analogies to extant equids indicate typical herd sizes of 10-20 individuals, enabling coordinated escape from predators via speed and agility. Hipparion maintained a lifestyle, with limb adaptations supporting efficient locomotion across varied terrains. Trackway evidence from related hipparionins, such as Eurygnathohippus, reveals gaits including running walks at speeds of 2.2-2.4 m/s, while anatomical features like gracile metapodials indicate capabilities for faster bursts during predator evasion. These traits, inferred from postcranial fossils, underscore a reliance on speed for survival in open habitats. Lifespan estimates for Hipparion derive from dental and histology, revealing growth patterns slower than in modern Equus. Cyclical growth marks in teeth and bones suggest individuals reached skeletal maturity around 3 years, with potential longevity of 9-15 years based on age-at-death profiles from assemblages. For instance, Hipparion concudense specimens indicate older adults surviving to about 10 years, while population data from Neohipparion show average adult lifespans of around 8 years among survivors past infancy. Reproductive strategies in Hipparion paralleled those of modern equids, with a period of approximately 11 months, similar to modern equids. Fossil assemblages imply seasonal breeding, as evidenced by migration patterns in Neohipparion populations that aligned birthing with wet seasons for resource availability. Dental eruption timings, such as third molars at around 3 years marking , further support annual reproductive cycles tied to environmental cues.

Evolution

Origins

Hipparion and its close relatives within the tribe Hipparionini originated in during the middle to early , deriving from the three-toed equid Merychippus through the intermediate form Cormohipparion. This ancestral lineage, particularly Cormohipparion occidentale and related , emerged around 12–11 million years ago in the early , representing a key evolutionary step in the subfamily. The development of transitional traits, such as a well-defined nasomaxillary preorbital fossa—a shallow depression on the facial bone anterior to the eye orbit—and moderately teeth with crown heights of 45–50 mm, distinguished these early hipparions from their browsing predecessors and adapted them for more abrasive, grass-based diets in expanding grasslands. These features evolved progressively from the lower-crowned and less specialized cranial morphology of Merychippus, marking the onset of greater ecological specialization among equids. The fossil record indicates that proto-hipparion forms, akin to early Cormohipparion, first appeared in approximately 15 million years ago during the Barstovian land mammal age, but true Hipparion sensu lato materialized around 12 million years ago in the same region. The initial emergence in the occurred shortly thereafter, with the earliest records dating to 11.4–11.0 million years ago in , exemplified by Hippotherium primigenium in the Vienna Basin and Cormohipparion sinapensis at sites like Sinap Tepe in . This migration, facilitated by lowered sea levels during the early Tortonian supercycle, represents a pivotal dispersal event from across the , initiating the global radiation of hipparions. Ancient DNA analyses from Pleistocene equid remains have bolstered phylogenetic understanding, confirming Hipparion's placement within a distinct hippidiform separate from the lineage ancestral to modern Equus. Studies since 2009, examining mitochondrial sequences from South American hippidion fossils, reveal that these forms belong to the Equus , diverging from North American ancestors around 3–4 million years ago, underscoring the polyphyletic nature of late equid evolution and the independent trajectory of hipparionins.

Biogeography and diversification

Hipparions originated in North America and undertook a major migration across the Bering land bridge into Eurasia approximately 11 million years ago (Ma) during the early late Miocene, with the genus Cormohipparion representing the initial dispersers recorded at sites like the Potwar Plateau in Pakistan and Sinap Tepe in Turkey around 10.8 Ma. From Eurasia, hipparions further dispersed southward into Africa via the Levant corridor during the late Miocene, exemplified by lineages such as Eurygnathohippus appearing in North African deposits by the Turolian stage (MN12, ~9–7.3 Ma). Notably, there is no fossil evidence of hipparions reaching South America, limiting their global distribution to the Northern Hemisphere and adjacent regions. Major fossil localities underscore this biogeographic expansion, with classic European sites like Pikermi and in yielding abundant Hipparion remains from Turolian deposits (~8–7 Ma), revealing diverse cranial and postcranial morphologies indicative of adaptive variation. In , the Siwalik Hills of and have produced extensive Sivalhippus and Hipparion assemblages from Miocene-Pliocene sediments, documenting ongoing evolution in subtropical environments. African records are highlighted by in , where Pliocene footprints and skeletal elements of Hipparion sp. date to ~3.5 Ma, marking one of the southernmost extensions of the lineage. Overall, more than 60 hipparion species across 10 genera have been described from Miocene to deposits worldwide, reflecting a broad paleobiogeographic footprint. Diversification accelerated during the , particularly in MN11–MN12 (~8.9–6.8 Ma), coinciding with global climatic cooling and the expansion of C4 grasslands, which drove niche partitioning among hipparion lineages—such as increased hypsodonty in open-grassland adapted forms versus more browsing-oriented species. This radiation is linked to gradients across , fostering provincialism with distinct bioprovinces emerging in , the , and by the Turolian. Recent research reinforces these patterns, with 2021 analyses correlating hipparion diversification peaks to aridification and monsoon shifts that enhanced grassland habitats across the . A 2023 review highlights holdouts in Western Eurasia, where species like Plesiohipparion longipes and Proboscidipparion crassum persisted into the at sites such as Villarroya in and Kvabebi in Georgia, representing the final refugia before regional extirpation.

Extinction

Hipparion lineages disappeared from the fossil record in a staggered manner across continents, with the fully extinct by the Middle Pleistocene. In , Neohipparion , representing the terminal hipparion radiation, went extinct around the Hemphillian-Blancan boundary approximately 5 million years ago (Ma), marking an early regional end to three-toed equids on the continent. In , last occurrences cluster between 2.5 and 1.0 Ma, with hipparions vanishing from and around 2.5 Ma, persisting longer in eastern regions like until about 1.0 Ma. African records show a delayed pattern, with North African populations (e.g., Eurygnathohippus ) ending at 1.7 Ma and sub-Saharan forms surviving until 0.6–0.4 Ma. Recent analyses highlight prolonged survival in Mediterranean refugia, such as the and , where like Plesiohipparion rocinantis endured into the around 2.1 Ma, coexisting briefly with incoming Equus. The followed a north-to-south across the , correlating with intensifying since the and superimposed Milankovitch-scale climatic oscillations that disrupted habitats. Declining atmospheric CO2 levels, which fell progressively from the onward, contributed to reduced extent by favoring shifts in vegetation toward more arid, fragmented landscapes less suitable for hipparion grazing niches. , driven by these aridity increases and seasonality, isolated populations and limited dispersal, exacerbating regional declines. Ecological competition from monodactyl equids, particularly the radiation of Equus starting around 2.6 Ma in , played a key role in displacing hipparions, as the more efficient single-toed forms outcompeted them in open grasslands. These environmental pressures operated well before significant expansion, confirming purely climatic and biotic drivers without anthropogenic influence. By the , hipparions had been entirely replaced by advanced equids adapted to cooler, more variable conditions.

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  35. https://www.tdx.cat/bitstream/handle/10803/670137/goo1de1.pdf?sequence=1&isAllowed=y
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  41. https://doi.org/10.1127/njgpa/2022/1037
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