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Lagomorpha

Lagomorpha is an order of herbivorous mammals that includes pikas, hares, and rabbits, encompassing approximately 99 extant species divided into two families: Ochotonidae (pikas, with about 34 species) and Leporidae (rabbits and hares, with about 65 species).^[1]^[2] These small to medium-sized animals are characterized by continuously growing incisors with enamel on both the anterior and posterior surfaces, a distinctive feature that sets them apart from rodents, which have enamel only on the front; lagomorphs possess four upper incisors (two pairs, with the second pair smaller and often peg-like), a diastema separating the incisors from the cheek teeth, and a fenestrated skull with open areas behind the orbits.^[3] Their dental formula is typically 2/1, 0/0, 3/2, 3/3, with unrooted, hypsodont (high-crowned) teeth adapted for grinding fibrous vegetation.^[3] Lagomorphs originated in Asia during the Paleocene-Eocene epochs, with the earliest definitive fossils dating to the Early Eocene in regions like western India and northern China,^[4] and they underwent significant diversification starting in the Oligocene, particularly in Central Asia.^[5] Over 50 extinct species are known from Oligocene to Pliocene deposits in Mongolia alone, reflecting a peak in diversity before a decline in the Late Pliocene; modern lineages include primitive ochotonids and more derived leporids, which dispersed globally during the Miocene and Pliocene.^[5] Today, lagomorphs inhabit diverse environments from equatorial forests to Arctic tundras and high-altitude meadows up to 5,000 meters, occurring natively on all continents except Antarctica and Australia (though rabbits have been introduced to Australia and New Zealand).^[1] Pikas are typically small (70–300 grams), egg-shaped, and adapted to rocky talus or meadow habitats where they exhibit vocal communication and varying social structures, while hares (2–5 kilograms) feature long ears and legs for speed in open terrains, and rabbits display diverse forms including burrowing species and the domesticated European rabbit (Oryctolagus cuniculus).^[1] Ecologically, lagomorphs serve as key prey for predators, contribute to ecosystem engineering through grazing and burrowing, act as indicators of climate change (especially pikas), and hold cultural and economic significance in hunting, fur, and meat production.^[1]

Taxonomy and nomenclature

Etymology

The term "Lagomorpha" derives from the Ancient Greek words lagōs (λαγώς), meaning "hare," and morphḗ (μορφή), meaning "form" or "shape," thus referring to animals with a hare-like form. This name was coined in 1855 by the German naturalist Johann Friedrich von Brandt as a subordinal designation within Rodentia to group the hares and rabbits based on their shared dental and morphological traits, particularly the duplicated upper incisors.^[6] Historically, lagomorphs were long confused with rodents due to superficial similarities in gnawing dentition and herbivorous habits, leading to their initial inclusion in the order Rodentia since the 18th century; Brandt's proposal marked an early effort to distinguish them, though full separation into a distinct order occurred later in 1912 by James W. Gidley, who emphasized their unique evolutionary lineage.^[7] This taxonomic shift highlighted anatomical differences, such as the lagomorphs' second pair of smaller upper incisors positioned behind the primary pair, absent in rodents.^[6] Common names for lagomorph groups have evolved linguistically across cultures, reflecting regional encounters and onomatopoeia. "Hare" traces to Proto-Germanic *hasô, via Old English hara, denoting the wild, long-eared species of the genus Lepus. "Rabbit," originally referring to the young of hares or the European species Oryctolagus cuniculus, entered English in the 14th century from Old French rabot or Walloon robète, of uncertain origin but possibly imitative of the animal's burrowing habits. For pikas (Ochotonidae), the name derives from Tungusic piika or Mongolian ogotona (оготно), mimicking their high-pitched calls, and was adopted into English in the 19th century as European explorers documented Asian species.^[8]

Taxonomic history and phylogeny

The taxonomic history of Lagomorpha begins with Carl Linnaeus, who in 1758 classified hares (genus Lepus) within the order Rodentia, based on superficial similarities in their gnawing dentition and overall morphology. Pikas (genus Ochotona) were not scientifically described until 1795 by Heinrich Friedrich Link.^[6] This grouping persisted for over a century, as early naturalists often treated lagomorphs as a subgroup of rodents due to shared ecological roles and anatomical features like continuously growing incisors. However, in 1855, Johann Friedrich von Brandt proposed the order Lagomorpha to distinguish hares and pikas from rodents, emphasizing their unique duplicidentate dentition—featuring two pairs of upper incisors instead of one—as a defining trait derived from the Greek words lagos (hare) and morphē (form).^[6] This separation gained traction in the late 19th and early 20th centuries through comparative anatomy studies, which highlighted additional differences in skull structure, jaw mechanics, and reproductive systems, leading to formal recognition of Lagomorpha as a distinct order by the mid-20th century.^[6] In modern taxonomy, Lagomorpha is recognized as an order within the superorder Euarchontoglires, where it forms the clade Glires alongside Rodentia, supported by molecular phylogenies that confirm their sister-group relationship based on shared genomic signatures and early divergences from other placental mammals. The order comprises two extant families: Ochotonidae (pikas), with a single genus Ochotona encompassing 34 species (as of 2024) primarily adapted to rocky, alpine environments; and Leporidae (rabbits and hares), which includes 11 genera and approximately 75 species (as of 2024), such as Lepus for hares, Oryctolagus for the European rabbit, and Sylvilagus for New World cottontails, reflecting diverse burrowing and cursorial lifestyles. There are no formal suborders within Lagomorpha; the two families represent the primary phylogenetic divisions, with Ochotonidae branching earlier as the outgroup to the more speciose Leporidae. Contemporary phylogenetic understanding relies heavily on molecular data, including mitochondrial and nuclear genes, which robustly affirm the monophyly of Lagomorpha with high bootstrap support in maximum likelihood trees. A 2024 analysis using a supermatrix of 17,942 nucleotide sites from 80 lagomorph taxa estimates the crown-group divergence of Lagomorpha at approximately 57.2 million years ago (95% highest posterior density: 52.9–57.2 Ma),^[9] aligning with Paleogene radiations shortly after the Cretaceous-Paleogene boundary. These molecular time-calibrated phylogenies, incorporating fossil constraints, further resolve intra-family relationships, such as the basal position of certain leporid genera, and underscore the order's evolutionary independence from rodents despite their historical taxonomic linkage.

Evolutionary history

Origins and early diversification

Stem lagomorphs within the larger Glires clade that also includes rodents trace their origins to the early Paleocene in Asia, approximately 60 million years ago, shortly after the Cretaceous-Paleogene extinction event.^[10] Early ancestors, such as the duplicidentate Mimotona species from localities in China, represent stem Glires with mixed rodent- and lagomorph-like mandibular features, indicating an initial phase of morphological experimentation within the clade.^[10] These basal forms likely inhabited forested environments, with evidence suggesting arboreal habits inferred from limb proportions and dental adaptations suited to a herbivorous diet.^[11] Crown Lagomorpha emerged in the early Eocene around 55 million years ago. By the Paleocene-Eocene boundary around 55 million years ago, lagomorph ancestors had diverged from rodents, marking the emergence of true stem lagomorphs in Asia, including taxa like Dawsonolagus from Inner Mongolia.^[12] A key adaptation during the Eocene was the development of duplicated upper incisors, characteristic of the Duplicidentata, which enhanced gnawing efficiency for processing tough vegetation and distinguished lagomorphs from their simplicidentate rodent relatives.^[11] This period saw a transition from predominantly arboreal lifestyles to more terrestrial forms, driven by evolving locomotor traits such as elongated hindlimbs in some lineages, allowing exploitation of ground-level foraging opportunities.^[11] Initial diversification accelerated in the middle to late Eocene, with stem lagomorphs radiating across Asia and eventually reaching North America via Beringian land bridges.^[12] By the late Eocene, the clade had diverged toward the two extant families: Ochotonidae (pikas and relatives, represented early by forms like Desmatolagus) and Leporidae (rabbits and hares, with precursors like Shamolagus), reflecting ecological partitioning between alpine and open-habitat niches.^[12] This branching was facilitated by post-Cretaceous environmental shifts, including the Paleocene-Eocene Thermal Maximum warming event around 56 million years ago, which promoted forest expansion and herbaceous plant proliferation, providing favorable conditions for herbivore radiation in Asia.^[12] The oldest definitive lagomorph fossils include small ankle bones from Early Eocene deposits in western India and the complete skeleton of Gomphos elkema from China, dating to approximately 55 million years ago.^[13]^[14]

Fossil record and key events

The fossil record of Lagomorpha extends back to the early Eocene, with the earliest known specimens from Asia, such as Gomphos, and later middle Eocene forms like the genus Gobiolagus from the Shara Murun Formation at Ula Usu in Inner Mongolia, China.^[15] This genus, comprising at least six species, represents one of the basal lagomorphs and highlights Central Asia as a key center for early diversification, with additional middle Eocene finds from sites like Erden Obo further refining the morphology of primitive forms.^[16] In North America, the genus Palaeolagus, exemplified by P. haydeni, appears prominently in late Eocene to early Oligocene deposits, such as those in the Brule Formation of Nebraska, providing abundant cranial and postcranial evidence of early leporid-like adaptations and marking the continent's initial lagomorph presence.^[17] A pivotal event in lagomorph evolution was the Miocene radiation of the family Leporidae, which saw a peak in diversity during the Miocene-Pliocene transition, driven by ecological expansions across Eurasia and North America.^[18] This radiation involved the spread of leporids from North American origins into Asia by the late Miocene, as evidenced by the "Leporid Datum"—a biotic marker reflecting the widespread invasion and abundance of the family around 10-7 million years ago.^[19] Later, during the Pleistocene, climate shifts associated with glacial-interglacial cycles contributed to extinctions of several lagomorph lineages, particularly in North America, where megafaunal turnover and habitat fragmentation led to dietary shifts in surviving species toward C4 grasses, underscoring the vulnerability of smaller herbivores to environmental instability.^[20] Recent discoveries post-2020 have enhanced understanding of lagomorph evolution, including new Asian fossils from middle Eocene sites that bolster evidence for the divergence between leporids and ochotonids around 40-50 million years ago, aligning fossil data with molecular estimates placing crown Lagomorpha at approximately 57 million years ago.^[21] A 2022 study on cranial evolution in leporids analyzed fossil specimens to reveal unique adaptations like enlarged auditory bullae, which emerged during the Miocene radiation and facilitated diversification into open habitats.^[22] These findings also address gaps in the record, notably the sparse representation of lagomorphs in Africa; while historical data show limited pre-Pleistocene fossils, recent excavations have yielded new species, such as Pronolagus from early Pleistocene sites in Angola, indicating greater southern African diversity than previously recognized.^[23]

Physical characteristics

General morphology and anatomy

Lagomorphs are small to medium-sized herbivorous mammals characterized by a quadrupedal body plan with cursorial limbs adapted for swift terrestrial locomotion. Body mass varies widely across the order, ranging from about 100 g in the smallest pikas to up to 7 kg in larger hares, with head-body lengths typically spanning 125–750 mm. Their pelage is dense and soft, often exhibiting countershading for camouflage, and they possess a short or rudimentary tail, except in pikas where it is absent. These structural features support their primarily herbivorous lifestyle, enabling efficient foraging and predator evasion in diverse environments.^[24]^[25]^[26] A defining anatomical trait of lagomorphs is their dentition, which features a unique dental formula of I 2/1, C 0/0, P 3/2, M 2–3/3, resulting in 26–28 teeth total. Unlike rodents, they have two upper incisors per side—the primary incisor is large and chisel-like, while the secondary (peg) incisor sits directly behind it, aiding in precise gnawing of tough plant material. Canines are absent, creating a prominent diastema between the incisors and cheek teeth, and all post-incisor teeth are hypsodont (high-crowned) and ever-growing, with transverse enamel ridges on the occlusal surfaces for grinding fibrous vegetation. The mandibular symphysis is unfused, allowing independent lateral movement of the jaw halves for optimal occlusion. This specialized dentition is crucial for their hindgut fermentation-based digestion.^[24]^[27]^[28] Skeletally, lagomorphs exhibit adaptations for agility and speed, including elongated hind limbs with an extended femur, tibia, and fibula that facilitate powerful leaps, often exceeding 3 m in distance. The spine is notably flexible, with additional lumbar vertebrae compared to many mammals, enhancing maneuverability during rapid directional changes. The skull is fenestrated, particularly in the maxillary region, reducing weight while maintaining structural integrity, and the palate is short. Laterally positioned eyes provide a nearly panoramic field of view, spanning almost 360 degrees, which is essential for detecting threats from multiple angles.^[29]^[24]^[30] Sensory systems in lagomorphs are finely tuned for survival in predator-rich habitats. Hearing is acute, with sensitivity profiles similar to modern rabbits, enabling detection of low-frequency sounds from approaching dangers; the auditory bullae are inflated for enhanced sound localization. Olfaction is well-developed, supported by a complex nasal turbinal skeleton that increases the surface area for odorant detection, aiding in foraging, social communication, and predator avoidance. Vibrissae, or whiskers, are prominent on the muzzle and serve as tactile sensors for navigating low-light or obstructed environments, providing feedback on nearby objects and textures.^[31]^[17]^[24]

Unique traits compared to other mammals

Lagomorphs possess a distinctive dentition that sets them apart from other mammals, particularly rodents. Unlike rodents, which have a single pair of continuously growing upper incisors, lagomorphs feature two pairs: large central incisors flanked by smaller, peg-like second incisors positioned directly behind them. These peg teeth play a crucial role in stabilizing the lower incisors during occlusion and aiding in the precise shearing of fibrous vegetation, enhancing feeding efficiency in herbivorous diets.^[32]^[33] This dual-incisor arrangement is unique to the order Lagomorpha and contributes to their specialized oral morphology, distinct from the simpler gnawing apparatus of rodents.^[34] For thermoregulation, lagomorphs employ specialized fur and ear morphologies adapted to diverse environments. Their dense, multilayered fur provides superior insulation against cold, with guard hairs and underfur trapping air for thermal retention, a trait more pronounced than in many similarly sized mammals lacking such profuse pelage. In species like hares, elongated ears serve as efficient radiators, facilitating heat dissipation through vascular networks that dilate in warm conditions to increase surface area for convective cooling, while vasoconstriction conserves heat in colder climates—a mechanism particularly vital in arid or open habitats.^[30]^[35]

Variations among families

The two families within Lagomorpha, Ochotonidae (pikas) and Leporidae (hares and rabbits), exhibit notable morphological and physiological variations that reflect their distinct evolutionary paths. Ochotonids are generally smaller in body size, typically ranging from 15 to 25 cm in length and weighing 100 to 200 g, compared to the larger leporids, which can reach up to 70 cm and several kilograms.^[18] This size disparity influences their overall build, with pikas featuring rounded ears, an absence of an external tail, and dense fur that provides insulation.^[36] In contrast, leporids possess elongated ears and hind legs adapted for rapid locomotion, a visible tail often tipped in black in certain lineages, and pronounced sexual dimorphism where females are typically larger than males to support higher reproductive demands.^[37]^[38] Physiologically, these families diverge in metabolic strategies suited to their lifestyles. Pikas maintain higher basal metabolic rates, enabling efficient thermoregulation in cooler, high-altitude settings through elevated energy expenditure for heat production.^[39] Conversely, leporids are specialized for endurance running, with enhanced cardiovascular and muscular adaptations that prioritize sustained speed over intense bursts, facilitated by their longer limbs and lighter skeletal proportions.^[29] A 2021 study on lagomorph morphology demonstrated significant expansion of leporid cranial morphospace following divergence from ochotonids, highlighting increased disparity in skull shape linked to locomotor and dietary specializations.^[11] Despite these differences, both families share core lagomorph traits, such as the duplicated upper incisors for gnawing.^[18]

Diversity

Pikas (Ochotonidae)

Pikas belong to the family Ochotonidae, which consists of a single extant genus, Ochotona, encompassing approximately 32 species distributed primarily across Asia and North America.^[40]^[41] These small, tailless mammals, commonly known as "rock rabbits" or "conies," are characterized by their rounded ears, short limbs, and soft fur, typically measuring 15–25 cm in body length and weighing 100–300 g.^[36] Unlike their leporid relatives, pikas exhibit a more compact body form suited to navigating rocky terrains, with dense pelage that provides insulation against harsh alpine conditions. A defining behavioral adaptation of pikas is their hay-piling, or "haying," where individuals harvest vegetation during the short summer months, drying it in the sun before storing it in central haypiles within their home ranges for winter consumption, as they do not hibernate.^[42] This stockpiling can amount to several kilograms per individual, ensuring survival through periods of snow cover when fresh forage is unavailable.^[43] Pikas also construct extensive burrow systems in talus slopes—loose accumulations of rocks and boulders—using these interstices for shelter, nesting, and escape from predators.^[44] These habitats provide thermal stability, buffering against extreme temperature fluctuations in high-elevation environments. Communication among pikas relies heavily on vocalizations, including high-pitched bleats, whistles, and yodel-like calls that function in territorial defense, alarm signaling, and individual recognition, often echoing across rocky slopes to maintain social spacing.^[45] Adults produce a repertoire of at least nine distinct call types, with acoustic variation aiding in mate attraction and kin identification within family groups.^[45] The diversity within Ochotona reflects adaptations to varied montane niches, with species exhibiting subtle morphological differences such as ear size and fur coloration correlated with local climates. The American pika (Ochotona princeps), the sole North American endemic, occupies talus fields in the Rocky Mountains and Sierra Nevada, from elevations of 1,500–4,000 m, where it demonstrates resilience to cold but vulnerability to heat stress.^[46] In contrast, Asian species like the plateau pika (Ochotona curzoniae) thrive in the high-altitude meadows of the Qinghai-Tibetan Plateau at 3,000–5,000 m, serving as a keystone species by engineering burrow networks that enhance soil aeration and biodiversity.^[47] Recent taxonomic investigations in the 2020s, leveraging genomic data, have affirmed the monophyly of the genus Ochotona and prompted minor revisions, such as the recognition of additional lineages within species complexes and the description of two new species, Ochotona galunglaensis and another, from the Himalayan region in 2025.^[48]^[41] These studies underscore the genus's evolutionary cohesion while highlighting ongoing diversification driven by geographic isolation.

Hares and rabbits (Leporidae)

The family Leporidae, commonly known as hares and rabbits, comprises 11 genera and approximately 64 species distributed worldwide, excluding Antarctica and Australia (where some have been introduced). This family represents the majority of lagomorph diversity, with hares belonging primarily to the genus Lepus (32 species) and rabbits encompassing other genera such as Oryctolagus and Sylvilagus.^[1] A key distinction between hares and rabbits lies in their reproductive strategies: hare young (leverets) are precocial, born fully furred with open eyes and capable of limited mobility shortly after birth, while rabbit young (kits) are altricial, born hairless, blind, and helpless in burrows or nests.^[49] These differences reflect adaptations to their respective lifestyles, with hares relying on speed and camouflage in open habitats rather than extensive sheltering. Hares in the genus Lepus are typically solitary and adapted for rapid escape from predators, achieving speeds up to 70 km/h in bursts, as exemplified by the European hare (Lepus europaeus), which inhabits grasslands and farmlands across Europe and Asia.^[50] Their precocial development enables leverets to disperse quickly after birth, reducing vulnerability without the need for prolonged parental care. Unlike the rock-dwelling pikas of the family Ochotonidae, leporids are predominantly terrestrial, with elongated hind limbs specialized for leaping and bounding locomotion. Recent studies on cranial evolution highlight unique leporid adaptations, such as an intracranial joint at the occipital region and a fenestrated rostrum, which enhance structural integrity and reduce weight during high-speed cursorial activities.^[51] Rabbits, in contrast, often exhibit more social tendencies and dependence on burrowing, with species like the European rabbit (Oryctolagus cuniculus) forming colonies in extensive underground warrens that serve as protective networks for altricial offspring.^[52] Native to the Iberian Peninsula, O. cuniculus has a long history of human association, with evidence of managed populations in walled enclosures dating to Roman times, laying the foundation for modern domestication.^[53] Similarly, cottontail rabbits in the genus Sylvilagus, widespread in the Americas, give birth in shallow burrows or ground nests, emphasizing their altricial strategy while utilizing vegetative cover for concealment. These traits underscore the leporids' evolutionary divergence within Lagomorpha, prioritizing evasion through either velocity or communal refuge.

Distribution and habitats

Global range

Lagomorphs are native to every continent except Australia and Antarctica, with natural populations spanning North and South America, Europe, Asia, and Africa, though they are absent from southern South America and most oceanic islands.^[24] Human-mediated introductions have extended their range to Australia, New Zealand, and various islands, where species like the European rabbit (Oryctolagus cuniculus) have become established and, in some cases, invasive.^[54] The order's distribution reflects a combination of ancient dispersals and more recent anthropogenic influences, resulting in a near-cosmopolitan presence today. The family Ochotonidae (pikas) exhibits a more restricted range, primarily across Asia—from the Himalayas to eastern Siberia—and western North America, with limited occurrences in Eastern Europe.^[11] In contrast, the family Leporidae (rabbits and hares) is more widespread, achieving a cosmopolitan distribution with the highest species diversity concentrated in the Americas and Eurasia; for instance, the genus Lepus includes 32 species distributed nearly worldwide, while Sylvilagus with 17 species is confined to the Americas.^[55] Asia is a major center of diversity for lagomorphs, underscoring the continent's role in their evolutionary history. Historical range expansions have shaped modern distributions, including post-glacial migrations that allowed species to recolonize northern latitudes in Europe and North America following the Last Glacial Maximum.^[56] Human introductions have further facilitated spread, such as the translocation of the European rabbit from its native Iberian Peninsula to other parts of Europe by the Romans around the 1st century AD, and later to distant regions like Australia in the 19th century.^[54] Endemic hotspots include the Ethiopian Highlands, home to species like the Ethiopian highland hare (Lepus starcki), which is restricted to the Afroalpine zones of central Ethiopia.^[57]

Habitat preferences and adaptations

Lagomorphs exhibit diverse habitat preferences shaped by their family affiliations, with each demonstrating specialized physiological and morphological adaptations to environmental challenges. Pikas (family Ochotonidae) primarily inhabit high-altitude rocky terrains, such as talus slopes and boulder fields above the tree line, as well as tundra-like alpine meadows in cool, moist ecosystems. These environments provide thermal stability through rock insulation and access to vegetation for foraging. To cope with harsh winters, pikas remain active year-round without entering torpor or hibernation, relying instead on thick insulating fur, a high metabolic rate for heat generation, and behavioral adaptations like caching "hay piles" of dried vegetation to sustain them through snow-covered periods.^[58]^[59] Hares (family Leporidae) favor open habitats including grasslands, shrublands, and deserts, where their speed and vigilance offer protection from predators. In seasonal environments, species like the snowshoe hare (Lepus americanus) undergo biannual molts, shifting from brown summer fur for camouflage in dry vegetation to white winter pelage against snow cover, enhancing survival by reducing detection. This fur color polymorphism has evolved convergently in multiple hare lineages to match variable snow durations.^[60]^[61] Rabbits (also Leporidae) prefer more structured habitats such as forests, meadows, and scrublands, often utilizing burrows for shelter. They construct extensive underground warrens—networks of tunnels with chambers for nesting and escape—that provide protection from predators and extreme weather, allowing populations to thrive in areas with dense cover. Burrowing behavior is particularly pronounced in species like the European rabbit (Oryctolagus cuniculus), facilitating communal living and resource defense.^[62] Recent research highlights how climate variability influences lagomorph camouflage adaptations, particularly in hares and jackrabbits. A 2023 study on white-tailed jackrabbits (Lepus townsendii) revealed that genetic variation in seasonal pelage color tracks historical snow cover patterns, with lighter winter coats predominant in snowy regions and darker ones in milder areas; populations with higher genetic diversity in color genes may better adapt to future reductions in seasonal snow due to climate change.^[63]

Ecology and biology

Diet and digestion

Lagomorphs are strictly herbivorous, with diets centered on foliage such as grasses, herbs, bark, and twigs.^[53] Members of the family Ochotonidae (pikas) primarily forage on alpine and meadow plants, collecting and drying them into haypiles during summer for winter consumption, which can sustain them for up to 350 days in some populations.^[64] In contrast, species in the family Leporidae (hares and rabbits) graze predominantly on fresh green vegetation when available, relying on immediate foraging rather than extensive storage.^[65] Lagomorphs employ hindgut fermentation as their primary digestive mechanism, facilitated by an enlarged cecum that hosts microbial communities breaking down complex plant fibers.^[66] This process produces volatile fatty acids for energy, but to compensate for nutrient losses post-foregut digestion, lagomorphs form soft cecotropes—nutrient-dense fecal pellets containing B vitamins, proteins, and fermentation byproducts—that are re-ingested through cecotrophy.^[66] Pikas exhibit a variant of this behavior, sometimes storing cecotropes for later consumption alongside direct ingestion.^[67] Enzymatic digestion in lagomorphs relies on microbial rather than endogenous enzymes, with high cellulase activity in the cecum enabling cellulose breakdown absent in their own physiology.^[68] Gut morphology varies among families: rabbits possess relatively longer gastrointestinal tracts and slower digesta passage rates than hares, enhancing fiber digestibility and nitrogen retention, while hares' faster transit supports rapid intake aligned with their cursorial habits.^[69] ^[70] Seasonal dietary shifts pose significant nutritional challenges, particularly in resource-poor habitats. During winter, leporids transition to low-quality foods like bark and twigs, which provide limited protein and energy, potentially leading to mass loss if twig diameter exceeds optimal sizes for digestion.^[71] Pikas mitigate these constraints through haypile reserves, selecting summer forage with lower phenolic content for immediate use and higher-fiber plants for storage, though extreme conditions can still strain their metabolic demands.^[65]

Reproduction and development

Lagomorphs exhibit a polyestrous reproductive strategy, with females capable of multiple breeding cycles within a single season, often producing several litters annually to maximize reproductive output in variable environments.^[72] Ovulation in lagomorphs is induced rather than spontaneous, triggered by copulation or mechanical stimulation, which ensures fertilization efficiency and is a key adaptation shared across the order.^[73] Litter sizes typically range from 1 to 12 offspring, varying by species and environmental conditions, with larger litters more common in rabbits than in pikas or hares.^[74] Reproductive patterns differ markedly among lagomorph families. In pikas (Ochotonidae), breeding pairs are often monogamous, with females producing 2-4 altricial young per litter—born hairless, blind, and helpless—typically in two litters per year within rock crevices or burrows.^[75] Hares (Leporidae) give birth to precocial young in shallow ground depressions known as forms; these offspring are furred, eyes open, and mobile shortly after birth, enabling rapid independence with minimal parental investment.^[76] In contrast, rabbits (also Leporidae) produce altricial kits in elaborate underground burrows, where the young remain dependent for several weeks post-birth.^[77] Gestation periods in lagomorphs are relatively short, lasting 25-31 days in rabbits and pikas, though slightly longer (up to 42 days) in some hares, facilitating quick turnover of generations.^[74] Sexual maturity is attained early, generally between 3 and 8 months of age, allowing individuals to breed in their first year and contributing to the order's high reproductive potential.^[37] Parental care varies significantly, reflecting developmental differences. Hares provide minimal care, with mothers nursing leverets briefly once daily and leaving them concealed to avoid predation, as the young forage independently soon after birth.^[78] Rabbits exhibit more structured care, including burrow construction and once-daily communal nursing bouts where multiple females in a warren may nurse litters collectively, enhancing survival in social groups.^[79] Pikas show intermediate investment, with both parents defending territories and the female nursing altricial young for about three to four weeks until weaning.^[75]

Behavior and sociality

Lagomorphs display a diverse spectrum of social structures that reflect adaptations to their environments and predation pressures. Hares, such as the European hare (Lepus europaeus), are predominantly solitary, maintaining individual territories and interacting primarily during brief mating encounters, with occasional loose aggregations at food sources in high-density areas. In contrast, rabbits like the European rabbit (Oryctolagus cuniculus) form colonial groups of 2–8 adults plus juveniles in complex warrens, featuring a rigid dominance hierarchy where males compete aggressively for territory defense and females protect nesting burrows. Pikas, including the American pika (Ochotona princeps), often live in pair-bonded family units, with mated pairs sharing territories year-round and subadults sometimes philopatric, remaining with parents into winter; colonial variants, such as steppe pikas (O. curzoniae), organize into larger family clusters with minimal overlap between units.^[80]^[53]^[62] Daily activity patterns in lagomorphs are typically crepuscular or nocturnal, aligning with reduced predator activity at dawn and dusk to facilitate foraging while minimizing exposure; for instance, snowshoe hares (Lepus americanus) and cottontail rabbits (Sylvilagus spp.) peak in activity during these twilight periods, though some pikas exhibit diurnal habits in cooler alpine zones. Alarm responses include foot-thumping, a rapid hind-limb percussion that transmits vibrations and alerts nearby individuals, as seen in rabbits and certain pikas like the Afghan pika (O. rufescens). Freezing postures—immobility with erect ears and flattened bodies—are a common initial reaction to potential threats, allowing camouflage in vegetation or rocky substrates before flight.^[81]^[62]^[82] Anti-predator behaviors emphasize evasion and detection, with lagomorphs relying on bursts of speed up to 45 mph (72 km/h) in hares and 35 mph (56 km/h) in rabbits, often executed in erratic zig-zag trajectories to disrupt predator pursuit angles and exploit terrain irregularities. Vigilance is heightened in open habitats, involving frequent pauses for scanning and ear orientation, as documented in cottontail rabbits where individuals adjust alertness based on perceived risk from visual or olfactory cues. Colonial rabbits may engage in group mobbing, collectively approaching and vocalizing at intruders to deter them, enhancing survival through shared vigilance in social settings.^[83]^[84]^[85] Communication among lagomorphs integrates vocal, chemical, and postural signals for territory maintenance and coordination. Vocalizations include distress screams and low grunts in hares and rabbits during agonistic encounters or capture, while alarm calls—sharp whistles or trills—prompt conspecific flight. Scent marking via submandibular or anal glands deposits pheromones to delineate boundaries, a practice universal across families and intensified during breeding. Pikas notably employ complex "songs," prolonged call sequences by males to advertise territories and attract mates, with studies indicating these vocalizations convey individual identity through unique spectral signatures, facilitating recognition in talus habitats.^[62]^[75]^[86]

Conservation

Threats and challenges

Lagomorph populations worldwide are confronting multiple anthropogenic and environmental pressures that exacerbate their vulnerability, with approximately 31% of the 91 known species assessed as threatened (Vulnerable, Endangered, or Critically Endangered) on the IUCN Red List as of 2025.^[87] These threats often compound, leading to population declines across diverse taxa, including pikas, hares, and rabbits. Habitat loss and degradation, primarily driven by agricultural expansion and urbanization, represent one of the most pervasive dangers to lagomorphs, fragmenting their preferred open grasslands, shrublands, and montane environments. In regions like Mexico and China, conversion of native habitats for intensive farming has severely impacted species such as the volcano rabbit (Romerolagus diazi), reducing available foraging areas and increasing exposure to predators. Urban sprawl further erodes these habitats, as seen in European hare (Lepus europaeus) populations in peri-urban zones where development displaces burrows and food sources. Climate change intensifies this issue for talus-dwelling pikas, particularly the American pika (Ochotona princeps), whose high-elevation habitats are shifting upward due to warming temperatures, resulting in local extirpations and an estimated 97% decline in suitable climate space in parts of the Sierra Nevada by mid-century under high-emission scenarios.^[88]^[89] Predation pressures have escalated through overharvesting and the introduction of non-native predators, disrupting natural population dynamics. Unsustainable hunting for meat, fur, and sport has depleted hare and rabbit stocks in parts of Africa and Asia, with species like the Cape hare (Lepus capensis) facing intensified poaching amid growing human demand.^[90] Invasive predators, such as feral cats and red foxes introduced to islands and mainland areas, prey heavily on vulnerable lagomorphs; for instance, in New Zealand and Australia, these exotics have contributed to sharp declines in introduced rabbit populations by targeting juveniles and exploiting altered landscapes.^[91] Diseases pose acute risks, with viral pathogens causing mass mortality events that can decimate local populations. Myxomatosis, a poxvirus intentionally introduced to control European rabbits (Oryctolagus cuniculus) in Australia in the 1950s, persists as a cyclical threat, leading to epizootics that kill up to 99% of susceptible individuals in naive populations. More recently, rabbit hemorrhagic disease virus 2 (RHDV2) has emerged as a global concern, with outbreaks in the 2020s causing widespread die-offs; in the southwestern United States, the virus was first detected in wild lagomorphs in March 2020 and has since spread across multiple states, infecting both wild and domestic rabbits with mortality rates exceeding 70% in affected areas.^[92] Emerging pathogens, including coinfections with myxoma virus, further compound these impacts, particularly in dense populations.^[93] Competition from invasive species adds another layer of challenge, as non-native lagomorphs and herbivores outcompete natives for resources in overlapping ranges. The invasive European rabbit, for example, alters vegetation structure in Iberian ecosystems, reducing forage availability for endemic Iberian lynx prey like the wild rabbit (Oryctolagus cuniculus—native form) and indirectly pressuring hare populations through resource depletion.^[94] Such interactions, combined with habitat modifications by invasives, heighten extinction risks for island-endemic species like the Sardinian pika (extinct) and contribute to broader biodiversity losses.^[95]

Status and protection efforts

Lagomorph conservation statuses vary widely across the order's 91 species, with the International Union for Conservation of Nature (IUCN) Red List classifying one species as critically endangered, 13 as endangered, and 14 as vulnerable as of 2025.^[87] The critically endangered riverine rabbit (Bunolagus monticularis) in South Africa exemplifies acute risk, with fewer than 250 breeding pairs remaining due to habitat loss.^[96] Endangered species include the hispid hare (Caprolagus hispidus) in India and Nepal, the Amami rabbit (Pentalagus furnessi) in Japan with around 3,000 individuals, and the volcano rabbit (Romerolagus diazi) in Mexico. Many critically endangered and endangered lagomorphs lack sufficient recent studies, highlighting ongoing gaps in monitoring and research.^[97] Several lagomorphs receive international protection through the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), with species like the hispid hare and volcano rabbit listed in Appendix I, prohibiting commercial trade to prevent further population declines.^[98] Pikas benefit from national park protections in regions like the United States, where the American pika (Ochotona princeps) is safeguarded in parks such as Rocky Mountain National Park through habitat monitoring and climate impact assessments.^[99] Conservation efforts are coordinated by organizations like the World Lagomorph Society, which funds research and projects for threatened species, including population surveys and habitat restoration initiatives.^[100] Reintroduction programs have targeted species such as the pygmy rabbit (Brachylagus idahoensis), with over 1,200 individuals released in Washington State since 2007, supported by captive breeding to enhance wild populations despite challenges like low survival rates.^[101] Disease management research focuses on rabbit hemorrhagic disease virus (RHDV2), with vaccines tested successfully in endangered riparian brush rabbits (Sylvilagus bachmani riparius), inducing protective antibodies without adverse effects.^[102] Recent advances from 2023 to 2025 include climate modeling to identify habitat corridors for species like the American pika, aiding connectivity amid warming temperatures.^[103] Genetic diversity studies for captive breeding, particularly in pygmy rabbits, use noninvasive fecal sampling to monitor inbreeding and guide releases, improving long-term viability.^[104] As of 2025, expanded RHDV2 vaccination programs have been implemented for wild populations in affected regions, showing initial success in reducing mortality rates.^[87]

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None
### Summary of IUCN Red List Categories for Lagomorph Species
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