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Pecora

Pecora is an infraorder of even-toed ungulate mammals (order Artiodactyla) within the suborder Ruminantia, encompassing the majority of living ruminants except for the primitive chevrotains (family Tragulidae). It includes five extant families: Antilocapridae (pronghorns), Bovidae (cattle, sheep, goats, and antelopes), Cervidae (deer), Giraffidae (four giraffe species and the okapi), and Moschidae (musk deer).^[1]^[2] Pecorans are characterized by a specialized four-chambered stomach that facilitates microbial fermentation of fibrous plant material, enabling efficient digestion through rumination (chewing the cud).^[3] Most species possess bony cranial appendages—such as true horns (permanent, keratin-covered cores in Bovidae), antlers (deciduous, branched structures in Cervidae), pronghorns (branched and shed annually in Antilocapridae), or ossicones (skin-covered projections in Giraffidae)—which emerge from the frontal bones and serve functions in intraspecific competition, defense, and sexual selection.^[3] Exceptions occur in the musk deer (Moschus spp.) and the Chinese water deer (Hydropotes inermis), which lack these structures.^[3] Dentally, they typically follow the formula I 0/4, C 0/1, P 3/2–3, M 3/3 (32–34 teeth), with no upper incisors and a gap (diastema) for food manipulation.^[3] The evolutionary origins of Pecora trace back to Eurasia in the late Eocene to early Oligocene, over 37 million years ago, with significant diversification during the Miocene epoch around 23–5 million years ago, driven by climatic changes and habitat expansion.^[4]^[5] Fossil records indicate an initial radiation in Asia and Europe, followed by migrations to Africa and the Americas, giving rise to stem groups like the palaeomerycids and climacoceratids before the emergence of modern families.^[6] Today, Pecora exhibit global distribution across diverse habitats from tundras to savannas, with approximately 200–210 species as of 2025; the Bovidae alone accounts for over 140 species in more than 50 genera, including economically vital domesticated forms like cows (Bos taurus) and sheep (Ovis aries).^[7] Their ecological roles range from herbivores shaping vegetation dynamics to keystone species in food webs, while many face threats from habitat loss and overhunting.^[8]

Evolutionary history

Origins and divergence

The suborder Ruminantia emerged during the Eocene epoch, with molecular estimates and fossil evidence indicating an origin around 50 million years ago in Asia, where early forms adapted to forested environments. The oldest known ruminant fossils, such as Archaeomeryx from the middle Eocene of Mongolia, date to approximately 44 million years ago and represent primitive, small-bodied artiodactyls with basic selenodont dentition.^[4] These early ruminants laid the foundation for the clade's subsequent radiation, characterized by the development of foregut fermentation as a key digestive innovation. Pecora, comprising the advanced ruminants including bovids, cervids, and giraffids, diverged from the sister clade Tragulina (represented by chevrotains and their relatives) within Ruminantia during the late Eocene. Phylogenetic analyses combining molecular and morphological data confirm this sister-group relationship, with Bayesian relaxed molecular clock methods estimating the divergence at 44.3–46.3 million years ago. This split marked a critical point in ruminant evolution, as Pecora began to exhibit traits like more complex cranial structures and enhanced hypsodonty, setting the stage for later ecological expansions. Alternative molecular timescales place the divergence slightly earlier, around 51.6 ± 4.9 million years ago in the early Eocene, highlighting some variability in clock calibrations but consistently supporting an Eocene origin.^[9] The initial diversification of Pecora accelerated in the Miocene (23–5 million years ago), coinciding with global climatic cooling after the Middle Miocene Climatic Optimum and the widespread expansion of C4 grasslands across Eurasia and Africa. This environmental shift, involving increased aridity and seasonality, drove adaptations for mixed browsing and grazing diets, enabling pecorans to exploit newly available open habitats more effectively than their traguline relatives.^[9] These changes facilitated the evolution of advanced ruminant digestion, with rumen microbial communities optimizing fiber breakdown in grasses. Early Miocene fossils like Prodremotherium (approximately 20 million years ago from Europe) exemplify this phase, displaying primitive cranial features such as elongated, straight nasal bones, short curved canines, and a relatively simple hornless skull, consistent with its position as a stem pecoran.^[10]

Fossil record

The fossil record of Pecora is relatively sparse during the Eocene and Oligocene epochs, with transitional forms providing key insights into the early evolution of this ruminant suborder. Stem pecorans, such as those belonging to the Gelocidae family, including the genus Gelocus, are documented from Oligocene deposits in Europe, representing primitive morphologies that bridge non-pecoran ruminants and crown-group Pecora.^[6] These early taxa exhibit basicranial features and dental characteristics transitional to pecoran specializations, such as elongated nasal bones and rudimentary pedicles that foreshadow the development of cranial appendages, though they lack fully formed horns.^[11] Fossils from sites like the Phosphorites du Quercy in France highlight this phase, where gelocids coexisted with other early ruminants during a period of mammalian turnover linked to climatic shifts.^[12] The Miocene marks a significant radiation of Pecora, with genera like Prodremotherium and Gelocus giving rise to forms resembling early cervids and bovids across Eurasia. Prodremotherium, known from early to middle Miocene localities in Europe and Asia, displays elongated limbs and dental adaptations indicative of browsing habits, evolving toward the more specialized morphologies seen in later pecorans.^[13] Key fossil sites, such as the Siwalik Group in Pakistan, yield diverse assemblages from this period, including primitive pecorans with mixed tragulid-like and advanced pecoran traits, documenting the diversification amid expanding grasslands.^[14] This radiation reflects adaptive responses to environmental changes, with taxa transitioning from forest-dwellers to open-habitat grazers. During the Pliocene and Pleistocene, the fossil record documents the diversification of modern pecoran families, with notable dispersals shaping continental faunas. The earliest Bovidae fossils appear around 18 million years ago in Eurasia, with genera such as Eotragus known from sites in Pakistan and Europe. The family dispersed to Africa during the middle Miocene, with early records from East African sites dating to approximately 16–18 million years ago, marking the initial radiation of horned ruminants in tropical environments.^[15] Cervidae underwent migrations to the Americas approximately 5 million years ago via the Bering land bridge, with the oldest North American records from late Miocene-Pliocene deposits like the Ellensburg Formation in Washington, USA, representing initial colonization by Eurasian lineages.^[16] These events contributed to the establishment of diverse pecoran communities across hemispheres. Extinct pecoran groups, such as Prolibytheridae from the early Miocene of North Africa and Pakistan, play a crucial role in elucidating the evolution of cranial appendages. Prolibytherium, characterized by bizarre, sexually dimorphic frontal structures—including butterfly-shaped ossicone-like projections in males and simpler forms in females—suggests basal homologies with later bovid horns and giraffid ossicones.^[17] Fossils from sites like Gebel Zelten in Libya reveal these appendages as multifunctional, potentially for display and combat, informing models of appendage diversification within Pecora.^[18]

Taxonomy and classification

Higher classification

Pecora constitutes an infraorder within the suborder Ruminantia of the order Artiodactyla, the even-toed ungulates, encompassing all ruminant lineages except the basal Tragulina (chevrotains).^[19]^[1] The infraorder is traditionally subdivided into two superfamilies: Cervoidea, which includes the families Cervidae (deer), Giraffidae (giraffes), and Moschidae (musk deer); and Bovoidea, comprising Bovidae (bovids such as cattle, antelopes, sheep, and goats) and Antilocapridae (pronghorns).^[1] The monophyly of Pecora is robustly supported by molecular evidence, including shared short interspersed nuclear element (SINE) insertions unique to this lineage, such as a specific retroposon integrated in the common ancestor of cows, sheep, deer, and giraffes. Within the broader order Artiodactyla, now recognized as part of the clade Cetartiodactyla (which also incorporates cetaceans), Pecora forms a sister group to Tylopoda (camels and relatives) in the ruminant-tylopod branch, with this combined clade diverging from the non-ruminant Suina (pigs and peccaries) approximately 55 million years ago during the early Eocene.^[20]^[21] Early 20th-century taxonomic schemes often separated Giraffoidea (giraffes and pronghorns) as a distinct superfamily from other pecorans, reflecting uncertainties in morphological character states like cranial appendages and dental features; however, cladistic analyses incorporating both morphological and molecular data in the 1990s confirmed the monophyletic structure of Pecora and resolved inter-superfamily relationships, integrating Giraffoidea within the current framework.^[11]

Families and diversity

The infraorder Pecora encompasses five extant families, representing a total of approximately 200 species of advanced ruminants.^[8] These families exhibit significant biodiversity, with species richness varying widely among them and concentrated in specific geographic regions. The Antilocapridae includes only one species, the pronghorn (Antilocapra americana), which is endemic to open habitats in North America.^[22] The Bovidae is the most species-rich family, comprising about 143 species such as antelopes, cattle, sheep, and goats, and accounting for roughly 70% of Pecora's total diversity.^[23] Bovids dominate in Africa and Asia, particularly in east African savannas where species richness peaks due to adaptive radiations into diverse ecological niches.^[24] The Cervidae contains approximately 50 species of deer, distributed widely across temperate zones of Eurasia, North America, and introduced elsewhere.^[25] The Giraffidae consists of five species: the northern giraffe (Giraffa camelopardalis), reticulated giraffe (G. reticulata), Masai giraffe (G. tippelskirchi), southern giraffe (G. giraffa), and the okapi (Okapia johnstoni), all restricted to African forests and savannas.^[26] Finally, the Moschidae includes seven species of musk deer (Moschus spp.), small-bodied forms inhabiting forested mountains of central and eastern Asia.^[27] Families within Pecora are distinguished primarily by their cranial appendages, which serve functions in defense, display, and combat, setting them apart from the Tragulina's complete lack of such structures.^[28] Bovids possess permanent, unbranched horns consisting of a bony core covered by a keratin sheath, present in both sexes in many species.^[29] In contrast, cervids bear deciduous antlers—branched, bony outgrowths shed and regrown annually, typically in males only.^[29] Antilocaprids feature forked pronghorns that are shed yearly, combining traits of both horns and antlers. Giraffids have skin-covered ossicones, while moschids lack horns or antlers altogether, relying instead on elongated upper canines as tusks.^[28] Recent taxonomic revisions, driven by molecular phylogenetic studies in the 2010s, have refined classifications within Pecora, notably in Cervidae where analyses of mitochondrial and nuclear DNA supported the splitting of the Capreolini tribe to better reflect evolutionary relationships among Old World deer.^[30] Additionally, in August 2025, the IUCN recognized four distinct giraffe species within Giraffidae, based on genetic, morphological, and ecological evidence, elevating former subspecies to full species status.^[31]

Anatomy and physiology

General morphology

Pecora exhibit a characteristic even-toed (cloven) hoof structure, where the weight is borne equally by the third and fourth digits, with the first digit reduced or absent and the second and fifth digits vestigial or lost. This adaptation supports their quadrupedal locomotion, with cursorial limbs featuring elongated metapodials and reduced side toes to enhance speed and endurance across diverse terrains. Body sizes vary widely within the group, ranging from approximately 7–18 kg in musk deer (Moschidae) to over 1,200 kg in adult male giraffes (Giraffidae), reflecting adaptations to different ecological niches.^[32]^[33]^[34] Most Pecora possess cranial appendages arising from the frontal bones, serving functions in defense, display, and intraspecific competition. In Bovidae, true horns are permanent structures with a bony core covered by a keratin sheath that is not shed; in Antilocapridae, pronghorns have similar structures but the keratin sheath is shed annually, while Cervidae feature antlers that are branched, deciduous bony growths shed annually after the breeding season. Giraffidae display ossicones, which are skin-covered bony protuberances present in both sexes, and Moschidae lack such appendages entirely. Sexual dimorphism is pronounced, particularly in body size and the development of these appendages, with males typically larger and more ornamented than females.^[35]^[36] The adult dental formula of Pecora is typically I 0/4, C 0/1, P 3/2–3, M 3/3, lacking upper incisors and canines, which facilitates browsing and grazing. The molars are hypsodont, featuring high crowns with complex folding enamel ridges suited for grinding tough, fibrous vegetation. Sensory adaptations include large eyes positioned laterally for a broad field of view, aiding crepuscular and nocturnal vigilance against predators, and a well-developed vomeronasal (Jacobson's) organ that enhances acute olfaction for detecting pheromones and food sources.^[37]^[38]

Digestive system

Pecora possess a specialized four-chambered stomach that enables efficient fermentation of fibrous plant material, distinguishing them from Tragulina, which have a three-chambered stomach lacking a fully developed omasum.^[39] The rumen, the largest chamber acting as a fermentation vat, holds ingested material where microbes initiate breakdown; the reticulum mixes and softens the contents while aiding in regurgitation; the omasum absorbs water and volatile fatty acids; and the abomasum functions as the true stomach, secreting hydrochloric acid and enzymes for protein digestion similar to monogastrics.^[40] This multi-chambered structure allows Pecora to derive nutrients from cellulose-rich diets that other herbivores cannot efficiently process.^[41] The digestive process relies on microbial symbiosis in the rumen, where diverse communities of bacteria and archaea degrade complex plant polysaccharides, particularly cellulose, into simpler compounds.^[42] These microbes ferment the substrates anaerobically, producing volatile fatty acids (VFAs) such as acetate, propionate, and butyrate as primary end products, which are absorbed through the rumen wall to provide energy to the host.^[43] Protozoa and fungi also contribute by enhancing fiber breakdown and preventing bacterial overgrowth.^[43] Rumination, or cud-chewing, further optimizes digestion by regurgitating partially fermented boluses from the rumen into the mouth for re-mastication, increasing surface area for microbial access and promoting additional breakdown of tough plant fibers.^[40] This cyclic process, involving reticulum contractions to propel material upward, occurs for several hours daily and is essential for maximizing nutrient extraction from low-quality forage.^[40] The system achieves 50-70% digestibility of fibrous plants, with grass fiber often reaching 60-70% and legume fiber 40-50%, depending on plant type and lignin content.^[44] Adaptations include rhythmic forestomach contractions, occurring at rates up to 1-2 per minute in the rumen, which mix contents, expel gas, and facilitate digesta movement.^[45] These contractions maintain optimal pH (around 6.0-7.0) for microbial activity and prevent digestive stasis.^[46] Variations exist among pecoran families; for instance, giraffids exhibit smaller rumens relative to body size compared to grazing ruminants, an adaptation suited to browsing on higher-quality, less fibrous leaves that require less fermentation volume.^[47] VFA production from ruminal fermentation supplies approximately 70% of the animal's energy needs, underscoring the efficiency of this symbiotic process.^[48]

Distribution and habitats

Geographic distribution

Pecora, the infraorder encompassing advanced ruminants such as deer, cattle, and antelopes, are native to every continent except Australia and Antarctica, with their distribution shaped by both natural evolutionary processes and human influences.^[49] The highest species diversity occurs in Africa, particularly within the family Bovidae, which dominates savannas and grasslands, while Eurasia serves as a major hotspot for both Bovidae and Cervidae, reflecting ancient radiations across the Old World.^[50]^[51] The family Bovidae exhibits the broadest range among Pecora, spanning Africa, Asia, Europe, and extending into North America through species like bison and mountain goats.^[50]^[7] Cervidae, including deer and elk, are primarily distributed across the Americas and Eurasia, with species adapted to temperate forests and tundras from North America to Southeast Asia.^[52] In contrast, Giraffidae is strictly endemic to sub-Saharan Africa, where giraffes and okapis occupy fragmented savanna and forest habitats.^[53] Antilocapridae, represented solely by the pronghorn, is confined to open plains and deserts of western North America.^[22] Moschidae, the musk deer, are restricted to mountainous regions of central and eastern Asia, from the Himalayas to parts of China and Vietnam.^[54] Human activities have altered Pecora distributions through introductions and extirpations. Reindeer (Rangifer tarandus), a cervid, have been introduced by humans to Arctic regions beyond their native Eurasian and North American ranges, including Alaska from Siberian stock in the late 19th century.^[55] Similarly, axis deer (Axis axis) were introduced to Hawaii, establishing feral populations that now impact local ecosystems.^[56] Historical migrations, such as those of cervids across Pleistocene land bridges like Beringia, facilitated the colonization of the Americas from Eurasian ancestors.^[57] Conversely, overhunting has led to extirpations of deer species, including large-scale local extinctions of red deer (Cervus elaphus) populations across parts of Europe during the 19th and early 20th centuries.^[58]

Habitat preferences

Pecora exhibit diverse habitat preferences aligned with their ecological roles and family affiliations. Members of the Bovidae family, such as antelopes and cattle, predominantly favor open grasslands and savannas, where expansive areas support grazing on herbaceous vegetation.^[8] In contrast, Cervidae, including deer and elk, prefer forested woodlands and ecotones between forests and grasslands, environments conducive to browsing on leaves, twigs, and shrubs.^[25] Giraffidae, represented by giraffes, occupy semi-arid savannas and open woodlands rich in tall acacia trees, while certain bovid antelopes, like gazelles, adapt to arid deserts and montane regions with sparse vegetation.^[53]^[59] These species span broad altitudinal gradients, from sea level to elevations exceeding 5,000 meters. For example, the Tibetan antelope (Pantholops hodgsonii) inhabits high-altitude alpine steppes and cold deserts on the Tibetan Plateau at 3,700–5,500 meters, where low precipitation and extreme cold prevail.^[60]^[61] Water requirements vary significantly; many pecorans derive hydration from forage, but giraffes exemplify low dependency, obtaining most moisture from foliage and rarely drinking, which enables persistence in water-scarce savannas.^[33] Specialized adaptations facilitate exploitation of these habitats. Forest-adapted cervids possess pelage with disruptive patterns, such as spots on fawns, that mimic dappled sunlight filtering through the canopy for camouflage against predators. Giraffes' extended necks, reaching up to 2.4 meters, allow access to high acacia browse in savannas, minimizing competition with shorter herbivores.^[62] Numerous pecorans, particularly bovids in dynamic ecosystems like the Serengeti, undertake seasonal migrations to follow rainfall-driven forage availability, traveling to ungrazed patches during dry periods.^[8] Habitat loss severely threatens Pecora survival. Deforestation fragments dense woodlands essential for musk deer (Moschus spp.), reducing cover and increasing vulnerability to poaching and predation.^[63] Similarly, conversion of grasslands to cropland for agriculture diminishes foraging areas for grazing bovids, exacerbating population declines across open habitats.^[64]

Behavior and ecology

Pecora exhibit a wide range of social structures, from solitary lifestyles in species like musk deer and okapi to large, dynamic herds exceeding 100 individuals in wildebeest. In cervids such as deer, social groups often form matriarchal family units with multiple generations, while bachelor groups of young males are common outside breeding seasons. Bovids frequently display harem systems where territorial males maintain groups of females and offspring, as seen in impala populations, with similar systems in pronghorn (Antilocapridae). Nursery herds provide collective protection for young.^[65]^[65]^[65] Communication among Pecora involves vocalizations, scent marking, and visual displays to convey information about identity, territory, and threats. Vocal signals include low-frequency grunts in cervids like caribou for maintaining contact within groups and alarm snorts in antelopes such as Thomson's gazelle to alert others to predators. Scent marking is prevalent, with cervids using preorbital glands to deposit individualized odors on vegetation during aggressive or affiliative interactions, while bovids employ interdigital glands for similar spacing and recognition purposes. Visual displays, such as stotting—a stiff-legged bounding leap—in gazelles and oribi, signal fitness to predators or alert conspecifics to danger.^[65]^[66]^[65] Territoriality is prominent in many Pecora, particularly among males who defend ranges using cranial appendages through behaviors like sparring in pronghorns, where individuals clash horns to establish dominance. Alloparenting, though rare, occurs in some bovids, such as gaur where non-maternal females nurse unrelated calves, and in wildebeest nursery herds where group members assist in vigilance and care for young. Human activities have significantly altered social behaviors in domesticated forms like cattle, where intensive farming disrupts natural gregariousness and induces stress responses during handling, leading to manipulated herding patterns that deviate from wild bovid dynamics.^[65]^[67]^[65]

Diet and feeding

Pecorans exhibit primarily herbivorous diets, relying on plant material as their main food source, with distinct feeding categories based on species adaptations. Grazers, such as bison (Bison bison), predominantly consume graminoids and grasses, selectively cropping short vegetation in open grasslands to maximize nutrient intake from fibrous forages.^[8] Browsers, exemplified by giraffes (Giraffa camelopardalis), target leaves, twigs, and shrubs from trees and bushes, often accessing higher foliage unavailable to other herbivores.^[8] Intermediate or mixed feeders, like deer (e.g., Odocoileus spp.), combine grasses, forbs, and browse, adjusting based on availability to balance fiber and protein needs.^[8] Foraging techniques in Pecora are specialized to optimize energy acquisition while minimizing risks. Giraffes employ selective browsing with their prehensile tongues—up to 45 cm long—and mobile lips to strip leaves from thorny acacias, avoiding spines and targeting nutrient-rich parts.^[68] In contrast, grazing species like bison use herd-based strategies, moving collectively across landscapes to lightly crop grasses, which reduces individual predation exposure and promotes efficient patch use.^[69] Across Pecora, daily dry matter intake typically ranges from 2% to 3% of body weight, supporting high-fiber digestion and maintaining metabolic demands.^[70] Seasonal variations profoundly influence Pecoran diets and behaviors, driving adaptive shifts to ensure nutritional sufficiency. Many species undertake migrations to follow fresh growth pulses, as seen in wildebeest (Connochaetes spp.), which travel vast distances across the Serengeti-Mara ecosystem in response to rainfall-triggered grass regrowth, synchronizing with calving to exploit protein-rich vegetation.^[71] In temperate or harsh winters, fallback foods become critical; for instance, deer resort to browsing bark, twigs, and woody stems when herbaceous plants are unavailable, providing essential but lower-quality energy.^[72] Rumen microbial communities adapt dynamically to these diet changes, altering volatile fatty acid profiles—for example, increasing propionate production in grain-fed domestic ruminants to enhance energy yield from starch-rich feeds.^[73] Intraspecific and interguild competition shapes feeding ecology, particularly for shared resources. Within grazing niches, Pecorans compete with equids (e.g., zebras and horses) for high-quality grasses, where equids' faster intake rates can displace ruminants from optimal patches, influencing coexistence through niche partitioning.^[74] Nutritional bottlenecks, such as sodium deficiency in low-mineral habitats like certain inland pastures, further constrain diets, leading to reduced intake and performance unless supplemented naturally via salt licks or geophagy.^[75] These factors underscore the interplay between foraging strategies and environmental pressures in maintaining Pecoran nutritional ecology.

Reproduction and life history

Mating systems

Pecora exhibit predominantly polygynous mating systems, where a single male mates with multiple females, a pattern driven by sexual dimorphism and resource availability. This includes resource-defense polygyny, in which territorial males defend areas rich in forage or water to attract females, as seen in antelopes such as the waterbuck (Kobus ellipsiprymnus). Female-defense strategies involve males herding or tending groups of females, exemplified by bison (Bison bison), where males form temporary harems during the breeding season. Lekking occurs in some species, particularly gazelles and antelopes like the kob (Kobus kob) and topi (Damaliscus lunatus), where males aggregate in display arenas to perform courtship rituals without defending resources, allowing females to select mates based on displays.^[76] Courtship in Pecora typically involves elaborate displays using sexual appendages, pheromonal signals, and synchronization with female estrus cycles, which last 18-24 days in most species. Male deer engage in antler clashes during the rut to establish dominance and attract females, while sheep and goats feature horn-locking battles among rams to resolve competition. Pheromones released via urine or glandular secretions play a key role in signaling receptivity, particularly during the short estrus phase of 12-36 hours. These behaviors are influenced by social hierarchies, as detailed in discussions of social behavior.^[76]^[77]^[78] Sexual selection in Pecora favors larger males with superior horns or antlers, which confer advantages in male-male combat and female choice, leading to pronounced dimorphism across families like Cervidae and Bovidae. In deer and antelopes, victorious males secure more mating opportunities, enhancing their reproductive success through traits that signal genetic quality. Infanticide by incoming males occurs in some bovids, such as bighorn sheep (Ovis canadensis), to hasten the return of females to estrus by eliminating non-kin offspring, thereby accelerating the male's breeding window.^[76]^[79] Domestication has significantly altered natural mating systems in Pecora, particularly in cattle (Bos taurus), where artificial insemination bypasses polygynous competition and allows selective breeding for desirable traits. This technique, widely adopted since the mid-20th century, reduces disease transmission and enables use of superior sires across herds but disrupts traditional harem or territorial dynamics.^[80]

Development and lifespan

Pecorans exhibit viviparous reproduction with a synepitheliochorial placenta, where embryonic development begins in the oviduct following fertilization. In representative Bovidae species such as cattle (Bos taurus), the zygote reaches the 16-cell stage by day 4 and enters the uterus, forming a blastocyst by day 7 and hatching around days 9–10. The conceptus then transitions from spherical to ovoid (days 12–13), tubular (days 14–15), and finally filamentous (days 16–17), elongating to up to 150 mm to enhance nutrient exchange with uterine histotroph. In sheep (Ovis aries), a close relative, the morula enters the uterus by days 3–4, becomes a blastocyst by day 6, hatches by days 8–9, and reaches the filamentous stage by days 12–17, extending to 25 cm. This elongation is driven by trophectoderm proliferation and is crucial for implantation, which begins around days 16–20 in cattle and days 16–18 in sheep. Maternal recognition of pregnancy occurs via interferon tau secretion from the trophectoderm (days 13–17 in cattle, days 10–21 in sheep), preventing luteolysis and sustaining the corpus luteum.^[81] In Cervidae, such as red deer (Cervus elaphus), early embryonic development mirrors Bovidae patterns but includes embryonic diapause in some species like roe deer (Capreolus capreolus), where the blastocyst remains dormant for 4–5 months before resuming growth. Without diapause, as in mule deer (Odocoileus hemionus), the embryo develops continuously, with the abomasum appearing at 22% of gestation and gastric structures forming by mid-gestation. Giraffidae, including giraffes (Giraffa camelopardalis), show similar initial stages but with extended timelines due to prolonged gestation.^[82]^[83] Gestation lengths in Pecora vary significantly by family and body size, reflecting adaptations to ecological niches. In Bovidae, periods range from 120–150 days in small duikers to 300–330 days in African buffalo (Syncerus caffer), with domestic cattle at approximately 280 days, sheep and goats at 150 days, and pronghorn (Antilocapra americana, Antilocapridae) at 250 days. Cervidae gestations average 200 days, as in white-tailed deer (Odocoileus virginianus). Giraffidae exhibit notably longer periods of 425 days in giraffes and about 450 days in okapi (Okapia johnstoni), correlating with larger body masses and slower intrauterine growth rates (scaling as body mass^0.13). Shorter gestations in non-giraffid Pecorans (<300 days) enable higher reproductive rates and seasonal breeding, a key innovation distinguishing them from Tragulina.^[84]^[85]^[86] Birth typically results in precocial young capable of standing shortly after delivery, though giraffe calves drop from heights up to 2 meters. Postnatal development involves rapid rumen maturation; in calves and lambs, microbial colonization begins within hours, with functional fermentation by 4–6 weeks, supported by milk rich in volatiles like butyrate. Weaning occurs at 2–6 months, depending on species. Sexual maturity is reached earlier in females (average 22 months) than males (28 months) across Pecora, with variations: smaller Bovidae like goats mature at 6–12 months, while larger cervids like red deer reach maturity at 16–24 months for females and 24–36 months for males. Cranial appendages, such as horns or antlers, often develop around puberty, signaling maturity.^[81]^[87] Lifespans in Pecora scale with body mass (as mass^0.15) and vary widely, influenced by predation, habitat, and captivity. In Bovidae, wild American bison (Bison bison) live up to 33 years, African buffalo to 29.5 years, and domestic cattle average 20 years in captivity. Cervidae show 10 years wild for white-tailed deer but up to 26.8 years for red deer. Giraffidae have extended longevities, with giraffes reaching 36 years wild or captive and okapi up to 33 years in captivity, exceeding expectations for their size due to low metabolic rates. Overall, wild lifespans are shorter than captive ones, with males often dying younger due to sexual dimorphism and competition.^[88]^[86]

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[PDF] The Ruminantia (Mammalia, Cetartiodactyla) from the Oligocene to ...
ordered from the Early Oligocene to the Early Miocene. I Introduction ... pecoran genera Gelocus and Prodremotherium in having a deep external ...
[14]
[PDF] oligocene and early miocene ruminants - Palaeontologia Electronica
The slightly younger site Z 127 has three species associated with the primitive pecoran Bugtimeryx pilgrimi. In contrast, tragulids apparently do not make ...
[15]
[PDF] The fossil record and evolution of Bovidae: State of the field
18Ma, Pakistan), and an unnamed species from the Vihowa Formation. (ca.18Ma, Pakistan) have been tentatively pro- posed as early or stem Bovinae (Solounias and.
[16]
[PDF] A new early occurrence of Cervidae in North America from the ...
We here describe a new cervid occurrence from the Miocene-Pliocene Ellensburg Formation in Central Washington. This new cervid occurrence is below a date of 5.0 ...Missing: migration Bering
[17]
Sexual dimorphism of the frontal appendages of the early Miocene ...
Prolibytherium magnieri Arambourg, 1961, is a bizarre-looking pecoran discovered in lower Miocene strata from Libya and Egypt (Gebel Zelten and Wadi Moghara ...Missing: pecora | Show results with:pecora
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Systematics and Evolution of the Miocene Three-Horned ...
Dec 2, 2015 · This phylogenetic reconstruction establishes an evolutionary hypothesis that implies a basal homology of the appendages of Prolibytherium and ...
[19]
A genomic view with mitochondrial 16S rRNA and cytochrome b gene
Tragulina is represented by a single extant family, Tragulidae, which includes three genera (Grubb, 1993). The other infra order, Pecora, comprises five living ...
[20]
Pattern and timing of diversification of Cetartiodactyla (Mammalia ...
The MMCO was directly followed by a drastic global cooling, from 15 to 13 Ma, marking a transition from a greenhouse world to an “icehouse-world” [99,117].
[21]
Phylogenetic relationships of artiodactyls and cetaceans as ...
Cetartiodactyla includes five major lineages: Ruminantia, Tylopoda, Suina ... Calculation of divergence dates suggests that the Cetartiodactyla could have ...
[22]
Antilocapridae (pronghorn antelope) - Animal Diversity Web
A single modern species is found in the family Antilocapridae, but the group has a fossil record going back to the Miocene.
[23]
Family Bovidae - Cattle, antelopes, and goats - Ultimate Ungulate
Aug 19, 2024 · The Bovidae are the most diverse group of living ungulates with 143 Recent species (over 55% of modern ungulates).<|control11|><|separator|>
[24]
Bovidae (antelopes, cattle, gazelles, goats, sheep, and relatives)
Bovid species richness is highest in the savannah of east Africa and the family has radiated to fill an enormous variety of ecological niches resulting in a ...
[25]
Cervidae (deer) | INFORMATION - Animal Diversity Web
The family Cervidae , commonly referred to as "the deer family", consists of 23 genera containing 47 species, and includes three subfamilies: Capriolinae ( ...
[26]
https://giraffeconservation.org/state-of-giraffe/
[27]
https://inaturalist.org/taxa/42140-Moschidae
[28]
Molecular and Morphological Phylogenies of Ruminantia and the ...
The evolutionary radiation of Pecora occurred at the Early/Late Oligocene transition, and Pecoran families diversified and dispersed rapidly during the Early ...
[29]
Horns and Antlers - Animal Diversity Web
Antlers are branched, bony structures shed annually, while horns are permanent, unbranched, with a bony core and keratin sheath.Missing: Pecora | Show results with:Pecora
[30]
The systematics of the Cervidae: a total evidence approach - PeerJ
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Siberian musk deer - Ultimate Ungulate
Weight: 11-18 kg / 24-40 lb. The overall colouration of animals varies from a light yellowish brown to almost black, although a dark brown is most common. The ...
[32]
Giraffa camelopardalis (giraffe) - Animal Diversity Web
At birth, giraffe calves are 2 m tall from the ground to the shoulders. Newborn giraffes weigh 50 to 55 kg. Both male and female giraffes have a spotted coat.
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How the even‐toed ungulate vertebral column works
Cursorial form is an adaptation to life in open landscapes, characterized by maximum speeds and endurance. Its distinctive features are a shallower trajectory ( ...
[34]
Evolution of ruminant headgear: a review - PMC - NIH
The horns, ossicones and antlers of ruminants are familiar and diverse examples of cranial appendages. We collectively term ruminant cranial appendages ...
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Jul 6, 2011 · The horns, ossicones and antlers of ruminants are familiar and diverse examples of cranial appendages. We collectively term ruminant cranial ...
[36]
Eruption of posterior teeth in the maxilla and mandible for age ...
The following dental formula is almost universal in Pecora. Pecora have three incisors, a canine, three premolars, and three molars in a quadrant of the ...
[37]
Key innovations in ruminant evolution: a paleontological perspective
Oct 21, 2013 · Then, Hoplitomerycidae could be placed somewhere at the basal divergence of Pecora (Mazza 2013) and their horn-like appendages could have been ...
[38]
Fig. 1. Schematic representation of the stomach of tragulids (top) and...
The rumen, the first compartment of the ruminants' digestive tract, is a fermentation chamber that hosts a diverse and dynamic microbial community referred to ...
[39]
The ruminant digestive system - University of Minnesota Extension
Ruminant stomachs have four compartments: the rumen, the reticulum, the omasum and the abomasum. Rumen microbes ferment feed and produce volatile fatty acids, ...
[40]
Elucidating the Evolutionary Relationships among Bos taurus ... - NIH
Like other ruminants, Bos taurus has four stomach-like compartments, as opposed to the single stomach in many other mammals. These four chambers allow digestion ...
[41]
Rumen Microbial Predictors for Short-Chain Fatty Acid Levels and ...
Oct 31, 2022 · In the rumen, microbial fermentation enables the conversion of plant fibers into small molecule products, particularly short-chain fatty acids, ...
[42]
International Symposium on Ruminant Physiology: Rumen fungi ...
Bacteria, protozoa, and fungi in the rumen produce enzymes that degrade and ferment plant cell carbohydrates, proteins, and lipids to volatile fatty acids.Missing: Pecora | Show results with:Pecora
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Plant limitations to fiber digestion and utilization - PubMed
Ruminants digest 40-50% of legume fiber and 60-70% of grass fiber. Some fiber cannot be digested no matter how long it remains in the rumen.
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Rumen motility in sheep and cattle given different diets
Mean rates of contraction were similar for sheep and cows, with A sequences averaging 1.W/min and B sequences averaging 0.62/min. Sheep and cows differed in.
[45]
Microbial and Animal Limitations to Fiber Digestion and Utilization
Four major factors regulate ruminant fiber digestion: 1) plant structure and composition, which regulate bacterial access to nutrients; 2) nature of the ...Fibrolytic Microorganisms · Microbial Adhesion And... · Enhancement Of Fiber...<|separator|>
[46]
Relationships between body mass and combined intestine mass ( ),...
When compared to grazing species of a similar body mass, a giraffe's rumen is small (Mitchell et al., 2015) , emphasising their behavioral adaptations for ...
[47]
Relationships between rumen volatile fatty acid concentrations and ...
Introduction. Volatile fatty acids (VFA) account for 60–70% of metabolizable energy supply in ruminants (Armentano, 1992, Van Soest, 1982) ...
[48]
Bovid | African & Asian Antelopes, Wild Cattle & Goats | Britannica
During the Pleistocene Ice Age, while most of the Eurasian tropical savanna fauna became extinct, Africa remained the main refuge of Plio-Pleistocene mammals.Missing: Pecora Bering
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Pecora - Variety of Life
Jul 24, 2018 · Contains: Giraffoidea, Bovidae, Cervidae, Palaeomerycidae. The Pecora are the clade containing most modern ruminants except for the chevrotains.
[50]
Bovidae - an overview | ScienceDirect Topics
Bovids are found across all of mainland Africa and in 30 countries in Europe, the Middle East, and Asia. Four subfamilies exist only in the African continent ...
[51]
Cervidae - deer, elk, moose - New Hampshire PBS
There are 55 species in this family, and they are found in all parts of the world, except for Sub-Saharan Africa, Australia, and Antarctica.
[52]
Natural History: Giraffe - Center for Biological Diversity
Range: Giraffes are found in fragmented habitats scattered throughout sub-Saharan Africa. Breeding: Giraffe pregnancy usually lasts about 15 months, with two- ...
[53]
Moschidae - an overview | ScienceDirect Topics
The four species of musk deer in the Moschidae are found in central Asia, the Himalayan plateau, and east through parts of China and Vietnam. A primitive deer, ...
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Reindeer Introgression and the Population Genetics of Caribou in ...
May 19, 2014 · Reindeer were introduced into Alaska from domestic Siberian herds in 1891 in an attempt to introduce an alternate food source and ...
[55]
Removing Hawaii's illegally introduced axis deer
The helicopter pilot agreed to provide 500 hours of flight time to locate and eradicate the Hawaii Island deer population in restitution.<|separator|>
[56]
Pleistocene megafauna from eastern Beringia: Paleoecological and ...
Late Pleistocene eastern Beringia is a model paleo-ecosystem for the study of potential and realized species interactions within a diverse mammalian fauna.
[57]
Population dynamics of a natural red deer population over 200 ...
This short period led to massive reductions in red deer population sizes and large‐scale extinctions of red deer populations due to poaching and unregulated ...
[58]
Effect of locomotor preference on the evolution of mitochondrial ...
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[59]
Pantholops hodgsonii (chiru) | INFORMATION - Animal Diversity Web
Chiru are most often found along the alpine steppe in northwest Tibet and China, where annual precipitation is less than 16 inches and elevations are between ...
[60]
The Tibetan Antelope Population Depends on Lakes on the Tibetan ...
Nov 22, 2023 · The Tibetan antelope is a wild animal unique to the Tibetan Plateau, mainly distributed across 3700–5500 m alpine desert grassland and plateau ...
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How did the giraffe get its long neck? | Penn State University
May 17, 2016 · "To achieve their extraordinary length, giraffe cervical vertebrae and leg bones have evolved to be greatly extended," Cavener said. "At least ...
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Conservation status and causes of decline of musk deer (Moschus ...
As well as deforestation, forest fires may have a substantial impact upon musk deer populations, by degrading their habitats. As stated earlier, the musk deer ...
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Beef and Bison Can Help in Fight Against Climate Change
Jul 20, 2020 · Due to market forces, many of these lands are at risk of being converted to cropland including corn and soybeans. When grasslands are plowed ...
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[PDF] The Behaviour of Ungulates and its relation to management
51. 52. 53. The social organization of the roan antelope Hippotragus equinus and its influence on the spatial distribution of herds in the Kruger National.
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Pre-orbital gland opening during aggressive interactions in rusa ...
The opening of the preorbital gland in deer serves as a visual communication and has been linked to a wide variety of behavioural situations.
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Sep 28, 2025 · Allonursing is the nursing of non-filial offspring by lactating females. It is a rare form of alloparental care and remains virtually ...
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Getting Started With Bison Ranching - SDSU Extension
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[PDF] Dry Matter Intake for Beef Cattle
Dry matter intake of diets developed with a high percentage of grains may only be 1 - 1.5 percent of body weight. This type of diet is limit-fed. Cattle fed ...
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Most people think that the Wildebeest Migration only takes place between July and October, but it's actually an ever-moving, circular migration with various but ...
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How to manage deer damage on trees and other plants
Food is scarce during winter and deer will eat any available vegetation including dried leaves, sedges, grasses, mushrooms and woody plants.
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Effects of High-Grain Diet on Performance, Ruminal Fermentation ...
Aug 30, 2024 · In other ruminants, evidence suggests that long-term feeding of high-grain diets can change rumen fermentation patterns and microbial ...
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a test of the nutritional model of equid/bovid competition ... - PubMed
Data presented here demonstrate that compared to similarly sized grazing ruminants (bovids), hind-gut fermenters (equids) have higher rates of food intake ...
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Sodium deficiency of grazing cattle in Papua New Guinea
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Evolution of ungulate mating systems: Integrating social and ...
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Oestrous cycles in Bos taurus cattle - PubMed
The oestrous cycle in cattle lasts for 18-24 days. It consists of a luteal phase (14-18 days) and a follicular phase (4-6 days).
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Reproductive Physiology of Sheep - Management and Nutrition
Ewes are seasonally polyestrous, cycling every 16–17 days during the breeding season. · The duration of estrus (~30 hours) is influenced by the breed and age of ...
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Infanticide as Sexual Conflict: Coevolution of Male Strategies and ...
The original sexual selection hypothesis arguing that infanticide improves male mating success by accelerating the return of females to fertilizable condition ...
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Artificial Insemination in Cattle: Efficiency, Opportunities and ...
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Deer Development - UNSW Embryology
Dec 24, 2019 · The average prenatal development period for deer (Mule) is 200 days. Deer are typically a uniparental species, where the offspring (fawn) is only cared for by ...
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Artificial Reproductive Technology (ART) Applied to Female Cervids ...
Oct 11, 2021 · Diapause is a period in early neonatal development in some species during which an embryo is suspended at the blastocyst stage, and the ...
[83]
Table: Approximate Gestation Periods-Merck Veterinary Manual
Gestation period is 58–72 d from breeding at unknown stage of estrus; from day of ovulation (which can be determined by progesterone or luteinizing hormone ...
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Old World Ruminant Morphophysiology, Life History, and Fossil ...
The omasum of pecoran ruminants (which is absent in tragulids) and shorter gesta- tion periods in non giraffid crown pecorans (as opposed to giraffids) ...
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[PDF] A long life among ruminants: giraffids and other special cases
Giraffids (giraffe and okapi) have high longevities, long gestation periods, and low intra-uterine growth rates. Anoa and some small ruminants also have high ...
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Sexual maturity and shape development in cranial appendages of ...
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Longevity Records: Life Spans of Mammals, Birds, Amphibians ...
American bison can live up to 33 years, while Addax can live up to 25.3 years. Pronghorn can live up to 11.8 years.