Fact-checked by Grok 2 weeks ago

Elephas

Elephas is a genus of elephants in the family Elephantidae, subfamily Elephantinae, characterized by high-crowned molars adapted for grazing and comprising one extant species, the Asian elephant (Elephas maximus), along with numerous extinct species that thrived during the Pliocene and Pleistocene epochs.^[1] The genus originated in Africa approximately 3 to 4 million years ago during the Pliocene, from where it dispersed to Asia in the mid-Pliocene, giving rise to the lineage leading to the modern Asian elephant, and underwent a second dispersal in the late Pliocene to Asia and Europe, resulting in additional extinct lineages.^[1]^[2] The Asian elephant (E. maximus), the sole surviving member of the genus, is the largest terrestrial mammal in Asia, with adult males reaching shoulder heights of 240–300 cm and weights of 3,500–6,000 kg, while females are smaller at 195–240 cm and 2,000–3,500 kg; it features gray skin sparsely covered in hair, a trunk ending in a single finger-like projection for grasping, and in males, long curved tusks, though females and calves of both sexes are typically tuskless.^[3] This species inhabits diverse tropical habitats including grasslands, dry and wet deciduous forests, and scrublands across South and Southeast Asia, from elevations of sea level to 3,000 m, though populations have been fragmented by habitat loss.^[3] Recognized subspecies include the Sri Lankan (E. m. maximus), mainland Indian (E. m. indicus), and Sumatran (E. m. sumatranus) elephants, with the latter critically endangered.^[3] Fossil evidence indicates that Elephas once had a broader distribution, with species present in Africa until about 11,500 years ago and in Europe until the end of the Pleistocene, including at least ten recognized extinct species from the middle Pliocene onward, such as early forms from the Ekora beds in Kenya.^[1]^[2] The genus is closely related to the extinct mammoths (Mammuthus) and the African elephants (Loxodonta), sharing a common ancestry within Elephantidae, and genomic studies confirm Elephas as a distinct lineage endemic to Asia in modern times, with extinct forms contributing to understanding elephantid evolution and diversification.^[1]^[4] Today, the Asian elephant is classified as endangered by the IUCN, with a wild population estimated at 41,410–52,410 individuals as of 2021, though a 2025 census in India (home to ~60% of the global population) reported 22,446 individuals, suggesting further decline; it faces threats from poaching, habitat fragmentation, and human-elephant conflict.^[5]^[6]

Physical Description

Anatomy and Morphology

Elephas species display the quintessential proboscidean body plan, characterized by a robust, barrel-shaped torso supported by four thick, columnar legs that function as pillar-like structures to distribute the animal's substantial mass and facilitate movement across varied terrains. These legs feature straight alignment of bones, with minimal flexion at joints, enabling efficient weight-bearing while minimizing energy expenditure during locomotion. The trunk, an elongated fusion of the nose and upper lip, serves as a versatile appendage comprising approximately 40,000 muscle fascicles organized into horizontal and vertical bands, culminating in a single finger-like extension at the tip for grasping and manipulating objects with precision.^[7]^[8]^[9] Prominent among the external features are the large, fan-shaped ears, which are richly vascularized and act as primary thermoregulatory organs by increasing surface area for convective and radiative heat loss, particularly in tropical habitats. The skeletal framework includes a distinctive dome-shaped skull in extant species like Elephas maximus, with a compressed rostrocaudal profile and elevated occipital region that accommodates the attachment of powerful neck muscles. Tusks, modified elongated upper incisors, emerge from the premaxillae and exhibit species-specific curvature—straighter and more downward-directed in Elephas compared to the outward-flaring form in related genera—serving roles in foraging, defense, and intraspecific display.^[10]^[11]^[12] Dentally, Elephas is equipped with hypsodont molars featuring a lamellar structure of tightly packed, enamel-covered plates that progressively wear to form grinding surfaces ideal for processing abrasive vegetation; these molars possess a higher total number of plates than those in Mammuthus, reflecting adaptations to diverse dietary demands within the genus. The skin of living Elephas maximus is notably thick—up to several centimeters in places—and deeply wrinkled, enhancing flexibility and creating folds that trap moisture to aid in evaporative cooling, while sparse, coarse hairs provide minimal insulation.^[11]^[13]^[14]

Size and Variations

The Asian elephant (Elephas maximus), the sole extant species in the genus Elephas, exhibits considerable size variation influenced by sex and regional populations. Adult males typically reach a shoulder height of 2.4–3.0 m and a body length of 5.5–6.4 m, with weights ranging from 3,500 to 6,000 kg, while females are smaller, attaining shoulder heights of 1.95–2.4 m, similar body lengths, and weights of 2,000–3,500 kg.^[3]^[15] Sexual dimorphism is pronounced, with males averaging about 30–40% larger in linear dimensions and up to 50% heavier than females, supporting their role in intra-sexual competition.^[3] This size disparity is facilitated by anatomical adaptations such as columnar legs, which provide structural support for the greater body mass in males.^[3] Tusks in E. maximus are sexually dimorphic and primarily occur in males, consisting of elongated upper incisors that average 2–3 m in length and can weigh up to 45 kg per tusk in exceptional cases.^[16]^[17] Females often lack prominent tusks, though some populations exhibit small tushes (underdeveloped tusks) protruding less than 5 cm, and tusklessness is common in certain groups due to selective pressures.^[16]^[3] Morphological variations are evident among the recognized subspecies of E. maximus. The Sri Lankan elephant (E. m. maximus) is the largest, with males achieving shoulder heights up to 3.5 m and weights exceeding 5,000 kg, featuring darker skin and larger ears compared to continental forms.^[18] In contrast, the Sumatran elephant (E. m. sumatranus) is notably smaller, with shoulder heights of 2–3.2 m and weights of 2,000–4,000 kg, possessing relatively larger ears and an additional pair of ribs.^[18]^[15] The mainland Indian elephant (E. m. indicus) occupies intermediate sizes.^[18]

Distribution and Habitat

Current Range

The Asian elephant (Elephas maximus), the sole extant species in the genus Elephas, is endemic to South and Southeast Asia, where it occupies fragmented habitats across 13 range countries: Bangladesh, Bhutan, Cambodia, China, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Sri Lanka, Thailand, and Vietnam.^[19]^[20] India hosts the largest population, estimated at 22,446 individuals (95% CI: 18,255–26,645) as of the 2025 synchronized census using DNA-based methods, representing over 60% of the global total.^[21]^[22] Smaller but significant populations persist in Sri Lanka (around 6,000–7,000 as of 2022–2025 estimates), Myanmar (estimated 1,000–1,850), and Thailand (about 3,000–4,000), while numbers in countries like China, Nepal, Bhutan, and Bangladesh are critically low, often fewer than 500 each.^[23]^[24] In Indonesia, populations are concentrated on Sumatra and Borneo, with fewer than 1,800 on Sumatra alone.^[23] The global wild population is estimated at 40,000–50,000 individuals as of 2024, though this figure is approximate due to challenges in surveying dense forests and ongoing declines, with recent national censuses indicating potential downward revisions.^[20]^[19] These populations are highly fragmented, with over 70% occurring outside protected areas, primarily due to habitat loss from agriculture, infrastructure development, and urbanization, leading to increased human-elephant conflict.^[19] Asian elephants prefer habitats with access to water, including tropical and subtropical moist broadleaf forests, dry deciduous forests, grasslands, scrublands, and even agricultural landscapes where they raid crops.^[19] They are adaptable to elevations up to 3,000 meters but require areas rich in browse and foraging opportunities.^[23] Three main subspecies are recognized based on geographic isolation and morphological differences: E. m. indicus (Indian elephant), distributed across mainland Asia from India to Indochina; E. m. maximus (Sri Lankan elephant), confined to Sri Lanka; and E. m. sumatranus (Sumatran elephant), found exclusively on Sumatra, Indonesia.^[23] The Bornean elephant (E. m. borneensis) is sometimes treated as a fourth subspecies, limited to northern Borneo in Malaysia and Indonesia, with a population of around 1,000.^[23]

Fossil Distribution

The genus Elephas originated in sub-Saharan Africa during the Pliocene epoch, approximately 3 to 4 million years ago, evolving from earlier elephantid ancestors at the Miocene-Pliocene boundary. Early fossils, such as those of Elephas ekorensis, have been recovered from East African sites like the Ekora Formation in Kenya, dating to the early to mid-Pliocene (approximately 5–4.2 million years ago). From Africa, the genus dispersed northward, migrating into Eurasia around 3–4 million years ago via land bridges across the Levant, marking the beginning of its expansion into Asian and European ecosystems.^[2]^[25]^[26] Fossil evidence of Elephas in Asia is abundant, particularly from the Indian subcontinent, where the Siwalik Group's Upper Siwalik deposits (Pliocene to early Pleistocene) have yielded extensive remains of primitive species, including Elephas hysudricus, indicating a broad distribution across northern India and Pakistan. In China, early Elephas fossils from Late Pliocene to Pleistocene contexts, such as those in the Nihewan Basin, document the genus's northward spread into East Asia. The Middle East, especially the Levant region, preserves key migration-route sites; for instance, Middle Pleistocene remains of Elephas cf. hysudricus from Ma'ayan Baruch in Israel and 'Ain Soda in Jordan highlight the corridor between Africa and Asia.^[27]^[28]^[29] Extinct Elephas species exhibited diverse ranges tied to Pleistocene environments. Elephas hysudricus, an early form ancestral to modern Asian elephants, was widespread in the Indian subcontinent during the Pliocene, with fossils concentrated in the Siwalik foothills. The dwarf species Elephas falconeri (now often classified under Palaeoloxodon) occupied Mediterranean islands like Sicily, Malta, and Crete during the Middle to Late Pleistocene, adapting to insular conditions through extreme size reduction. In North Africa, Elephas iolensis ranged across savanna habitats in the Late Pleistocene, representing a terminal African branch of the genus before its regional extinction.^[29]^[30]^[31] By the Late Pleistocene, approximately 12,000 years ago, Elephas species had become widespread across Europe, Asia, and parts of Africa but underwent rapid extinction, particularly in Europe and North Africa, driven by post-glacial climatic shifts and human overhunting. This event eliminated all continental Elephas populations outside South and Southeast Asia, where the sole surviving species, E. maximus, endured amid contracting habitats—a stark contrast to the genus's earlier expansive range.^[32]^[1]

Behavior and Ecology

Diet and Foraging

Elephas species, exemplified by the living Asian elephant (Elephas maximus), are strictly herbivorous megaherbivores that consume a diverse array of vegetation to meet their high energetic demands.^[33] Their diet primarily consists of browse such as leaves, twigs, bark, and fruits, supplemented by grazing on grasses, herbs, and sedges, with over 100 plant species recorded in some populations.^[34] Adults ingest approximately 100–200 kg of plant material daily, representing about 1.5–3% of their body weight, to compensate for the low digestibility of fibrous foods.^[35] This substantial intake supports their large size and activity levels, with foraging occupying 12–19 hours per day.^[36] Foraging strategies in Elephas rely on specialized anatomical adaptations for efficient food acquisition. The prehensile trunk serves as the primary tool for plucking leaves, stripping bark, and grasping grasses, enabling precise manipulation without teeth.^[16] Tusks, present mainly in males, assist in uprooting plants, debarking trees, and digging for roots or water during scarcity.^[34] Individuals travel 5–20 km daily in search of food patches, adjusting paths based on resource availability and covering larger distances in fragmented habitats.^[37] Dietary habits exhibit pronounced seasonal variations, particularly in tropical monsoon environments. During wet seasons, E. maximus favors nutrient-rich grasses and herbs, which are abundant and support higher protein intake.^[38] In dry seasons, they shift toward browse like leaves and bark from deciduous trees, as grasses desiccate, necessitating longer foraging bouts near reliable water sources.^[36] Water requirements are substantial, with adults consuming 80–200 liters daily—roughly 2–3% of body weight—to aid digestion and thermoregulation, often drinking multiple times per day when available.^[39] The digestive physiology of Elephas is adapted for processing cellulose-rich forage through hindgut fermentation. As monogastric hindgut fermenters, they rely on microbial communities in the enlarged cecum and colon to break down fibrous material, extracting volatile fatty acids for energy, though overall digestibility remains low at 40–50%.^[40] Young calves engage in coprophagy, consuming adult feces to inoculate their gut with essential microbes, facilitating the establishment of a functional fermentative flora for plant digestion.^[41] As keystone species, Elephas profoundly influence forest ecosystems through their foraging activities. By felling trees and browsing selectively, they create gaps that promote understory regeneration and habitat heterogeneity, enhancing plant diversity.^[42] Additionally, they disperse seeds of large-fruited plants over distances up to 5 km via endozoochory, with intact seeds passing through the gut to germinate in nutrient-enriched dung piles, sustaining tropical forest composition.^[43] Elephas species, particularly the Asian elephant (Elephas maximus), exhibit a matriarchal social structure where females form stable family units led by the oldest and most experienced female, known as the matriarch. These herds typically consist of 10 to 20 related females and their offspring, including immature males, providing protection and support through cooperative behaviors. Adult males, upon reaching maturity around 14-15 years, generally leave the family unit to live solitarily or join loose bachelor groups of 2-5 individuals, occasionally forming larger temporary aggregations during resource abundance.^[16]^[44] Communication within these groups is multifaceted, relying on vocal, tactile, visual, and chemical signals to maintain bonds and coordinate activities. Low-frequency infrasonic rumbles, produced through the larynx and often amplified by the trunk, allow individuals to convey information over distances up to 10 km, facilitating reunions in dense habitats. Tactile interactions, such as trunk touches and entwining, reinforce social affiliations, while visual cues like ear flapping signal emotions or warnings; the trunk's dexterity enables precise contact, as adapted from its anatomical versatility. Chemical signals from temporal gland secretions, especially prominent in males, provide olfactory cues about reproductive status and identity, detectable by others in the vicinity.^[45]^[46]^[47] Social cohesion is further strengthened by allomothering, where non-maternal females in the herd assist in calf protection, nursing, and guidance, significantly improving juvenile survival rates by distributing caregiving responsibilities. In males, periods of musth—a hormonal surge marked by elevated testosterone—induce heightened aggression and dominance displays, such as charging or trunk swinging, which can alter group interactions and prioritize mating opportunities over typical hierarchies. Group dynamics often follow a fission-fusion pattern, with herds splitting and reforming based on resource availability, leading to temporary large aggregations at water sources; conflicts are resolved through dominance displays and ritualized postures rather than lethal fights, maintaining overall stability.^[48]^[49] Increasing overlap between human settlements and elephant ranges has intensified conflicts, manifesting in crop-raiding behaviors where herds enter agricultural areas to forage on nutrient-rich crops like rice and sugarcane, resulting in economic losses and retaliatory actions against elephants. These interactions disrupt traditional social structures, as herds adapt by becoming more nocturnal or evasive, yet they underscore the need for habitat corridors to mitigate such pressures.^[50]^[51]

Reproduction and Life Cycle

Mating and Reproduction

The genus Elephas exhibits a promiscuous or polyandrous mating system, in which females typically mate with multiple males during estrus to maximize genetic diversity and reproductive success.^[52] Males, often solitary or in loose bachelor groups outside of musth periods, roam widely in search of receptive females, detecting estrus through pheromones released via urine and temporal gland secretions.^[53] Competition among males intensifies during musth, a periodic physiological state marked by elevated testosterone levels, aggressive displays such as ear flapping and trunk curling, and dominance contests involving tusk thrusts, charges, and vocalizations to secure mating priority.^[54] Larger, older males in peak musth generally dominate these interactions, though female choice plays a role in final mate selection based on male vigor and health signals.^[55] Breeding in Elephas species is largely aseasonal but shows peaks in certain regions tied to environmental cues like resource availability, with conceptions often occurring during the dry season when day lengths increase and forage quality supports gestation demands.^[56] Females reach sexual maturity between 10 and 15 years of age, after which they enter ovarian cycles lasting about 16 weeks, though ovulation is induced by male presence.^[57] The gestation period lasts 18 to 22 months, the longest of any terrestrial mammal, resulting in the birth of a single calf in nearly all cases, with twins occurring in approximately 1% of pregnancies and often leading to reduced survival rates.^[58] Interbirth intervals average 4 to 5 years, reflecting the high energetic costs of reproduction and extended lactation periods.^[59] Reproductive success in Elephas is influenced by social dynamics, particularly the presence of experienced matriarchs in family groups, which enhances calf survival through protective behaviors, resource access, and predator avoidance strategies—young mothers' calves, for instance, face up to eight times higher mortality without nearby grandmothers.^[48] This low fecundity, characterized by infrequent breeding and long generational times, heightens population vulnerability to threats like habitat loss and poaching, as even modest adult mortality can prevent recovery.^[60]

Growth and Development

Asian elephant calves (Elephas maximus) are born after a gestation period of approximately 22 months, typically weighing around 100 kg and standing about 1 meter tall at the shoulder.^[3]^[61] These newborns can stand and walk within minutes to hours of birth, though they remain highly dependent on their mothers for the initial years.^[61] During the neonatal stage, calves rely exclusively on mother's milk, consuming 2-3 gallons daily, which supports rapid initial growth and development of motor skills, including trunk coordination that strengthens over the first few months.^[3]^[61] Calves begin supplementing their diet with solid vegetation around 4 months of age but continue nursing frequently until about 2 years, with weaning typically occurring between 3 and 5 years as they transition to full foraging independence.^[3]^[62] Growth is particularly rapid in the early years, with calves reaching sub-adult size by around 10 years, though sexual maturity arrives earlier for females at 10-15 years and for males at 10-15 years, with males achieving full physical maturity later, between 25 and 30 years.^[63]^[3] This extended growth trajectory allows for the development of massive body size, with continued weight gain into adulthood, especially in males.^[64] Parental care is intensive and communal within matriarchal herds, where the mother provides primary nursing and protection, supplemented by allomothering from other adult and adolescent females who comfort, guard, and assist calves against predators such as tigers.^[65]^[66] Calves learn essential foraging techniques, social interactions, and navigation skills through close observation and imitation of herd members during these formative years.^[66] In the wild, Asian elephants have a lifespan of 60-70 years, though individuals in captivity may live longer, up to 80 years or more, due to reduced predation and veterinary care.^[3]^[67] Senescence in older individuals is often marked by the progressive wear of their six sets of molars, which eventually leads to inability to chew vegetation effectively, resulting in malnutrition and starvation as a common cause of death.^[68]^[69] Calf mortality is notably high, ranging from 25-50% in the first few years, primarily due to predation, infectious diseases, or maternal abandonment, with risks amplified if the mother dies early, increasing first-year mortality up to tenfold.^[70]^[71] Despite this, surviving calves benefit from herd support, which enhances long-term development and integration into social structures.^[72]

Evolutionary History

Origins and Phylogeny

The genus Elephas originated in Africa during the early Pliocene, approximately 5 million years ago, deriving from early elephantid ancestors such as Primelephas, which exhibited gomphothere-like dental features adapted to forested environments.^[73]^[74] The earliest species widely attributed to Elephas, E. ekorensis, appeared in East Africa around 5–4.2 million years ago during the early to mid-Pliocene, marking the initial diversification within sub-Saharan habitats.^[75] This African origin reflects the broader evolution of Elephantinae from proboscidean lineages that had already adapted to browsing diets in tropical settings. Phylogenetically, Elephas occupies a sister position to Mammuthus (mammoths) within the family Elephantidae, with their divergence estimated at 5–6 million years ago based on integrated molecular clocks from mitochondrial DNA (mtDNA) and fossil calibrations.^[75]^[76] The split from the African elephant genus Loxodonta occurred earlier, around 7–7.6 million years ago, establishing three primary lineages within Elephantidae that radiated from a common African ancestor.^[75] mtDNA analyses further confirm that the Elephas–Mammuthus clade forms a monophyletic group basal to Loxodonta, with the Asian elephant (E. maximus) representing the extant sister to the extinct mammoth branch.^[75] Evidence of ancient hybridization, particularly between Elephas or related forms and Palaeoloxodon (straight-tusked elephants), is supported by genomic data from fossils showing gene flow events, including up to 10% mammoth ancestry in some Eurasian Palaeoloxodon populations.^[4] Recent genomic studies, such as those from 2018, refine these relationships, highlighting admixture within Elephantinae lineages.^[4] Key divergence events facilitated the genus's expansion beyond Africa. The split from Loxodonta coincided with late Miocene climatic shifts, while the Eurasian radiation of Elephas followed the Messinian Salinity Crisis (ending ~5.33 million years ago), which temporarily exposed land bridges across the Mediterranean, enabling migrations into Europe and Asia by early Pliocene species like Palaeoloxodon recki.^[77]^[75] By the late Pliocene (~3.7 million years ago), Elephas had dispersed widely into Asian habitats via these routes.^[78] The evolutionary timeline of Elephas spans the Pliocene origins in African forests, diversification with multiple migrations and adaptations to open woodlands, and a peak in species diversity during the Pleistocene, when forms like E. namadicus occupied vast Eurasian ranges before widespread extinctions.^[4]^[78] This trajectory underscores the genus's role in proboscidean adaptability amid global environmental changes.^[76]

Key Extinct Species and Adaptations

The genus Elephas displayed key evolutionary adaptations that enabled its extinct species to thrive in diverse Pleistocene and Pliocene environments, including progressive dental modifications for grinding tougher, more abrasive vegetation as habitats shifted from forests to open woodlands, and pronounced size variations among mainland populations that likely served as a deterrent against large carnivores.^[79]^[4] These traits reflect responses to climatic fluctuations and ecological pressures, with molars evolving higher crowns and more complex enamel folding to handle fibrous plants, while body size often scaled with resource availability and predation risks.^[80] Elephas hysudricus represents an early member of the genus from the Pliocene of India, particularly the Siwalik hills, where its primitive molars with fewer ridges were suited to the softer vegetation of forested floodplains along ancient river systems like the Gangetic plain.^[28] These dental features indicate an adaptation to a mixed browsing-grazing diet in humid, subtropical environments dominated by woodlands and swamps, marking a transitional form in the lineage leading to later Elephas species.^[81] Fossils from the Pinjor Formation highlight its role as a basal representative, with remains suggesting a body size intermediate between earlier proboscideans and more derived forms.^[82] Note: Species such as Palaeoloxodon antiquus (straight-tusked elephant), P. falconeri (dwarf elephant), and P. jolensis (North African form) were formerly classified under Elephas but are now recognized in the related genus Palaeoloxodon, reflecting refinements in elephantid taxonomy based on morphological and genomic evidence.^[4]

Taxonomy and Classification

Genus Definition

Elephas is a genus of elephants in the family Elephantidae, subfamily Elephantinae, distinguished by its high-crowned molars featuring transverse ridges adapted for grinding vegetation, primarily distributed in Asia among extant members, and teeth that are less hypsodont relative to those of mammoths in the genus Mammuthus.^[11] As the type genus of Elephantinae, it represents advanced proboscideans with craniodental specializations for diverse habitats, originating from late Miocene ancestors in Africa before dispersing to Eurasia.^[83] The genus was first formally described by Carl Linnaeus in 1758, who established Elephas maximus as the type species based on the Asian elephant, providing the foundational binomial nomenclature for the group.^[83] In 1847, Hugh Falconer and Proby Cautley expanded the taxonomic scope of Elephas to incorporate fossil material, distinguishing it from other proboscideans through early analyses of dental and skeletal remains, which facilitated the inclusion of extinct species.^[83]^[84] Diagnostic criteria for Elephas include a skull with a high, domed cranium, prominent occipital region where the occipital plane faces downward, and parallel premaxillaries supporting downward-directed tusks.^[11] The dental formula comprises 1/0 incisors (upper tusks only), 0/0 canines, 3/3 premolars, and 3/3 molars, though molars replace sequentially across five sets per quadrant over the animal's life.^[11] Genetic markers, such as a 6.56% sequence divergence in mitochondrial DNA, clearly separate Elephas from the African genus Loxodonta.^[85] Historically, Elephas faced nomenclatural confusion with Palaeoloxodon, often treated as a subgenus due to overlapping dental traits like lamellar molars, but modern analyses resolve this through cranial distinctions—such as the pronounced parieto-occipital crest in Palaeoloxodon—and genomic evidence placing the latter closer to Loxodonta.^[86] These differences confirm Elephas as a distinct lineage within Elephantinae.^[83]

Living and Extinct Species

The genus Elephas is represented by a single living species and approximately 4–6 extinct species, primarily known from Pleistocene and earlier fossil records.^[2] The sole extant species is Elephas maximus, the Asian elephant, which inhabits forested and grassland ecosystems across South and Southeast Asia and is characterized by its highly social matriarchal groups and adaptation to diverse vegetation through browsing and grazing.^[25] E. maximus is considered monotypic at the species level but includes four recognized subspecies: E. m. indicus (continental or Indian elephant, distributed across mainland Asia), E. m. maximus (Sri Lankan elephant, endemic to Sri Lanka), E. m. sumatranus (Sumatran elephant, restricted to Sumatra, Indonesia), and E. m. borneensis (Bornean elephant, restricted to Borneo); the Sumatran and Bornean subspecies are critically endangered.^[25]^[87] The validity of these subspecies has been supported by genomic analyses in the 2020s, revealing distinct evolutionary lineages with limited gene flow among populations.^[88] No new species within Elephas have been described since 2000, reflecting stable taxonomic consensus based on morphological and genetic evidence.^[4] The IUCN assesses E. maximus as Endangered, with a declining population due to habitat fragmentation, human-elephant conflict, and ivory poaching. Extinct species of Elephas are mostly confined to the Pleistocene epoch (2.58 million to 11,700 years ago), with fossils indicating a once-widespread distribution across Eurasia and Africa; most non-E. maximus lineages became extinct by the late Pleistocene.^[89] Key extinct taxa currently classified under Elephas include E. hysudricus from the Late Pleistocene of South Asia, considered a direct ancestor to E. maximus with transitional hypsodont molars;^[25] E. platycephalus from the Early Pleistocene of East Africa, displaying broad skull proportions and early Elephantidae traits;^[89] E. ekorensis from the middle Pliocene of Kenya; E. hysudrindicus from the Pleistocene of Indonesia; and E. beyeri (doubtful) from the Pleistocene of the Philippines.^[89] Note that several species formerly placed in Elephas (e.g., E. antiquus, E. falconeri, E. naumanni) have been reclassified into other genera such as Palaeoloxodon based on recent phylogenetic studies.

References

[1]
The Elephantidae
Around 11,500 years ago, Elephas became extinct in Africa, and some species of Elephas disappeared from Europe. All members of the family became extinct in ...
[2]
[PDF] Asian Elephant (Elephas maximus) - - Clark Science Center
Jun 18, 1982 · Ten fossil species of Elephas were recognized; the earliest is from the middle Pliocene Ekora beds, southeastern Turkana, Kenya, formed about ...
[3]
Elephas maximus (Asiatic elephant) - Animal Diversity Web
Elephas maximus is one of the largest terrestrial organisms and is considered a megafauna species. Asian elephants have gray skin that is covered with hair.
[4]
A comprehensive genomic history of extinct and living elephants
Feb 26, 2018 · We report a total of 14 genomes, including 2 from the American mastodon, which is an extinct elephantid relative, and 12 spanning all three extant and three ...
[5]
Shape variation in the limb long bones of modern elephants reveals ...
In this study, we investigate the shape variation of the six limb long bones among the modern elephants, Elephas maximus and Loxodonta africana.
[6]
The functional anatomy of elephant trunk whiskers - Nature
Jun 8, 2023 · Trunk musculature consists of ~40,000 muscles, which compares to only 600–700 muscles in the human body. The trunk musculature is innervated by ...
[7]
The sensorineural specializations of the trunk tip (finger) of the Asian ...
Background: The dorsal extension of the tip of the trunk of Asian elephants (Elephas maximus), often referred to as "the finger," possesses remarkable ...
[8]
Elephant Thermoregulation: Anatomy, Physiology & Behavior
Sep 22, 2022 · Elephants use different thermoregulatory mechanisms that depend on the anatomical and morphological characteristics of the species. The crevices ...
[9]
Qualitative Comparison of the Cranio‐Dental Osteology of the Extant ...
Dec 28, 2009 · The overall shape of the cranium is very different in the two genera. The Loxodonta cranium is much more rounded in profile than Elephas, which ...<|control11|><|separator|>
[10]
Of tusks and trunks: A review of craniofacial evolutionary anatomy in ...
Oct 8, 2024 · Their crania are rostrocaudally compressed in profile, with that of Elephas being especially more so due to the more rounded morphology in that ...
[11]
[PDF] MAMMALIAN SPECIES NO. 182, pp. is, 4 ñgs. - ElephaS maximUS ...
Jun 18, 1982 · Elephas maximus has more lamellae and wid- er upper and lower third molars than extinct species of Elephas. (MagHo, 1973). The Asian elephant ...
[12]
Elephants develop wrinkles through both form and function - PMC
The coloration of the trunk and skin texture differs between species, with Asian elephant trunk skin looking lighter, having pinkish pigmentation and smoother ...
[13]
Physical Characteristics - Asian Elephant (Elephas maximus) Fact ...
Sep 9, 2025 · Body smaller than African elephant · Ears smaller · Forehead flatter · Back convex or more level to ground · Head is highest point of Asian ...
[14]
Asian elephant | Smithsonian's National Zoo and Conservation ...
Asian elephant skin is gray, but parts sometimes lack color, especially on and around the ears, forehead and trunk. This de-pigmentation is believed to be ...
[15]
About Asian Elephant | WWF India
An Asian elephant bull can attain a height of 11 feet and a weight of over six tonnes. Record tusks have measured over 8ft (240cm) in length ... males and join ...
[16]
Evolution and taxonomy of Asian elephants | Natural History Museum
Elephants from Sri Lanka: are the largest; have the darkest skin colour; have the largest ears; are most prone to pink depigmentation of the skin on the face, ...Project Summary · Fossil Comparisons · About Borneo Elephants<|separator|>
[17]
First tracks of newborn straight-tusked elephants (Palaeoloxodon ...
Sep 16, 2021 · Based on well-preserved skeletons, estimates of maximum shoulder height varies from 300 to 420 cm and body mass from 4.5–5.5 to 13 tonnes for ...
[18]
A new partial skeleton of Elephas (Palaeoloxodon) antiquus ...
Based on extant and extinct elephant specimens, the Amyntaio's elephant estimated height at the shoulder is 3.5 m and its weight close to 9 tonnes. A CT scan ...
[19]
Asian Elephant | WWF - World Wildlife Fund
They inhabit dry to wet forest and grassland habitats in 13 range countries spanning South and Southeast Asia. While they have preferred forage plants, Asian ...
[20]
The state of the world's elephant populations - Our World in Data
Sep 10, 2024 · For the Asian species, this is in the range of 40,000 to 50,000. ... Asian elephant (Elephas maximus). There are fewer estimates of Asian ...
[21]
Population & Conservation Status - Asian Elephant (Elephas ...
Sep 9, 2025 · Population estimates · Fewer than 1,800 individuals · Once widespread; now only remain in small, severely fragmented populations · 70% of potential ...
[22]
Asian Elephant (Elephas maximus) Fact Sheet: Taxonomy & History
elephants. Genus: Elephas Linnaeus, 1758. Species: Elephas maximus Linnaeus, 1758 — Asian elephant, Indian elephant.
[23]
[PDF] Isotopic evidence of foraging ecology of Asian elephant (Elephas ...
Fossil Elephas is scarce in Eurasia during the Middle Pleistocene, except for several fossils reported from India (Kundal and Kundal, 2011) and the Middle East ...
[24]
Fossil distribution of Elephas species in south and southeast Asia...
Fossil distribution of Elephas species in south and southeast Asia (dotted lines indicate fossils that are rare or of doubtful age) and a schematic of the ...
[25]
New fossil remains of Elephas from the southern Levant
New elephant fossils are described from archaeological sites in Israel and Jordan. · The remains are 500–200,000 years old and are of the lineage of the Asian ...
[26]
Range-wide mtDNA phylogeography yields insights into the origins ...
It therefore appears that both clades had their origin in E. hysudricus. Elephas hysudricus fossils have been recorded widely, from the Pinjor horizon of ...
[27]
Report Estimating the dwarfing rate of an extinct Sicilian elephant
Feb 18, 2021 · We use combined molecular and fossil evidence to define the minimum and maximum rate of dwarfing in an extinct Mediterranean dwarf elephant from Puntali Cave ( ...
[28]
Reanalysis of African Elephas recki: implications for time, space and ...
The African Elephas lineage, consisting of E. ekorensis, E. recki, and E. iolensis, is an important faunal component of Plio-Pleistocene African ...
[29]
Global late Quaternary megafauna extinctions linked to humans, not ...
Jul 22, 2014 · We show that the severity of extinction is strongly tied to hominin palaeobiogeography, with at most a weak, Eurasia-specific link to climate change.
[30]
Determining the Dietary Preferences of Wild Asian Elephants ... - NIH
Elephas maximus are considered mega-herbivores with tremendous dietary requirements. Sukumar (2006) stated that elephants are able to consume 10% of their body ...
[31]
Asian Elephant (Elephas maximus) Fact Sheet: Diet & Feeding
Sep 9, 2025 · Feed as browsers (eat shrubs and trees) and grazers (eat grass) · Eat a wide variety of trees, shrubs, climbing plants (e.g., vines), herbs, and ...
[32]
Asian elephants' picky diet helps shape their forest home, study shows
Aug 9, 2023 · Adult Asian elephants can forage for 19 hours and consume up to 180 kilograms (400 pounds) of plant material per day, roughly half of which is “ ...
[33]
Feeding preferences of the Asian elephant (Elephas maximus) in ...
Nov 17, 2016 · This study investigates the feeding preferences of the Asian elephant in Parsa Wildlife Reserve (PWR) and Chitwan National Park (CNP), Nepal.
[34]
[PDF] Locomotion in captive Asian elephants (Elephas maximus)
Oct 31, 2014 · The mean daily distance travelled by captive Asian elephants was 9.05±0.61 km, with a range of 6.21–15.00 km, compared to 5-10 km for wild ...
[35]
Seasonal variation of health in Asian elephants - PMC
Dec 29, 2020 · This influences their protein intake, since in the dry season, elephants ingest low levels of protein (browse period), and in wet seasons, they ...
[36]
[PDF] Climate Influences on Human - Elephant Conflict in Sri Lanka
Most elephants consume 100 - 150 kilograms (220 - 330 pounds) of food and 80 - 160 liters (20 - 40 gallons) of water per day. The Asian elephant is adapted to ...
[37]
(PDF) Characteristics of gut microbiota in captive Asian elephants ...
Dec 19, 2023 · The Asian elephant is a typical hindgut fermenter that depends on trillions of gut microbiota within its gastrointestinal tract to ferment food ...
[38]
(PDF) Characteristics of Gut Microbiota in Captive Asian Elephants ...
Jun 28, 2025 · ... coprophagia events had an impact on the foal microbiome which promoted the establishment. of suitable gut microbial ora in young animals ...<|control11|><|separator|>
[39]
The impact on local forest ecosystem by elephants - ResearchGate
Aug 6, 2025 · The positive impacts include: improved seed dispersal and germination; creating forest gaps and maintaining community diversity; and enriching ...
[40]
Seed dispersal potential of Asian elephants - ScienceDirect.com
We examined the effects of elephant fruit consumption on potential seed dispersal using the example of a tree species with mega-faunal characteristics, ...
[41]
Elephants are socially complex - ElephantVoices
A matriarch, usually the oldest and most respected female, leads each family. Families may consist of as few as two or as many as 50 or more individuals. The ...
[42]
Vocalizations of wild Asian elephants (Elephas maximus)
Nov 5, 2009 · Intermittent touching, trunk twining with conspecifics ... Rumbles were the only call type in the repertoire with infrasonic components.
[43]
Asian Elephant - Columbus Zoo
Size: Height ranges between 7 to 10 feet, with males generally larger than females. Weight: 7,000 to 13,000 pounds, with females tending to weigh less than ...
[44]
The Chemical Ecology of Elephants: 21st Century Additions to ... - NIH
Males of both species dribble urine, secrete an odorous, viscous fluid from the temporal gland (i.e., temporal gland secretions, TGS), can act aggressively, ...
[45]
Nearby grandmother enhances calf survival and reproduction in ...
Jun 10, 2016 · In line with this, in Asian elephants, some observations suggest that mothers with suckling calves never permitted other calves to suckle them ...
[46]
Social Behavior and Group Formation in Male Asian Elephants ... - NIH
May 8, 2022 · Although younger male elephants are virtually always subordinate to older males [2], musth status seems to outweigh age-based dominance ...
[47]
Human-Elephant Conflict: A Review of Current Management ...
Contrary to Asia's focus on domestication, the culling of crop raiding elephants or those that kill humans has been regularly practiced in Africa to manage ...Abstract · Introduction · Causes and Consequences of... · Conceptual Model for...
[48]
The efficacy of interventions to protect crops from raiding elephants
Human retaliation to elephant crop raiding and other motivations of illegal killing can have negative population-level consequences on elephant demography ( ...
[49]
(PDF) Interplay of male traits, male mating strategies and female ...
Aug 9, 2025 · Furthermore, these results help to explain the evolution of musth as a sexual strategy: female elephants will often breed with multiple males ...
[50]
Reproduction & Development - Asian Elephant (Elephas maximus ...
Sep 9, 2025 · Gestation · Approximately 20 to 23 months, on average (Corse 1799; Dittrich 1966; Eisenberg 1980; Sukumar 2003; Meyer et al. 2004; Wittemyer 2011 ...
[51]
Defining progressive stages of male sexual behavior in in-situ and ...
Male elephants undergo a unique sexual state called “musth” that further complicates in-situ and ex-situ management strategies.
[52]
https://www.researchgate.net/publication/274713140_Interplay_of_male_traits_male_mating_strategies_and_female_mate_choice_in_the_Asian_elephant_Elephas_maximus
[53]
Seasonality of reproduction in Asian elephants Elephas maximus ...
Jul 31, 2018 · Semi-captive Asian elephants mainly bred in the dry season when body condition was lowest and day-lengths were increasing.
[54]
[PDF] Behavioral and Hormonal Indicators in the Asian Elephant
Females reach sexual maturity in their early teens, with captive females remaining reproductive into their thirties (Taylor & Poole 1998; Wiese 2000), while ...<|control11|><|separator|>
[55]
Foetal age determination and development in elephants - PMC - NIH
Their gestation length is the by far longest of all mammals (623–729 days in the Asian and 640–673 days in the African elephant; Sukumar 2003; Meyer et al. 2004) ...Missing: sources | Show results with:sources
[56]
Key reproductive insights of captive female asian elephants ...
The average age at first calving was 5447 ± 1344 days (14 years 11 months 7 days), while the average inter-calving interval was 1543 ± 771 days (4 years 2 ...
[57]
[PDF] Demography of captive Asian elephants (Elephas maximus) in ...
The average fecundity of the captive population (0.095 calves/adult female/year) is substantially lower than that for a wild population (0.21 calves/adult ...
[58]
[PDF] Asian Elephant Elephas maximus Asian or African? - Denver Zoo
Interestingly, Elephas maximus is a monotypic genus, meaning the Asian elephant is the only species within its genus. Class: Mammalia. Order: Proboscidea.Missing: taxonomy | Show results with:taxonomy<|control11|><|separator|>
[59]
Elephants are long-lived - ElephantVoices
By age three, calves can be fully weaned although they typically suckle for longer. Elephant calves are also dependent on their mothers and other family ...
[60]
Asiatic elephant (Elephas maximus) longevity, ageing, and life history
Although both sexes may become sexually mature at about 9 years of age, males do not normally reproduce until they are about 15 years old. Full size is attained ...Missing: height mortality parental care
[61]
[PDF] Sex-specific body mass ageing trajectories in adult Asian elephants
Apr 29, 2021 · We show sex-specific ageing trajectories: adult males gained weight up to the age of 48 years old, followed by a decrease in body mass until ...Missing: maturity | Show results with:maturity
[62]
68-2-438.pdf
This report describes allomaternal caretaking behavior in cow-calf groups of known lineage in the Asian elephant herd at the Washington Park. Zoo in Portland, ...Missing: weaning | Show results with:weaning
[63]
Observations on the social behaviour of free ranging groups of tame ...
Aug 7, 2025 · In elephant society reproductively inactive females, often nulliparous adolescents, play an important role in care giving through allomothering, ...
[64]
Age estimation of captive Asian elephants (Elephas maximus ...
Dec 11, 2023 · The average lifespan of Asian elephants is between 50–70 years but some have been known to survive for more than 80 years. DNA was extracted ...Missing: trajectory | Show results with:trajectory
[65]
Elephant body mass cyclicity suggests effect of molar progression ...
Aug 4, 2025 · As teeth are worn and migrate mesially toward the front of the jaw, fragments of the front teeth either fall out or are swallowed to make room ...
[66]
Age Determination in Elephants - Whole Earth Education
Jul 1, 2024 · When very little of the last teeth is left, the elephant cannot chew its food properly, it loses condition and finally dies of starvation. So ...
[67]
https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0294994
[68]
Short-term and delayed effects of mother death on calf mortality in ...
Aug 20, 2015 · Calves had a 10-fold higher mortality risk in their first year if they lost their mother, but this decreased with age to only a 1.1-fold higher risk in the ...Missing: neonatal stage
[69]
Evolutionary significance of maternal kinship in a long-lived mammal
Here, we measured the effects of maternal sister presence on female reproduction across a lifespan in a semi-captive population of Asian elephants. We report ...
[70]
Evolution - SANParks
It is thought that modern African and Indian elephants may have originated in Asia but the earliest fossils of the Proboscidea order have been found in Africa.<|separator|>
[71]
The Elephantinae
Loxodonta, Elephas and Mammuthus originated in Africa during the Pliocene (about 3 4 million years ago). Loxodonta remained in Africa while Elephas dispersed ...Missing: timeline | Show results with:timeline
[72]
Proboscidean Mitogenomics: Chronology and Mode of Elephant ...
Blue indicates a Mammuthus–Elephas sister group relationship, green indicates a Mammuthus–Loxodonta sister group relationship, and yellow indicates an Elephas– ...
[73]
Mammoth and Elephant Phylogenetic Relationships - NIH
Mar 29, 2007 · Morphologically, mammoths are considered sister to Asian elephants, but DNA studies show varying results. The study suggests using American ...
[74]
The impacts of the Messinian Salinity Crisis on the biogeography of ...
The aim of this study was to explore how the Messinian Salinity Crisis could influence the distribution and migration routes of the ancient Mediterranean ...
[75]
New Phylogenetic Analysis of the Family Elephantidae Based on ...
The elephant genus Elephas originated in Africa in the Miocene (around 7 million years ago) and spread to Asia during the Pliocene (around 3.7 million years ...
[76]
Elephant teeth: How they evolved to cope with climate change ...
Oct 16, 2023 · But they were ill-suited for tackling large quantities of tough, abrasive vegetation. Scientists theorized that two major factors drove the ...
[77]
Teeth of Past and Present Elephants: Microstructure ... - AGU Journals
Apr 7, 2021 · We report for the first time on the enamel microstructure data of the Indian elephantiformes Anancus, Stegodon, Elephas, and Palaeoloxodon<|control11|><|separator|>
[78]
[PDF] The straight-tusked elephant (Palaeoloxodon antiquus) in ...
African elephants can weigh up to 6 tonnes, with a shoulder height of up to 4 metres, while females weigh up to 3.2 tonnes and have a shoulder height of 2.6 ...
[79]
The presence and extinction of Elephas antiquus Falconer and ...
We discuss seven radiocarbon dates from remains of the straight-tusked elephant, Elephas antiquus Falconer and Cautley, 1847, from The Netherlands.
[80]
[PDF] India at the cross-roads of human evolution - Sci-Hub
Oct 26, 2009 · Our aim here is to broadly review the palaeoanthropo- logical records of India (with a few examples from Pakistan) from the Miocene onwards, ...
[81]
(PDF) Early Pleistocene Mammalian Faunas of India and Evidence ...
Jan 31, 2015 · In India, Early Pleistocene mammals have been found in Karewa Group of Kashmir Valley and the Pinjor Formation (Upper Siwalik Subgroup) of ...
[82]
Sicilian dwarf elephant image buy Uchytel
This island-bound elephant was an example of insular dwarfism, with an adult male specimen measured 150-180 cm in shoulder height and weighed about 1000 kg, ...
[83]
Mediterranean dwarf elephant (Palaeoloxodon falconeri)
Palaeoloxodon (Elephas) antiquus ancestors most likely reached the Mediterranean islands of Malta and Sicily from North Africa or possibly, from northern ...
[84]
[PDF] Late Pleistocene and Holocene mammal extinctions on continental ...
Jan 3, 2016 · spread African elephant lineages, Elephas iolensis was founded by Pomel. (1895) on the basis of a lower molar from Beausejour Farm in Algeria ...
[85]
Genomics Reveals Distinct Evolutionary Lineages in Asian Elephants
Aug 18, 2025 · We identify three distinct genetic clusters: Borneo, Sumatra, and Mainland, along with a fourth separation of Sri Lanka from the Mainland.
[86]
The latest straight-tusked elephants ( Palaeoloxodon)? “Wild ...
Aug 6, 2025 · ... Elephas maximus to Palaeoloxodon sp., and proposed that this extinct proboscidean also survived until around 3000 years ago in China. Many ...<|control11|><|separator|>
[87]
Elephas antiquus in Greece: New finds and a reappraisal of older ...
Dec 6, 2011 · ... height around 3.7 m at the shoulder and body mass around 9.0 tonnes. The good state of preservation of the MAR-1 bones allows the ...
[88]
The genus Elephas, member of the family of elephants
The genus Elephas, belongs to the family of elephants, elephantidae, and was named by Linnaeus 1758. Genus and species. Elephas - (genus). †Elephas antiquus - ( ...