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Proboscidea
Temporal range: Middle Paleocene-Holocene 60.0–0 Ma
The three living elephant species: Asian elephant, Elephas maximus (top right), African bush elephant, Loxodonta africana (left) and African forest elephant, Loxodonta cyclotis (bottom right)
Skeleton of the early proboscidean Moeritherium
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Mirorder: Tethytheria
Order: Proboscidea
Illiger, 1811
Subclades

Proboscidea (/ˌproʊbəˈsɪdiə/; from Latin proboscis, from Ancient Greek προβοσκίς (proboskís) 'elephant's trunk') is a taxonomic order of Afrotheria paenungulate mammals described by J. Illiger in 1811. It encompasses the elephants (family Elephantidae) and their extinct relatives.[1] Three living species of elephant are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant.

Extinct members of Proboscidea include the deinotheres, mastodons, gomphotheres, amebelodonts and stegodonts. The family Elephantidae also contains several extinct groups, including mammoths and Palaeoloxodon. Proboscideans include some of the largest known land mammals, with the elephant Palaeoloxodon namadicus and mastodon "Mammut" borsoni suggested to have body masses surpassing 16 tonnes (35,000 lb), rivalling or exceeding paraceratheres, the otherwise largest known land mammals in size.[2] The largest living proboscidean is the African bush elephant, with a recorded maximum size of 4 meters (13.1 feet) at the shoulder and a weight of 10.4 tonnes (11.5 short tons).[2] In addition to their enormous size, later proboscideans are distinguished by tusks and long, muscular trunks, which were less developed or absent in early proboscideans.

Evolution

[edit]

Over 180 extinct members of Proboscidea have been described.[3] The earliest members of Proboscidea like Eritherium are known from the Paleocene of Africa, around 60 million years ago, the earliest proboscideans were much smaller than living elephants, with Eritherium having a body mass of around 3–8 kilograms (6.6–17.6 lb).[4] By the late Eocene, some members of Proboscidea like Barytherium had reached considerable size, with an estimated mass of around 2 tonnes,[2] while others like Moeritherium are suggested to have been semi-aquatic.[5]

A major event in proboscidean evolution was the collision of Afro-Arabia with Eurasia, during the Early Miocene, around 18-19 million years ago allowing proboscideans to disperse from their African homeland across Eurasia, and later, around 16-15 million years ago into North America across the Bering Land Bridge. Proboscidean groups prominent during the Miocene include the deinotheres, along with the more advanced elephantimorphs, including mammutids (mastodons), gomphotheres, amebelodontids (which includes the "shovel tuskers" like Platybelodon), choerolophodontids and stegodontids.[6] Around 10 million years ago, the earliest members of the family Elephantidae emerged in Africa, having originated from gomphotheres.[7] The Late Miocene saw major climatic changes, which resulted in the decline and extinction of many proboscidean groups such as amebelodontids and choerolophodontids.[6] The earliest members of modern genera of Elephantidae appeared during the latest Miocene-early Pliocene around 6-5 million years ago. The elephantid genera Elephas (which includes the living Asian elephant) and Mammuthus (mammoths) migrated out of Africa during the late Pliocene, around 3.6 to 3.2 million years ago.[8]

Over the course of the Early Pleistocene, all non-elephantid probobscideans outside of the Americas became extinct (including mammutids, gomphotheres and deinotheres), with the exception of Stegodon.[6] Gomphotheres dispersed into South America during this era as part of the Great American interchange,[9] and mammoths migrating into North America around 1.5 million years ago.[10] At the end of the Early Pleistocene, around 800,000 years ago the elephantid genus Palaeoloxodon dispersed outside of Africa, becoming widely distributed in Eurasia.[11] By the beginning of the Late Pleistocene, proboscideans were represented by around 23 species. Proboscideans underwent a dramatic decline during the Late Pleistocene as part of the Late Pleistocene megafauna extinctions, with all remaining non-elephantid proboscideans (including Stegodon, mastodons, and the American gomphotheres Cuvieronius and Notiomastodon) and Palaeoloxodon becoming extinct, with mammoths only surviving in relict populations on islands around the Bering Strait into the Holocene, with their latest survival being on Wrangel Island around 4,000 years ago.[6][12]

The following cladogram is based on endocasts.[13]

Proboscidea
"plesielephantiforms"
"mastodonts"

Morphology

[edit]
Gallery of Proboscidean skeletons

Over the course of their evolution, proboscideans experienced a significant increase in body size. Some members of the families Deinotheriidae, Mammutidae, Stegodontidae and Elephantidae are thought to have exceeded modern elephants in size, with shoulder heights over 4 metres (13 ft) and masses over 10 tonnes (22,000 lb), with average fully grown males of the mammutid "Mammut" borsoni having an estimated body mass of 16 tonnes (35,000 lb), making it one the largest and perhaps the largest land mammal ever, with a fragmentary specimen of the Indian elephant species Palaeoloxodon namadicus only known from a partial femur being speculatively estimated in the same study to have possibly reached a body mass of 22 tonnes (49,000 lb).[2] As with other megaherbivores, including the extinct sauropod dinosaurs, the large size of proboscideans likely developed to allow them to survive on vegetation with low nutritional value.[14] Their limbs grew longer and the feet shorter and broader.[15] The feet were originally plantigrade and developed into a digitigrade stance with cushion pads and the sesamoid bone providing support, with this change developing around the common ancestor of Deinotheriidae and Elephantiformes.[16] Members of Elephantiformes and Deinotheriidae have retracted nasal regions of the skull indicating the development of a trunk.[17][18]

The skull grew larger, especially the cranium, while the neck shortened to provide better support for the skull. The increase in size led to the development and elongation of the mobile trunk to provide reach. The number of premolars, incisors and canines decreased. The cheek teeth (molars and premolars) became larger and more specialised.[15] In Elephantiformes, the second upper incisor and lower incisor were transformed into ever growing tusks on the upper and lower jaws,[19][20] while in Deinotheriidae there are only tusks on the lower jaw.[18] The tusks are proportionally heavy for their size, being primarily composed of dentine. In primitive proboscideans, a band of enamel covers part of the tusk surface, though in many later groups including modern elephants the band is lost, with elephants only having enamel on the tusk tips of juveniles. The upper tusks were initially modest in size, but from the Late Miocene onwards proboscideans developed increasingly large tusks, with the longest ever recorded tusk being 5.02 metres (16.5 ft) long belonging to the mammutid "Mammut" borsoni found in Greece, with some mammoth tusks likely weighing over 200 kilograms (440 lb). The lower tusks are generally smaller than the upper tusks, but could grow to large sizes in some species, like in Deinotherium (which lacks upper tusks), where they could grow over 1.5 metres (4.9 ft) long, the amebelodontid Konobelodon has lower tusks 1.61 metres (5.3 ft) long, with the longest lower tusks ever recorded being from the primitive elephantid Stegotetrabelodon which are around 2.2 metres (7.2 ft) long.[21]

The molar teeth changed from being replaced vertically as in other mammals to being replaced horizontally in the clade Elephantimorpha.[22] While early Elephantimorpha generally had lower jaws with an elongated mandibular symphysis at the front of the jaw with well developed lower tusks/incisors, from the Late Miocene onwards, many groups convergently developed brevirostrine (shortened) lower jaws with vestigial or no lower tusks.[23][24] Elephantids are distinguished from other proboscideans by a major shift in the molar morphology to parallel lophs rather than the cusps of earlier proboscideans, allowing them to become higher crowned (hypsodont) and more efficient in consuming grass.[25]

Dwarfism

[edit]
Main article: Dwarf elephant
Size comparison of the dwarf elephant Palaeoloxodon falconeri from the Pleistocene of Sicily and Malta to a human

Several species of proboscideans lived on islands and experienced insular dwarfism. This occurred primarily during the Pleistocene, when some elephant populations became isolated by fluctuating sea levels, although dwarf elephants did exist earlier in the Pliocene. These elephants likely grew smaller on islands due to a lack of large or viable predator populations and limited resources. By contrast, small mammals such as rodents develop gigantism in these conditions. Dwarf proboscideans are known to have lived in Indonesia, the Channel Islands of California, and several islands of the Mediterranean.[26]

Elephas celebensis of Sulawesi is believed to have descended from Elephas planifrons. Elephas falconeri of Malta and Sicily was only 1 m (3 ft), and had probably evolved from the straight-tusked elephant. Other descendants of the straight-tusked elephant existed in Cyprus. Dwarf elephants of uncertain descent lived in Crete, Cyclades and Dodecanese, while dwarf mammoths are known to have lived in Sardinia.[26] The Columbian mammoth colonised the Channel Islands and evolved into the pygmy mammoth. This species reached a height of 1.2–1.8 m (4–6 ft) and weighed 200–2,000 kg (440–4,410 lb). A population of small woolly mammoths survived on Wrangel Island as recently as 4,000 years ago.[26] After their discovery in 1993, they were considered dwarf mammoths.[27] This classification has been re-evaluated and since the Second International Mammoth Conference in 1999, these animals are no longer considered to be true "dwarf mammoths".[28]

Ecology

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It has been suggested that members of Elephantimorpha, including mammutids,[29] gomphotheres,[30] and stegodontids,[31] lived in herds like modern elephants. Analysis of remains of the American mastodon (Mammut americanum) suggest that like modern elephants, that herds consisted of females and juveniles and that adult males lived solitarily or in small groups, and that adult males periodically engaged in fights with other males during periods similar to musth found in living elephants. These traits are suggested to be inherited from the last common ancestor of elephantimorphs,[29] with musth-like behaviour also suggested to have occurred in gomphotheres.[32] All elephantimorphs are suggested to have been capable of communication via infrasound, as found in living elephants.[33] Deinotheres may have also lived in herds, based on tracks found in the Late Miocene of Romania.[34] Over the course of the Neogene and Pleistocene, various members of Elephantida shifted from a browse-dominated diet towards mixed feeding or grazing.[35]

Classification

[edit]

Below is a taxonomy of proboscidean genera as of 2019.[36][37][38][39]

References

[edit]

Bibliography

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Proboscidea

View on Grokipedia
from Grokipedia
Proboscidea is an order of eutherian mammals within the clade , encompassing the three extant species of elephants—Loxodonta africana (African savanna elephant), Loxodonta cyclotis (African forest elephant), and Elephas maximus (Asian elephant)—along with over 160 extinct relatives including mammoths, mastodons, and gomphotheres. Members of this order are distinguished by their elongated, muscular (trunk) used for feeding, manipulation, and respiration; elongated upper incisors forming prominent tusks; massive crania; and pillar-like limbs supporting enormous body masses often exceeding several tons. The evolutionary history of Proboscidea traces back approximately 60 million years to the late Paleocene or early Eocene in , where the earliest known forms, such as Phosphatherium and Numidotherium, emerged as small, pig-sized herbivores within the broader group, which also includes , sirenians, and aardvarks. Over the era, proboscideans underwent significant diversification, radiating into at least 10 families, 42 genera, and around 175 and , adapting to diverse habitats across , , and the through migrations facilitated by land bridges. Key evolutionary trends included increasing body size (), development of complex grinding molars for processing abrasive vegetation, and specialized trunks for browsing and grazing, with many lineages achieving ecological dominance as megaherbivores. Proboscideans reached their peak diversity during the and epochs, with forms like gomphotheres and deinotheres spreading globally and influencing ecosystems through their foraging and roles. However, most lineages went extinct during the , likely due to , habitat loss, and human hunting pressures, leaving only the family today. Modern elephants, the sole survivors, exhibit remarkable intelligence, complex social structures, and long gestations of up to 22 months, underscoring their unique adaptations. Recent phylogenetic analyses, incorporating total evidence and , have clarified interfamily relationships, supporting a basal position for early African forms and confirming 's with mammoths and mastodons as derived groups.

Taxonomy and Definition

Etymology and Scope

The order Proboscidea was established by German zoologist Johann Karl Wilhelm Illiger in 1811 to encompass living and extinct elephants (Elephantidae) and their relatives, such as the American mastodon. The name derives from the New Latin Proboscidea, rooted in the Greek proboškís (προβοσκίς), meaning "elephant's trunk" or "one with a trunk," highlighting the defining elongated nasal appendage central to the group's identity. This etymology underscores the order's focus on mammals characterized by a prehensile proboscis adapted for feeding, manipulation, and social behaviors. Proboscidea belongs to the mammalian superorder Afrotheria, a clade of primarily African-origin mammals that also includes orders like Hyracoidea (hyraxes) and shares a common ancestry with Sirenia (sirenians). The scope of Proboscidea extends to all known trunk-bearing herbivorous mammals, from primitive Paleogene forms to the three extant elephant species—African bush elephant (Loxodonta africana), African forest elephant (L. cyclotis), and Asian elephant (Elephas maximus)—representing a lineage that originated in Africa and dispersed globally. Approximately 175 species and subspecies have been recognized across 10 families, with the vast majority extinct, illustrating the order's historical diversity and adaptive radiation as large-bodied herbivores. Key diagnostic traits of proboscideans include the elongated, muscular (trunk) formed by fusion and extension of the and upper lip; pillar-like, columnar limbs supporting immense body mass; and high-crowned () molars with complex folding for grinding abrasive . These features evolved to facilitate a herbivorous lifestyle in varied terrestrial environments. Early taxonomic classifications underwent significant revisions; for instance, in 1834, grouped proboscideans with sirenians under "gravigrades," a cohort later refined as , before molecular and morphological evidence separated them into distinct orders within .

Phylogenetic Relationships

Proboscidea belongs to the superorder , a clade of placental mammals that also encompasses paenungulates such as (Hyracoidea) and (Sirenia), along with other African-endemic groups like elephant shrews, tenrecs, golden moles, aardvarks, and sengis. This placement emerged from molecular phylogenetic analyses in the late 1990s, which revealed unexpected affinities among morphologically disparate African mammals through comparisons of nuclear and sequences. Subsequent genomic studies have robustly confirmed Afrotheria's , with high bootstrap support across large-scale phylogenomic datasets, highlighting shared insertions and protein sequence signatures as additional molecular evidence. Morphological synapomorphies supporting are limited due to extensive adaptive divergence, but key features include the testicond condition (intra-abdominal testes without descent into a ), an elevated number of thoracolumbar vertebrae (typically 20 or more), and certain dental patterns such as the pseudohypocone on upper molars, which reflects early stages of lophodonty.00223-8) Within , forms part of the subclade alongside Hyracoidea and , where morphological traits like a continuous navicular facet on the astragalus in the ankle joint provide additional support for this grouping, though these are more pronounced in paenungulates. Debates persist regarding the exact branching order within , with early molecular studies yielding conflicting topologies—such as Proboscidea sister to or to Hyracoidea—but 2020s phylogenomic analyses using whole-genome data reveal an almost perfect , with gene trees supporting various resolutions and only 8-10% of genes providing statistically significant signal to reject the hypothesis of unresolved branching. This configuration aligns with some fossil-calibrated trees, emphasizing rapid diversification. Fossil evidence, including early afrotherian-like remains from , corroborates molecular estimates of an initial afrotherian divergence around 80 million years ago in the , prior to the end-Cretaceous extinction, when was isolated as a landmass favoring endemic radiations.

Major Taxonomic Groups

The order Proboscidea comprises 10 families, encompassing 175 and classified across 42 genera, with only 3 extant today. This classification, as outlined in seminal taxonomic work, reflects a hierarchical structure that includes both early diverging and advanced lineages, with Proboscidea nested within the Afrotherian of placental mammals. Recent paleontological revisions in the 2020s have refined genus-level distinctions, such as the separation of from based on cranial morphology and molecular data from . The basal family Moeritheriidae, known from the Eocene epoch, represents one of the earliest proboscidean radiations and includes the genus as its type, a small-bodied form that anchors the family's nomenclatural history. Deinotheriidae is distinguished by genera like , featuring unique tusks curving downward from the lower jaw, with type specimens from and deposits establishing its monotypic status within the family. Gomphotheriidae encompasses diverse forms from the , including genera such as and Anancus, with ongoing taxonomic adjustments elevating certain subfamilies to reflect phylogenetic branching. Mammutidae, commonly referred to as mastodons, includes key genera like and , with the family's nomenclature tracing back to early 20th-century descriptions based on North American fossils. The family , established by in 1821 through analysis of extant morphology, houses modern elephants and their close extinct relatives, such as mammoths; it comprises two extant genera—Loxodonta (African elephants, including L. africana and L. cyclotis) and (, E. maximus)—along with extinct ones like Mammuthus () and . The remaining families—Numidotheriidae, Barytheriidae, Palaeomastodontidae, Phiomiidae, and —include genera such as Numidotherium, Barytherium, , Phiomia, and , respectively, contributing to the order's total of over 170 extinct species.

Evolutionary History

Origins in the Paleocene

The earliest known proboscideans emerged in during the late , approximately 60 million years ago, evolving from endemic ungulate-like mammals within the superorder . These proto-proboscideans represent the basal radiation of the order in a period shortly after the Cretaceous-Paleogene extinction, with fossils indicating small-bodied forms adapted to forested or environments. The order Proboscidea is nested within based on both molecular and morphological evidence, underscoring its African continental origins. A key early genus is Phosphatherium escuilliei, a small, semi-aquatic proboscidean dating to 59–56 million years ago, discovered in the Ouled Abdoun phosphate basin of . Standing less than one meter at the and weighing around 10–15 kg, Phosphatherium featured bunodont teeth suited for soft and enlarged, procumbent upper incisors that served as precursors to tusks, alongside evidence of a short, fleshy proboscis-like structure indicated by the retracted position of the nasal opening. This exemplifies the primitive morphology of early proboscideans, bridging them to later forms through shared dental and cranial traits. During the Eocene, proboscideans underwent an initial radiation, notably with the Moeritheriidae family around 37–35 million years ago, known from abundant fossils in the Fayum Depression of . These semi-aquatic, hippopotamus-like forms reached up to 200 kg in body mass and inhabited marshy, riverine habitats, as evidenced by stable carbon and oxygen analyses of their showing a diet of aquatic plants and an amphibious lifestyle. Moeritheriids displayed further elongation of the incisors as proto-tusks and adaptations for wading, such as shortened limbs, marking key evolutionary steps toward more specialized proboscidean traits. By the late Eocene, early proboscidean dispersals had reached , broadening their distribution beyond as suggested by tentative dental fossils from southern .

Diversification During the Miocene

The epoch (23–5 million years ago) marked the peak of proboscidean diversity, with the order undergoing a major that resulted in numerous genera and species across multiple continents. This expansion began with migrations into around 22 million years ago, facilitated by climatic changes and land connections, leading to widespread distribution. By the middle to , proboscideans had reached via the , further diversifying into new ecological niches. Overall, proboscidean taxonomic richness reached its zenith during this period, with estimates of around 175 species and recognized across the order's history, many originating or peaking in the . Key families that diversified during the included Gomphotheriidae, , and the earliest members of . Gomphotheriidae, often featuring shovel-like tusks adapted for browsing or grazing vegetation, underwent rapid and became dominant in and , with genera like exemplifying this radiation. , characterized by unique downward-curving lower tusks used for stripping bark from trees, originated in the but achieved greater abundance and geographic spread in the across and . Early , ancestors to modern elephants, emerged in the late Miocene, initially in , with forms showing preliminary adaptations toward more open habitats. Adaptive innovations during the drove this diversification, particularly shifts in diet and body size. Many proboscideans transitioned from browsing on soft forest vegetation to on tougher grasses, accompanied by the evolution of high-crowned () molars for processing abrasive plants, as seen in middle fossils from containing grass phytoliths. Body sizes increased significantly, with like reaching shoulder heights of about 3 meters and weights of 4–5 tons, enabling exploitation of diverse habitats from woodlands to emerging grasslands. These changes coincided with and the spread of C4 grasslands, promoting niche partitioning among sympatric . Important fossil sites from the Miocene include the Bugti Hills in Pakistan, which preserve early Miocene proboscideans documenting initial migrations from Africa to Asia, and the Samburu Hills in northern Kenya, yielding late Miocene assemblages with diverse gomphotheres and deinotheres. Recent discoveries in the 2020s, such as refined ecomorphological analyses from East African sites, have clarified migration routes and dietary transitions, showing how biogeographic expansion beyond Africa fueled functional diversity.

Quaternary Extinctions and Survivors

The period, spanning from approximately 2.6 million years ago to the present, marked a profound decline in proboscidean diversity, with the vast majority of genera—estimated at over 90%—becoming extinct, leaving only the modern elephants as survivors. This mass extinction primarily affected iconic groups such as mammoths (Mammuthus spp.), American mastodons (Mammut americanum), and various gomphotheres (e.g., and ), which had thrived across , , and during the Pleistocene. of fossil remains indicates that the peak of these extinctions occurred between 12,000 and 10,000 years ago, coinciding with the transition from the Pleistocene to the epoch. records from cores further corroborate this timeline, revealing rapid shifts in from open grasslands to closed forests, which altered habitats critical for these large herbivores. Multiple interacting factors contributed to this late-Quaternary collapse, including at the end of the , intensified human hunting pressures, and consequent habitat fragmentation. The termination of Ice Age conditions around 11,700 years ago led to warmer temperatures and altered patterns, reducing the extent of steppe-tundra ecosystems that supported proboscidean populations. activities, particularly by Paleoindian groups, played a significant role, as evidenced by sites in where fluted spear points have been found embedded in and bones, indicating targeted hunting of proboscideans around 13,000–11,000 years ago. Habitat loss exacerbated these pressures, with expanding human populations and megafaunal overhunting leading to ecosystem destabilization, as supported by overlapping radiocarbon chronologies of human arrivals and proboscidean disappearances across continents. While debates persist on the relative weights of climatic versus anthropogenic drivers, integrated evidence from isotopic analysis and archaeological contexts points to a synergistic effect driving the extinctions. Amid this widespread die-off, only two genera within the family Elephantidae persisted: Loxodonta (African elephants) in and Elephas (Asian elephants) in South and , retreating to isolated refugia where fragmented forests and savannas provided relative sanctuary from hunting and climatic extremes. Recent genomic studies from the 2020s, analyzing whole-genome sequences from wild populations, reveal that these survivors endured severe population bottlenecks during the , resulting in critically low —such as only two mitochondrial haplotypes in some groups—and heightened vulnerability to depression.01177-1) Island populations occasionally delayed local extinctions; for instance, dwarfed woolly mammoths (Mammuthus primigenius) on in the survived in isolation until approximately 4,000 years ago, as determined by of over 100 bone and ivory samples spanning 7,000–4,000 years . These insular endemics, adapted to a resource-limited environment post-land bridge submersion around 12,000 years ago, represent one of the latest known holdouts of non-elephantid proboscideans.

Anatomy and Morphology

Overall Body Plan

Proboscideans exhibit a massive, herbivorous body build characterized by pillar-like limbs that provide robust support for their enormous weight, which in extinct forms could reach up to 14 tons. Their is barrel-shaped to accommodate a large digestive system for processing fibrous , while the head is positioned low on the shoulders to maintain balance under the pull of heavy cranial structures. This graviportal architecture, with straight, columnar legs resembling pillars, evolved to distribute the animal's mass efficiently during movement and standing. The skin of proboscideans is notably thick, measuring up to 4 cm in extant elephants, offering protection against environmental hazards and aiding in thermoregulation. In modern elephants, the skin is sparsely haired, but extinct species like the woolly mammoth possessed a denser coat for insulation in colder climates. Ear size varies for thermoregulatory purposes, with African elephants featuring large, flap-like ears that facilitate heat dissipation through increased surface area and blood vessel exposure. Locomotion in proboscideans is adapted to their size via a pillar-legged , enabling a top speed of approximately 25 km/h in bursts despite their bulk. Skeletal adaptations include fused carpals and tarsals in the wrists and ankles, which enhance stability and reduce flexibility to better withstand compressive forces during weight-bearing. Sexual dimorphism is pronounced in proboscideans, with males generally larger and displaying more robust skeletal features compared to females, as evidenced by comparisons between remains and living . This size disparity, often exceeding 30% in body mass, likely relates to male-male competition for mates.

Trunk, Tusks, and Dentition

The trunk, or , of proboscideans is a multifunctional formed by the fusion of the and upper , functioning as a muscular hydrostat capable of precise movements without skeletal support. Composed of approximately 90,000 muscle fascicles arranged in longitudinal, radial, and transverse orientations, the trunk enables elongation, shortening, bending, and constriction through antagonistic muscle contractions. In extant , it reaches lengths of up to 2 meters, with the distal tip exhibiting exceptional dexterity due to miniaturized fascicles as small as 10 cubic micrometers in volume, allowing fingertip-like grasping—African possess two such protrusions, while Asian have one. This structure facilitates object manipulation, such as plucking vegetation or wielding tools, as well as drinking by siphoning water or dust into the via generated by dilation. Additionally, the trunk serves in communication through tactile interactions and trunk-to-trunk contact for social bonding, and it integrates sensory functions, including touch via specialized mechanoreceptors at the tip. The olfactory capabilities of the trunk surpass those of dogs, supported by around 2,000 functional genes—more than twice the approximately 800 in canines—enabling detection of scents over several kilometers for locating , , or kin. Tusks in proboscideans are elongated upper incisors that grow continuously throughout life, primarily composed of (commonly termed ) overlaid by a thin layer of and an initial enamel cap that wears away early. In extinct species like woolly mammoths, tusks could exceed 3 meters in length, curving outward to aid in by uprooting , defense against predators, and intraspecific displays for dominance or . Modern retain similar functions, using tusks to dig for minerals or strip bark, though intense poaching pressure has driven evolutionary shifts toward tusklessness in some populations, particularly females in regions like , where genetic mutations on the linked to tusk development have increased in frequency due to selective survival advantages. Proboscidean dentition features a unique horizontal replacement system, with molars migrating forward from the back of the as wear down from , allowing up to six sets per lifetime in . These molars are lophodont, with transverse ridges adapted for grinding tough material, evolving from the bunodont (cusped) teeth of basal proboscideans like Phosphatherium to the highly (high-crowned) forms in later grazers such as mammoths, enhancing durability against silica-rich grasses. This serial replacement and morphological progression reflect adaptations to increasingly diets over proboscidean evolution.

Size Variations and Dwarfism

Proboscideans exhibit a remarkable range in body size across their evolutionary history, from diminutive basal forms to colossal later species. Early proboscideans, such as Phosphatherium and from the Eocene, were small, pig-sized herbivores weighing approximately 10 to 200 kg and standing less than 1 meter at the shoulder. In contrast, the largest known proboscideans, including from the Pleistocene of , achieved shoulder heights of up to 4.3 meters and body masses estimated at 10 to 14 tons, representing the peak of terrestrial mammalian . Modern elephants, the sole surviving proboscideans, are considerably smaller; African bush elephants (Loxodonta africana) typically weigh 2 to 6 tons with shoulder heights of 3 to 4 meters, while Asian elephants ( maximus) average 2 to 5 tons and 2.5 to 3 meters. This size variation follows allometric scaling principles, where body mass increases cubically with linear dimensions like shoulder height, influencing locomotion, , and resource requirements. Growth in proboscideans is characterized by rapid juvenile development supported by nutrient-rich maternal , enabling calves to achieve substantial size early in life. Elephant contains high levels of protein, typically 2.5% to 4.7% in early , along with elevated fat content, which fuels accelerated somatic growth and prepares offspring for around 2 to 4 years of age. is reached at 10 to 15 years, with females often maturing earlier (around 10 to 12 years in African elephants) than males (12 to 15 years), marking the transition to reproductive adulthood after a prolonged period of dependency. evidence from tusk cross-sections reveals incremental growth rings, analogous to rings, that record annual or seasonal deposition rates, providing insights into individual , diet, and age at death in extinct species like mammoths and mastodons. Insular dwarfism represents a striking in proboscideans isolated on islands, where body size reductions occur as part of the broader "island rule" evolutionary pattern, driven by limited resources and reduced predation pressure rather than climatic factors like . On Mediterranean islands such as and , evolved to a shoulder height of about 1 meter and a body mass of 200 to 300 kg, roughly 1% the size of its mainland ancestor antiquus. Similarly, on California's , the Mammuthus exilis diminished to approximately 1.7 meters at the shoulder and 700 to 800 kg, a fraction of the 6- to 10-ton from which it descended, with some estimates suggesting even smaller individuals approaching 200 kg. These reductions likely enhanced survival in resource-scarce environments by lowering metabolic demands and accelerating maturation rates. Recent genomic analyses in the , including comparisons of from insular and mainland proboscideans, have begun to identify potential genetic mechanisms underlying size reduction, such as alterations in pathways, though full causal links remain under investigation.

Ecology and Biology

Habitats and Distribution

Proboscideans achieved a during the through the periods, with fossil evidence spanning diverse environments from tundras to tropical forests across all continents except and . Early forms originated in around 55 million years ago and dispersed to via land bridges approximately 18-19 million years ago, later reaching around 5-6 million years ago through and even extending to . Iconic cold-adapted like the (Mammuthus primigenius) inhabited northern high latitudes, including , , and parts of , thriving in steppe-tundra ecosystems up to the during the Pleistocene. Today, the three extant proboscidean species occupy fragmented ranges in and . African elephants (Loxodonta africana and L. cyclotis) are distributed across in savanna woodlands, grasslands, and tropical forests, with a total estimated range of approximately 5.9 million km², though much of this is under pressure from human activity. Asian elephants ( maximus) inhabit dry and wet forests, grasslands, and shrublands across 13 countries from through to , covering about 500,000 km²—roughly 5% of their historical extent. Proboscideans exhibit strong preferences tied to resource availability, particularly , which they require in substantial volumes due to their large body size and thermoregulatory needs. Both extant species drink 100-200 liters of daily, often accessing permanent sources like rivers or waterholes to prevent , and they avoid arid interiors without reliable moisture. Asian elephants demonstrate notable elevational flexibility, typically ranging from to 3,000 meters but occasionally ascending higher into Himalayan to evade or follow seasonal . Climate variability profoundly influences proboscidean distribution through seasonal migration patterns synchronized with wet and dry cycles. In ecosystems, African elephants shift ranges to exploit flush of grasses and during wet seasons, contracting to riverine areas in dry periods, with movements covering hundreds of kilometers in response to rainfall. Similarly, Asian elephants in monsoon-driven habitats migrate altitudinally or laterally to track and , though ongoing fragmentation has reduced these patterns. Recent tracking data from the indicate a roughly 50% contraction in suitable range since 1900, driven by habitat loss and isolation into smaller patches, limiting adaptive responses to shifts.

Diet, Foraging, and Physiology

Proboscideans are strictly herbivorous, consuming a diverse array of plant material to meet their high energetic demands. Extant species, such as the African savanna elephant (Loxodonta africana), ingest approximately 100–300 kg of vegetation daily, depending on body size, availability, and season. Dietary preferences vary by habitat and : forest-dwelling proboscideans like the (Loxodonta cyclotis) primarily browse on twigs, leaves, bark, and fruits in wooded environments, while inhabitants favor grasses and sedges as . This dichotomy is reflected in dental adaptations, with molars in grazing lineages evolving high-crowned () structures and complex enamel folding to withstand abrasion from silica phytoliths in grasses, a trait that emerged prominently in the as open habitats expanded. Foraging strategies in proboscideans rely heavily on the multifunctional trunk, which enables precise manipulation for plucking leaves, stripping bark, or uprooting grasses without using teeth, allowing selective feeding on nutrient-rich parts of . Individuals typically travel 10–50 km per day in search of patches, covering extensive ranges to compensate for the low nutritional quality of their diet. Digestion occurs via in the enlarged and colon, where a diverse gut —dominated by such as Fibrobacter and —breaks down and other fibrous components into volatile fatty acids for energy extraction. This microbial is essential, as proboscideans lack the rumination of smaller herbivores, achieving moderate efficiency (around 40–50% for ) through prolonged retention times of up to 60 hours. Physiological adaptations support survival on this low-energy diet and in arid or variable climates. Proboscideans exhibit a approximately 50% lower than expected for their body mass based on interspecific scaling laws, reducing daily expenditure and enabling sustained with minimal fat reserves. is facilitated by temporary storage in the trunk (up to 10 liters) and pharyngeal pouches near the , which hold 3–5 liters for short-term hydration between bouts every 2–3 days. involves large, vascularized ears that dissipate heat through and , potentially lowering core temperature by 3–5°C, supplemented by baths that provide evaporative cooling and UV protection. Reproduction is adapted to long lifespans, with lasting about 22 months—the longest among mammals—and twinning occurring in less than 1% of births due to limited uterine capacity and maternal resources. Evolutionary transitions in diet are evident from fossil records, particularly in the when climatic cooling and promoted C4 grasslands. Early proboscideans relied on C3 browse, but lineages like deinotheres and early elephantids shifted toward C4 grasses, as indicated by increasing δ¹³C values in and microwear patterns showing coarser scratches from abrasive silica. These changes, dated to around 7–5 million years ago in , drove dental innovations such as ever-growing molars, enhancing processing of tougher and facilitating diversification into open biomes.

Social Behavior and Reproduction

Proboscideans, particularly elephants, exhibit complex social structures characterized by matriarchal family units. These herds typically consist of 8 to 100 related females and their offspring, led by the oldest female, known as the matriarch, who guides the group in decision-making for movement, foraging routes, and responses to threats. Male proboscideans, in contrast, leave their natal herds at maturity and live solitarily or in loose, temporary associations with other males, often forming bachelor groups without the stable bonds seen in female kin groups. Allomothering is a common practice within these herds, where non-maternal females, such as aunts or older sisters, provide care to calves, including protection, grooming, and teaching, which enhances calf survival rates especially for young or inexperienced mothers. Communication among proboscideans is multimodal, facilitating coordination over varying distances and contexts. They produce infrasonic rumbles with fundamental frequencies of 14-24 Hz, which can travel up to several kilometers in open habitats, allowing herds to maintain contact or signal alarms. Close-range interactions involve tactile signals like trunk touches to mouths, genitals, or bodies for and reassurance, while chemical cues from temporal gland secretions convey information on reproductive status, identity, and emotional states such as aggression during . Reproductive strategies in proboscideans are polygynous, with males competing for access to females through displays and combat, often heightened during —a periodic state of elevated testosterone that induces aggression and attraction via temporal gland secretions and urine dribbling. Females reach around 10-12 years, with lasting 18-22 months, the longest among mammals, followed by calving intervals of 4-5 years to allow for extended maternal investment. Proboscideans have lifespans of 60-70 years in the wild, during which learned behaviors—such as migration routes and social norms—are transmitted matrilineally from matriarchs to and kin, preserving cultural across generations. Evidence from fossil bone beds and trackways suggests that extinct proboscideans like mammoths maintained similar matriarchal structures, with assemblages often dominated by female and juvenile remains indicative of family groups traveling together. In extant , observations in the have documented mourning-like rituals, where investigate, touch, and linger over deceased kin or even unfamiliar carcasses, appearing subdued and covering bodies with vegetation, reflecting deep social bonds.

Conservation and Human Impact

Extant Species Status

The extant species of Proboscidea are the African savanna elephant (Loxodonta africana), the African forest elephant (L. cyclotis), and the Asian elephant (Elephas maximus). The African savanna elephant is classified as Endangered on the IUCN Red List, with a population estimated at approximately 352,000 individuals as of the 2016 Great Elephant Census, which covered 93% of the savanna range and remains a key reference for recent assessments. Recent total African elephant estimates indicate around 415,000-540,000 individuals as of 2024, with savanna elephants comprising the majority. The African forest elephant is recognized as a distinct species and listed as Critically Endangered, with current estimates placing its population at fewer than 100,000 individuals, concentrated mainly in Central African forests like Gabon, where recent surveys indicate around 95,000. The Asian elephant is also Endangered, with a wild population ranging from 40,000 to 50,000 individuals across its fragmented range in Asia. Genetic health across these species is compromised by and small sizes, resulting in reduced diversity and risks of , which manifests as decreased fitness, higher juvenile mortality, and lower reproductive success. Studies in the 2020s utilizing single nucleotide polymorphisms (SNPs) from whole-genome sequencing have revealed that Asian elephants display higher levels of and lower heterozygosity compared to African elephants, attributable to their smaller overall numbers and more isolated subpopulations. For instance, fragmented Asian populations show elevated homozygosity in key genomic regions, exacerbating vulnerability to environmental stressors. In African elephants, while populations maintain relatively higher diversity due to larger group sizes, elephants exhibit notable declines in in isolated Central African groups. Subspecies variation further highlights status disparities. The African forest elephant (L. cyclotis) represents a distinct lineage with specialized adaptations to dense forests, but its Critically Endangered status underscores severe population isolation. Among Asian elephant subspecies, the Sri Lankan elephant (E. m. maximus) is the largest, with adult males averaging up to 5,500 kg and noted for its darker pigmentation and robust build; its population stands at approximately 11,000 individuals based on 2024 national census estimates finalized in 2025, comprising about 22% of the total count. Accurate census and health monitoring rely on advanced non-invasive techniques. GPS collars enable real-time tracking of movement patterns and home ranges in both African and Asian populations, providing data on group dynamics and habitat use. Complementarily, dung DNA analysis allows for genetic identification of individuals without disturbance, facilitating population estimates, kinship analysis, and detection of inbreeding in remote areas like African forests. These methods have improved census precision, revealing previously undercounted groups and supporting targeted genetic management.

Threats and Extinction Risks

Poaching for ivory remains a primary threat to African elephants (Loxodonta africana), with an estimated 100,000 individuals killed across the continent between 2010 and 2012 due to surging illegal trade demands. This selective pressure has driven rapid evolutionary changes, including an increase in tuskless females to approximately 30% in certain populations, such as in Mozambique's Gorongosa National Park, where poachers disproportionately target tusked individuals. Similar poaching incidents affect Asian elephants (Elephas maximus), though at lower intensities, exacerbating vulnerability through targeted removal of large-tusked males. Habitat loss from and has fragmented over 60% of suitable ranges in since 1700, confining populations to isolated patches that limit and increase isolation. In , comparable pressures have reduced distributions by about 30% in recent decades, further compounded by human- conflicts. These conflicts are acute in , where crop-raiding result in approximately 600 human deaths and 150 deaths annually, based on 2020-2024 data, often from retaliatory killings or accidental encounters. Climate change intensifies these risks by prolonging droughts that diminish water and forage availability, leading to and heightened mortality among . Models project that rising temperatures could cause up to 40% loss for African in regions like by 2050, forcing greater overlap with human areas and amplifying conflict. Diseases such as pose additional dangers, with outbreaks killing over 100 in in 2019 amid drought-stressed conditions that facilitate spore ingestion. Competition with for grazing resources further strains , particularly in fragmented landscapes where encroaches on habitats. In , incidents in wild highlight the role of interfaces in disease transmission. The interplay of these threats—poaching, habitat fragmentation, climate-induced scarcity, and disease—cumulatively elevates extinction risks for both elephant species, potentially leading to functional extinction where surviving populations lose ecological roles. Low genetic diversity, arising from isolation, compounds this vulnerability by reducing adaptive capacity.

Conservation Efforts

Conservation efforts for proboscideans, particularly elephants, encompass a range of international legal frameworks, protected area designations, monitoring programs, and innovative initiatives aimed at safeguarding populations and habitats. The Convention on International Trade in Endangered Species of Wild Fauna and Flora () listed all elephant populations in Appendix I in 1989, effectively banning international commercial trade in to curb . National parks play a crucial role in habitat ; for instance, in serves as a key conservation area for African , supporting long-term monitoring and of approximately 1,870 individuals as of early 2025 through collaborative efforts between researchers and local communities. Similarly, in provides essential safeguards for Asian within its diverse , contributing to broader as a with a century-long history of conservation management. Key initiatives include the Monitoring the Illegal Killing of Elephants (MIKE) program, established in 1997, which operates a site-based system across and to track trends in illegal killings, build management capacity, and inform policy decisions through data collection from protected areas. In , rewilding projects focus on securing elephant corridors to facilitate movement and genetic connectivity; the national Elephant Corridors of India report identifies 150 such pathways, with efforts like the Wildlife Trust of India's Right of Passage project rehabilitating conflict-affected areas and restoring connectivity spanning over 100 kilometers in critical landscapes. Notable successes demonstrate the impact of community-driven approaches and technology. In , community conservancies have facilitated a significant rebound in numbers, growing from approximately 7,600 in 1995 to 25,664 by 2023 through sustainable and benefit-sharing models that empower local stakeholders. technologies, such as drones equipped with AI for real-time detection, have enhanced patrol efficiency in African reserves; programs like Air Shepherd have conducted over 4,000 missions to deter threats and monitor elephant movements in regions like and . As of 2025, the IUCN Species Survival Commission released updated African Elephant Status Reports (December 2024 for forest elephants, June 2025 for savanna elephants), confirming Endangered and Critically Endangered statuses respectively while highlighting stable but vulnerable populations and the need for enhanced transboundary efforts. Looking to the future, transboundary collaborations address habitat fragmentation by linking ecosystems across borders. The Kavango-Zambezi Transfrontier Conservation Area (KAZA TFCA), spanning about 520,000 km² across Angola, Botswana, Namibia, Zambia, and Zimbabwe, integrates 36 protected areas to support the world's largest contiguous elephant population through joint management and anti-poaching strategies. In the 2020s, emphasis has shifted toward human-wildlife coexistence, with compensation schemes reimbursing communities for losses incurred from elephant interactions, as outlined in global guidelines promoting sustainable livelihoods alongside conservation.

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