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Notiomastodon

Notiomastodon is an extinct genus of proboscideans in the family Gomphotheriidae, consisting of large, herbivorous mammals closely related to modern elephants that inhabited South America exclusively during the Pleistocene epoch, from approximately 2.5 million years ago until their extinction around 12,000 years ago.^[1] The sole species, Notiomastodon platensis (originally described as Mastodon platensis by Ameghino in 1888 and later reassigned), was a robust, lowland gomphothere characterized by a short, tall skull, upper tusks that were curved to nearly straight and enamel-free in adults, and lower tusks that were small or absent.^[1] Adults typically reached a shoulder height of about 2.8 meters and a body mass of around 6.5 tonnes for large individuals, making them among the largest terrestrial mammals of their time in the region.^[2] Phylogenetically, Notiomastodon represents a derived lineage within Gomphotheriidae with a debated position; analyses based on ancient DNA place it as the sister taxon to Elephantidae (the family including modern elephants and mammoths), with a divergence estimated at approximately 13.5 million years ago during the Miocene, while proteomic studies suggest a closer affinity to Mammutidae.^[3]^[4] This genus migrated to South America via the Great American Biotic Interchange around 2.5 million years ago and became widespread, with fossils documented across a broad latitudinal range from Ecuador and Colombia in the north to central Chile and Argentina's Pampean Region in the south, spanning over 140 localities.^[1] It did not overlap in distribution with the related gomphothere Cuvieronius hyodon, suggesting ecological partitioning.^[1] Notiomastodon platensis was adapted to savannahs and xerophytic pasture habitats in warm to temperate climates, functioning as a mixed feeder with a preference for grazing on grasses and shrubs, as indicated by dental microwear, enamel isotopes, and coprolite analyses.^[1] Evidence also points to frugivory, with fruit remains found in dental calculus from late Pleistocene specimens.^[5] The species' extinction coincided with the end-Pleistocene megafaunal die-off, likely driven by a combination of rapid climate change toward cooler, drier conditions and early human hunting pressures, as suggested by cut-marked bones and dated remains from sites in Chile, Colombia, and Venezuela dating to 13,000–11,000 years ago.^[1] In some areas, such as the Pampas, populations declined around 20,000 years ago prior to human arrival in the region, though overall extinction occurred before 12,000 years ago.^[1]

Research history

Early discoveries

The initial scientific interest in South American proboscidean fossils emerged during Alexander von Humboldt's expedition to the region between 1799 and 1804, where he collected two teeth resembling those of mastodons from Andean localities, including one from the Imbabura Volcano near Quito, Ecuador, and another from Chile.^[6] These specimens were presented to the French naturalist Georges Cuvier, who formally described and illustrated them in 1806 as Mastodon andium (a left lower second molar) and Mastodon humboldtii (an incomplete lower first molar or deciduous fourth premolar), marking the first European recognition of extinct elephant-like mammals from the continent.^[6] Cuvier's work highlighted the distinctiveness of these remains from North American mastodons, laying the groundwork for further paleontological exploration in South America.^[1] In the late 19th century, Argentine paleontologist Florentino Ameghino advanced the study through extensive collections from the Pampas region, formally naming Mastodon platensis in 1888 based on dental and skeletal fragments from Argentine sites, including molars and tusks that suggested a robust, South American-adapted proboscidean.^[1] Ameghino's description, published in Rápidas diagnosis de mamíferos fósiles nuevos de la República Argentina, emphasized the species' differences from northern forms and contributed to identifying multiple Mastodon taxa in the region, though many were later synonymized.^[7] By the early 20th century, Ángel Cabrera refined the taxonomy in 1929 by establishing the genus Notiomastodon for South American proboscideans, designating Notiomastodon platensis as the type species to distinguish it from North American mastodons, with the type locality at Playa del Barco in Monte Hermoso, Buenos Aires Province, Argentina, based on a mandible and tusk exhibiting a lateral enamel band.^[7] This reclassification addressed prior confusions in Ameghino's nomenclature.^[1] Concurrent excavations bolstered these findings; in Argentina, Cabrera's work at Playa del Barco yielded partial skeletal elements, while in Brazil, Gerald Waring's 1912 dig at Pedra Vermelha in Bahia uncovered isolated teeth and postcranial bones described as Mastodon waringi in 1920, later attributed to Notiomastodon.^[8] These efforts revealed Notiomastodon's widespread distribution and provided foundational material for understanding its anatomy.^[8]

Taxonomic debates

The taxonomic history of Notiomastodon is characterized by persistent confusion with the North American genus Stegomastodon, stemming from overlapping morphological traits such as similar dental crown patterns and limb bone robusticity. In the mid-20th century, researchers like Hoffstetter (1952) proposed treating Notiomastodon as a subgenus of Stegomastodon to account for these similarities, reflecting broader uncertainties in distinguishing South American proboscidean taxa. This led to proposals for synonymy, notably by Prado et al. (2005), who argued that Notiomastodon represented a junior synonym of Stegomastodon based on shared postcranial adaptations and stratigraphic correlations across the Americas.^[9]^[10] During the 1990s, efforts to refine classifications highlighted ecological distinctions, with Haplomastodon emerging as a proposed separate genus for highland-adapted forms, while Notiomastodon was retained for lowland species. This separation relied on differences in dental morphology—such as simpler, less folded enamel in Haplomastodon molars suited to abrasive high-altitude vegetation—and limb proportions, where Haplomastodon exhibited shorter, more robust metapodials indicative of montane locomotion. These revisions aimed to address prior lumping of highland and lowland fossils under unified taxa.^[11] A notable controversy arose with the 2007 description of Amahuacatherium peruvium as a late Miocene gomphothere from Peruvian Amazonia, implying an earlier migration of proboscideans to South America than previously thought. However, reexamination of the type fossils in 2015 revealed stratigraphic inconsistencies, re-dating the material to the Late Pleistocene and identifying morphological affinities with Notiomastodon platensis, including comparable tusk curvature and molar loph(id) structure, leading to its synonymization.^[12]^[13] These debates culminated in the 2012 taxonomic revision by Mothé et al., which analyzed over 90 South American specimens and affirmed Notiomastodon platensis as the sole valid species for lowland gomphotheres, dismissing multiple synonyms and emphasizing diagnostic cranial and dental traits to streamline classification. Phylogenetic analyses briefly supported this resolution by placing Notiomastodon as distinct from highland forms.^[14]

Recent research

Recent research on Notiomastodon since 2010 has incorporated advanced analytical techniques to elucidate its paleoecology, taxonomy, and temporal distribution. A pivotal 2018 multiproxy study analyzed stable isotopes, dental microwear, and dental calculus from molars of Notiomastodon platensis across multiple South American sites, revealing a predominantly leaf-browsing diet in closed forest habitats.^[15] This approach refined understandings of its ecological niche, showing adaptations to mixed C3-C4 vegetation environments during the Late Pleistocene.^[15] Taxonomic clarifications have also advanced, with a 2022 review in Historical Biology synthesizing morphological data from over 140 localities to reaffirm Notiomastodon as the sole lowland gomphothere genus in South America.^[1] The study invalidated previous referrals of South American fossils to Stegomastodon, reassigning them to N. platensis based on diagnostic cranial and dental features, thus resolving long-standing nomenclatural ambiguities and emphasizing its adaptation to warm, temperate savannas and xerophytic pastures.^[1] New fossil discoveries have expanded the known geographic range. In 2024, remains from north-central Chile (31°S–36°S) documented N. platensis in Mediterranean ecosystems during the Late Pleistocene, indicating behavioral flexibility in exploiting sclerophyllous forests and seasonal wetlands amid arid conditions.^[16] These findings, supported by sedimentological context, highlight its broad latitudinal tolerance from 31°S to 42°S.^[16] Dating techniques have further refined its chronology. A 2025 electron spin resonance (ESR) analysis of specimens from northwestern Venezuela yielded ages ranging from 560 ± 40 ka to 47 ± 7 ka, providing the oldest numeric ages for the species and extending its known temporal range back into the Middle Pleistocene, confirming presence from the Ensenadan to Lujanian stages.^[17] This updates prior estimates of its early distribution in northern South America. Paleopathological investigations provide insights into health stressors. A 2025 study of mandibular remains from Argentine sites (Córdoba province) identified chronic osteomyelitis in multiple individuals, characterized by bone remodeling, abscesses, and periosteal reactions, likely triggered by bacterial infections from environmental factors such as drought-induced fodder scarcity or injury during foraging.^[18] These pathologies suggest population-level vulnerabilities to Late Pleistocene ecological pressures, including habitat fragmentation.^[18]

Taxonomy and phylogeny

Classification

Notiomastodon is classified within the kingdom Animalia, phylum Chordata, class Mammalia, order Proboscidea, superfamily Elephantioidea, family Gomphotheriidae, and subfamily Cuvieroniinae.^[19] The genus Notiomastodon was erected by Cabrera in 1929, with the type species N. platensis (originally described as Mastodon platensis by Ameghino in 1888), based on a holotype mandible from the Pleistocene of Buenos Aires Province, Argentina.^[20]^[21] The genus is monotypic, containing only N. platensis, with no accepted subspecies; proposals for regional variants based on morphological differences were rejected in recent analyses due to high intraspecific variation and overlapping distributions.^[1] Taxonomic revisions from 2015 to 2022 have stabilized the classification, absorbing South American Quaternary forms previously assigned to Stegomastodon (e.g., S. platensis) and declaring Haplomastodon a nomen dubium, unifying them under Notiomastodon platensis to reflect a single widespread lowland species.^[20]^[1]^[21] Notiomastodon is distinguished from the Mammutidae (true mastodons) by its shorter mandible and possession of upper tusks only, lacking the prominent lower tusks typical of many mammutids.^[20] It differs from the Elephantidae (modern elephants and mammoths) in having gomphothere-like molars: low-crowned, trilophodont cheek teeth with bunodont cusps adapted for mixed browsing-grazing, rather than the high-crowned, multilophodont molars of elephantids suited for abrasive grasses.^[22]^[23]

Phylogenetic position

Notiomastodon is traditionally placed within the extinct family Gomphotheriidae, specifically in the subfamily Cuvieroniinae, as part of the trilophodont brevirostrine gomphotheres based on morphological cladistic analyses of cranial and postcranial features. A 2016 cladistic study of South American proboscideans, incorporating 48 morphological characters from 14 taxa, recovered Notiomastodon as closely related to North American Rhynchotherium and South American Cuvieronius, though not directly sister to the latter; instead, Rhynchotherium and Cuvieronius formed a clade nearer to Notiomastodon than to other gomphotheres like Stegomastodon. Key synapomorphies supporting this positioning include an elongated premaxilla that supports upturned and slightly diverging upper tusks, reduced or absent lower tusks (incisors), a shortened brevirostrine mandibular symphysis, and tetralophodont molars with bunodont cusps adapted for mixed browsing-grazing diets.^[24] This morphological framework posits Notiomastodon as derived from North American Rhynchotherium-like ancestors that dispersed southward during the Great American Biotic Interchange approximately 3 million years ago, leading to its radiation across South American lowlands during the Pleistocene. However, the subfamily Cuvieroniinae, which includes Notiomastodon alongside Cuvieronius, is characterized by these derived tusk and dental traits that distinguish it from earlier, more longirostrine gomphotheres.^[13] A 2022 phylogenetic update using Bayesian total-evidence methods, integrating ancient DNA from Notiomastodon platensis collagen with 406 morphological characters across 25 proboscidean taxa, challenges the traditional gomphothere placement by positioning Notiomastodon as the sister taxon to the crown Elephantidae (modern elephants and mammoths), with a divergence estimated at 13.5 million years ago (95% HPD: 16.2–11.1 Ma). This analysis renders Gomphotheriidae paraphyletic, with Notiomastodon clustering closely with Stegomastodon (divergence ~2.5 Ma) within a broader trilophodont clade that excludes Mammutidae, and confirms its North American origins via the GABI around 2.5–3.5 Ma based on calibrated fossil constraints. The study attributes this revised position to palaeogenetic evidence resolving long-standing morphological ambiguities in proboscidean relationships, and this phylogeny remains the most recent consensus as of 2025.^[25]

Evolutionary origins

Notiomastodon originated from North American gomphotheriid ancestors that migrated southward during the Great American Biotic Interchange, coinciding with the final closure of the Isthmus of Panama around 3 million years ago (Ma) at the Pliocene–Pleistocene transition. This dispersal event allowed proboscideans and other northern taxa to cross into South America, marking a pivotal phase in intercontinental faunal exchange. The earliest undisputed South American proboscidean fossil, a fragmentary vertebra likely belonging to an early representative of the lineage leading to Notiomastodon, comes from the middle section of the Uquía Formation in northwestern Argentina and dates to approximately 2.5 Ma.^[13]^[26]^[27] Following this initial incursion, Notiomastodon underwent diversification primarily in northern South America, particularly in regions of present-day Colombia and Venezuela, during the early Pleistocene around 1.8 Ma. Here, the genus adapted to tropical lowland ecosystems, developing traits suited to forested and open habitats with abundant vegetation. Fossil evidence from these areas indicates rapid establishment of populations, with dental and postcranial remains showing morphological consistency suggestive of early evolutionary experimentation in feeding and locomotion strategies.^[28]^[1] By the Middle Pleistocene, Notiomastodon had radiated southward, reaching the eastern foothills of the Andes and the expansive pampas of central Argentina, occupying a broad latitudinal gradient from equatorial lowlands to temperate grasslands. Unlike its highland relative Cuvieronius, Notiomastodon dominated lowland niches.^[29] Throughout its temporal and geographic range, spanning from the early Pleistocene to the early Holocene, Notiomastodon persisted as a single species, N. platensis, with no substantive evidence for speciation despite fluctuating climates and habitat shifts. Comprehensive taxonomic revisions of extensive fossil assemblages confirm morphological unity across sites, attributing variations to ecophenotypic plasticity rather than distinct lineages.^[13]^[28]

Description

Size and build

Notiomastodon displayed a robust build suited to bearing its considerable weight, with pillar-like limbs analogous to those in modern elephants for structural support, though featuring a comparatively narrower pelvis.^[30]^[31] Adult individuals typically measured 2.5–3.0 m in shoulder height and 4.5–5.5 m in body length, with estimated body masses of 3.15–6.5 tonnes calculated via regressions from femoral circumference.^[30]^[31] Sexual dimorphism was evident, with males larger than females and possessing longer tusks.^[32] Juveniles exhibited shoulder heights of about 1 m, achieving full adult dimensions by 15–20 years through gradual ontogenetic growth.^[30]^[32]

Skull and dentition

The skull of Notiomastodon platensis was characterized by a relatively short and tall elephantoid morphology, featuring a high-domed cranium, large forehead, and horizontally oriented rostrum formed by the premaxilla, which provided support for the upper tusks.^[24]^[6] The rostrum showed a slightly developed incisive fossa, and the upper tusk alveoli diverged slightly, accommodating the robust tusks.^[24] The upper tusks were straight to slightly curved, with a circular to oval cross-section, and lacked enamel bands in adults, though juveniles exhibited enamel covering or bands; lengths varied, with the lectotype measuring 150 cm.^[24]^[6] Lower tusks were absent, consistent with the gomphotheriid condition.^[6] The mandible was brevirostrine with a rounded profile and reduced symphysis that was not strongly downturned; the ascending ramus was long and nearly perpendicular to the mandibular body.^[6] The dentition included intermediate molariforms that were trilophodont and last molariforms that were tetra- or pentalophodont, with 35–82 cusps forming bunolophodont patterns suitable for grinding vegetation; enamel was thick (5–6 mm in some teeth) and complex, often producing trefoil wear patterns.^[24]^[33] Third molars, for example, measured up to 220 mm in crown length and exhibited 4½–5½ lophs with 45–60 cusps.^[6]^[33] The teeth were hypsodont and followed the typical proboscidean pattern of horizontal succession from milk to permanent molars.^[24]

Postcranial skeleton

The postcranial skeleton of Notiomastodon was adapted for supporting its massive body mass, estimated at up to 6.5 tonnes in large individuals, with features emphasizing graviportal locomotion suited to terrestrial environments.^[30] The axial skeleton provided robust trunk support, while the appendicular skeleton featured pillar-like limbs for weight-bearing efficiency. The vertebral column included 7 cervical vertebrae, typical of proboscideans, facilitating neck flexibility for browsing. Preserved specimens reveal thoracic vertebrae with broad, robust centra and tall neural spines for muscle attachment, contributing to overall spinal stability; lumbar vertebrae exhibit similar robustness to anchor hindlimb muscles. Although complete formulas are rare in the fossil record, comparisons with related gomphotheriids suggest 18–19 thoracic and 5 lumbar vertebrae, enabling a slightly arched trunk posture that distributed the weight of the head and proboscis.^[34]^[35] The pectoral girdle consisted of a broad scapula with a large glenoid cavity for shoulder joint stability during locomotion. This structure supported the forelimbs, which were slightly shorter than the hindlimbs overall, reflecting adaptations for weight distribution in a graviportal posture. The humerus measured approximately 83.5 cm in length in large specimens, robust and cylindrical with expanded articular ends for load-bearing, while the radius and ulna emphasized vertical pillar support over speed. Metapodials were short and massive, functioning as weight-bearing columns rather than levers.^[35]^[30]^[31] The pelvic girdle was relatively narrow, with flared ilia providing attachment for powerful hindlimb muscles. The femur ranged up to approximately 109.5 cm in length in large specimens, longer than the humerus, with a straight shaft and robust trochanters for gluteal muscle leverage. The tibia was proportionally shorter (around 71 cm). Metatarsals were similarly pillar-like and compact, adapted for graviportal support.^[35]^[30]^[31] Foot structure featured five toes on the manus and pes, with weight distributed across padded digital pads and a central carpal/tarsal pad, ideal for traversing soft or uneven terrain such as grasslands and forests. Carpals and tarsals were blocky and interlocking for stability, with metacarpals and metatarsals fused in function to form a solid platform beneath the body weight. Trackway evidence corroborates this upright limb posture, indicating efficient weight transfer during movement.^[35]

Paleobiology

Diet and feeding ecology

Notiomastodon platensis exhibited a mixed feeding strategy, consuming both C3 plants such as trees and shrubs (indicative of browsing) and C4 grasses (indicative of grazing), with opportunistic adaptations to local vegetation availability. Stable carbon isotope analyses (δ¹³C values ranging from -20‰ to -10‰) from tooth enamel and tusks across South American sites confirm this herbivorous diet, reflecting a blend of forest and open grassland resources that varied seasonally and annually.^[36]^[37] Regional dietary differences are evident, with populations in the Andean highlands showing a greater emphasis on browsing C3 vegetation like leaves and woody elements, while those in the Argentine Pampas leaned toward grazing on C4 grasses in more open, grassy environments. Dental microwear texture analysis supports this flexibility, revealing low to moderate scratch and pit densities consistent with a generalist diet incorporating both abrasive grasses and softer browse.^[22]^[38] A 2025 multiproxy study of Chilean fossils from central regions provides direct evidence of frugivory, with dental microwear on 96 teeth indicating occasional consumption of fleshy fruits, complemented by starch residues and plant microfossils from dental calculus suggesting intake of fruits like those from the Chilean palm (Jubaea chilensis). This role underscores its ecological importance, as the loss of such megafaunal dispersers has contributed to ongoing declines in fruit-dependent plant populations in modern South American forests.^[39] The species' trunk likely aided in manipulating foliage, stripping bark, and excavating roots or tubers, behaviors inferred from tusk morphology and wear patterns analogous to those in extant proboscideans. Molar wear patterns, characterized by heavy abrasion from silica-rich plants, further indicate an abrasive diet that included gritty soils and tough vegetation, with the bunodont teeth enabling efficient grinding of diverse plant matter.^[22]

Locomotion and ichnofossils

Notiomastodon displayed a graviportal gait typical of large proboscideans, characterized by columnar limb posture and synchronized fore- and hindlimb movements inferred from its skeletal proportions, which featured relatively shorter forelimbs (82% of hindlimb length on average) compared to modern elephants. This build supported a slow, stable walking pace suited to its body mass of around 4–5 tons, with a maximum estimated speed of approximately 20 km/h based on biomechanical models derived from comparable proboscidean limb scaling.^[30]^[40] Ichnofossils attributed to Notiomastodon, primarily from the Late Pleistocene Pehuen Co site in the Argentine Pampas, preserve trackways that indicate travel in family groups, with multiple parallel paths suggesting coordinated movement among individuals of varying sizes. These trackways exhibit stride lengths of 1.5–2 m and foot impressions measuring 40–50 cm in width, reflecting the animal's sub-circular pes and manus prints with five forward-pointing nail impressions on the forefoot. A 2015 study analyzing these ichnofossils dated the site to approximately 12,000 years ago, highlighting the preservation of over 300 proboscidean prints in aeolian dune sediments overlying coastal lagoons.^[41]^[42] Trackway patterns at Pehuen Co and similar Pampas sites show evidence of wallowing in shallow mudflats, with clustered, irregular impressions indicating pauses for dust or water bathing, alongside linear foraging paths through vegetated wetlands. Notiomastodon lacked morphological adaptations for climbing, such as elongated metapodials or flexible joints, confining its locomotion to flat, lowland terrains and precluding uphill or rugged mobility.^[41]^[40] Compared to modern elephants, Notiomastodon footprints exhibit similar overall morphology, including rounded heels and distinct pad impressions, but feature a narrower trackway gauge due to its lighter, more compact build and proportionally shorter, thicker limbs that reduced lateral sway during movement.^[43]^[30] Notiomastodon exhibited a social organization akin to that of modern elephants, forming matriarchal family groups comprising 5–15 individuals, primarily females and their young. This structure is inferred from fossil bone beds preserving multiple individuals across age classes, including the notable assemblage of 47 skeletons from a karstic cave at Águas de Araxá in Minas Gerais, Brazil, where both adults and juveniles were represented, suggesting a group mortality event.^[44] Adult males dispersed from family units after reaching maturity, adopting a solitary lifestyle or forming transient bachelor herds. Indications of musth-like aggression in males, characterized by elevated testosterone levels leading to heightened territoriality and combat, are supported by tusk wear patterns and growth interruptions observed in proboscidean fossils, including gomphotheres, which mirror annual hormonal cycles in extant elephants.^[45] Reproductive biology in Notiomastodon paralleled that of other proboscideans, with females undergoing a gestation period of approximately 22 months and typically producing a single calf per birth. Sexual maturity was achieved between 8 and 10 years, contributing to a lifespan estimated at 40–60 years based on dental eruption sequences and tusk growth annuli.^[45] Seasonal migrations likely occurred, driven by habitat shifts such as floodplain fluctuations, as evidenced by isotopic signatures in teeth indicating movement between foraging areas; trackways of grouped individuals provide additional support for herding during these travels.^[45]

Paleopathology

Paleopathological evidence from Notiomastodon fossils reveals a range of skeletal and dental conditions, reflecting the physical demands and environmental challenges faced by this proboscidean. Osteoarthritis, characterized by degenerative changes in joint surfaces and vertebral bodies, has been documented in approximately 10% of adult specimens, primarily attributed to chronic mechanical stress from supporting their massive body weight during locomotion and foraging.^[46] These lesions, often involving osteophyte formation and joint space narrowing, indicate long-term wear in weight-bearing structures like the spine and limbs.^[34] Infections such as osteomyelitis and associated abscesses are evident in cranial remains, particularly the mandibles. A 2025 Argentine study analyzed two Notiomastodon individuals from Pleistocene deposits in Córdoba and Santiago del Estero Provinces, identifying chronic osteomyelitis with extensive bony proliferation, periosteal reactions, and abscess cavities that likely impaired mastication and overall health.^[47] These pathologies suggest bacterial invasion, possibly secondary to dental trauma or systemic infection, and highlight vulnerabilities in the feeding apparatus. Traumatic injuries are also preserved, point to survivable traumas potentially caused by falls during movement over uneven terrain or interactions with predators such as large carnivores.^[34] Dental remains frequently show enamel hypoplasia, manifesting as linear defects on tooth crowns, which signals episodes of nutritional stress during early development. These disruptions in enamel formation likely stemmed from periodic resource scarcity or illness, compounded by abrasive wear from a gritty diet.

Distribution and paleoecology

Geographic range

Notiomastodon platensis inhabited lowland regions across much of South America during the Pleistocene, with its core range extending from northern Colombia and Ecuador southward through the Andean foothills, Amazon Basin, Chaco region, and Pampas to central Argentina and southern Chile.^[48]^[1] The species is known from over 140 localities continent-wide, primarily in open savanna and woodland environments below 1,500 meters elevation, though brief highland occurrences are excluded as they pertain to related taxa like Cuvieronius.^[1] The northern limit of N. platensis reached the Venezuelan llanos and Caribbean coastal areas of Colombia and Venezuela, with confirmed fossils from sites such as El Breal de Orocual in Monagas State, Venezuela, and the Cauca Valley in Colombia.^[49]^[50] In the south, the distribution extended to approximately 42°S, including recent findings along the Chilean coast at 36°S in north-central regions like Tagua Tagua and La Campana National Park. Key fossil sites include the Tarija Valley in Bolivia, where multiple specimens, including tusks and mandibular remains, have been recovered from the Pleistocene sediments of the Tarija Formation, contributing to over 100 associated megafaunal records in the basin.^[51]^[52] In Argentina, abundant material comes from the Pampean region, with notable assemblages from the Ituzaingó Formation exposures in Entre Ríos Province.^[1] Brazil's São Paulo Basin yields more than 160 remains, including teeth and postcranial elements from sites like Águas de Araxá in nearby Minas Gerais, highlighting the species' prevalence in southeastern lowlands.^[53] The temporal span of N. platensis is the Pleistocene, from approximately 2.5 Ma to 0.01 Ma, with the earliest records in the Ensenadan stage (early Pleistocene); no verified pre-Pliocene records; the earliest South American proboscidean arrivals date to around 2.5 Ma in Colombia and northwestern Argentina, but these predate the genus.^[48]^[26]

Habitat and environmental associations

Notiomastodon platensis primarily inhabited lowland environments across South America during the Pleistocene, favoring savannas, open woodlands, and riverine forests, particularly during interglacial periods when warmer conditions expanded these habitats.^[29] These ecosystems provided a mix of browse and grasses, with the species showing adaptations to seasonal flooding, as evidenced by concentrations of fossils in flood-related deposits. Such adaptations likely included behavioral flexibility in foraging and movement to exploit nutrient-rich floodplains following wet seasons.^[50] The species was associated with warm-temperate climates, characterized by mean annual temperatures ranging from 15–25°C, supporting diverse vegetation including C3-dominated plants.^[48] Pollen records from Andean foothill sites indicate the presence of gallery forests along rivers, with humid, wooded corridors persisting amid surrounding drier landscapes.^[54] These forests, rich in trees and shrubs, aligned with the proboscidean's browsing habits, as confirmed by microfossils in dental calculus.^[55] Recent evidence from north-central Chile (31°S–36°S) dated to the Late Pleistocene reveals Notiomastodon platensis occupied Mediterranean-like ecosystems, including shrublands with pronounced dry summers and humid winters, demonstrating greater environmental tolerance than previously recognized.^[16] This broader adaptability is supported by stable isotope data indicating foraging in semi-arid, xerophytic vegetation under cooler and wetter conditions than modern analogs.^[56] Fossils of Notiomastodon are commonly preserved in fluvial deposits and loess sequences, reflecting a preference for wetland margins and riverine settings conducive to foraging on soft vegetation and minerals.^[57] These sedimentary contexts, often linked to seasonal water availability, underscore the species' reliance on dynamic, moisture-influenced landscapes for sustenance.^[50]

Interactions with contemporaneous fauna

Notiomastodon platensis coexisted with a diverse array of Pleistocene megafauna across South America, particularly in open habitats such as grasslands, savannas, and woodland edges of the Pampas, Patagonia, and Andean foothills. This community included large herbivores like the ground sloth Megatherium americanum, which functioned as a bulk feeder in similar landscapes, the extinct horse Hippidion principale adapted to grassy environments, and other taxa such as Toxodon platensis and camelids like Hemiauchenia. Carnivores, notably the saber-toothed cat Smilodon populator, were also present, sharing these ecosystems and contributing to a complex trophic structure.^[58] Ecological niche partitioning likely minimized direct competition among these herbivores, with Notiomastodon exhibiting a mixed feeding strategy that emphasized browsing on C3 plants such as leaves, shrubs, and fruits in forested or semi-closed habitats, distinct from the grazing habits of equids like Hippidion and Equus neogeus on C4-dominated grasslands. This dietary flexibility allowed Notiomastodon to occupy a unique role in the proboscidean guild, as it represented the primary large proboscidean in southern South America following the Great American Biotic Interchange, with limited overlap from northern taxa like Cuvieronius in more equatorial regions. Additionally, Notiomastodon's frugivory facilitated seed dispersal for large-fruited plants, including palms (Arecaceae) and legumes (Fabaceae), promoting forest regeneration and plant diversity in its range.^[59]^[60]^[39] Predation and scavenging interactions involved carnivores targeting Notiomastodon, particularly juveniles or weakened individuals, as evidenced by tooth marks on bones from sites like Pilauco in Chilean Patagonia, attributable to large felids such as Smilodon populator. These traces indicate scavenging of exposed carcasses rather than active predation, suggesting opportunistic feeding behaviors within the megafaunal community. No other proboscideans directly competed in this guild, reinforcing Notiomastodon's ecological dominance until the late Pleistocene.^[61]^[58] The extinction of Notiomastodon contributed to a trophic cascade, disrupting seed dispersal services essential for forest regeneration; a 2025 multiproxy analysis of fossils from central Chile revealed that its extinction increased the extinction risk for megafaunal fruit plants dependent on proboscidean seed dispersal, many of which are now threatened, highlighting ongoing ecosystem vulnerabilities. Human arrival in South America around 15,000 years ago disrupted these biotic interactions by altering predation dynamics and habitat use.^[39]

Extinction and human relations

Timing and causes of extinction

The extinction of Notiomastodon occurred at the Pleistocene-Holocene boundary into the Early Holocene, with the latest reliable records dating to approximately 11,000 years BP (calibrated), aligning synchronously with the broader wave of South American megafaunal extinctions that eliminated over 80% of large mammal species by around 10,000 BP.^[62]^[63] Recent radiocarbon dating of fossils from sites across southern South America, including Brazil and Argentina, indicates a sharp population decline beginning around 12,900 calibrated years BP, with near-complete disappearance by 10,900 calibrated years BP; a 2024 study from Minas Gerais, Brazil, confirms dates up to ~17,000 cal BP but supports no survival beyond ~11,000 cal BP.^[62]^[64] This timing coincides with the Pleistocene-Holocene transition, marked by abrupt environmental shifts that affected grassland-dominated ecosystems preferred by Notiomastodon.^[22] Primary causes of extinction included climate-driven changes following the Younger Dryas period (approximately 12,900–11,700 BP), characterized by cooling and subsequent warming that led to habitat fragmentation and grassland retraction between 12,400 and 11,500 calibrated years BP.^[62] Stable isotope analyses (δ¹³C and δ¹⁸O) from Notiomastodon enamel reveal dietary stress from increasing aridification around 15,000 BP, with shifts toward more C₄-dominated diets indicating reduced availability of preferred mixed C₃/C₄ vegetation in lowland mid-latitudes.^[22] Microwear patterns on teeth from Brazilian sites further support this, showing signs of abrasive feeding on tougher, scarcer browse during dry phases, which likely concentrated populations near dwindling water sources and contributed to mass mortality events.^[22] Notiomastodon found no effective refugia in Andean highlands, having withdrawn from these elevations by the Last Glacial Maximum (ca. 20,000–16,000 BP) due to unsuitable cold, arid conditions.^[1] Human overhunting amplified these climatic pressures, as evidenced by species distribution modeling from 2021 that demonstrates high spatiotemporal overlap between Notiomastodon ranges and early human sites equipped with Fishtail projectile points after 13,000 calibrated years BP.^[62] The legacy of this extinction persists in modern South American ecosystems, where 2025 multiproxy fossil analyses confirm Notiomastodon as a key long-distance seed disperser for large-fruited plants like the Chilean palm (Jubaea chilensis); its loss has contributed to ongoing forest degradation, with 40% of such plant species now endangered due to disrupted dispersal.^[39]

Evidence of human interactions

The earliest evidence of temporal overlap between humans and Notiomastodon in South America dates to around 15,000 years before present (BP), coinciding with the arrival of humans as indicated by the Monte Verde site in southern Chile, which is radiocarbon dated to approximately 14,500 calibrated years BP.^[65] Notiomastodon persisted across much of the continent until around 10,900–11,000 cal BP, allowing for several millennia of coexistence during the late Pleistocene.^[62] Stable isotope analyses of Notiomastodon remains reveal dietary and habitat preferences for open woodlands and grasslands, environments that isotopic signatures from early human sites suggest were also frequented by Paleoindians, indicating potential spatial overlap in resource use. Direct evidence of hunting includes an organic-type artifact (possibly wood, bone, horn, or antler) embedded in the rostral region of a juvenile Notiomastodon platensis skull from the Tanque Grande site in the Lagoa Santa Karst, Minas Gerais, Brazil, dated to the late Pleistocene.^[66] Taphonomic analysis of the specimen shows the artifact penetrated deeply, consistent with a projectile impact, and associated bone weathering suggests the animal died shortly after the injury, marking the first confirmed case of human-killed proboscidean in South America.^[66] At the Taguatagua 3 site in central Chile, dated to around 12,800–12,400 cal BP, multiple Notiomastodon bones exhibit cut marks from stone tools, indicative of butchery activities such as defleshing and disarticulation.^[67] Archaeological site associations further support interactions, as seen at Taima-Taima in northwestern Venezuela, where El Jobo complex stone tools, including unifacial points and scrapers, were found in direct stratigraphic association with butchered remains of a juvenile Notiomastodon, radiocarbon dated to at least 13,000 BP.^[68] The tools show use-wear patterns consistent with processing large mammal carcasses, and the mastodon bones display percussion fractures and cut marks aligned with marrow extraction and hide removal.^[68] Possible cultural representations appear in late Pleistocene rock art from the Serranía de la Lindosa in the Colombian Amazon, dated to around 12,600 BP, where some panels depict proboscideans interpreted as gomphotheres resembling Notiomastodon based on trunk, tusk, and body morphology, potentially illustrating hunting scenes or environmental observations.^[69] However, these identifications remain tentative, as stylistic and chronological ambiguities prevent definitive confirmation. No evidence exists for domestication or sustained management of Notiomastodon by humans, with all interactions appearing opportunistic and focused on hunting.^[62]

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