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Ground pangolin

The ground pangolin (Smutsia temminckii), also known as Temminck's ground pangolin or Cape pangolin, is a medium-sized, scaly mammal belonging to the order Pholidota, endemic to sub-Saharan Africa from southern Sudan and Chad southward to northern South Africa.^[1] It measures 40–70 cm in head-body length with a tail of similar length, its body almost entirely covered in overlapping keratinous scales that form a protective armor, leaving only the undersides, face, and inner limbs unscaled.^[2] Lacking teeth, it possesses a long, sticky tongue—up to 25 cm extended—for capturing prey, and strong claws adapted for digging burrows and tearing open ant and termite nests.^[2] This solitary, nocturnal species inhabits a range of dry habitats including savannas, grasslands, and open woodlands up to 1,700 m elevation, avoiding dense forests and arid karroid regions.^[1] Its diet consists exclusively of ants and termites, which it locates using keen senses of smell and hearing, consuming them in large quantities—up to 70,000 ants per night—while ingesting grit to aid digestion.^[2] Behaviorally, it travels bipedally with forelimbs tucked, defends itself by rolling into an impenetrable ball, and maintains home ranges of about 6 km², creating temporary burrows for shelter.^[1] Reproduction occurs year-round, with females giving birth to a single offspring after a gestation of 105–140 days; the young clings to the mother's back for 2–3 months before independence.^[2] Classified as Vulnerable by the IUCN Red List, ground pangolin populations are declining due to illegal poaching for scales and meat used in traditional medicine and bushmeat trade, habitat loss from agricultural expansion, and non-targeted threats like electrocution on electrified fences (estimated 377–1,028 deaths annually in South Africa) and road collisions.^[1] With low reproductive rates—one offspring every 1–2 years—and longevity exceeding 20 years in the wild, the species' slow recovery exacerbates vulnerability to these anthropogenic pressures, though protected areas offer some refuge.^[1]

Taxonomy and evolution

Classification and nomenclature

The ground pangolin was first described scientifically by South African physician and zoologist Johan Smuts in 1832, under the binomial Manis temminckii, in his work Dissertatio zoologica, enumerationem mammalium capensium continens.^[3] The species epithet temminckii honors Dutch zoologist Coenraad Jacob Temminck (1778–1858), director of the National Museum of Natural History in Leiden, who contributed to early mammalian systematics. The genus name Smutsia derives from the describer Smuts, reflecting taxonomic convention for honoring the original author in sub-Saharan African mammals.^[4] Initially classified within the broad genus Manis encompassing all pangolins, S. temminckii was reclassified into the distinct genus Smutsia in the 2010s following molecular phylogenetic analyses of mitochondrial and nuclear DNA, which resolved three monophyletic genera within Manidae: Manis (Asian species), Phataginus (arboreal African species), and Smutsia (terrestrial African species including the ground and giant pangolins).^[4]^[5] These studies, incorporating complete mitogenomes and multi-locus datasets, demonstrated deep genetic divergence between African terrestrial pangolins and other lineages, supporting the separation based on shared morphological traits like larger body size and ground-dwelling adaptations, rather than prior lumping driven by limited pre-genomic data. No subspecies are currently recognized for S. temminckii, owing to insufficient morphological or genetic evidence for intraspecific variation across its range.^[4] Common names for the species include Temminck's ground pangolin, Cape pangolin, and steppe pangolin, reflecting its terrestrial habits and southern African distribution; the term "pangolin" itself originates from the Malay pengguling ("one who rolls up"), alluding to the animal's defensive behavior but not specific to nomenclature.^[6]

Phylogenetic relationships

The ground pangolin (Smutsia temminckii) is classified within the subfamily Smutsiinae of the family Manidae, order Pholidota, alongside the giant pangolin (S. gigantea) and the African tree pangolins of genus Phataginus. Phylogenetic analyses of mitochondrial genomes affirm the monophyly of Smutsiinae as a distinct African clade, basal to the Asian subfamily Maninae, with robust support from concatenated gene sequences showing divergence in key morphological and molecular synapomorphies.^[7]^[5] Molecular dating using Bayesian relaxed clock models estimates the split between African (Smutsiinae) and Asian (Maninae) pangolin lineages at approximately 38 million years ago, during the late Eocene to early Oligocene, consistent across genomic datasets from multiple species.^[8]^[9] This deep divergence predates the Oligocene-Miocene boundary and reflects vicariance driven by continental drift and paleoclimatic shifts, rather than recent dispersal.^[4] The fossil record of Pholidota reveals a sparse history, with the earliest definitive remains from the middle Eocene of Europe (e.g., Eomanis and Pholidocercus), indicating a Laurasian origin for the order prior to the African-Asian split.^[10] Miocene fossils, such as partial humeri from localities like the French Quercy phosphorites (ca. 20-15 million years ago), represent early manids potentially ancestral to modern subfamilies, though direct Smutsiinae precursors remain elusive due to poor preservation of scaleless juvenile or dental elements.^[11] Recent genomic sequencing (2016–2023) reinforces the monophyly of Smutsia within Smutsiinae, revealing low intraspecific genetic diversity in S. temminckii populations, which supports recognition of evolutionarily significant units for conservation amid poaching pressures.^[5]^[4] These studies also position Pholidota as sister to Carnivora within Ferae, underscoring convergent scale armor with distantly related xenarthrans rather than shared ancestry.^[12]

Morphology and physiology

External features and scales

The ground pangolin (Smutsia temminckii, formerly Manis temminckii) exhibits a distinctive external morphology characterized by a head-body length ranging from 35 to 90 cm and a tail length of 30 to 60 cm, with adults typically weighing 7 to 18 kg, though individuals exceeding 20 kg have been recorded.^[13] ^[2] The body is elongated and cylindrical, covered dorsally and laterally by large, overlapping scales composed of keratin, formed from agglutinated hairs, measuring 2 to 5 cm in length and exhibiting a yellow-brown coloration.^[13] These scales encase the head, back, sides, and tail, leaving the snout, chin, throat, sides of the face, and belly unprotected by scales and instead covered in softer, sparsely haired skin.^[2] The scales serve a primary defensive role, interlocking to form a rigid armor when the animal curls into a ball during threats, as observed in field encounters where predators fail to penetrate the scaled exterior.^[13] Scales are periodically shed and regrown throughout the animal's life to accommodate body growth and repair damage, with replacement occurring continuously rather than in discrete annual cycles.^[14] The forelimbs are equipped with five digits, featuring three enlarged, sickle-shaped claws on the middle toes, up to 4 cm long, adapted for excavation, while the hindlimbs have shorter claws.^[13] The tail, tapering to a point, aids in balance and can grasp objects weakly, with scales extending along its length for added protection.^[2] Sexual dimorphism is minimal, with males averaging slightly larger in body mass and length than females, though overlapping size ranges preclude reliable external distinction without measurement.^[2] Scale color may vary subtly from light brown to darker tones influenced by environmental dust accumulation in arid habitats, but the underlying keratin structure remains consistent.^[13] The overall scaled covering constitutes a significant portion of the body surface, estimated to protect approximately the dorsal half, enhancing survival against predation through mechanical resistance verified in biomechanical studies of scale overlap and hardness.^[15]

Internal anatomy and sensory adaptations

The ground pangolin lacks teeth entirely, relying instead on a specialized digestive system adapted for consuming ants and termites. Its tongue, which can extend up to approximately 30 cm, is anchored to an elongated hyoid apparatus that allows protrusion deep into nests, coated in viscous saliva to capture prey.^[16] The stomach functions as a muscular gizzard, where ingested abrasive grit from soil aids in grinding the chitinous exoskeletons of insects, facilitating enzymatic breakdown in the absence of mastication.^[17] Sensory adaptations prioritize chemoreception and audition over vision, suited to a nocturnal, fossorial lifestyle. Vision is poor, with small eyes lacking advanced acuity, though low-light sensitivity may be enhanced by a retinal tapetum lucidum in some pholidotans. Olfaction is acute, supported by enlarged olfactory bulbs and vomeronasal organs that detect pheromonal and prey cues from afar. Hearing is similarly developed, enabling detection of subtle substrate vibrations from insect activity.^[6]^[18]^[19] Physiological efficiencies include a basal metabolic rate roughly half that predicted for mammals of comparable mass, conserving energy from a low-nutrient, formicivorous diet. Water requirements are minimal, met primarily through prey moisture and metabolic water, with conservation achieved via behavioral aridity avoidance rather than specialized renal or osmoregulatory mechanisms.^[20]^[21]

Habitat and distribution

Geographic range

The ground pangolin (Smutsia temminckii) inhabits sub-Saharan Africa, with its current distribution forming an arc from northeastern Chad and Sudan eastward to Ethiopia and Somalia, southward through Kenya, Tanzania, and Zambia, and westward to Angola, extending to the Cape provinces of South Africa.^[2]^[13] This range encompasses fragmented populations across Sahel margins, savannas, and semi-arid zones, often confined to protected areas in regions like Zimbabwe and South Africa's northern provinces.^[22]^[23] Historical records suggest a broader extent prior to colonial-era and modern habitat alterations, including agricultural expansion that fragmented suitable landscapes.^[24] Surveys from the 2020s, such as those in Tanzania's Ruaha landscape and Hwange National Park, document ongoing range contractions linked to land conversion, with verified sightings increasingly limited to remnant savanna patches.^[25]^[22] The species avoids equatorial rainforests and dense forest mosaics, showing a macro-scale preference for drier biomes; extralimital vagrant occurrences beyond this core distribution remain undocumented in recent data.^[2]^[1]

Environmental preferences and adaptability

The ground pangolin (Smutsia temminckii) selects habitats characterized by open savannas, woodlands, and floodplains featuring friable, sandy or loamy soils that facilitate burrowing for refuge and foraging access to subterranean ant and termite nests.^[22] These soil types, often associated with active termite mounds in mopane-dominated vegetation, provide structural integrity for extensive burrow networks while allowing efficient excavation with the species' powerful foreclaws.^[22] Radio-tracking data from central Namibia indicate opportunistic burrow selection, with individuals utilizing both self-dug earthen burrows and pre-existing ones from aardvarks or other excavators, prioritizing proximity to prey colonies over strict habitat typology. In semi-arid environments like the Kalahari, where seasonal droughts prevail, the species demonstrates adaptability through behavioral adjustments rather than rigid habitat fidelity; radio-collared individuals in arid South African populations exhibited no significant selectivity among available shrubveld and dune habitats, exploiting varied terrain as long as invertebrate prey remains viable.^[26] Burrow systems serve a critical thermoregulatory function, shielding against diurnal heat exceeding 40°C and nocturnal cools, with individuals retreating underground during peak temperatures to maintain body temperatures averaging 34–36°C under resource-plentiful conditions.^[21] During prey scarcity induced by dry seasons, pangolins relax body temperature precision, tolerating fluctuations up to 38.2°C maxima and 28.9°C minima, coupled with shifts to diurnal foraging when nocturnal ant activity diminishes, thereby conserving energy without free water intake by deriving hydration solely from prey.^[21] ^[27] This resilience extends to anthropogenic landscapes, where ground pangolins persist in farmlands and fragmented areas provided termite and ant abundances persist, underscoring causal reliance on edaphic factors like soil friability over vegetative density; dense forests are avoided due to compacted soils hindering mobility and burrow stability.^[22] Empirical tracking reveals core burrow reuse for estivation-like dormancy during extreme dry periods, minimizing metabolic demands until monsoon rains restore prey populations, though prolonged droughts may elevate mortality risks via reduced foraging efficiency.

Behavioral ecology

Activity patterns and locomotion

The ground pangolin (Smutsia temminckii) exhibits predominantly nocturnal activity patterns, emerging from burrows shortly after sunset to forage and returning before dawn, with mean active periods of approximately 5.7 hours per night ranging from 1 to 12 hours depending on individual and environmental factors.^[26] Activity onset varies seasonally, with increased diurnal tendencies observed during winter months in arid regions such as the Kalahari, where cooler temperatures may facilitate daytime movement, contrasting with more strictly nocturnal behavior in mesic habitats.^[28] Emergence times show no significant correlation with lunar cycles, indicating that moonlight illumination does not substantially alter their temporal rhythms in studied populations.^[26] Individuals maintain solitary home ranges that vary by sex, age, and habitat, with males typically occupying larger areas of 6 to 14 km² that may overlap multiple female ranges of 0.17 to 23.4 km², while seasonal resource availability influences range size and utilization.^[29]^[30] Locomotion in ground pangolins is characterized by a quadrupedal gait involving deliberate, shuffling steps on all fours, supported by powerful forelimbs adapted for digging burrows and excavating prey with elongated claws.^[31] Under stress, they can adopt a bipedal stance for brief defensive or rapid movements, achieving speeds of up to 5 km/h, though sustained quadrupedal travel remains slow and methodical to conserve energy in their armored form.^[32] Ground pangolins (Smutsia temminckii) exhibit a predominantly solitary social structure, with adults interacting minimally except during brief mating encounters or when females are rearing dependent offspring.^[33]^[2] Field observations indicate no formation of packs, social hierarchies, or cooperative groups among conspecifics in the wild, reflecting their asocial nature and low population densities in suitable habitats.^[2] Territorial boundaries are maintained primarily through olfactory cues, with individuals scent-marking using secretions from enlarged anal glands, urine, and feces to signal occupancy and deter intruders.^[34]^[2] Males and females may have overlapping home ranges in resource-variable environments, but direct confrontations are rare, and intraspecific aggression appears limited to defensive responses rather than territorial disputes, consistent with abundant prey reducing competition.^[34]^[2] Communication relies heavily on chemosensory signals due to the species' acute sense of smell, with visual, tactile, and acoustic modalities playing secondary roles.^[2] Vocalizations are minimal, lacking true vocal cords; observed sounds include hissing during disturbance and huffing or chuffing for short-range signaling, though their role in conspecific interactions remains undocumented in field studies.^[34] Tactile contact occurs opportunistically during mother-offspring associations or mating.^[2] The primary social bond forms between mothers and single offspring, which ride on the mother's tail or back starting around one month post-birth and achieve independence after 3–4 months, after which the young disperses to establish its own territory.^[29] This brief dependency period underscores the species' low investment in prolonged familial ties beyond nutritional support.

Diet and foraging behavior

The ground pangolin (Smutsia temminckii) maintains a strictly myrmecophagous diet consisting exclusively of ants (Formicidae) and termites (Isoptera), with no consumption of plant matter, bees, wasps, or other insect orders. Ants predominate, comprising approximately 96% of dietary occurrences across sampled prey species, while termites form a minor portion; pangolins selectively target 15 ant and 5 termite species out of available options, favoring those with large, accessible colonies such as Anoplolepis custodiens (ants) and Trinervitermes (termites).^[35]^[36] This selectivity reflects preferences for prey with high biomass availability rather than random foraging, enabling efficient energy acquisition from subterranean and arboreal nests.^[35] Foraging occurs nocturnally or crepuscularly, primarily guided by acute olfaction to detect volatile cues from insect colonies, as vision is limited; the elongated, prehensile tongue—anchored near the pelvis and extensible up to 40 cm—then extracts prey via adhesive saliva, without reliance on teeth for mastication. Small stones are ingested to aid trituration in the glandular stomach, where enzymatic digestion breaks down the keratinous exoskeletons. Daily consumption supports metabolic demands, with individuals capable of ingesting thousands of insects per bout, though precise biomass varies with prey density.^[37]^[38] Dietary composition exhibits seasonal flexibility to counter prey scarcity, particularly in semi-arid habitats; during winter dry periods, pangolins shift toward energy-dense species like certain ants and termites, extending foraging bouts to maintain intake despite reduced overall availability. Such adaptations mitigate nutritional shortfalls but do not fully compensate for prolonged resource limitations.^[39]^[39]

Reproduction and lifecycle

Mating and breeding

The ground pangolin (Smutsia temminckii) employs a polygynous mating system, in which individual males mate with multiple females, often competing aggressively through physical confrontations to secure access to receptive females.^[2] Males and females are otherwise solitary and do not form lasting pair bonds, with encounters limited to brief mating periods.^[2]^[40] Breeding occurs year-round across much of the species' range, though regional observations suggest possible peaks aligned with seasonal conditions such as increased rainfall or warmer months; for instance, in Namibia, mating activity has been noted from September to December.^[2]^[41] Due to their nocturnal and elusive habits, detailed courtship behaviors remain poorly documented, but as with other pangolins, olfactory cues likely play a key role in mate detection given the species' reliance on acute sense of smell for navigation and resource location.^[2]^[6] Gestation lasts approximately 105-140 days, typically resulting in the birth of a single offspring, though twins are occasionally reported in rare cases.^[42]^[1]^[2] This low fecundity contributes to slow population recovery, as females produce only one young annually under optimal conditions.^[41]

Development and parental care

Ground pangolin neonates, typically a single offspring, are born in underground burrows or shelters after a gestation period of approximately 139 days, with birth weights ranging from 200 to 500 grams.^[2] At birth, the young possess soft, pliable scales that gradually harden and darken over the first few weeks, providing increasing protection.^[43] Females provide exclusive parental care, nursing the offspring in the burrow for the initial 2–4 weeks while abstaining from foraging to minimize disturbance.^[2] Once mobile, the juvenile clings to the mother's back or tail during nocturnal foraging excursions, allowing it to accompany her without impeding locomotion.^[2] This carrying behavior persists until the young reaches approximately 3 kg, nearing subadult size. Weaning begins around 3 months of age, as juveniles start supplementing milk with ants and termites scavenged from the mother's foraging sites.^[2] Full independence follows by about 2 years, when the offspring disperses to establish its own territory, coinciding with attainment of sexual maturity.^[6]^[40] In response to threats, mothers curl into a tight defensive ball, enveloping the juvenile beneath their armored body to deter predators.^[44] Juvenile survival appears high under maternal protection, contingent on evasion of predation, though empirical mortality rates remain undocumented due to limited field observations.^[2]

Ecological role

Trophic interactions

The ground pangolin (Smutsia temminckii) occupies a mid-trophic level as a specialized myrmecophage, preying on ants and termites with minimal competition from other mammals due to its unique morphological adaptations, such as elongated claws and a protrusible tongue, which limit overlap with generalist insectivores like aardvarks or birds.^[2] This dietary niche reduces intraspecific and interspecific competition, positioning it as a low-density consumer in savanna and woodland food webs.^[45] As prey, ground pangolins encounter few natural predators, primarily large carnivores including lions (Panthera leo), spotted hyenas (Crocuta crocuta), and leopards (Panthera pardus), which can occasionally breach their defenses through sustained attacks or crushing bites.^[45] Verified predation events are rare, attributed to the pangolin's nocturnal habits and effective countermeasures, though non-fatal interactions—such as predators removing tracking devices—indicate opportunistic encounters with leopards and hyenas in regions like South Africa's Kalahari.^[46] Humans act as apex predators via direct hunting, rendering scales ineffective against tools or firearms.^[2] Primary defenses include rapid curling into a spherical ball, with overlapping keratin scales forming an impenetrable barrier that protects soft underparts, head, and limbs; this posture is supplemented by powerful tail lashes capable of injuring assailants and emission of a musky, noxious secretion from anal glands to deter approach.^[2] These adaptations confer high efficacy against smaller or less persistent predators, with natural mortality from predation appearing low based on limited field observations, but they offer negligible protection against human exploitation or large felids employing prolonged grappling.^[45]

Ecosystem services

The ground pangolin (Smutsia temminckii) contributes to ecosystem services mainly by preying on ants and termites, its exclusive diet, which curbs populations of these invertebrates known to inflict substantial damage on crops and timber in agricultural and forested areas.^[1] ^[38] Termites, in particular, represent a major agricultural pest across sub-Saharan Africa, where ground pangolin foraging reduces infestation risks without reliance on chemical interventions.^[47] Burrowing for food sources aerates compacted soils, improving permeability and microbial activity, while the digestion and deposition of insect remains aids nutrient redistribution, enhancing soil fertility in savanna and woodland habitats.^[48] ^[49] These activities position the species as an ecosystem engineer, with indirect benefits extending to vegetation health through moderated insect herbivory and termite mound dynamics.^[50] Potential disservices, such as abandoned burrows posing hazards to livestock or altering microhabitats, appear minimal and lack quantitative assessment in available studies.^[51]

Conservation and threats

Current status and population data

The ground pangolin (Smutsia temminckii) is classified as Vulnerable on the IUCN Red List, based on a 2019 assessment that continues to reflect its status amid ongoing data limitations as of 2025 evaluations. In June 2025, the U.S. Fish and Wildlife Service proposed listing it as Endangered under the Endangered Species Act, recognizing severe pressures on its persistence wherever found.^[52] Population size remains poorly quantified, with no reliable global census available; coarse estimates suggest fewer than 100,000 mature individuals across its range, though these derive from fragmented local data rather than comprehensive surveys.^[42] Overall trends indicate declines, inferred from sparse field observations and indirect indicators like habitat contraction, with generation times estimated at approximately nine years supporting projections of 30-40% reductions over recent decades.^[53] Effective monitoring is constrained by the species' nocturnal habits, solitary lifestyle, and cryptic burrowing, which yield low detection rates in standard surveys. Camera-trap deployments in the 2020s have captured occasional individuals in protected savanna and grassland sites, suggesting localized stability where enforcement is strong, but such efforts underscore persistent gaps in coverage across its extensive sub-Saharan distribution from Senegal to South Africa.^[54]^[55]

Primary threats: habitat and exploitation

Habitat loss through agricultural expansion threatens ground pangolin populations by converting savanna and grassland habitats essential for foraging and burrowing. Cultivated land across Africa expanded by 60 million hectares from 2000 to 2020, with significant conversions from grasslands in central and southern regions overlapping the species' range.^[56] In southern Africa, including areas like KwaZulu-Natal Province, crop agriculture directly fragments pangolin habitats, reducing available ant and termite prey resources. Illegal poaching for scales, primarily destined for traditional Chinese medicine markets in Asia, represents the dominant exploitation pressure. Reported seizures of African pangolin scales exceeded 287,000 kg across 130 incidents involving 15 African countries from 2016 onward, with Nigeria serving as a key transit hub.^[57] ^[58] In South Africa, authorities arrested 679 individuals linked to pangolin trade between 2016 and 2024, reflecting sustained trafficking volumes.^[59] Temminck's ground pangolin scales, composed of keratin akin to fingernails or rhino horn, drive this trade despite lacking empirical medicinal value.^[60] Local bushmeat hunting affects ground pangolins but remains secondary to scale poaching in scope and impact, particularly outside West African locales where meat commands higher local value than scales.^[61] These pressures interact synergistically: expanding road networks fragment habitats, facilitate poacher access to remote savannas, and elevate roadkill risks for nocturnal pangolins.^[62] ^[63] Concurrent climate-driven drying in African savannas exacerbates habitat degradation by shifting vegetation patterns and diminishing invertebrate prey density.^[64]

Conservation measures and outcomes

In 2016, all pangolin species, including the ground pangolin (Smutsia temminckii), were listed in Appendix I of the Convention on International Trade in Endangered Species (CITES), effectively banning commercial international trade to curb poaching driven by demand for scales and meat.^[65] Nationally, the species receives protection under frameworks like South Africa's Threatened or Protected Species (TOPS) regulations, which classify it as endangered and prohibit unlicensed possession or trade.^[1] Protected areas such as Kruger National Park implement habitat safeguards alongside rehabilitation protocols, involving nightly foraging simulations for rescued individuals prior to release to enhance survival post-captivity.^[66] Anti-poaching patrols, often in collaboration with law enforcement, have intensified in South Africa during the 2020s, leading to increased arrests and localized reductions in trafficking attempts within monitored regions.^[67] Captive breeding and reintroduction programs face significant hurdles, with high neonatal mortality rates—often exceeding 90% in early attempts—and challenges in replicating wild diets contributing to limited viable offspring.^[68] Initiatives like the Phinda Pangolin Reintroduction Project have achieved modest progress by translocating and monitoring individuals to re-establish populations in historically depleted areas, providing a potential breeding nucleus.^[69] Outcomes remain mixed, as illegal trade endures despite prohibitions; global pangolin scale seizures declined by 84% from 2019 peaks to 2024 levels, partly attributable to pandemic disruptions in supply chains and enhanced enforcement, yet these intercepts represent only a fraction of total volume due to underreporting and adaptive smuggling tactics.^[70]^[71] In South Africa, patrol efforts correlate with fewer detected local trade incidents in the early 2020s compared to pre-2019 surges, but broader persistence of demand-driven poaching underscores enforcement gaps and the need for supply-chain interventions.^[72] Reintroduction successes are site-specific and small-scale, failing to offset inferred population pressures from ongoing exploitation.^[73]

Human interactions

Traditional uses and cultural significance

In southern African communities, particularly among tribal groups in South Africa, scales and bones of the ground pangolin (Smutsia temminckii) are employed in traditional healing practices to address ailments including rheumatism, headaches, back pains, and swollen limbs.^[74] ^[24] The flesh serves as bushmeat and is regarded as a delicacy in some locales, contributing to local protein consumption.^[74] ^[75] Parts of the animal, including live specimens, feature in muti rituals, where they are incorporated into spiritual or protective practices by traditional healers.^[75] Ethnographic surveys document scales being ascribed properties to ward off evil spirits and provide protection, with such beliefs persisting in rural areas.^[76] Ground pangolins hold cultural value in certain communities as symbols of prosperity, believed to ensure good rains, harvests, and overall well-being when ritually handled.^[53] These uses trace to pre-colonial traditions but have gained economic prominence in impoverished rural regions, such as those within the species' distributional range in South Africa, where harvesting supports livelihoods amid limited alternatives.^[24]^[74]

Scientific evaluation of trade claims and policy debates

Pangolin scales, composed primarily of keratin—a fibrous protein identical to that in human fingernails and hair—exhibit no pharmacological activity or medicinal efficacy beyond placebo effects in traditional applications.^[77] Multiple peer-reviewed analyses from the 2010s and 2020s, including systematic reviews of traditional Chinese medicine claims, confirm the absence of bioactive compounds supporting purported benefits such as treating rheumatism or stimulating blood flow.^[78]^[79] Demand persists due to entrenched cultural beliefs rather than empirical validation, with scales removed from official pharmacopeias in regions like China for lacking scientific substantiation.^[77] Policy debates surrounding pangolin trade, including for the ground pangolin (Smutsia temminckii), center on the efficacy of absolute bans versus regulated alternatives. International commercial trade bans under CITES Appendix I, implemented in 2017 following the 2016 CoP17 decision, have failed to eradicate demand, as evidenced by persistent black market seizures exceeding hundreds of thousands of animals annually between 2016 and 2019, alongside rising per-kilogram prices surpassing $500 in some markets.^[80]^[59] Critics argue these prohibitions exacerbate illegal trafficking by driving activities underground without addressing root cultural drivers, potentially inflating scarcity perceptions and poaching incentives, while enforcement challenges in source countries like those in southern Africa compound non-compliance.^[81]^[82] Proponents of regulated trade advocate for sustainable harvesting or captive breeding to generate revenue for communities, undercutting poachers through legal supply chains, as explored in feasibility studies on pangolin farming.^[83] Such approaches draw on property rights frameworks, positing that incentivizing local stewardship via controlled quotas or ranching—rather than top-down bans—could align economic interests with conservation, particularly for species like the ground pangolin where baseline population data remains imprecise, with mature estimates varying widely from 7,000 to 32,000 individuals due to sampling gaps and reliance on indirect indicators.^[1]^[84] Opponents, often from conservation NGOs, counter that biological constraints like the species' slow reproduction and cryptic habits render farming unviable at scale, risking laundering of wild-sourced products and accelerating local extirpations amid uncertain decline rates.^[83] These tensions highlight how sparse, model-dependent population metrics may overestimate threats, fueling precautionary bans that overlook potential for evidence-based management.^[1]

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Dec 4, 2024 · Unlike olfaction and vision which have been followed with interest for pangolins, their auditory sensation is still mysterious, and our results ...
[21]
Energetics suggest cause for even further conservation concern for ...
Sep 13, 2019 · The measured field metabolic rates for Temminck's ground pangolins also suggest a slow life history. Ignoring the unusually high and low values ...
[22]
Temminck's pangolins relax the precision of body temperature ...
Aug 28, 2023 · Low 24-h minimum body temperatures presumably allowed pangolins to conserve energy by reducing metabolism, as explained by the Q10 effect in ...
[23]
Ecology, status, and distribution of Temminck's pangolin (Smutsia ...
Feb 26, 2024 · Temminck's pangolin, Smutsia temminckii (Smuts, 1832), faces conservation threats of poaching and trafficking in Zimbabwe.Missing: original | Show results with:original
[24]
Distribution records for Temminck's Ground Pangolin (Smutsia...
It occurs in the Northern Cape, North West, Limpopo, Mpumalanga and KwaZulu- Natal provinces (Figure 1). They are thought to be locally extinct in the Free ...
[25]
[PDF] ethnozoological survey of the - African Pangolin Working Group
Historically, the range of the Temminck's ground pangolin in South Africa is believed to cover eight of the nine provinces with its occurrence being ...
[26]
Local Ecological Knowledge Insights Into the Distribution and ...
Aug 16, 2025 · Temminck's pangolin (Smutsia temminckii) is believed to be widespread across various ecosystems in Tanzania. In contrast, the white‐bellied (or ...
[27]
[PDF] Home range, habitat selection and activity patterns of an arid-zone ...
All previous behavioural studies of Temminck's ground pangolins (Smutsia temminckii) have focused on populations in mesic regions.Missing: molting dimorphism
[28]
10 Amazing Pangolin Facts | Working with Wildlife
Furthermore, while ground pangolins do drink water when available, they do not require water and are able to obtain sufficient moisture from their prey alone.
[29]
[PDF] Home range, habitat selection and activity patterns of an arid-zone ...
The time at which activity commenced varied seasonally as well as among individuals, with an increase in diurnal activity during winter. Young pangolins also ...
[30]
Temminck's pangolin - African Pangolin Working Group
Home ranges vary in size from 600–1 400 hectares (6–14 km2),and are smaller for younger animals. Some studies have found that a male's home range overlaps the ...
[31]
Home range size and distribution in a wild population of Cape ...
Aug 7, 2025 · In Zimbabwe, home ranges are estimated to be between 0.17 -23.4 km², with males having the largest ranges and temporarily overlapping several ...<|separator|>
[32]
Osteology and Radiological Anatomy of the Thoracic Limbs of ...
Dec 11, 2017 · Temminck's ground pangolin is the only pangolin present in South Africa. It is a myrmecophagous mammal with a bipedal gait.
[33]
A novel description of the Vastus lateralis morphology of the ...
Mar 31, 2022 · The Temminck's ground pangolin (Manis temminckii, also called Smutsia temminckii) is a unique terrestrial mammal covered with keratinous scales ...
[34]
Mpala Live! Field Guide: Pangolin | MpalaLive
Social Structure. Pangolins are solitary except when mating or when females are rearing their young. · Communication. Ground pangolins communicate through scents ...Missing: organization | Show results with:organization
[35]
What is a pangolin - African Pangolin Working Group
Pangolins make very few sounds (as they have no vocal chords). Different species may create varying sounds such as 'chuffing' through inhalation or exhalation ...Missing: communication | Show results with:communication
[36]
Ecological factors affecting the feeding behaviour of pangolins ...
Feb 28, 2006 · Five termite and 15 ant species were preyed on by pangolins. Six of these species constituted 97% of the diet while ants formed 96% of the diet.
[37]
Ecological factors affecting the feeding behaviour of pangolins ...
S. temminckii was found to selectively feed on 30 ant and 10 termite species, out of the 50 potential prey species, preferring an ant species, Anoplolepis ...
[38]
Investigating the use of sensory information to detect and track prey ...
Jun 17, 2020 · Behavioural anecdotes of pangolins using their elongated snouts and tongues to forage seem to support the idea that they rely on olfaction and ...
[39]
[PDF] Ecological factors affecting the feeding behaviour of pangolins ...
Cape pangolins Manis temminckii feed exclusively on ants and termites and they have several morphological adaptations that enable them to gain access to the ...
[40]
Do seasonal dietary shifts by Temminck's pangolins compensate for ...
Pangolins responded to winter resource scarcity by increasing the proportion of consumed Trinervitermes termites, but the shift did not prevent low energy ...Missing: rate | Show results with:rate
[41]
Pangolin Fact Sheet | Blog | Nature - PBS
Mar 25, 2020 · The gestation period differs based on species but typically ranges between 70 to 140 days. While African pangolin females give birth to a single ...
[42]
Pangolin Project Update - The AfriCat Foundation
Females give birth to one pup per year after a gestation period of about 140 days. Marked breeding seasons have been noted across their range. In Namibia, a ...
[43]
Everything you need to know about Temminck's pangolins - Born Free
Temminck's pangolins are classed as 'Vulnerable' by the IUCN, threatened by human infrastructure, poaching and habitat loss. Found in low-lying regions, living ...Missing: behavior | Show results with:behavior
[44]
Pangopups: All About Baby Pangolins
Sep 1, 2021 · At birth, pangolins measure just six inches and weigh 3/4 of a pound. During the first days of life, pangolin mothers protect their young in ...
[45]
Pangolin Protection - Global Conservation Force
A pangolin's best defense is to roll in a ball and “hope for the best” – using its overlapping scales as protection against a predators bite or grasp.Missing: maternal juveniles
[46]
Manis temminckii (Ground pangolin) - biodiversity explorer
Predators, parasites and commensals. Few natural predators, only hyaena and lion are able to penetrate the scale defense. Reproduction. After a gestation ...
[47]
Non‐fatal removal of ground pangolin (Smutsia temminckii Smuts ...
Oct 25, 2023 · The occurrence of both fatal and non-fatal interactions between pangolins and predators gives an indication of predator activity in the study ...Missing: records | Show results with:records
[48]
The pangolin puzzle: scratching the surface
Jun 3, 2020 · By preying heavily on termites, pangolins influence their abundance and community structure in a way that minimizes these losses and thereby ...
[49]
pangolins: ecosystem gardeners worth conserving
Feb 16, 2023 · Greenwood adds that pangolins help to regulate populations of termites and ants by way of their voracious appetites. Some estimates hold that ...Missing: Manis temminckii control
[50]
Pangolins: Natural Pest Controllers and Soil Caretakers
Nov 1, 2011 · Pangolins provide pest control by eating insects and improve soil by burrowing and aerating it, mixing and improving nutrient quality.
[51]
To Save Pangolins: A Nutritional Perspective - PMC - PubMed Central
Nov 14, 2022 · Swart et al. [17] found that Smutsia temminckii feeds on 15 kinds of ants and 5 kinds of termites without an obvious preference. The literature ...
[52]
[PDF] management plan for the community-based pangolin conservation ...
5.1 Protecting pangolins in community forests also ensures the preservation of broader ecosystem services, such as soil aeration and pest control, which.<|separator|>
[53]
Endangered Species Status for Seven Species of Pangolin
Jun 17, 2025 · Home ranges have been estimated to be 96 hectares (ha) (0.37 square miles (mi2 )) for males and 24.4 ha (0.09 mi2 ) for females across various ...
[54]
Save This Species: Temminck's Pangolin - The Revelator
Sep 12, 2025 · Species name: Temminck's pangolin (Smutsia temminckii), also known as the ground, Cape, or steppe pangolin. · IUCN Red List status: Vulnerable.
[55]
Pangolins in global camera trap data: Implications for ecological ...
Our results suggest that monitoring pangolins over space or time remains very challenging with camera traps. At coarse scales, we have shown that it is ...
[56]
[PDF] Methods for monitoring populations of pangolins (Pholidota: Manidae)
Equally, if and how levels of activity and circadian patterns differ. Page 17. 16 between species, by season, the lunar phase, and climatic conditions, remains ...
[57]
(PDF) Spatial-temporal patterns of cultivated land expansion and ...
Aug 6, 2025 · Results indicated that the net cultivated land area increased by 6.00 × 10⁷ ha from 2000 to 2020. Over 62% of cultivated land remained stable, ...
[58]
Scaly trade: Analyses of the media reports of pangolin (Pholidota ...
We report 130 seizure incidents accounting for 287,272 kg of African pangolin scales involving 15 African and seven Asian countries. Overall, Nigeria and the ...
[59]
Genomic analyses reveal poaching hotspots and illegal trade in ...
Dec 14, 2023 · Network mapping of African pangolin seizures identified Nigeria as the highest-volume transit hub in Africa, where traffickers amass pangolin ...
[60]
https://www.unodc.org/documents/data-and-analysis/wildlife/2020/WWLC20_Chapter_4_Pangolin.pdf
[61]
[PDF] PANGOLIN SCALES - Unodc
Pangolin scales are used medicinally, especially in traditional Chinese medicine, and are the main product of most recent seizures, mainly from Africa.Missing: 2016-2023 | Show results with:2016-2023
[62]
In southeast Nigeria, pangolins hunted for meat, not scales, study finds
Sep 10, 2025 · Since a pangolin's meat fetched 3-4 times as much money as its scales in the local market, there was no financial incentive to trade the scales.
[63]
Threats to Pangolins
The building of new roads, does not only fragment habitat, it also creates inroads into previously inaccessible areas, with few legislated controls and ...Missing: climate | Show results with:climate
[64]
A Review of the Anthropogenic Threats Faced by Temminck's ...
Aug 7, 2025 · Throughout its range, Temminck's ground pangolin, Smutsia temminckii, is becoming increasingly threatened, predominantly as a result of ...
[65]
Extinction of Pangolins - Climate Change Writers
Pangolins are becoming increasingly vulnerable to extinction as our forests diminish and temperatures rise. This is due to the loss of their habitats and food ...Missing: roads | Show results with:roads
[66]
[PDF] CoP17 Inf. 59 - CITES
Oct 5, 2016 · For the full IUCN Red List account for this species including distribution map, see here. Distribution: Ranges from Guinea, Sierra Leone and ...
[67]
Back to the Wild – Our Stories – Scientific Services - SANParks
Feb 24, 2024 · The pangolin rehabilitation protocol requires walking the animals nightly for five hours in an area with sufficient ants to feed on. Pangolins ...
[68]
https://pmc.ncbi.nlm.nih.gov/articles/PMC4490220/
[69]
Captive breeding of pangolins: current status, problems and future ...
Jun 8, 2015 · The winner will mate with the female pangolin, with the mating period generally lasting three to five days. Female pangolins have two to five ...<|separator|>
[70]
Pangolin Reintroduction Project - andBeyond
This groundbreaking, collaborative project to reintroduce the Temminck's ground pangolin to a region where it has been locally extinct.
[71]
Global trafficking of pangolin scales, elephant ivory plummets post ...
May 2, 2025 · By 2024, the volume of seized pangolin scales and ivory had declined by 84 per cent and 74 per cent respectively compared to 2019 levels. “It is ...
[72]
Lack of data and reporting gaps hamper global efforts to protect ...
Aug 27, 2025 · Visit the IUCN Congress website for the full list of motions. Notes to editors. The full “Report on the conservation status of pangolins (Manis ...
[73]
[PDF] An analysis of pangolin scale and ivory trafficking, 2015-2024
For pangolin scales, this trend is driven by an uptick in multi-ton land-based and stockpile seizures in Asia and Africa since 2023. For ivory, the trend is ...
[74]
The Return of the Pangolin | Reversing a Local Extinction - andBeyond
The aim is to help re-establish a new and healthy population of Temminck's ground pangolin, provide a breeding nucleus from which to create further ...A Conservation First · Like-Minded Conservationists · A Slow RehabilitationMissing: programs | Show results with:programs
[75]
Ethnozoological Survey of Traditional Uses of Temminck's Ground ...
Aug 7, 2025 · ... muti, headaches, back aches/pains, swollen feet/. legs, foot aches ... pangolins, Manis temminckii. South African Journal of Wildlife ...
[76]
Meet the Species: Cape Pangolin
Feb 16, 2012 · The Cape Pangolin (Smutsia temminckii) is one of the larger and more terrestrial of Africa's four pangolin species.Missing: subspecies | Show results with:subspecies<|control11|><|separator|>
[77]
(PDF) Symbolism, myth and ritual in Africa and Asia - ResearchGate
Temminck's pangolins are used as sacrifices to predict rainfall and pangolin products are often used as protection and good luck charms to ward off bad luck ( ...
[78]
Evidence for the medicinal value of Squama Manitis (pangolin scale)
There is no reliable evidence that Squama Manitis has special medicinal value. The removal of Squama Manitis from Pharmacopoeia is rational.
[79]
Pangolin scales are not a party drug, say researchers
Aug 9, 2019 · It has no scientifically proven medicinal value. Jacobs was doing related research when she first heard the rumors about pangolin scales ...Missing: studies | Show results with:studies<|separator|>
[80]
The Use and Prescription of Pangolin in Traditional Vietnamese ...
Feb 22, 2021 · Further, there was the acknowledgement that there is no scientific evidence for any medicinal properties of pangolin scales: 'The knowledge ...
[81]
United States Finds China's Pangolin Trade Undermines Wildlife ...
Sep 12, 2023 · The US Department of the Interior has announced that China's failure to halt its trade in endangered pangolins diminishes the effectiveness of an international ...Missing: black | Show results with:black
[82]
Are wildlife trade bans backfiring? - Knowable Magazine
Nov 13, 2020 · Legal trade in Asian pangolins dropped to near-zero in the 13 years after a CITES agreement to not export wild-caught pangolins from their ...<|separator|>
[83]
Curbing the trade in pangolin scales in China by revealing ... - Nature
Jan 21, 2025 · The illegal trading in pangolin scales and meat in China differ significantly. Pangolin scales originate primarily from Africa and are ...<|control11|><|separator|>
[84]
Evaluating the feasibility of pangolin farming and its potential ...
Pangolins are threatened by overexploitation for local and international use. They are subject to an international commercial trade ban, and are also the ...Missing: arguments | Show results with:arguments
[85]
Population structure and demographic history of two highly-trafficked ...
Sep 27, 2024 · Pangolin's population genetic structure can be used to trace the geographic origins of trafficked scales, but substantial sampling gaps across ...