African Pangolins Research Articles

Population structure and demographic history of two highly-trafficked species of pangolin in the Congo Basin.

African pangolins are hunted for their meat and for use in local traditional medicine, as well as for their scales, which are trafficked internationally, especially to growing Asian markets. Pangolin's population genetic structure can be used to trace the geographic origins of trafficked scales, but substantial sampling gaps across pangolins' ranges hinder these efforts. In this study, we documented population structure and dynamics in the two species of African pangolin, the white-bellied pangolin (Phataginus tricuspis) and the giant pangolin (Smutsia gigantea) in the underexplored Republic of Congo. Using the mitochondrial control region and two nuclear markers (beta-fibrinogen and titin), we identified high genetic diversity in both species. We document a distinct mitochondrial lineage of the white-bellied pangolin, which was most likely shaped by river barriers together with dynamics of forest refugia related to the climatic shifts during the Pleistocene. We detected population growth in the white-bellied pangolin coinciding with a dry period during the Pleistocene, suggesting some ability for this typically forest-dwelling species to persist under diverse environmental conditions. Using landscape genetics, we found all but one of the pangolins we sampled at bush meat markets originated locally. A single individual appeared to have been imported to Congo from Cameroon. These findings significantly contribute to our understanding of pangolin population biology and local trade dynamics. In addition, our data from a previously unstudied part of pangolins' ranges will help us to better understand international wildlife trafficking patterns and to target conservation and protection strategies for these highly vulnerable species.

Ants (Hymenoptera: Formicidae) and termites (Blattodea: Termitoidae) in the diet of wild white-bellied pangolin (Phataginus tricuspis) in forest-savanna habitats of Cameroon.

The white-bellied pangolin Phataginus tricuspis (Rafinesque 1821) is a semiarboreal species occurring in tropical sub-Saharan Africa. It is the world's most trafficked African pangolin species based on volumes recorded in seizures. Reintroduction of confiscated live pangolins and ex-situ rearing are being explored worldwide as a conservation action. However, the husbandry of seized animals is challenging as the diet of the white-bellied pangolin is poorly known and little studied. We analyzed the stomach contents of dead white-bellied pangolins from two forest-savanna protected areas. Stomach content samples from 13 white-bellied pangolin specimens contained ~165,000 Arthropoda, mostly Hymenoptera (60.34%) and Blattodea (39.66%). Overall, we identified 39 termite and 105 ant species consumed as prey by pangolins. Individual pangolins examined had fed on a maximum of 31 ant species and 13 termite species. The termite and ant species richness varied significantly across the pangolins'last consumed meal. We recorded 24 ant genera dominated by Crematogaster (relative importance [RI] = 17.28). Out of 18 termite genera recorded, the genus Pseudacanthotermes (RI = 17.21) was the most important prey. Ten ant species were preferentially eaten by white-bellied pangolin, with Crematogaster acis being the most common prey species. Four species of termite were most frequently eaten with Pseudacanthotermes militaris being the most abundant. The mean abundance of ants and termites varied among pangolin individuals. The season did not influence the mean abundance of termites eaten by pangolin individuals. However, ant abundance in stomach contents was significantly higher in the dry season. An improved understanding of pangolin feeding behavior and prey selection may help inform conservation husbandry efforts. For example, nutritional analysis of the food eaten by wild pangolins can guide the development of nutritional diets for captive pangolins.

Women and urban wildmeat trafficking in the Republic of Congo

As a result of declining biodiversity and increasing rates of urbanization, the illegal urban wildmeat trade is projected to become an integral sector of the wildlife crime industry. Adequate assessment of urban wildmeat trafficking requires investigation into the roles and behaviors of individuals who engage in wildlife crimes. However, akin to much of wildlife crime literature, women's engagement within the urban wildmeat trade have received little investigation. The objectives of our investigation were to (1) explore relationships between women and wildlife products across the supply chain, and (2) determine whether a significant relationship exists between women and specific wildlife products. Through systematic social observations, we evaluate the gendered dimensions of urban wildmeat trafficking in the Republic of Congo between the urban centers of Brazzaville and Pointe-Noire. We place particular emphasis on species of conservation concern, namely great apes, African pangolins, and dwarf crocodiles. Results indicate that there are gendered variations at the species and the geographic level, indicating that women are sourcing and sending their products to different locations than men, and that women are specializing in trade of different species. We attest that urban wildmeat trafficking prevention strategies implement a gender-aware approach due to the unique ways that individuals engage with the industry and how that engagement is gendered.

Conservation genetics of the white-bellied pangolin in West Africa: A story of lineage admixture, declining demography, and wide sourcing by urban bushmeat markets.

During the last 40 years, the volumes of African pangolins feeding the illegal wildlife trade have dramatically increased. We conducted a conservation genetics survey of the most traded African species, the white bellied pangolin (WBP; Phataginus tricuspis), across three West African countries including Guinea, Côte d'Ivoire, and Ghana. Our study combining mitochondrial DNA sequencing and microsatellite genotyping is the first to reveal a wide pattern of admixture between two of the six mitochondrial lineages as previously delimited within WBP. We found a signature of isolation by distance but a lack of population genetic structuring, supporting the idea that WBP may have underestimated dispersal abilities. Levels of genetic diversity were low in West African lineages (WAfr and Gha) compared to Central Africa, reinforcing the picture of genetic pauperization shared by West African WBP. We observed a 85%-98% decline in the effective population size of WBP occurring c. 3200 to 400 ya, with current numbers (520-590) at the lower end of the conservative thresholds for minimum viable population size. The microsatellite markers were powerful enough to differentiate between individuals and identify replicated samples, confirming the utility of this approach in tracing the pangolin trade. Genetic diversity estimates confirmed that Yopougon, the main bushmeat market from Abidjan (Côte d'Ivoire), was fed by a large trade network as confirmed by vendors reporting 10 different sources situated 62-459 km away from the market. We conclude that WBP distributed in the Upper Guinean Block should be considered a single management unit of high conservation concern, as impacted by genetic diversity erosion, drastic decline in effective population size, and wide range sourcing for feeding urban bushmeat markets. Given the genetic admixture pattern detected within WBP from West Africa, we advocate for a multi-locus strategy to trace the international trade of the species.

Genomic analyses reveal poaching hotspots and illegal trade in pangolins from Africa to Asia.

The white-bellied pangolin (Phataginus tricuspis) is the world's most trafficked mammal and is at risk of extinction. Reducing the illegal wildlife trade requires an understanding of its origins. Using a genomic approach for tracing confiscations and analyzing 111 samples collected from known geographic localities in Africa and 643 seized scales from Asia between 2012 and 2018, we found that poaching pressures shifted over time from West to Central Africa. Recently, Cameroon's southern border has emerged as a site of intense poaching. Using data from seizures representing nearly 1 million African pangolins, we identified Nigeria as one important hub for trafficking, where scales are amassed and transshipped to markets in Asia. This origin-to-destination approach offers new opportunities to disrupt the illegal wildlife trade and to guide anti-trafficking measures.

Uncovering the magnitude of African pangolin poaching with extensive nanopore DNA genotyping of seized scales.

Trade in pangolins is illegal, and yet tons of their scales and products are seized at various ports. These large seizures are challenging to process and comprehensively genotype for upstream provenance tracing and species identification for prosecution. We implemented a scalable DNA barcoding pipeline in which rapid DNA extraction and MinION sequencing were used to genotype a substantial proportion of pangolin scales subsampled from 2 record shipments seized in Singapore in 2019 (37.5t). We used reference sequences to match the scales to phylogeographical regions of origin. In total, we identified 2346 cytochrome b (cytb) barcodes of white-bellied (Phataginus tricuspis) (from 1091 scales), black-bellied (Phataginus tetradactyla) (227 scales), and giant (Smutsia gigantea) (1028 scales) pangolins. Haplotype diversity was higher for P. tricuspis scales (121 haplotypes, 66 novel) than that for P. tetradactyla (22 haplotypes, 15 novel) and S. gigantea (25 haplotypes, 21 novel) scales. Of the novel haplotypes, 74.2% were likely from western and west-central Africa, suggesting potential resurgence of poaching and newly exploited populations in these regions. Our results illustrate the utility of extensively subsampling large seizures and outline an efficient molecular approach for rapid genetic screening that should be accessible to most forensic laboratories and enforcement agencies.

Pharyngeal Communities and Antimicrobial Resistance in Pangolins in Gabon.

Wildlife can be a reservoir and source of zoonotic pathogens for humans. For instance, pangolins were considered one of the potential animal reservoirs of SARS-CoV-2. The aim of this study was to assess the prevalence of antimicrobial-resistant species (e.g., extended-spectrum β-lactamase [ESBL]-producing Enterobacterales) and Staphylococcus aureus-related complex and to describe the bacterial community in wild Gabonese pangolins. The pharyngeal colonization of pangolins sold in Gabon (n = 89, 2021 to 2022) was analyzed using culture media selective for ESBL-producing Enterobacterales, S. aureus-related complex, Gram-positive bacteria and nonfermenters. Phylogenetic analyses of ESBL-producing Enterobacterales was done using core-genome multilocus sequence typing (cgMLST) and compared with publicly available genomes. Patterns of cooccurring species were detected by network analysis. Of the 439 bacterial isolates, the majority of species belonged to the genus Pseudomonas (n = 170), followed by Stenotrophomonas (n = 113) and Achromobacter (n = 37). Three Klebsiella pneumoniae isolates and one Escherichia coli isolate were ESBL-producers, which clustered with human isolates from Nigeria (MLST sequence type 1788 [ST1788]) and Gabon (ST38), respectively. Network analysis revealed a frequent cooccurrence of Stenotrophomonas maltophilia with Pseudomonas putida and Pseudomonas aeruginosa. In conclusion, pangolins can be colonized with human-related ESBL-producing K. pneumoniae and E. coli. Unlike in other African wildlife, S. aureus-related complex was not detected in pangolins. IMPORTANCE There is an ongoing debate if pangolins are a relevant reservoir for viruses such as SARS-CoV-2. Here, we wanted to know if African pangolins are colonized with bacteria that are relevant for human health. A wildlife reservoir of antimicrobial resistance would be of medical relevance in regions were consumption of so-called bushmeat is common. In 89 pangolins, we found three ESBL-producing Klebsiella pneumoniae strains and one ESBL-producing Escherichia coli strains, which were closely related to isolates from humans in Africa. This points toward either a transmission between pangolins and humans or a common source from which both humans and pangolins became colonized.

Chromosome-length genome assemblies and cytogenomic analyses of pangolins reveal remarkable chromosome counts and plasticity.

We report the first chromosome-length genome assemblies for three species in the mammalian order Pholidota: the white-bellied, Chinese, and Sunda pangolins. Surprisingly, we observe extraordinary karyotypic plasticity within this order and, in female white-bellied pangolins, the largest number of chromosomes reported in a Laurasiatherian mammal: 2n = 114. We perform the first karyotype analysis of an African pangolin and report a Y-autosome fusion in white-bellied pangolins, resulting in 2n = 113 for males. We employ a novel strategy to confirm the fusion and identify the autosome involved by finding the pseudoautosomal region (PAR) in the female genome assembly and analyzing the 3D contact frequency between PAR sequences and the rest of the genome in male and female white-bellied pangolins. Analyses of genetic variability show that white-bellied pangolins have intermediate levels of genome-wide heterozygosity relative to Chinese and Sunda pangolins, consistent with two moderate declines of historical effective population size. Our results reveal a remarkable feature of pangolin genome biology and highlight the need for further studies of these unique and endangered mammals.

Compromised bilaterality in the small African pangolin (Phataginus tricuspis) – An expression of or compensation for developmental errors; asymmetry, elliptical Fourier analyses and modularity

ABSTRACT We present a postnatal evaluation of skull developmental signaling in small African pangolin emphasizing structural, and cognitive trend in ontogeny for assessment of developmental instability, proper identification and classification, forty digitally processed skulls and foramen magnum from different geo-locations were assessed for asymmetry, foramen magnum shape and modularity using geometric and Elliptical Fourier analyses. Multivariate analysis of regression demonstrated low (p < 0.5) but directional fluctuating asymmetry (F1539 = 3.4045, F882 = 3.2665, dorsal and ventral views). Allometric trajectories followed rostrocaudal, caudolateral directions; Intercepts for shape/size predictions were parallel. Mahalanobis distances between centroids (2.42) were significant (p < 0.01). The variance-covariance matrix in ontogeny lies between 0.0017 and 0.56. Foramen magnum outline descriptors by incremental harmonics revealed first 4 effective PCs defined 96.98% of shape properties and 3.02% constituted finerdetails. 74.1% accuracy decline after size factor elimination. Modulation PCA of covariance matrix and asymmetry component was 88.38% and 7.48% (PC1 and 2), respectively, the variance % predicted 10.08%. The studied samples confirmed ‘handedness’ and fluctuating asymmetries. Foramen magnum shape assumed priority over size in ontogeny with profound asymmetry (from the 5th harmonic), suggestive of compensations to lateralization in neural integration modules.

Optimizing DNA Analyses of Pangolin Products from the Trade for Species Identification in Vietnam

To support law enforcement related to illegal wildlife trafficking in Vietnam, DNA analyses have been widely used to identify species based on products seized from the trade. Pangolins are one of the most threatened mammal groups in Vietnam and across the world because they have been trafficked in huge numbers and considered the most traded mammals globally. Although the government and management agencies have implemented several deterrent measures to reduce the illicit trade of pangolins, a high number of violations with large seizures have still been detected in recent years in the country. Species identification of pangolin products is often challenging, as many products don’t have sufficient morphological diagnoses, especially their meat, hair, and scale. In this study, we present a protocol, which can be applied in laboratories in Vietnam, for extracting and amplifying a fragment of the mitochondrial cytochrome C oxidase subunit 1 (COI) from pangolin’s scales and hairs. This protocol was successfully employed to generate a fragment the COI gene from 10 confiscated pangolin samples. The results show for the first time that three samples originated from an African pangolin species and highlight the importance of applying molecular approaches to combating wildlife trafficking in Vietnam.&#x0D;

Continued availability and sale of pangolins in a major urban bushmeat market in Cameroon despite national bans and the COVID-19 outbreak.

Continued availability and sale of pangolins in a major urban bushmeat market in Cameroon despite national bans and the COVID-19 outbreak.

Can DNA help trace the local trade of pangolins? Conservation genetics of white-bellied pangolins from the Dahomey Gap (West Africa)

BackgroundAfrican pangolins are currently experiencing unprecedented levels of harvesting, feeding both local demands and the illegal international trade. So far, the lack of knowledge on the population genetics of African pangolins has hampered any attempts at assessing their demographic status and tracing their trade at the local scale. We conducted a pioneer study on the genetic tracing of the African pangolin trade in the Dahomey Gap (DG). We sequenced and genotyped 189 white-bellied pangolins from 18 forests and 12 wildlife markets using one mitochondrial fragment and 20 microsatellite loci.ResultsTree-based assignment procedure showed that the pangolin trade is endemic to the DG region, as it was strictly fed by the the Dahomey Gap lineage (DGL). DGL populations were characterized by low levels of genetic diversity, an overall absence of equilibrium, important inbreeding levels, and lack of geographic structure. We identified a 92–98% decline in DGL effective population size 200–500 ya—concomitant with major political transformations along the ‘Slave Coast’—leading to contemporaneous estimates being inferior to minimum viable population size (< 500). Genetic tracing suggested that wildlife markets from the DG sourced pangolins through the entire DGL range. Our loci provided the necessary power to distinguish among all the genotyped pangolins, tracing the dispatch of a same individual on the markets and within local communities. We developed an approach combining rarefaction analysis of private allele frequencies with cross-validation of observed data that traced five traded pangolins to their forest origin, c. 200–300 km away from the markets.ConclusionsAlthough the genetic toolkit that we designed from traditional markers can prove helpful to trace the illegal trade in pangolins, our tracing ability was limited by the lack of population structure within the DGL. Given the deleterious combination of genetic, demographic, and trade-related factors affecting DGL populations, the conservation status of white-bellied pangolins in the DG should be urgently re-evaluated.

The scale of Nigeria's involvement in the trans-national illegal pangolin trade: Temporal and spatial patterns and the effectiveness of wildlife trade regulations

The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) prohibits commercial trans-national trade in pangolin specimens. However, African pangolins are continually trafficked to Asia for traditional medicine, with Nigeria considered a key hub. Using reported Nigeria-linked pangolin seizure data and interviews with Nigerian law enforcement officials, we a) characterised Nigeria's involvement in global pangolin trafficking January 2010–September 2021, particularly observing trafficking trends after pangolin's CITES Appendix I listing; b) estimated the minimum number of pangolins whose scales are in Nigeria-linked seizures January 2010–September 2021, and; c) assessed ongoing efforts within Nigeria to curb pangolin trafficking. Nigeria-linked seizures involved 190,407 kg of pangolin derivatives (99.9% scales) from a minimum of 799,343 pangolins (95% confidence interval; 625,944-996,353) of four species (see caveats in Methods). All shipments confiscated in transit were destined for Asia, with a rapid increase in the mass of maritime shipments over time. Furthermore, stockpiling of pangolin derivatives for overseas shipment is perhaps a prominent trafficking model in Nigeria. Nigeria's law enforcement efforts improved from 2017, the same year Nigeria apparently began playing a hub role. The impact of pangolin's CITES Appendix I listing on pangolin trafficking was unclear, as the marked rise in seizures from 2017 when the listing became effective, coincided with improvements in Nigerian law enforcement efforts. COVID-19–induced travel restrictions likely reduced trafficking activities in 2020 but activities may have fully resumed in 2021. This study provides new information to inform effective enforcement and policy formulation efforts to protect African pangolins.

Distribution of Temminck's pangolin (Smutsia temminckii) in South Africa, with evaluation of questionable historical and contemporary occurrence records

AbstractTemminck's pangolin (Smutsia temminckii) is a threatened mammal found in southern and eastern Africa. It is the most widely distributed African pangolin species yet is listed as Vulnerable on the IUCN Red List due to a projected population decline of 30%–40% within the next three generations. This species' secretive, predominantly nocturnal behaviour and generally low population densities make it difficult to directly assess its distribution, and thus, alternative methods are required. Furthermore, many historical and some contemporary records of this species exist from outside of its accepted range, suggesting that the distribution may be greater than currently accepted, although the validity of these records has never been assessed. We used MaxEnt modelling to evaluate the most likely distribution of S. temminckii in South Africa and compared the past and present distribution of the species, as well as assessed the probable validity of the questionable records. Our results suggest that one of the questionable historical museum records is probably valid, while the other may refer to an erroneous collecting locality. At least some contemporary sightings which were originally deemed questionable are probably valid, while others should be treated as dubious until supporting information to validate their authenticity becomes available.

Molecular Genetics Supports a Potential Fifth Asian Pangolin Species (Mammalia, Pholidota, Manis).

Recently, two mitochondrial haplotypes, H4 and H8, of Manis sp. were found in two seizures in Hong Kong that do not correspond to Manis javanica, Manis pentadactyla or Manis crassicaudata of Asian pangolin species or any African pangolin species. It was proposed that both haplotypes derived from Manis culionensis, an unknown lineage of M. javanica, or a thus far unidentified Asian pangolin species (Manis sp.). To further investigate these three hypotheses, we used two mitochondrial genes of all eight known extant pangolin species and conducted phylogenetic tree reconstructions, divergence time estimation, and species delimitation analyses. All analyses consistently confirmed that these two haplotypes of Manis sp. constitute a distinct lineage, potentially representing a fifth Asian pangolin species, which originated around the Late Miocene to Early Pliocene (6.95 [4.64-9.85] million years ago). Our study provides genetic support for a potential fifth Asian pangolin species and helps to better understand species diversity of Asian pangolins, which is urgently needed for effective conservation work.

Facial and Body Morphometric Indices of the Arboreal African Pangolin ‎‎(Manis tricuspis): Anthropometric Tools in Describing the Pangolin

Body and craniofacial measurements are essential tools in identifying species and characterising their external traits. Body surface measurements were obtained from six African pangolins (Manistricuspis) – 3 males and 3 females. Facial and body measurements obtained from the pangolins revealed that the females had higher averages for body weight, body length and girth. The dimensions of the head showed that the males had a wider zygoma. Also, the manus and arm (humerus) in the females were longer (27.52±3.80 mm and 59.69±3.54 mm respectively) than in the males (26.95±3.35 mm and 50.68±10.22 mm respectively), while the lengths of the arm and the leg (all its components) were longer in the males. Data obtained from this study may provide basic information in assessing and comparing other species of the pangolin, and find application in archaeological, forensic and comparative anatomy of wildlife.

Understanding Local Ecological Knowledge, Ethnozoology, and Public Opinion to Improve Pangolin Conservation in the Center and East Regions of Cameroon

African pangolins are exceptionally difficult to monitor in the wild, which means that assessing threats and impacts to these species also remains a challenge. Local ecological knowledge (LEK) surveys have been recommended as an important tool to collect data for small elusive species. We carried out community-based interviews in 20 villages surrounding Mpem et Djim and Deng-Deng National Parks to gather information on local ecological and traditional medicine knowledge, perceived population status, and the level of conservation awareness amongst local people concerning giant pangolin ( Smutsia gigantea), white-bellied pangolin ( Phataginus tricuspis), and black-bellied pangolin ( Phataginus tetradactyla). The arboreal black-bellied pangolin was least known by local people, and along with the giant pangolin was the least recently observed species. The white-bellied pangolin was reported to be the most recently observed species by most people. Local people reported seeing giant pangolins most around forest and savannah burrows and white-bellied pangolins in forests, commonly crossing dead fallen trees. Limited traditional or cultural uses of pangolins were reported, and most respondents were aware of threats to pangolins. Respondents showed great interest in pangolin conservation, suggesting various strategies for conservation activities, such as trade controls and provisioning of alternative sources of food. We show that working with local communities can provide useful information for pangolin conservation and management efforts within protected areas.

Genetic identification of African pangolins and their origin in illegal trade

To track the illegal pangolin trade from Africa to Asia, we analyzed 1800 DNA samples from 30 seizures of African pangolin scales in Hong Kong during the period 2012–2016. We concluded that all four African pangolin species were present in trade, and that the white-bellied pangolin (Phataginus tricuspis) appeared most frequently (88.5%) in our samples. All six previously described phylogeographic lineages originating from the entire distribution range of P. tricuspis were found in the seizures, and the western central African lineage alone accounted for 67.1% of the samples of this species. Confirmed by modelling data, high DNA haplotype richness was present in most of the pangolin scale seizures, including those contained in small air parcels and large-volume sea shipments. Results suggest that African pangolins were hunted across large areas of their natural range and then delivered to a small number of trade transit hubs. Our study illustrates the utility of genetic analysis for characterizing the illegal pangolin trade and identifying the geographic origin of poaching hotspots.

Isolation and characterization of 30 STRs in Temminck’s ground pangolin (Smutsia temminckii) and potential for cross amplification in other African species

Temminck's ground pangolin (Smutsia temminckii) is one of four species of pangolin, endemic to Africa. Two of the African pangolin species are listed as vulnerable and two are listed as endangered on the International Union for Conservation of Nature Red List of Threatened Species due to their ongoing exploitation for traditional medicine and bushmeat. In this study, we developed 30 species-specific short-tandem repeats (STRs) in Temminck's ground pangolin using next-generation sequencing. The markers were also optimized for crossamplification in other African species. All the markers amplified successfully in Temminck's ground pangolin with allelic polymorphisms observed in 87% of the markers in giant pangolin (S. gigantea) whereas 60% of the markers were amplified polymorphic loci in both whitebellied pangolin (Phataginus tricuspis) and black-bellied pangolin (P. tetradactyla). Analysis of diversity estimates showed moderate levels of variability in Temminck's ground pangolin (Na = 5; Ho = 0.559), giant pangolin (Na = 4.909; Ho = 0.514) and white-bellied pangolin (Na= 2.686; Ho = 0.541) with lower values being observed in black-bellied pangolin (Na = 3; Ho = 0.242). This study provides data of the first available STR markers which was amplified in all four African pangolin species that can now be used in conservation genetic and evolutionary aspects of population histories.

Skull shape variation in extant pangolins (Pholidota: Manidae): allometric patterns and systematic implications

Abstract Pangolins are among the most endangered groups of mammals, comprising eight extant species delineated into three genera. Despite several studies dedicated to their skeletal anatomy, the potential taxonomic insight from cranial morphological variation in extant Pholidota is yet to be assessed with modern geometric morphometric methods. We present the first comprehensive study on the cranial morphology of extant pangolins and discuss its implications for the taxonomy and evolution of the group. We performed landmark-based morphometric analyses on 241 museum specimens to describe the variation in skull shape in seven of the eight extant species. Our analyses revealed genus- and species-level morphological discrimination, with Asian species (Manis spp.) being grouped together, whereas African pangolins present distinct skull shapes between small (Phataginus spp.) and large (Smutsia spp.) species. Analyses of allometry also identified a set of traits whose allometric trajectories distinguish Asian from African specimens. Finally, we uncovered intraspecific variation in skull shape in white-bellied pangolins (Phataginus tricuspis) that partly corroborates recent DNA-based differentiation among biogeographically distinct populations. Overall, our results shed light on the morphological diversity of the skull of these enigmatic myrmecophagous mammals and confirm the genus-level classification and cryptic diversity within the white-bellied pangolin revealed by molecular phylogenetics.