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Taxonomic revision of the pampas cat Leopardus colocola complex (Carnivora: Felidae): an integrative approach
Abstract
The pampas cat Leopardus colocola has been subject to conflicting classifications over the years. Currently, one polytypic species with seven subspecies is recognized, but integrative taxonomic study for this debated group has never been done. Here, we combine the broadest morphological coverage of the pampas cat to date with molecular data and ecological niche models to clarify its species composition and test the validity of recently proposed subspecies. The multiple lines of evidence derived from morphology, molecular, biogeography and climatic niche datasets converged on the recognition of five monotypic species: L. braccatus, L. colocola, L. garleppi (including thomasi, budini, steinbachi, crespoi and wolffsohni as synonyms), L. munoai and L. pajeros (including crucina as synonym). These five species are morphologically diagnosable based on skin and skull traits, have evolved in distinct climatic niche spaces and were recovered in molecular species delimitation. Contrary to previous taxonomic arrangements, we do not recognize subspecies in pampas cats. To objectively define the two most controversial species, we designate neotypes for L. colocola and L. pajeros. The diversification of pampas cats is associated with Middle Pleistocene glaciations, but additional genetic samples from the central Andean region are still needed to conclusively reconstruct its evolutionary history.
... This ecoregion presents great importance to botanical and animal studies due to its biodiversity with different areas being highlighted as important to conservation (Bilenca and Miñarro 2004;Loyola et al. 2009). Of particular concern is Leopardus munoai, an endemic felid at high risk of extinction due to those landscape changes (Nascimento et al. 2021;Tirelli et al. 2021). Indeed, land use change emerges as the primary threat to the survival of 91% of small carnivores facing extinction, with 85% speci cally attributing this threat to agricultural activities (Marneweck et al. 2021). ...
Understanding how species respond to changes in their environment is crucial for effective conservation efforts, particularly in vulnerable ecoregions like the Uruguayan Savannah in Brazil and Uruguay. Here, we focused on two often overlooked species, the South American coati (Nasua nasua) and Molina's hog-nosed skunk (Conepaptus chinga), deploying 79 cameras across 15 areas in both the Brazilian and Uruguayan sectors of the ecoregion. Using occupancy models, we investigated the influence of different land cover types (forests, grasslands, farmland, and silviculture) and human density on the presence of these species. We tested the activity pattern uniformity for each species. We looked at the landscape transition history. Furthermore, we also generated innovative occupancy maps to better understand and guide policies and actions for these species. The coati occupancy exhibited a positive relation with forest areas but a negative correlation with grasslands and crop farming areas (p<0.05). The skunk presented a positive response to grassland and crop farming areas but a negative response to forest areas (p<0.05). The South American coati occupancy was estimated at approximately 0.141 (0.041 – 0.420), while for Molina’s hog-nosed skunks, it was 0.377 (0.200 – 0.610). Despite the coati's use of open areas, it demonstrated a stronger association with natural forests than altered landscapes. In contrast, Molina's hog-nosed skunk displayed adaptability, persisting in altered environments. In conclusion, our findings underscore the urgency of prioritizing conservation efforts for coatis, while highlighting the skunk's resilience to landscape alterations. This knowledge can guide targeted conservation plans for these species in threatened ecoregions.
... Brazil is home for the largest number of mammal species (Rylands et al. 2000;Paglia et al. 2012), which includes approximately ten wild felid species (Reis et al. 2006;Trigo et al. 2013aTrigo et al. , 2013bTrigo et al. , 2013cNascimento et al. 2020). Their populations have been declining in recent years due to many factors such as illegal hunting, habitat fragmentation, and human-wildlife competition (Nowell and Jackson 1996;Morato et al. 2016) being considered threatened within the Brazilian territory (MMA-Ministry of the Environment 2022) and included in conservation programs in their natural habitats (Gasparini- Morato et al. 2021). ...
Among the species described within the Onchocercidae family, Dirofilaria immitis is regarded as the most common worldwide, causing severe and often fatal conditions in dogs, cats, and occasionally humans. Dirofilaria spp. are vectored by mosquitoes, simulids, and culicoids, with their epidemiology dependent on the geographical distribution of competent vectors. Eight species of Dirofilaria have been reported so far in Brazil, of which six parasitize non-human primates, deer, procyonids, and marsupials. Here, we investigated the occurrence of Onchocercidae in wild felids (i.e., Panthera onca, Puma concolor, Herpailurus yagouaroundi, Leopardus geoffroyi, Leopardus guttulus, Leopardus pardalis, Leopardus wiedii, Leopardus munoai) from different locations in Brazil. Overall, 82 samples (n = 63 blood; n = 19 tissues) were molecularly screened for cytochrome c oxidase subunit-1 (cox1) gene. Four (i.e., 4.8%) wild felid samples were positive, and at BLAST analysis, the obtained sequences showed varying percentage of nucleotide identity with the genera Brugia (i.e., 87–88%), Setaria (i.e., 89%), and D. immitis (i.e., 94.4%). Phylogenetic analyses clustered sequences obtained into three distinct clades, one with D. immitis and the remaining two with other Onchocercidae spp. Data herein obtained highlight the need for a more comprehensive understanding of the diversity and biology of Onchocercidae in South America in order to assess the potential impact that these species may have for domestic and wild animals, as well as humans.
... Scientists should engage in discussions about their day-to-day operations of scientific research and conservation that will help make their science more inclusive, welcoming, and just. There have been similar initiatives for wild cats; for example, to push for the common names (1) guiña instead of the kodkod for L. guigna (Molina, 1782)the latter is indeed a local name referring to another species, the colocola or the Pampas cat L. colocola (Sanderson 2010, Kitchener et al. 2017, Nascimento et al. 2021, and (2) the Andean cat L. jacobita, previously also referred to as the Andean mountain cat, to replace the name Jacobita's cat (Cornalia, 1865). We invite scientists to consider local names of the cat species in their studies. ...
We present a new update of the official list of mammals of Ecuador. We follow a taxonomic order for the supra-generic categories, but alphabetical for the lower levels. We incorporate and point out the taxonomic changes that have occurred since the last version published in December 2023. For the elaboration of this list we kept a constant review of the scientific publications generated and the taxonomic changes occurred. The current version of the list of mammals of Ecuador includes 471 native species belonging to 13 orders, 53 families and 214 genera; among the main changes is the addition of two species to the Ecuadorian fauna (Cyttarops alecto and Lontra annectens); also noteworthy is the revalidation of the genus Pudella (instead of Pudu), the synonymization of the genus Vicugna with Lama and the taxonomic changes of other species (Saimiri macrodon, Oecomys galvez, O. paricola, Myotis osculatii, M. oxyotus, Leopardus pardinoides, and Lontra longicaudis). The mammalian orders with the highest species richness in Ecuador are Chiroptera (182), Rodentia (134), Artiodactyla (41) and Carnivora (37). We also document 27 additional species that are expected or potential for the Ecuadorian fauna. In addition, we list the subspecies (143) that correspond to the mammals of Ecuador, endemic species (62), extinct (3), recorded in the Ecuadorian Antarctic zone (11) and those introduced into the country (17).
Recently, the tiger-cat species complex was split into Leopardus tigrinus and Leopardus guttulus, along with other proposed schemes. We performed a detailed analysis integrating ecological modeling, biogeography, and phenotype of the four originally recognized subspecies-tigrinus, oncilla, pardinoides, guttulus-and presented a new multidimensional niche depiction of the species. Species distribution models used > 1400 records from museums and photographs, all checked for species accuracy. Morphological data were obtained from institutional/personal archives. Spotting patterns were established by integrating museum and photographic/camera-trap records. Principal component analysis showed three clearly distinct groups, with the Central American specimens (oncilla) clustering entirely within those of the Andes, namely the pardinoides group of the cloud forests of the southern Central-American and Andean mountain chains (clouded tiger-cat); the tigrinus group of the savannas of the Guiana Shield and central/northeastern Brazil (savanna tiger-cat); and the guttulus group in the lowland forests of the Atlantic Forest domain (Atlantic Forest tiger-cat). This scheme is supported by recent genetic analyses. All species displayed different spotting patterns, with some significant differences in body measurements/proportions. The new distribution presented alarming reductions from the historic range of − 50.4% to − 68.2%. This multidimensional approach revealed a new species of the elusive and threatened tiger-cat complex.
The taxonomic status and interspecies relationships of hares (genus Lepus ) in Xinjiang, China, remain ambiguous owing to extremely similar external morphologies, introgressive hybridization between species and a lack of molecular genetic records. In this study, a total of 154 hare specimens were used to clarify their taxonomic status. Based on multiple lines of evidence, including observations and comparisons of external morphological characteristics, skull morphometrics and genome‐wide single nucleotide polymorphism (SNP) datasets, four hare species were recognised. Lepus yarkandensis and Lepus timidus , which have obvious morphological characteristics, were identified as two distinct species, as previously reported. Combining the integrated morphological and molecular datasets with the relevant literature, we proposed suggestions on the classification of the more controversial hares. Due to molecular and morphological similarities, hares in northern and central Xinjiang were divided into two subspecies of Lepus tolai , namely, Lepus tolai lehmanni and Lepus tolai centrasiaticus . Despite similar external morphological characters, hares in the southwest Pamir Plateau and its surroundings can be distinguished from L. tolai by differences in skull morphology and genetic divergence and were, therefore, considered a subspecies of Lepus tibetanus , Lepus tibetanus pamirensis . Furthermore, we found that hybridization and introgression occurred between sympatric or parapatric hares, especially between L. yarkandensis and L. tibetanus pamirensis , L. yarkandensis and L. tolai centrasiaticus , and L. tolai lehmanni and L. timidus .
This chapter provides an overview of the knowledge on mammals in the Campos Sulinos. To date, 134 native species have been recorded, of which eleven are endemic. Through a topical analysis of the literature produced so far, we present research priorities on several topics and subregions. Currently, there is a reasonable body of knowledge on small mammal and carnivore population genetics and habitat selection, whereas other groups and topics are still poorly studied, such as the ecology of armadillos. Most of the knowledge comes from studies conducted in the west and south of the Brazilian Pampa, while studies in the highland grasslands and in the northwest and center of the Pampa have been scarce. Hybridization, recently described species, and adaptations for living in open environments are discussed, with an emphasis on the need to conserve species and ecosystem services in the region. The future systematic planning of mammal conservation in the Campos Sulinos depends on filling knowledge gaps and developing taxonomically and spatially more comprehensive research, as well as on a better connection between researchers and decision-makers/stakeholders.
Even in the genomics era, the phylogeny of Neotropical small felids comprised in the genus Leopardus remains contentious. We used whole-genome resequencing data to construct a time-calibrated consensus phylogeny of this group, quantify phylogenomic discordance, test for inter-species introgression, and assess patterns of genetic diversity and demographic history. We infer that the Leopardus radiation started in the Early Pliocene as an initial speciation burst, followed by another in its subgenus Oncifelis during the Early Pleistocene. Our findings challenge the long-held notion that ocelot (Leopardus pardalis) and margay (L. wiedii) are sister species, and instead indicate that margay is most closely related to the enigmatic Andean cat (L. jacobita), whose whole-genome data are reported here for the first time. In addition, we found that the newly sampled Andean tiger cat (L. tigrinus pardinoides) population from Colombia associates closely with Central American tiger cats (L. tigrinus oncilla). Genealogical discordance was largely attributable to incomplete lineage sorting, yet was augmented by strong gene flow between ocelot and the ancestral branch of Oncifelis, as well as between Geoffroy’s cat (L. geoffroyi) and southern tiger cat (L. guttulus). Contrasting demographic trajectories have led to disparate levels of current genomic diversity, with a nearly tenfold difference in heterozygosity between Andean cat and ocelot, spanning the entire range of variability found in extant felids. Our analyses improved our understanding of the speciation history and diversity patterns in this felid radiation, and highlight the benefits for phylogenomic inference of embracing the many heterogeneous signals scattered across the genome.
The Pantanal Cat, Leopardus colocola braccatus , presents a broad distribution over several biomes in South America. However, the distribution of the species is based on a limited number of occurrences, and confirmed records of the species are scarce. Here, we present 2 confirmed records of Leopardus colocola braccatus from the state of São Paulo, southeast Brazil. Both records are juvenile individuals, 1 recorded in the municipality of Castilho and the other in Araçatuba. These São Paulo records expand the distribution of this uncommon felid to the most populous state of Brazil. This species is currently Data Deficient on the São Paulo Red List, and these new records may help with future assessments of the conservation status of this felid.
The pampas cat is a small felid that occurs in open habitats throughout much of South America. Previous studies have revealed intriguing patterns of morphological differentiation and genetic structure among its populations, as well as molecular evidence for hybridization with the closely related L. tigrinus. Here we report phylogeographic analyses encompassing most of its distribution (focusing particularly on Brazilian specimens, which had been poorly sampled in previous studies), using a novel dataset comprising 2,143 bp of the mitogenome, along with previously reported mtDNA sequences. Our data revealed strong population strutucture and supported a west-to-east colonization process in this species' history. We detected two population expansion events, one older (ca. 200 thousand years ago [kya]) in western South America and another more recent (ca. 60-50 kya) in eastern areas, coinciding with the expansion of savanna environments in Brazil. Analyses including L. tigrinus individuals bearing introgressed mtDNA from L. colocola showed a complete lack of shared haplotypes between species, indicating that their hybridization was ancient. Finally, we observed a close relationship between Brazilian/Uruguayan L. colocola haplotypes and those sampled in L. tigrinus, indicating that their hybridization was likely related to the demographic expansion of L. colocola into eastern South America.
Dasypus is the most speciose genus of the order Cingulata, including approximately 40% of known living armadillos. Nine species are currently recognized, although comprehensive analyses of the entire genus have never been done. Our aim is to revise the taxonomy of the long-nosed armadillos and properly define the taxa. We examined 2126 specimens of Dasypus preserved in 39 different museum collections, including 17 type specimens. Three complementary methods were applied to explore morphological datasets both qualitatively and quantitatively. Qualitative morphological variation in discrete characters was assessed by direct observations of specimens. Linear morphometric variation was based on external data and cranial measurements of 887 adult skulls. The shape and size of the skull was abstracted through two-dimensional geometric morphometric analyses of dorsal, lateral and ventral views of respectively 421, 211, and 220 adult specimens. Our results converge on the recognition of eight living species (D. beniensis, D. kappleri, D. mazzai, D. novemcinctus, D. pastasae, D. pilosus, D. sabanicola, and D. septemcinctus), and three subspecies of D. septemcinctus (D. s. septemcinctus, D. s. hybridus, and a new subspecies from Cordoba described here). Information on type material, diagnosis, distribution, and taxonomic comments for each taxon are provided. We designate a lectotype for D. novemcinctus; and a neotype for Loricatus hybridus (= D. septemcinctus hybridus).
The Pampas cat Leopardus colocolo occurs in a variety of habitats from northern Ecuador to southern Argentina, yet the species has been poorly studied. There is scant information about its northern distribution or about populations in desert and dry forest. We aimed to determine the presence of the Pampas cat in the Sechura Desert and seasonally dry forest of north-western Peru and south-western Ecuador, identify threats to the species, and describe people's perceptions of it. Using 32 camera traps and compiling confirmed and unpublished records, we mapped the species’ distribution and identified 12 new localities, three in the Sechura Desert and nine in the dry forest. The first records of the Pampas cat in the Ecuadorian dry forest are reported from La Ceiba Natural Reserve and Jorupe Reserve; the northernmost record in the dry forest is from Cerros de Amotape National Park, Peru. In 56 semi-structured interviews with local people we found that most of them (76.8%) did not know the species; 61.5% of those who knew the species had a neutral perception and did not think the cat affected their personal activities. Here we update the northern distribution of the Pampas cat, describe threats to the species in arid ecosystems, and highlight the need for further studies to identify other possible threats and mitigation methods.
The tigrina Leopardus tigrinus (Schreber, 1775) is a small-sized Neotropical spotted cat found from northern Argentina and southern Brazil to Costa Rica. Four subspecies are traditionally recognized: L. t. tigrinus (Schreber, 1775) from northern Brazil, the Guianas and eastern Venezuela; L. t. pardinoides (Gray, 1867) from western Venezuela, Colombia, Ecuador and Peru; L. t. guttulus (Hensel, 1872) from southern Brazil, Paraguay and northern Argentina; and L. t. oncillus (Thomas, 1903) from Costa Rica. We studied external and craniodental morphology in quantitative and qualitative terms from 250 specimens in order to clarify the taxonomic status of tigrina. Based on the characters analyzed in this study, we recognize three diagnosable morphogroups, each with a distinct geographic distribution: northern/northwest-ern/west (samples from northern Brazil, the Guianas, Venezuela, Colombia, Ecuador, Peru, northwestern Argentina and Costa Rica), eastern (samples from northeastern and central Bra-zil), and southern (samples from southern Brazil, Paraguay and northeastern Argentina). Taking into account the morphologic evidence presented here, supported by biogeographic data and molecular studies available, we recognize three full species for tigrinas: L. tigrinus (including the putative subspecies L. t. pardinoides and L. t. oncillus as junior synonyms) for northern/ northwestern/west group; L. emiliae (Thomas, 1914) for eastern group; and L. guttulus for southern group.
We created a new dataset of spatially interpolated monthly climate data for global land areas at a very high spatial resolution (approximately 1 km 2). We included monthly temperature (minimum, maximum and average), precipitation, solar radiation, vapour pressure and wind speed, aggregated across a target temporal range of 1970–2000, using data from between 9000 and 60 000 weather stations. Weather station data were interpolated using thin-plate splines with covariates including elevation, distance to the coast and three satellite-derived covariates: maximum and minimum land surface temperature as well as cloud cover, obtained with the MODIS satellite platform. Interpolation was done for 23 regions of varying size depending on station density. Satellite data improved prediction accuracy for temperature variables 5–15% (0.07–0.17 ∘ C), particularly for areas with a low station density, although prediction error remained high in such regions for all climate variables. Contributions of satellite covariates were mostly negligible for the other variables, although their importance varied by region. In contrast to the common approach to use a single model formulation for the entire world, we constructed the final product by selecting the best performing model for each region and variable. Global cross-validation correlations were ≥ 0.99 for temperature and humidity, 0.86 for precipitation and 0.76 for wind speed. The fact that most of our climate surface estimates were only marginally improved by use of satellite covariates highlights the importance having a dense, high-quality network of climate station data.
1. The current classification of the Felidae was reviewed by a panel of 22 experts divided into core, expert and
review groups, which make up the Cat Classification Task Force CCTF of the IUCN Cat Specialist Group.
2. The principal aim of the CCTF was to produce a consensus on a revised classification of the Felidae for use
by the IUCN.
3. Based on current published research, the CCTF has fully revised the classification of the Felidae at the level
of genus, species and subspecies.
4. A novel traffic-light system was developed to indicate certainty of each taxon based on morphological,
molecular, biogeographical and other evidence. A concordance of good evidence in the three principal
categories was required to strongly support the acceptance of a taxon.
5. Where disagreements exist among members of the CCTF, these have been highlighted in the accounts for
each species. Only further research will be able to answer the potential conflicts in existing data.
6. A total of 14 genera, 41 species and 77 subspecies is recognised by most members of the CCTF, which is
a considerable change from the classification proposed by Wozencraft (2005), the last major revision of the
Felidae.
7. Future areas of taxonomic research have been highlighted in order to answer current areas of uncertainty.
8. This classification of the Felidae will be reviewed every five years unless a major new piece of research
requires a more rapid revision for the conservation benefit of felid species at risk of extinction.
Juan Ignacio Molina (1740–1829) was a Jesuit priest born in Chile but forced to flee to Europe after his religious order was expelled from the Spanish Empire in 1767. He settled in Bologna, Italy, and began to write a natural history of his homeland, which was first published in Italian in 1782 (he is also known as Giovanni Ignazio Molina). He completed a second volume on the history of the people of Chile in 1786, and a version of the book was in the library of HMS Beagle. This two-volume English translation was published in 1809, and also includes notes from other explorers, including a 1791 account of Chile by Pedro González de Agüeros (1768–93) and a 1774 description of Patagonia by Thomas Falkner (1707–84). Volume 1 covers the natural history of Chile, including the territory's diverse climate, geology, plants, and wildlife.