Taxonomic Notes
Previously recognised as belonging to a monotypic genus Oreailurus, primarily based on the relative size difference in the skull's auditory chambers obtained from very few specimens (Cabrera 1940, Wozencraft 1993, Nowell and Jackson 1996, Yensen and Seymour 2000). Subsequently, based on genetic evidence, the Andean Cat was included in the genus Leopardus. The specific name, jacobita, is in honour of Jacobita Mantagazza (Cornalia 1865) and should not be declined to jacobitus as is sometimes seen in the literature (Sanderson 2015).
Justification
The Andean Cat has been categorised as Endangered under C2a(i) because the species population does not exceed 2,500 mature individuals, and there is an inferred continuing decline in the population, with less than 250 mature individuals in the largest subpopulation.
A reduction in the population of the Andean Cat is inferred based on several lines of evidence: a. the decrease in reports of live animal sightings from local people; b. the increase of other threats that lead to the direct death of individuals, such as roadkills, retaliatory killings and dog attacks. In addition, a population decrease is suspected due to the rapidly increase in mining activities in most of the distribution range of this felid, especially the high Andes.
Although formal population density estimates (using robust statistical methods) are only available for two sites (Reppucci et al. 2011, Huaranca et al. 2019), our conclusions are supported by a number of additional studies on the ecology and distribution of this felid and much anecdotal evidence collected over 20 years by a team of experts that is specifically dedicated to the investigation and conservation of the Andean Cat across its entire range, the Andean Cat Alliance. Furthermore, the low population number is fully congruent with the results of recent studies that found very low genetic and genomic diversity (Cossios et al. 2012, Lescroart et al. 2023).
A further reason of concern for the conservation of this species is that most of the Andean Cat habitat is subject to major threats due to the expansion of the agricultural frontier, inadequate livestock management, and water extraction (Villaroel et al. 2014, Huaranca et al. 2022), as well as extensive water and soil transformation and contamination from a growing mining and petroleum industry (Walker et al. 2013, Chavez et al. 2023). While mining is already widely distributed on the Andean Cat's known distribution and is affecting the species both directly and indirectly, most of nearby unexploited areas are affected by requests or permits for mining exploration and exploitation. In addition, some areas, particularly in Bolivia and Peru, are subject to illegal mining. Besides this, the most suitable area (core area) for the Andean Cat population lying in the central part of its distribution, is located within the so called “lithium triangle”, and is now threatened by the growing lithium demand for batteries. Although the areas for lithium extraction are situated in salt flats, which are not the most suitable habitat for Andean Cats, possible impacts could arise from the growing demand of lithium in the surrounding areas, besides the increase of road construction and usage, urbanisation, chemical spillage, and contamination of surface and ground waters.
Furthermore, models show that global climate change will be a great threat for Andean Cats throughout most of its range. The modification of the habitat due to changes in climatic patterns is causing the species to shift their range to higher altitudes, ultimately resulting in a decrease in the available habitat for the Andean Cat therefore reducing the size of its total geographic distribution (Bennett et al. 2017). Andean Cats require large home ranges to subsist (average size: 51.2 km2, N=4, Villalba et al. 2009, Tellaeche 2015), and the low density of the species reduces the chances of meetings between individuals of opposite genders in the mating season. The fast degradation of the regions surrounding the protected areas may turn those areas in unsuitable places for the species, acting as barriers for dispersal and mating. If corridors are not created urgently, the protected areas existing in Andean Cat habitat will be insufficient to ensure proper habitat connectivity.
Considering the constant increase of the mentioned threats, the natural fragmentation of the habitat and the augmented fragmentation due to habitat loss and degradation, a continued decline of the population number is inferred.
Geographic Range Information
The Andean Cat was rarely recorded until the late 1990s, and its range was thought to be limited to southern Peru, southwest Bolivia, northwest Argentina, and northern Chile (Yensen and Seymour 2000). There were also few sightings (Scrocchi and Halloy 1986, Sanderson 1999) and museum specimens (Garcia-Perea 2002) of this species. However, the number of distribution records has grown significantly in the past 20 years, mainly because of the work of the Andean Cat Alliance (
www.gatoandino.org), a group of expert researchers that works in the four countries where the species live.
The Andean Cat has a patchy distribution due to a specialisation for naturally fragmented rocky habitats and most presence records are from the central Andes above 3,600 m in Argentina, Bolivia, Chile and Peru. However, it has also been found at lower elevations, as low as 1,800 m in the Andes of central-southern Argentina (Sorli
et al. 2006) and 1,150 m in central Chile (Segura
et al. 2021). More recent records of the Andean Cat in Argentina and Chile extend its distribution range to the south and outside the Andes, in proximity of the deciduous temperate forest of central Chile in Radal Siete Tazas (CONAF 2023) and of the Patagonian steppe and scrub habitats in Argentina, at elevations as low as 650 m (Martinez
et al. 2008, Novaro
et al. 2010).
The known northern and southern distribution limits for the Andean Cat are respectively, in central Peru (9°54' S), and central Argentina (38°23' S; Cossios
et al. 2007, Novaro
et al. 2010), but the new information gathered by researchers confirms that throughout this range, the Andean Cat subpopulations are patchily distributed. The results of modelling the Andean Cat distribution, considering four bioclimatic variables (annual mean and mean diurnal range temperature, annual precipitation, and coldest season winter), showed the presence of three biogeographic barriers that affect the species’ distribution. These include: i) In the north of Peru, coinciding with the Huancabamba depression, known as the “Peruvian low,” which defines the northern boundary of its distribution range. ii) In the centre of Bolivia, with the “Andean knee,” an extremely arid transition zone between the wet and dry Puna. This area has no records of Andean Cats. iii) In the south, with the “Arid Diagonal” in Chile and Argentina, also a hyper-arid zone with very few records of Andean Cats (Marino
et al. 2011). According to this model and despite the differences in elevation over the current geographic range, there are common climatic conditions that favour its presence, which are cold, dry and extreme diurnal variations in temperature (Marino
et al. 2011).
Population Information
Formal probabilistic estimates of population numbers for the Andean Cat have been conducted in only two sites that were selected because of the high quality of habitat available for the species (e.g., multiple Andean Cat records in the area, great abundance of prey, high terrain roughness, and rocky outcrops). In both cases spatially explicit capture recapture models (SECR, Royle et al. 2014) were applied to camera trap data. The first study obtained a density of 0.07 and 0.12 individuals/km2 in two consecutive years in northwestern Argentina (Reppucci et al. 2011) and the second study found a density of 0.064–0.069 individuals/km2 in central western Bolivia (Huaranca et al. 2019). A study in northern Chile reported a minimum density of 0.02 ind./km2 using genetic individual identification (Napolitano et al. 2008).
To supplement the formal information available on Andean Cat abundance, we compiled information of camera trap surveys carried out in different locations. Using those surveys, we estimated a minimum density for each area in terms of number of individuals detected/sampled area. To reduce the possibilities of overestimation or sub-estimation, we only used data from surveys that met the following criteria: (i) surveyed area greater than 100 km2 (approximately two Andean Cat home ranges), (ii) camera density greater than 0.07 camera traps/km2 (to avoid gaps where an individual has no possibility of being detected), and (iii) sampling effort greater than 900 camera trap-days (Llerena-Reátegui et al. 2022). After these filters were applied, there were eight surveys to perform the estimations (see Supplementary Information), including formal density estimations. For one area where population density was estimated both formally and non-formally, the two procedures produced similar results once filters were applied (Reppucci et al. 2011).
We grouped the surveys in three regions, since it is likely that abundance changes with latitude, being the more suitable habitats and higher densities in the centre of the species’ distribution (Brown 1984). The northern region ranges from central Peru to 21.5° of latitude South. The central region, considered the most suitable habitat for Andean Cats, ranges from 21.5° to 27.5° of latitude South. Finally, the southern region ranges from 27.5° of latitude South to the southernmost record in Argentina. The mean Andean Cat population densities for north, central and south regions were 0.048, 0.063 and 0.034 individuals/km2, respectively (see Supplementary Information). The mean density for all the published studies (Reppucci et al. 2011, Huaranca et al. 2019) and those compiled for this categorisation is 0.05 individuals/km2 (range: 0.012–0.120).
The Andean Cat distribution is patchy and strongly associated with steep rocky environments and these habitat patches are surrounded by large areas that are unsuitable for the species. One study found a value of around 1.4% of a total area of approximately 65,000 km2 to be suitable for the Andean Cat in northern Patagonia (Palacios et al. 2019). A second study carried out in northern Chile over 13,711 km2 estimated that 6.7% of the area was suitable for the species (Lagos et al. 2020). This second study was carried out in an area where Andean Cat density was expected to be relatively high, so it might overestimate the representativeness of suitable habitat at greater scale.
Considering the "Extant" area of 343,689.75 km2 (divided in 176,742.13 km2 in the north, 103,043.31 km2 in the centre and 63,904.32 km2 in the south; see Supplementary Information), for each region we used the mean population density found for that region and calculated the number of individuals for the 6.7% of the total area (being this the proportion of suitable habitat found by Lagos et al. (2020)). Although it is probable that the density for the remaining part of the area (93.3%) is close to 0, to avoid underestimating population numbers, we used the minimum density that we found in our compilation of surveys as explained above (0.01 individual/km2) for the rest of each area. It is necessary to remember that this local density estimation was obtained from good quality habitats, and therefore it likely overestimated the cat densities over much larger areas. As a result, we estimated the total population of Andean Cats at 4,354 individuals including adults and subadults. There is no information about population structure available, thus we used a default value of 50% for the proportion of mature individuals within the Andean Cat population (Nowell et al. 2007). This gave an estimate of only 2,177 mature individuals (see Supplementary Information).
Given the naturally fragmented arrangement of the Andean Cat's habitat, a great patch or group of close patches would be necessary to sustain a population of 250 mature individuals. According to our estimations, the patch of suitable continuous habitat large enough to hold at least 250 mature individuals should measure a minimum of 2,083 km2 (considering the highest known density estimation). A patch that has or exceeds the size required to sustain a population of 250 mature individuals has not been identified anywhere in the Andean Cat range. Therefore, there is no subpopulation numbering 250 or more mature individuals.
Given the uncertainty surrounding population estimates and the fact that they were obtained from areas that researchers identified as best habitat for the species, it is possible that we are overestimating population numbers (both at range and subpopulation level). The formal density estimates outlined above were obtained from two regions that are considered to have the most favourable conditions for Andean Cats (e.g., climate, habitat, topography and prey availability; Marino et al. 2011). Because of this, it is expected that Andean Cat densities in areas with less favourable conditions are much lower, as suggested by the additional studies reported here and that the ecological density might be even lower at landscape level. This hypothesis is supported by research done by Reppucci et al. (unpublished data) who extended their study site to the less suitable habitat surrounding the high-quality area and estimated that abundance decreased by 69%.
The genetic diversity of the Andean Cat has been found to be very low in mitochondrial genes and nuclear microsatellites across its range (Napolitano et al. 2008, Cossios et al. 2012). A study sampled 56 individuals across Perú, Bolivia and Argentina and found only five mitochondrial haplotypes (Cossios et al. 2012). In the same study, Andean Cat microsatellite diversity (allelic richness) was also very low compared with the congeneric species, the Pampas Cat Leopardus colocolo, which inhabits the same geographic areas. Similarly, the number of alleles was three to five times lower in Andean Cats than in Ocelots Leopardus pardalis, Margays Leopardus wiedii and Northern Tiger Cats L. tigrinus from Brazil (Cossios et al. 2012). These initial results led to the determination that the Andean Cat had two distinct Evolutionary Significant Units” (ESUs). The geographical separation between the northern and southern ESU concurs with the third biogeographic barrier, the South American Arid Diagonal and the separation between the two northern management units coincides with the transition zone between the wet and dry Puna (Cossíos et al. 2012).
A recent whole-genome study found that the Andean Cat has the lowest genomic diversity of the Leopardus genus, harbouring 10 times less genetic variation (heterozygosity) than the most diverse species evaluated (the Ocelot, Lescroart et al. 2023). Estimated heterozygosity was extremely low in the Andean Cat, comparable to the Iberian lynx (Lynx pardinus). Also, the lowest effective population size (Ne) of the Leopardus genus was observed in the Andean Cat, which appears to have been steadily declining in population size since at least the Last Interglacial (LIG; 130–116 thousand years Before Present). This study suggests long-term small population size as the cause of the observed low diversity (Lescroart et al. 2023).
Genetic diversity is associated with population size: high genetic diversity is related to large populations, while low genetic diversity to small populations. Therefore, low genetic diversity estimates complement camera trap low density estimates, and support that the Andean Cat has small population numbers across its range.
Habitat and Ecology Information
The Andean Cat is found primarily in rocky and steep terrains, in arid and sparsely vegetated areas of the High Andes above the timberline, and in scrub and steppe habitats within the Andean foothills of Central Argentina and Chile. In general, climatic conditions are extreme, with very low winter temperatures, large daily thermal variations all year, and low precipitations that determine the presence of adapted plants such as bunchgrasses, cushion plants, and low shrubs with small or resinous leaves (Cabrera and Willink 1973). The High Andean peatlands are known as “bofedales” or “vegas” and are an important resource for wildlife and domestic animals, including the Andean Cat (Villalba et al. 2004, Lagos et al. 2020, Huaranca et al. 2022). At the local scale, in two studies in Argentina, Andean Cats were most likely to use areas with a high proportion of rocks (Tellaeche 2015, Palacios 2019), and elevated terrain ruggedness (Tellaeche 2015), closer to water (vegas or small creeks), and close to areas populated by their main prey (Palacios 2019). In the north of Chile, the more important variables to predict Andean Cat presence were temperature, distance to peatlands and topographic position (Lagos et al. 2020).
According to the results of diet studies in Argentina, Bolivia and Chile, and compared with that of other sympatric carnivores, the Andean Cat occupies a narrower trophic niche with a wide overlap with the Pampas Cat (Walker et al. 2007, Marino et al. 2010). Rodents constitute the main prey item. Some studies identify Mountain Viscacha (Lagidium viscacia) as the more frequently consumed item (Napolitano 2008, Viscarra 2008), while in other areas, small rodents (mainly Phyllotis spp, Octodontomys gliroides, and Abrocoma cinerea) are more commonly consumed (Walker et al. 2007, Torrico 2009, Tellaeche 2010, Reppucci 2012). Among the Andean Cat prey items, the Mountain Viscacha stands out due to its significantly greater body mass, making a substantial contribution to the Andean Cat's biomass intake, despite the lower number of individuals consumed (Walker et al. 2007, Napolitano et al. 2008, Reppucci 2012).
In Central Chile, the consumption of the exotic and invasive species European Hare (Lepus europaeus) has also been recorded by camera trap (C. Sepúlveda pers. comm. 2023).
The Andean Cat is thought to be a solitary species, but may be seen in pairs or with a single kitten during the reproductive season. Estimation of reproduction dates are lacking, some kittens’ records on camera traps suggest a birth season from January to March (Segura in prep.) to February and July (Aravena et al. 2016). In Coquimbo´s Region, Central Chile, there are records of courtship vocalisations in October and January. In addition, in the month of August there was a record of a cub approximately four to six months old walking with her mother (C. Sepúlveda pers. comm. 2023). Data from several locations suggest that Andean cats deliver only one kitten per litter.
A wide camera trap-based study showed that Andean Cats are mostly active during dusk and night, but compared with other sympatric species (Culpeo Fox, Pampas Cat and Puma), the Andean Cat was the most diurnal species (Lucherini et al. 2009). A similar result was found for the first radio-collared Andean Cat in southern Bolivia, where activity peaks were recorded between 18:00 and 23:00 hours (Villalba et al. 2009). Reppucci (2012) found activity peaks in the first half of the night and first half of the day (including a peak at noon) for the north of Argentina, while Huaranca et al. (2019) found peaks after dusk and after midnight with scarce activity during the day.
Home range sizes of three Andean Cats through a telemetry study in Argentina ranged from 18.8 to 61.2 km2 for two females and 59 km2 for a single male monitored (Kernel 95%; Tellaeche 2015). Home ranges overlapped largely between the male and both females (between 52.8 and 100%). A single female Andean Cat that was radio-tracked between April and December 2004 in Bolivia, had a home range size of 65.5 km² (95% minimum convex polygon: Villalba et al. 2009). In an area with a high abundance of quality habitat for the species, it was found that Andean Cat home ranges exhibited extensive overlap between males and females and between sexes, as well as with the Pampas Cat (Tellaeche 2015).
A very large overlap has been found in the ecological niche of Andean and Pampas Cats in the High Andes, which spans the diet, distribution, activity patterns, space and habitat use (Walker et al. 2007, Lucherini et al. 2009, Reppucci 2012, Tellaeche 2015). In all cases the niche breadth for the Andean Cat was narrower and more restricted (Reppucci 2012, Tellaeche 2018). It cannot be excluded that intraguild predation and competition is an additional factor reducing the Andean Cat population densities (Lucherini and Luengos Vidal 2003, Reppucci et al. 2011).
Threats Information
The rapid increase in worldwide demand for metals and minerals, including lithium, and the rapid expansion of exploitation of shale oil and gas through hydraulic fracturing or fracking in northern Patagonia (Walker et al. 2013) are the most pressing threats to Andean Cat habitat. The impact of lithium exploration and mining in the core area of the distribution range of the Andean Cat has not been assessed yet but could have extremely negative impacts not only on the Andean Cat but also on other high Andean species. These activities not only change the disposition of the land and connectivity between patches, but more importantly they make an extensive use of water (Chavez et al. 2023) that is an important limiting factor on these arid ecosystems. In some areas, especially in the north of Chile extraction of underground and surface waters is seriously affecting suitable areas for the Andean Cat and its prey (Iriarte and Jaksic 2022, Iriarte et al. in prep.).
The expansion of the agricultural frontier and inadequate livestock management are also impacting the high Andean peatlands. These peatlands are crucial for the presence of the Andean Cat (Marino et al. 2010, Lagos et al. 2020) and are at risk not only due to extensive use but also as a result of the effects of global climate change (Marino et al. 2010). These factors will have a negative impact on the Andean Cat throughout most of its range by a decrease in the geographic distribution size (Bennett et al. 2017).
Poaching of Andean Cats is a complex phenomenon, and requires different interventions. Andean Cat killing by local people who consider them a predator of their small domestic livestock has been frequently reported particularly in some regions of Argentina, Chile and Peru (Lucherini and Merino 2008, Lucherini et al. 2016). In Argentina, high mortality rates due to hunting by local people have also been inferred both in the northwest (Tellaeche et al. unpublished data), and the northern Patagonia, where cats are being killed by herders in retaliation for predation by larger carnivores (Novaro et al. 2010; Rocío Palacios, unpublished data).
Hunting also occurs for bad management of pets. Dog attacks on livestock and wildlife are quite common in the Andes and Patagonia and local herders perceive a marked increase in recent times (Rocío Palacios, unpublished data), and even some feral dog populations have been recorded (Juan Carlos Huaranca, unpublished data). A second effect of domestic dogs and cats on wildlife is their potential to transmit diseases and parasites between the domestic and wild settings. This threat has not been corroborated as the few tested Andean Cat individuals returned negative results for the most common diseases (Napolitano et al. 2019), but there is an urgency for more research on this topic, since domestic animals visibly sick with zoonotic diseases have been detected in or around Andean Cat habitat (Huaranca and Palacios, unpublished data).
Since the first record of an Andean Cat road kill in 2018 several other vehicle-cat accidents (in most cases fatal for the cat) have been reported in different part of its distribution. This new threat has not been evaluated but it is important to keep monitoring it especially because of the increased extent of roads in the remote areas inhabited by Andean Cats related to mining exploration and exploitation.
A final indirect threat, that is global and difficult to assess and address, is the increased migration of local people to cities. This creates opportunities for extractive industries to advance, unchallenged, over the land. Additionally, when people come back from cities, they bring with them a whole set of tools to perform retaliatory killing against carnivores, such as traps, guns or poison, to prevent or in response to predation on livestock. The lack of job opportunities and reduction in the sense of belonging, combined with needs that are largely created by global media, facilitate this situation that indirectly affects all high Andean wildlife.
Although most of the threats have been identified at a general scale, including prime and secondary Andean Cat habitats, they cover a significant proportion of the species’ distribution. Nevertheless, a detailed assessment of the impact of the different threats on the species and its habitat is much needed.
Use and Trade Information
In Argentina, Bolivia, Chile and Perú, people of Aymara, and in some cases Quechua, origin have similar beliefs regarding the Andean Cat and Pampas Cat (both known as titi or osqollo). In some areas a common tradition is the use of a skin or a stuffed cat during ceremonies that people perform for marking their domestic livestock, mainly llamas or alpacas; the titi is considered a sacred animal related with abundance and fertility of the livestock or quality of crops (Giraldo-Jaramillo 2015). It is important to note that both the Andean Cat and Pampas Cat are part of these traditions and beliefs, and in general, are used indiscriminately. There are some local variations within and between countries and in some cases the Andean Cats (in some cases identified as a separate species but mostly together with Pampas Cats) are recognised as a special figure, with important influence in the cosmovision because of this felid’s relation with the mountainous areas, being considered the messenger with the local Apus, or mountain gods. The influence of western culture has resulted in a total or partial loss of the values of Andean cultures and the distortion of ancestral customs regarding the titi. Because of that, we do not believe the traditional use of the Andean Cat for ceremonies is currently affecting local populations, since its demand is reducing (Villalba et al. 2004). Beside this, the skins used for those ceremonies transfer through generations, so the real impact on Andean Cat subpopulations is expected to be low (Giraldo-Jaramillo 2015).
Conservation Actions Information
The Andean Cat Alliance (Alianza Gato Andino, or AGA) has worked to ensure the survival of the Andean Cat since 1999. AGA (
www.gatoandino.org) is the only organisation exclusively dedicated to the protection of this species and has been leading all the past and current interdisciplinary and cross-border initiatives to protect this wildcat in its four range countries (Peru, Bolivia, Chile, and Argentina). Its rareness and strong association with the high Andean ecosystem make this felid an ideal flagship species for conservation of this habitat, which is considered one of earth's last wild places.
AGA's activities are implemented throughout the four countries encompassing the distribution range of the Andean Cat, in the form of multinational programmes that have a sensible approach to the mitigation of the main threats that affect the species. With that in mind, the Alliance's work focuses on developing programs that diminish these threats globally, but are customised to each community they work with. These multinational conservation programs use flexible toolkits that adapt to the specific needs of each area and local community in order to find creative solutions that mitigate conservation threats. These programmes are also designed to work collaboratively, and the coordinators work together to have bigger impact programs can be summarised as follows:
The CATcrafts Program provides alternative income sources, reducing hunting by changing the perception and attitude towards Andean Cats, and indirectly reducing population reduction rates and habitat loss to extractive industries. The Conflict Mitigation Program facilitates coexistence and reduces killing of Andean Cats and other carnivores by using non-invasive tools to reduce conflicts with pumas/foxes. The Pawsitive Actions Program aims to reduce killing of Andean Cats and disease transmission by domestic dogs, while also improving human health. The In the Field 24/7 program investigates to fill the gaps in our knowledge of the species, particularly their ecological and genetic characteristics through camera trapping monitoring and genetic analysis of non-invasive samples to assess connectivity among populations. Finally, the Education Program transverses all of the above creating content and strategies that can be applied to all schools and communities across the Andean Cat habitat. AGA members from each range country work collaboratively to coordinate conservation, education, and research programs (see
https://gatoandino.org/programas/ for more information).
One of the main and major threats for Andean Cat conservation is large scale mining activities, and actions at a political and institutional level are urgently needed in order to minimise the impacts of this threat. Desiccation and/or contamination of peatlands and habitat destruction derived from these extensive activities should be reduced, through the limiting of mining activities in core habitat areas for the Andean Cat on one hand while making mining more sustainable in the entire Andean ecosystem.
An additional area where investigation and conservation efforts are recommended is the assessment and creation of awareness on the Ecological Services provided by Andean Cat subpopulations and, indirectly, the high Andes habitat they are primarily associated with, with emphasis on non-monetary and cultural values associated to native population cosmogony. The revitalisation of cultural traditions that involve the significant role the Andean Cat plays in Andean communities is necessary. These traditions encompass an inherent care and respect for both wildlife and the ecosystems in which communities reside, but they are being threatened by deculturisation and the influence of Western culture on Andean communities.