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Overview

Brief Summary

Biology

Despite the pampas cat being a fairly common species, there is little data available on its ecology (3). It has a diet primarily of small mammals, such as guinea pigs. They also prey on the eggs and chicks of ground-dwelling birds (3). Due to its ability to occupy a wide range of habitats, it is likely to eat any small vertebrate it can catch (4). It has even been reported to kill adult goats and raid domestic chicken houses (6). The pampas cat is thought to be a predominantly nocturnal and terrestrial species (3). They are adept climbers, though it is not clear whether they use this skill for hunting or for just escaping from predators (5). The little available information regarding the breeding behaviour of this species comes from the study of captive species; a female first reproduced at the age of two years, and gestation lasted for 80 to 85 days. The litter size varies from one to three, and the average life span is nine years, but some have lived for over 16 years (3) (6).
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Description

This little known South American cat can be found in a greater variety of habitats than any other cat on the continent. It looks like a robust domestic cat, with a broad face, amber eyes and distinctive pointy ears (4). The ears are black or grey on the back, with a silvery grey spot in the centre, and two conspicuous stripes run from the eyes, across the cheeks and meet beneath the throat (4) (6). The colour, pattern and texture of the pampas cat's coat varies considerably throughout its wide range. The background colour varies from yellowish white to greyish brown to silvery grey, and can be soft, short and vividly patterned, or long, coarse and virtually unmarked (4). In fact, these geographical differences are so pronounced it has been proposed that the pampas cat is actually three species. Genetic studies are underway to determine if this is correct (3). Generally, the forelimbs and hindlimbs have distinctive brown bands, and the short, bushy tail has fairly indistinct brown or black rings. Long hairs on the back, which can be up to 7 cm long, stand erect when the cat is nervous or frightened, creating an appearance of being much larger than they actually are. In Argentina and Chile, the pampas cat is known as the “gato pajero” or grass cat, pajero being the local name for pampas grass, one of the habitats in which it lives (4).
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Distribution

Range Description

The Pampas Cat, named after Argentine grasslands, ranges throughout most of Argentina and Uruguay beyond into the dry forests (chaco, cerrado) of Bolivia, Paraguay and Brazil, and north through the Andes mountain chain through Ecuador and possibly marginally into southwestern Colombia (Silveira 1995, Ruiz-Garcia et al. 2003, Nowell and Jackson 1996, Dotta et al. 2007).

The suggestion by Pereira et al. (2002) that the species can be considered extinct in the Pampas of central Argentina has been confirmed by more recent surveys.

The most recent information indicates that this species is rare throughout a very large portion of its distribution range, such as the Patagonia, Monte, Espinal, Yungas, and Mesopotamia in Argentina, the Pampas and Pantanal in Brazil, the dry forests of Bolivia, as well as in Uruguay and Peru.
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Pampas cats, Leopardus colocolo, have an expansive geographic range. In fact, they have been said to have a greater geographic range than any other South American cat. They are found in the forested slopes of the Andes in Ecuador, Peru, and Bolivia, the cloud forests of Chile, the Paraguayan chaco, open woodland areas of central, western, northeastern, and southern Brazil, the pampas of Argentina and Uruguay, and southern Patagonia. (IUCN, 1996; Silveira, 1995)

Biogeographic Regions: neotropical (Native )

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Range

Occurs in south-western South America, in Ecuador, Peru, Brazil, Bolivia, Chile, Paraguay, Uruguay, and Argentina (4).
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Physical Description

Morphology

The physical characteristics of L. colocolo vary across its range in South America. In the high Andes it is gray in color and has reddish stripes that are broken up into spots. In Argentina, the coat of L. colocolo is generally longer and yellow-brown in color with a muted pattern. Long fur is also typical to those individuals living in Brazil, but they tend to be rust colored with black bands on their yellow to orange sides and their lateral underparts.

A three month old male pampas cat from central Brazil that was brought into a zoo had the typical rusty color but also had very dark, irregular stripes over its entire body. By the time it had reached eight months of age, the dorsal and lateral striping had disappeared, and only the stripes on the limbs and underparts remained.

Ears of L. colocolo are large and more pointed than most other small, neotropical cats. Typical head and body length is 435-700 mm, tail length is 220-322 mm, and shoulder height is 300-350 mm. Average weight is 3 to 7 kg.

(IUCN, 1996; Wildlife On Easy Street, 2000; Silveira, 1995)

Range mass: 3 to 7 kg.

Range length: 435 to 700 mm.

Other Physical Features: endothermic ; bilateral symmetry

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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
The Pampas Cat has a wide distribution outside the moist forests of South America, being associated with more open habitats. It typically inhabits dry scrub and grassland, but can also be found in dry woodland as well as swampy wetland and rocky areas (Silveira 1995, Nowell and Jackson 1996, Pereira et al. 2002, Tellaeche 2015). Its prey includes small mammals as well as ground-dwelling birds (Nowell and Jackson 1996, Silveira et al. 2005). In the high Andes the diet is based on mountain viscacha and small rodents (Walker et al. 2007, Napolitano et al. 2008, Fajardo et al. 2014). In Brazil's Emas National Park, Pampas cats are primarily diurnal with some crepuscular and occasionally nocturnal activity. Home ranges (90% MCP) averaged 19.47 +/- 3.64 km (Silveira et al. 2005). In the High Andes 71% of camera trap photos were taken at night and more than 20% during the day (Lucherini et al. 2009) and average home range size (95% Kernel) was 14.9 km (Tellaeche 2015).

Systems
  • Terrestrial
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Just as the geographic range of the species varies widely, so does the habitat in which it is found. It can be found in open woodland or scrub thicket, cloud forest, cold, semi-arid desert regions, low-lying swamps, floodplains, and mountainous slopes. The only forest regions it has not been found to inhabit throughout its range are lowland tropical and temperate rain forests.(IUCN, 1996; Silveira, 1995)

Habitat Regions: temperate ; tropical

Terrestrial Biomes: desert or dune ; forest ; scrub forest ; mountains

Wetlands: marsh

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As its name suggests, the pampas cat is typically associated with the pampas, or open grasslands. However, it also occurs in thorn forest and scrub, open woodlands, cloud forests, floodplains, swampy areas and semi-arid cold deserts. It has been reported to occur up to altitudes of 4,800 metres (4). One habitat it does not occur in is lowland rainforest (3).
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Trophic Strategy

L. colocolo preys upon small mammals, such as guinea pigs, as well as ground-dwelling birds. It has been observed taking penguin eggs and chicks from nests. Pampas cats are known to take poultry in areas of human population. (IUCN, 1996; Silveira, 1995; Garman, 1997)

Animal Foods: birds; mammals; eggs

Primary Diet: carnivore (Eats terrestrial vertebrates, Eats eggs)

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Associations

As a predator, L. colocolo probably influences the population sizes of prey species.

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This species is poorly known, so information on predation and possible anti-predator adaptations is not available.

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Known prey organisms

Oncifelis colocolo preys on:
Aves
Mammalia

This list may not be complete but is based on published studies.
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Life History and Behavior

Behavior

Perception Channels: tactile ; chemical

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Life Cycle

See Reproduction.

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Life Expectancy

Longevity in this species has not been reported. However, other felid species of similar size typically live between 10 and 15 years. (Nowak, 1999)

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Lifespan, longevity, and ageing

Maximum longevity: 19.6 years (captivity) Observations: One wild born specimen was about 19.6 years of age when it died in captivity (Richard Weigl 2005).
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Reproduction

The mating system and behavior of this animal are not known.

L. colocolo in captivity in the northern hemisphere breeds period from April to July. Gestation is from 80 to 85 days, and 1 to 3 young are born per litter. Breeding season in the wild is unknown. (IUCN, 1996; Silveira, 1995)

Breeding season: Breeding occurs from April to July in pampas cats held in captivity in the Northern Hemisphere.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); viviparous

As in all mammals, the female provides the young with milk. Specific information on the parental care of this species is lacking, but it is likely that, as with other felids, the young are altricial. They are probably born in a den where they are cared for by their mother until they are able to accompany her on foraging trips.

Parental Investment: altricial ; female parental care

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2015

Assessor/s
Lucherini, M., Eizirik, E., de Oliveira, T., Pereira, J. & Wallace, R.

Reviewer/s
Nowell, K., Hunter, L., Schipper, J., Breitenmoser-Wrsten, C., Lanz, T. & Breitenmoser, U.

Contributor/s
Leite-Pitman, M.R.P.

Justification
Pampas Cat is generally rare or very rare (0.05-0.2 individuals/km) and localized throughout most of its range, and appears to be declining in several parts of its extent of occurrence because of extensive loss or reduction in quality of its habitat. Predation by dogs, hunting and road kills are additional threats. Population decline caused by loss of habitat is widespread and is a primary concern. Although little data is available on the rate of loss of natural habitats across the Pampas Cat range, we have some information from the Gran Chaco region that covers a large portion of Pampas Cats distribution range. The Gran Chaco forest extends over parts of northern Argentina, western Paraguay, eastern Bolivia, and western Brazil and occupies an approximate area of 1,141,000 km. Deforestation started in the 1970s and accelerated in 2002-2006. Following the global increase in commodity prices, it is likely that the rates of deforestation have maintained or increased in the last decade. Based on the data from the dry Chaco region, a portion of the whole Gran Chaco covering 790,000 km the yearly rate of transformation of forest into cropland (a habitat type where Pampas Cat do not occur) ranges from 0.63 to 1.75% (Clark et al. 2010). Thus, adopting a generation length of seven years (Pacifici et al. 2013) and assuming that transformed habitat becomes unavailable to the Pampas Cat, populations of this species could be suffering a 36.5% declination rate over three generations.

Another issue is that possibly the Pampas Cat should not be assessed as a single evolutionary unit. The taxonomy of this species is not yet resolved but the available evidence (morphological, genetic and ecological) indicates that there are clear differences among regional subpopulations, which have been proposed to represent distinct subspecies or even species. It is reasonable to consider the conservation status of these regional populations separately, because there may be little (or even no) historical and/or current gene flow among some of them, implying that limited genetic connectivity may further threaten the long term viability of the species as a whole.

Because of all the above and because the reasons for its rarity and population numbers are not known, the limited information available indicates Near Threatened as the most likely category (it almost qualifies for a threatened listing under criterion A2c). Nevertheless, it may qualify as Vulnerable in the near future and hence requires monitoring.

History
  • 2008
    Near Threatened (NT)
  • 2002
    Near Threatened (NT)
  • 1996
    Lower Risk/least concern (LR/lc)
  • 1994
    Indeterminate (I)
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L. colocolo is listed in CITES Appendix 2. Habitat destruction across their range is the major threat to this species. The pampas regions of Argentina and Uruguay have been heavily settled and grazed, which is suspected to have had a negative impact on pampas cat populations. Reduction of the prey base of L. colocolo is also a problem.

Trade of pampas cat pelts was ended in 1987. This species is listed as an endangered by the Instituto Brasileiro do Meio Ambiente e Recursos Naturais Renov·veis. L. colocolo is considered a rare species according to the rarity classification of Rabinowitz, because it is found in a widespread geographic range, is a habitat specialist, and occurs at low population densities.

(IUCN,1996; Garman, 1997)

US Migratory Bird Act: no special status

CITES: appendix ii

IUCN Red List of Threatened Species: near threatened

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Status

Classified as Near Threatened on the IUCN Red List (1) and listed on Appendix II of CITES (2).
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Population

Population
The Pampas Cat populations living in the High Andes and Puna eco-regions appear to be able to reach relatively high densities (0.74-0.78 individuals/km2) in the most productive habitat patches (Gardner et al. 2010). The population appears to be stable in these regions of Argentina and in slight decline in Bolivia and Peru because of decreasing habitat quality caused by increased anthropogenic modification.

The Pampas Cat is a naturally rare species in most of Patagonia, where records of its presence are scarce in comparison with those of other felids (e.g. Geoffroys Cat, Puma).

In the Argentine Mesopotamia (Corrientes province), a couple of records (road-killed animals) were obtained during the last decade, but almost no evidence was recorded through camera trapping for the Iber wetlands in spite of intensive trapping effort, thus suggesting that the species is rare in this region.

The Pampas Cat appears to be declining in central Argentina (Cordoba, San Luis), mainly due to habitat modification (i.e. soybean crop expansion), and is regionally extinct in the Pampas grasslands, where it only occurs in the southern and dry portion (Pereira et al. 2002 and recent unpublished surveys). In the Argentine Espinal, population numbers are low (0.11-0.17 individuals/ km2, lower than for the Geoffroys Cat, Caruso et al. 2012) and distribution is largely limited to grassland habitats that are threatened by human activities. Pampas cat are similarly rare in the adjacent southern part of the Monte eco-region (Pereira et al. 2011).

Pampas Cat records are very scarce in the Yungas eco-region of NW Argentina, where it is limited to the high-altitude grasslands (Di Bitetti et al. 2011).

A survey on small cats has found no records of Pampas Cats in the Bolivian Chaco dry forest (Cuellar et al. 2006).

In Brazil, the Pampas Cat inhabits open areas, such as the grasslands of the Pampas, the marshy Pantanal, and particularly the savannas of the Cerrado (Araujo Bagno 2004). The species is considered typically rare and with low population densities, typically of 0.010.05 individuals/km2 (or lower), throughout these regions, but may be relatively common in a few areas, usually protected, such as Emas National Parks in the Cerrado (Godoi et al. 2010), and Mirador State Park, where densities of 0.10.2 individuals/km2 are reached (Oliveira et al. 2010, Oliveira 2011).

In the Brazilian Pampas (southern portion of Rio Grande do Sul [RS] state), this felid is considered to occur only in well-preserved habitats areas (whose remaining area covers ca. 81,500 km), and is always rarer than Geoffroys Cat.

Habitat loss and population decline is expected to be 14% in the next 21 years or three generations (assuming the same generation length of seven years). Because of this, the species was considered Vulnerable (C1) in Brazil (Queirolo et al. 2013). The situation is thought to be the same in Uruguay.

An interesting point revealed by recent molecular studies is that the populations in RS state (Brazilian pampas) and Uruguay are genetically distinct from those present in the Brazilian Cerrado and also from Argentinean populations, indicating that they have been demographically isolated for a considerable period (Santos, Trigo and Eizirik, in preparation). Such a result indicates that this region constitutes a distinct conservation unit for this species, which should be taken into consideration when assessing its overall status.

In Peru, the Pampas Cat is generally uncommon or rare. A seemingly separate subpopulation of Pampas Cat occurs in the Tumbesian region of western Ecuador and northwestern Peru. Individuals here are small and well-marked compared with the Andean form. This form inhabits mainly forested habitats, favoring dense vegetation along permanent water-courses and even in mangroves. It also occurs in agricultural fields, especially sugar cane plantations and in sparse desert habitats with little vegetation along the coast where shorebirds congregate. In this region, the high level of human impacts and settlement has resulted in widespread forest loss and the Pampas Cat is now extremely rare, being primarily found in protected areas.

Most of its subpopulations, which could in fact end up being considered as genetically different species, would be Vulnerable individually.

Population Trend
Decreasing
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Threats

Major Threats
Habitat loss (to agricultural cropland) and degradation (by livestock grazing) is considered the major threat to this species throughout most of its range. Retaliatory killing for poultry depredation is also a threat, as are road kills.

In the High Andes of Bolivia and Peru the Pampas Cat is threatened by decreasing habitat quality caused by increased anthropogenic modification. In these regions, it is also actively persecuted by local people and often killed by dogs.

In Argentina, extensive habitat loss/modification due to the expansion of the agriculture frontier, mining, and oil extraction is the major threat.

The biggest threat to the species in Brazil is habitat loss, especially because of silviculture and agriculture expansion, as well as habitat conversion/degradation, mostly through fire. Given its rarity in Brazil, road kills are also considered a threat for some populations. In the Brazilian Pampas, predation by domestic dogs and hunting due to real or perceived conflicts, a problem that apparently has not been detected in the Cerrado yet (where it would be more incidental than widespread). A zone of hybridization between L. colocolo and L. tigrinus has been demonstrated by genetic analysis in central Brazil (Trigo et al. 2008).

In Peru, it is actively persecuted as it often kills chickens of local people, and is also impacted by dogs. Forest loss appears to be also an important threat.
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The pampas cat faces a number of threats, but due to the lack of information regarding the status of this cat in the wild, it is difficult to determine to what extent populations are being impacted. In the past they were hunted in large numbers for their pelts; 78,000 pelts were exported between 1976 and 1979, but international trade ceased in 1987 (3). Today, habitat destruction is believed to be the primary threat (4). A large percentage of pampas grassland in Argentina and Uruguay has been converted into agricultural land and heavily grazed, resulting in a reduction of available habitat and prey species. It is thought by some to be extinct in Uruguay, but probably persists there in very low numbers (3). Sport hunting is also considered to be a threat, but to what extent is not known. It is known that in central Chile hunters and their dogs are becoming more common, whilst dense cover for the pampas cat is becoming rarer (4).
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Management

Conservation Actions

Conservation Actions

Included on CITES Appendix II. The species is protected by national legislation across most of its range, with hunting prohibited in Argentina, Bolivia, Chile, Paraguay, and Peru. It occurs in 13 National Protected Areas in Argentina (Pereira et al. 2002), eight in Bolivia (Noss et al. 2010), 11 in Brazil (Queirolo et al. 2013), 75 in Chile (A. Iriarte in litt.), and 11 in Peru (Fajardo and Pacheco 2011).

The Pampas Cat is listed as Vulnerable by the national Red Lists of Argentina, Bolivia, and Brazil. In Peru is listed as Data Deficient.

Research into its ecology, distribution, taxonomy, and threats is urgently needed.

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Conservation

The pampas cat is listed on Appendix II of the Convention of International Trade in Endangered Species (CITES), which means that trade in this species should now be tightly controlled (2). Hunting of the pampas cat is prohibited in Argentina, Bolivia, Chile and Paraguay, and regulated in Peru, but there is no legal protection in place at present in Brazil or Ecuador (3). Conservation actions recommended by the IUCN/SSC Cat Specialist Group include determining whether the pampas cat is actually more than one species, and conducting research into its behaviour, ecology and distribution in the Argentinean pampas grasslands (3).
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Relevance to Humans and Ecosystems

Benefits

L. colocolo has been known to take domestic poultry as prey in areas where it lives near human habitation. (Garman, 1997)

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L. colocolo was formerly hunted across its range for its fur. (Garman, 1997)

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Wikipedia

Colocolo

For other uses, see Colocolo (disambiguation).

The colocolo (Leopardus colocolo) is a small spotted and striped cat native to the west Andean slope in central and northern Chile.[2] Until recently, it included the more widespread Pampas cat (L. pajeros) and Pantanal cat (L. braccatus),[1] and some maintain these as subspecies of the colocolo.[3][4] Confusingly, when these are treated as subspecies of the colocolo, the "combined" species is sometimes referred to as the Pampas cat.[5]

Taxonomy[edit]

Genus[edit]

Like most other small cats, the colocolo was formerly included in the genus Felis,[6] but together with Geoffroy's cat and the kodkod, some have placed it in Oncifelis instead.[7] Today, all major authorities place it in Leopardus.[1][3][4]

Species and subspecies[edit]

As traditionally defined, the colocolo occurs in the widest range of habitats of any small South American felid.[3] This, combined with distinct differences in pelage colour/pattern and cranial measurements, was the basis for splitting the Pantanal cat and Pampas cat from the colocolo.[2] Based on genetic divergence, the splits within the colocolo group are estimated to have occurred about 1.7 million years ago (Mya).[8] This divergence was lower than that found within Geoffroy's cat (about 2 Mya) or the oncilla (about 3.7 Mya; this very high divergence –far higher than other species in the genus– has resulted in some suggesting more than one species is involved in the oncilla).[8] Furthermore, the distribution pattern within the colocolo group based on genetics did not completely match that based on pelage colour/pattern and cranial measurements, and supported some of the traditional subspecific division rather than species division.[8] While the population in northern Chile has been placed in L. colocolo based on cranial measurements, genetics associate it with Pampas cats from Bolivia, and while the population in western Argentina has been placed in the Pampas cat based on pelage colour/pattern and cranial measurements, genetics associate it with the colocolo.[2][8] More recent genetic analysis also supports maintaining the Pantanal and Pampas cats as subspecies of the colocolo.[9]

When the Pantanal and Pampas cats are treated as a separate species, the colocolo has two subspecies: L. c. colocolo and L. c. wolffsohni.[1][2]

Description and habitat[edit]

The colocolo is a small, but heavy-set cat, only 56 to 67 cm (22 to 26 in) in body length, with a short 29 to 32 cm (11 to 13 in) tail, and weighing around 3 kg (6.6 lb) on average.[10] The two subspecies differ in their pelage colour and pattern:

  • L. c. colocolo (nominate): Reddish or dark grey with rusty-cinnamon stripes on the flanks and two stripes on each cheek, a cinnamon upper side of the ears with black edges and tips, four or five reddish rings on the tail (outer two are darker), dark brown stripes on the legs, black chest spots, and whitish underparts with rusty-ochraceous stripes.[2] It is found in central Chile in subtropical, xerophytic forests at altitudes of up to 1,800 m (5,900 ft).[2]
  • L. c. wolffsohni: Similar to nominate, but the flanks have large, reddish brown rosette-shaped spots with darker borders, the upper side of the ears are black with a greyish base and a small grey spot, usually eight rings are on the tail (of the same colour as the flank spots), and the stripes on the legs and spots/stripes on the underparts are very dark brown (almost black).[2] It is found in northern Chile in spiny shrublands and páramo.[2] Of two specimens, one was taken at an altitude between 2,000 and 4,000 m (6,600 and 13,100 ft), and the other at 4,100 m (13,500 ft).[2]

Externally, the colocolo differs from the Pantanal cat in its larger size, and pelage colour and pattern.[2] Some Pampas cats are as large as the colocolo, and some subspecies of the Pampas cats have the same pelage colour and pattern as colocolos of the subspecies L. c. wolffsohni.[2]

Behavior[edit]

Little is known about the colocolo's hunting and breeding habits; however, it is believed to prey mainly on small mammals and birds. Guinea pigs are thought to form a large part of its diet, along with viscachas and other rodents, and tinamous.[10] Though some have suggested it is chiefly nocturnal,[10] others suggest it is mainly diurnal.[9]

Litters are relatively small, usually consisting of only one or two kittens, and occasionally three. The kittens weigh around 130 g (4.6 oz) at birth.[10] The average lifespan is nine years, but some have lived for over 16 years.[11]

Status[edit]

The IUCN has only rated the "combined" species (including Pantanal and Pampas cats), in which case the colocolo is considered Near Threatened.[4] Whether the colocolo will receive a higher rating if the Pantanal and Pampas cats are treated as separate species is unclear, but it may be Endangered,[2] and its range is smaller than that of the kodkod, rated as Vulnerable by the IUCN.[12] However, unlike the kodkod, the colocolo is not associated with the highly threatened temperate rainforests found in the region.[2][12]

References[edit]

  1. ^ a b c d Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. p. 538. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  2. ^ a b c d e f g h i j k l m Garcia-Perea, R. (1994). The pampas cat group (Genus Lynchailurus Severertzov 1858) (Carnivora: Felidae), A systematic and biogeographic review. American Museum Novitates 3096: 1-35.
  3. ^ a b c Sunquist, M. E., & Sunquist, F. C. (2009). Colocolo (Leopardus colocolo). Pp. 146 in: Wildons, D. E., & Mittermeier, R. A. eds. (2009). Handbook of the Mammals of the World. Vol. 1. Lynx Edicions, Barcelona. ISBN 978-84-96553-49-1
  4. ^ a b c Pereira, J., Lucherini, M., de Oliveira, T., Eizirik, E., Acosta, G., Leite-Pitman, R. (2008). "Leopardus colocolo". IUCN Red List of Threatened Species. Version 2010.4. International Union for Conservation of Nature. Retrieved 5 January 2011. 
  5. ^ Novak, R. M., eds. (1999). Walker's Mammals of the World. Vol. 1. 6th edition. Johns Hopkins University Press, Baltimore. ISBN 0-8018-5789-9
  6. ^ Redford, K. H., & Eisenberg, J. F. (1992). Mammals of the Neotropics – The Southern Cone. Vol. 2. University of Chicago Press, Chicago. ISBN 0-226-70681-8
  7. ^ Wozencraft, C. (1993). Order Carnivora. Pp. 279-348 in: Wilson, D., & Reeder, D. eds. (1993). Mammal Species of the World. 2nd edition. Smithsonian Institution Press, Washington, D.C. ISBN 978-1-56098-217-3
  8. ^ a b c d Johnson, Slattery, Erizirik, Kim, Raymond, Bonacic, Cambre, Crawshaw, Nunes, Seuánez, Moreira, Seymour, Simon, Swanson, & O'Brien (1999). Disparate phylogeographic patterns of molecular genetic variation in four closely related South American small cat species. Molecular Ecology 8: S79–94
  9. ^ a b Macdonald, D., & Loveridge, A., eds. (2010). The Biology and Conservation of Wild Felids. Oxford University Press. ISBN 978-0-19-923445-5
  10. ^ a b c d Sunquist, Mel; Sunquist, Fiona (2002). Wild cats of the World. Chicago: University of Chicago Press. pp. 201–204. ISBN 0-226-77999-8. 
  11. ^ ARKive
  12. ^ a b Acosta, G. & Lucherini, M. (2008). "Leopardus guigna". IUCN Red List of Threatened Species. Version 2010.4. International Union for Conservation of Nature. Retrieved 5 January 2011. 
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