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Intraguild competition as a potential factor affecting the conservation of two endangered cats in Argentina.

Abstract

Although the kodkod, Oncifelis guigna, and Andean Mountain cat, Oreailurus jacobita, are the two cats in most immediate danger of extinction in Latin America, information on these felids is very scarce. As part of our effort to understand their current distribution, we collected data on the carnivore guild composition of these two small felids from 1998 to 2002. Sign search, in combination to genetic analysis of scat samples, enabled us to record the presence of the Andean Mountain cat, the similar-sized Pampas cat, Oncifelis colocolo, and the Culpeo fox, Pseudalopex culpaeus, at a site located at the high-altitude Argentina Andes. At this site, the abundance of O. colocolo and P. culpaeus appears greater than that of O. jacobita. At another site located at the Patagonia Mountain forest in Argentina, the kodkod, the slightly larger Geoffroy's cat, Oncifelis geoffroyi, and the Culpeo fox were live trapped. The population abundance of the kodkod in the more competitive guild of Argentina seems lower than what has been previously reported for the species in Chile. We suggest that intraguild competition may be an important factor affecting the present conservation status of the Andean Mountain cat and kodkod, and stress the need of a guild approach to the conservation of these endangered small carnivores.

Introduction

Argentina hosts all ten species of Neotropical felids, almost 28% of all the species in the world (Nowell and Jackson 1996). This impressive diversity of cats results in extensive range overlaps, which, in some regions, may involve up to eight species (Soler and Lucherini in press). The preservation of this felid diversity requires detailed knowledge about the true extent of sympatry and implies answering theoretical questions about niche overlap and segregation (Johnson et al. 1996).

The kodkod, Oncifelis guigna, and the Andean Mountain cat, Oreailurus jacobita, are the two cats that are in most immediate danger of extinction in Latin America and have been listed among the highest conservation priorities by the IUCN Cat Specialist Group (Nowell and Jackson 1996; Nowell 2002a) and as vulnerable in the Argentine Red List of mammals (Diaz and Ojeda 2000). Andean Mountain cats only inhabit the Puna, a high-altitude desert area of the Andes of Peru, Bolivia, Chile and Argentina (Oliveira 1994; Yensen and Seymour 2000). Although its natural history is still very poorly known, fragmentation, low local densities and decreasing prey populations are thought to be the causes of the endangered status of O. jacobita (Nowell and Jackson 1996). The main reasons for O. guigna's status are its extremely limited distribution and strong association to the Andean Patagonia forest of Southern Chile and Argentina, which is under strong human pressure (Nowell and Jackson 1996). Two recent studies have reported the first data on the natural history of two kodkod populations in Chile (Dunstone et al. 2002; Sanderson et al. 2002), while almost no information is available on the Argentine side of the Andes (Lucherini et al. 2000).

The population density of many carnivores has been found to be strongly correlated to prey biomass (Fuller and Sievert 2001; Carbone and Gittleman 2002). However, some of the variation in carnivore population density that is not explained by prey biomass may be due to the unique features of each population (Carbone and Gittleman 2002). Intraguild competition has been shown to be a widespread cause of mortality in carnivores (Palomares and Caro 1999), and one of the factors determining their abundance (Linnell and Strand 2000).

In this study, we report new data on the sympatry of O. jacobita and O. guigna with other carnivores in Argentina, and analyze the possible effects of intraguild competition on the conservation of these felids.

Study Sites

Data were collected from seven expeditions we conducted from 1998 to 2002 in the Andean Puna and Andean Patagonia forest. Four of these expeditions were carried out in the Puna, an arid, high-altitude (usually above 3500-4000 m a.s.l.) section of the Andes that extends 12,457,000 ha (Bertonatti and Corcuera 2000). This ecoregion is ranked regionally within the highest conservation priorities because of its outstanding biological value and vulnerable conservation status (Biodiversity Support Program 1995). We conducted three expeditions in the Patagonia Mountain forest in Argentina, a 6,604,000 ha southern temperate forest that has been ranked as Endangered and as a High Regional Priority (Biodiversity Support Program 1995; Bertonatti and Corcuera 2000). In the southern Argentine province of Chubut inside the Patagonia Mountain forest, the 263,000-hectare Los Alerces National Park (ANP) represents one of the largest protected areas in the southern temperate forest.

Data Collection

Because of the long-term nature of these projects and particularly, the great differences in the habitats where these cats live, we used an array of study techniques. The Puna survey areas were selected on the basis of previous reports and inteviews with local cattle breeders regarding the presence of Andean Mountain cats. Cat presence was assessed by sign search. In the survey areas, we intensively searched rocky areas, since, in this region, small cat scats are mainly found in shelters among rocks and boulders (Lucherini et al. 1999). All cat evidences were geo-positioned with a Geographic Position System Device (GPS). Cat scats were identified on the basis of their shape, texture, size, and, particularly, smell (Lucherini et al. 1999). Although none of these features alone is sufficient, their simultaneous use permits the correct identification of a sample (Lucherini et al. 1999). Fecal samples were collected in the Anconquija mountain range and Tucuman province in 1999, southwest of Jujuy province in 2001, and western Salta province in 2002. Each area was surveyed for 15 to 25 days. Collection methods are described in detail in Lucherini et al. (1999). For genetic analyses, all fresh feces were stored in paper bags and dried with silica-gel. The DNA of a sub-sample of each one of these scats was later extracted and amplified by polymerase chain reaction. The species were identified using a sequence-based analysis of the 16S rRNA mitochondrial gene (e.g. Johnson and O'Brien 1997).

Since 1998, the presence / absence of the kodkod has been studied in the Andean Patagonia forest, the only ecoregion of occurrence of the kodkod. Since 2000, presence/absence of small cats has been studied through live-trapping in ANP, which is the eastern-most part of the kodkod distribution (trapping procedures have been described in detail in Luengos Vidal et al. in press). Baited traps were set along trails crossing different habitats and checked three times a day. The total trapping effort was 1051 trap days. Cats were captured in box-traps and immobilized by intramuscular injection with tiletamine-zolazepam (Zoletil[R]) at a dosage suggested by Kreeger (1997). The combination of these two drugs has been widely recommended and its safety has been documented for a number of carnivores (Travaini and Delibes 1994; Lariviere and Messier 1996). All individuals were released at the capture site after their complete recovery. Trapping procedures have been described in detail in Luengos Vidal et al. (in press).

Results

Puna Carnivores

Twenty-five small cat fecal samples from the Puna were genetically analyzed, 14 of which were succesfully identified. One of the five fecal samples collected in 1999 from the Tucuman province, and eight of the eleven scats from Salta province were identified as belonging to Pampas cats. Four of the nine scats collected in May 2001 in the vicinity of the village of Coranzuli, Jujuy province, matched O. colocolo, while a scat found on a very steep rocky cliff at 3940 m a.s.l (about 23 [degrees] 04'30"S 66 [degrees] 16'W) matched O. jacobita. One of the Pampas cat's feces was collected close (2,000 m) to this O. jacobita scat. We also collected a cat skull at less than 2,900 m from the Andean Mountain cat scat that we later identified as a Pampas cat's skull, on the basis of Garcia Perea's (1994) craniometric review. Genetic analyses also confirmed the presence of the Culpeo fox (Pseudalopex culpaeus) in this area.

Patagonia Carnivores

Ten carnivores (two Patagonia skunks, Conepatus humboldtii, four Culpeo foxes and four small cats) were trapped in ANP. In October 2000, a kodkod cat was observed and photographed by a tourist about 500 m from Villa Futalaufquen, the main village of ANP. Based on this sighting we carried out carnivore live-trapping in the surroundings of Villa Futalaufquen from November 2001 to March 2002, and from September to December 2002. On December 18, 2001 at Puerto Limonao, 4,000 m from Villa Futalaufquen, a 2.7-kg, adult female cat was trapped (see Lucherini et al. 2002, for a photograph of this specimen). Its morphological characteristics (dark brown, heavily spotted coat; long and dense fur, thick tail with many black rings; comparatively large paws) mainly resembled those of a kodkod, but its size, larger than those previously reported (Sunquist and Sanderson 1998; Dunstone et al. 2002), and relatively long ears, resembled those of a Geoffroy's cat (Oncifelis geoffroyi). On March 6, 2002, another adult female cat was restrained in close proximity of Villa Futalaufquen (see Lucherini et al. 2002, for a photograph of this individual). The characteristics of this 3.2-kg cat were typical of the southern form of O. geoffroyi (Lucherini et al. 2001). On October 6, a 1.3-kg O. guigna (an adult female) was captured in the same area (42[degrees]53'S-71[degrees]37'30"W). Kodkod trapping efficiency (computed as the number of kodkods captured / 100 trap days) was 0.19 in Los Alerces, while trapping efficiency of all wild cats was 0.39. The DNA isolated from a sample of four cat scats (out of six collected in 2001 and 2002 in the proximity of Villa Futalaufquen) matched that of O. geoffroyi. Pumas (Puma concolor) were present but rare, while the likely occurrence of the lesser grison (Galictis cuja) has not been confirmed yet (Chehebar 2002).

Discussion

Puna Carnivores

Although the presence of skins of both the Andean Mountain cat and Pampas cat in the homes of villagers of the Andean Puna had been previously reported (Garcia Perea 2002, Walker and Novaro 2003), this is the first empirical evidence that these two cats can live sympatrically and at the same site.

Despite the scarcity of resources, the carnivore guild of the Puna comprises at least seven species. The body sizes of four of them (P. concolor, O. colocolo, P. culpaeus, and the South American gray fox, P. griseus) make them potential competitors of the Andean cat, while less niche overlap may be expected with the lesser grison, Galictis cuja, and the hog-nosed skunk, Conepatus spp. The information we gathered suggests that P. griseus uses more open and dry areas than the other carnivores, and a certain degree of spatial and habitat separation also occurs between cats, Culpeo foxes and pumas (Lucherini et al. 1999; Lucherini unpl. report). It also suggests that, at present, both the Culpeo fox and the Pampas cat are much more common than O. jacobita. Because of the new evidence that the Pampas cat may use the same areas as O. jacobita, we suspect that interspecific competition may affect the current status of the Andean Mountain cat.

The population density has been found to be strongly correlated to prey biomass in many carnivores (Fuller and Sievert 2001). One of the implications of intraguild competition is that human-caused changes to the main prey of one carnivore species are likely to have effects through the entire carnivore community. This may be the case with the Andean Mountain cat. O. jacobita is considered to be a conservation priority largely because it is thought to be dependent upon chinchillas (Chinchilla brevicaudata) and their less-valued relatives, the Mountain viscachas (Lagidium viscacia) (Nowell and Jackson 1996). Chinchillas have almost been exterminated in the wild because of their prized fur and are now listed as critically endangered both nationally (Diaz and Ojeda 2000) and internationally (IUCN 1976), while Mountain vizcachas colonies appear to be distributed very irregularly, separated by large unpopulated areas (Lucherini et al. unpubl. data). It is logical to expect that a decrease in the availability of O. jacobita main prey would cause a direct reduction in the numbers of the Andean Mountain cats. The indirect consequences of this alteration in prey populations are more difficult to predict. In the co-occurrence of two similar-sized, evolutionary-related, carnivores, we might expect that when a resource becomes limiting, intraguild competition increases strongly. The Pampas cat, because of its wide geographical and ecological distribution, is considered to be a more adaptable small felid (Oliveira 1994), and may be able to switch to alternative preys and would be competitively favoured with respect to the more specialized O. jacobita.

Patagonia Carnivores

Until now, the kodkod was thought not to occur in sympatry with the Geoffroy's cat in the Andean Patagonia forest (Nowell and Jackson 1996). Interspecific competition can take two forms: exploitation and interference. In many cases, in presence of larger species, small carnivores are affected by both types of competition and their density can be strongly reduced when compared to what might be expected on the sole basis of prey abundance (Linnell and Strand 2000). We suggest that this may be the case with the kodkod in Argentina. In the Chile Island of Chiloe, the largest carnivore coexisting with O. guigna is the small (three kg of body mass) Darwin fox, Pseudalopex fulvipes (Jimenez and McMahon in press). In the Laguna San Rafael area (Southern Chile), no other carnivore is present (Dunstone et al. 2002). Our results suggests that the kodkod lives in a larger and more competitive carnivore guild in Argentina, which can only be compared to that of the Chilean Queulat National Park (QNP). Dunstone et al. (2002) reported the occurrence of Puma concolor, Pseudalopex culpaeus, Mustela vison and Lyncodon patagonicus in QNP. Thus, the main difference between QNP and ANP, which are located at similar latitudes, is the presence of O. geoffroyi in the latter. If the co-occurrence of related carnivores results in competition as well as habitat and spatial segregation (as it has been shown in the case of P. culpaeus and P. griseus, Johnson and Franklin 1994), we would expect a lower kodkod population density at ANP than at QNP. Consistent with this prediction, the trapping efficiency in Queulat was 2.04 (Dunstone et al. 2002), more than 10 times that recorded in Los Alerces. Although other factors such as prey abundance (Carbone and Gittleman 2002) may affect carnivore population size, and taking into account that trapping efficiency is a very crude estimate of population abundance, this comparison suggests that the kodkod abundance is remarkably lower in our study site than in Chile. Furthermore, scat frequency seems to indicate that the Geoffroy's cat is more abundant than O. guigna at Los Alerces. Thus, as in the case of the Andean Mountain cat, our results support the hypothesis that intraguild competition may be an important factor affecting the conservation status of the kodkod.

Although they are generally accepted as two different species (Wozencraft 1993; O'Brien 1996), it has also been suggested that the kodkod might be a subspecies of the Geoffroy's cat (Nowell and Jackson 1996). The only available molecular genetic studies, based on an extremely small sample size, put these two cats in the same phylogenetic subgroup (Johnson and O'Brien 1997; Johnson et al. 1998). Since character displacement would be expected between similar species in the zone of sympatry (Dayan and Simberloff 1996), the capture of an individual with a mixture of traits of the two species stresses the need of data that will enable researchers to clarify the phylogenetic relatedness of O. guigna and O. geoffroyi.

General Conservation Implications

While the presence of other carnivores in the areas of occurrence of the Andean Mountain cat and the kodkod was previously known, our findings show that both of them live in close sympatry with related carnivores of similar size. Since the potential for competition between two species tends to increase with their relatedness and morphological similarity (Odum 1966), our results suggest that intraguild competition may be affecting the present status of O. jacobita and O. guigna.

The effective conservation of a species requires detailed knowledge of its present distribution, population status, ecological requirements, and genetic identity (Wilson 2000). Despite the increased attention they received in the last few years, O. jacobita and O. guigna are still among the species with a low research effort in comparison to their precarious conservation status (Nowell 2002b). Our results contribute to the reduction in this gap, which is especially wide in Argentina (Soler et al. in press).

The conservation of carnivores has traditionally been a species-by-species effort. However, it has been recently suggested that all carnivores are influenced to some degree by intraguild competition (Palomares and Caro 1999; Linnell and Strand 2000). In particular, intraguild competition can negatively affect the conservation status of extinction-prone carnivore populations (Creel et al. 2001). Consequently, a guild-based approach is more appropriate for conserving endangered carnivores when potential for competition is likely to exist. Our results emphasize the urgent need of this more complete approach to the conservation biology of endangered small carnivores, which combines the collection of autoecological data, and the simultaneous study of the interactions between guild members.

Competencia Intragrupal Como un Factor Potencial en la Conservacion de Dos Felinos en Peligro de Extincion en Argentina

Resumen

Aunque el kodkok, Oncifelis guigna, y el gato montes andino, Oreailurus jacobita, son los dos felinos en mayor riesgo de extincion en America Latina, la informacion referente a estas dos especies es muy escasa. Como parte de nuestros esfuerzos por entender su distribucion actual, de 1998 al 2002 colectamos datos sobre la composicion del grupo taxonomico de carnivoros al que estos dos pequenos felinos pertenecen. La busqueda de signos, en combinacion con analisis genetico de muestras fecales, nos permitio registrar la presencia del gato montes andino, del gato de las pampas, Oncifelis colocolo, y del zorro andino, Pseudalopex culpaeus, en un sitio a gran altitud localizado en los Andes argentinos. En este sitio, la abundancia de O. colocolo y P. culpaeus parece ser mayor que la de O. jacobita. En otro sitio localizado en el bosque de montana de la Patagonia en Argentina, el kodkod, el gato montes (de un tamano un poco mayor al del kodkod), Oncifelis geoffroyi, y el zorro andino fueron trampeados. La abundancia poblacional del kodkod en el grupo taxonomico mas competitivo en Argentina parece ser menor de lo que habia sido previamente reportado para esta especie en Chile. Sugerimos que la competencia intrataxonomica puede afectar en forma importante el estatus del gato montes andino y el kodkod, y que existe la necesidad de una nueva propuesta para la conservacion de estos dos pequenos carnivoros en peligro de extincion que tome en consideracion la competencia intragrupal.

La Concurrence Intraguilde Comme Facteur Potentiel Affectant la Conservation de Deux Chats en Peril en Argentine

Resume

Bien que le kodkod, Oncifelis guigna, et le chat de montagne andin, Oreailurus jacobita, soient les deux chats le plus en danger immediat de disparition en Amerique latine, l'information disponible sur ces felids est tres rare. En tant qu'element de notre effort de comprendre leur distribution courante, nous avons rassemble des donnees sur la composition de guilde carnivore de deux petits felids de 1998 a 2002. Recherche des Signes, en combination avec l'analyse genetique des echantillons de matieres fecales, nous avez permis d'enregistrer la presence du chat de montagne andin, du chat Pampas d'une taille semblable, Oncifelis colocolo, et du renard Culpeo, Pseudalopex culpaeus, dans un site dans l'haut-altitude des Andes argentines. Dans cet site, l'abondance O. colocolo et de P. culpaeus semble plus grande que du de O. jacobita. Dans un autre site dans la foret des montagnes patagones de l'Argentine, le kodkod, le chat Geoffroy Oncifelis geoffroyi (legerement plus grand que le kodkod), et le renard Culpeo etaient captures vivant. L'abondance de la population kodkod dans la guilde plus concurrentielle de l'Argentine semble plus bas que ce qui a ete precedemment rapporte pour les especes en Chili. Nous proposons que la concurrence entre-guilde puisse etre un facteur important affectant le statut actuel de conservation du chat de montagne andin et du kodkod, et soulignons le besoin d'une approche de guilde a la conservation de ces petites carnivores en peril.

Acknowledgments

The Patagonia office of the National park Agency (APN), and, specifically C. Chehebar, and the staff of Los Alerces and Campo de Los Alisos National Parks, and particularly, R. Neira and F. Nahuelpan, provided logistic support. We are grateful to D. Birochio, M.J. Merino, L. Soler, D. Castillo and M. Ciuccio for their essential contribution in data collection and E. Casanave for providing laboratory facilities. The veterinarians M. Uhart, P. Beldomenico and C. Baldi provided professional advice and support during the first captures at Los Alerces. A. Surgenor, S. Alarcon Farfan and two anonymous referees improved our previous drafts. E. Casanave (Universidad Nacional del Sur) provided laboratory facilities. The GECM's Andean Mountain cat and kodkod projects received the generous support of British Petroleum Conservation Programme, Cat Action Treasury and the Leonard X. Bosack and Bette M. Kruger Foundation, Cleveland Zoological Society, Denver Zoological Foundation, Idea Wild, ISEC Canada, La Torbiera Zoological Society, Wild About Cats. The Education Ministry of Jujuy province and Secretary of Environment of Salta Province endorsed the Andean Mountain cat project. APN endorsed the kodkod project and authorized all trapping and animal handling procedures.

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Mauro Lucherini

Estela Luengos Vidal

Mammal Behavioral Ecology Group (GECM)

Departamento de Biologia,

Bioquimica y Farmacia--UNS

San juan 670

8000 Bahia Blanca

Argentina

luengos@criba.edu.ar

Mauro Lucherini is Joint Researcher of the Argentine National Commission for Scientific and Technological Research (CONICET). In 1993, he earned a PhD in Zoology at Siena University in Italy. Since 1996, he has led the Mammal Behavioral Ecology Group (GECM), a team of Argentine biologists who work toward the understanding and conservation of neotropical mammals. The Mammal Behavioral Ecology Group has launched several projects on the Andean Mountain cat, kodkod, Geoffroy's cat and other little known carnivores in Argentina. Lucherini is part of the Cat and Canid Specialist Group of the World Conservation Union (IUCN) and has been appointed as coordinator of the multi-national Andean Mountain cat Conservation Committee (COCGA).

Estela M. Luengos Vidal has worked with Argentina carnivores since 1994 and is one of the founders of GECM. She is completing a Master's degree in Biology at the Universidad Nacional del Sur (UNS) studying the Pampas fox (Pseudalopex gymnocercus).
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Author:Lucherini, Mauro; Vidal, Estela Luengos
Publication:Endangered Species Update
Geographic Code:3ARGE
Date:Nov 1, 2003
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