FINAL REPORT. Submitted to: Prepared by: JAVIER A. PEREIRA. Gatos del Monte Project Association for the Conservation and Study of Nature (ACEN)
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1 FINAL REPORT ECOLOGY AND CONSERVATION OF SYMPATRIC PAMPAS CAT AND GEOFFROY S CAT IN AN ENDEMIC ECOREGION OF CENTRAL ARGENTINA Submitted to: THE RUFFORD FOUNDATION Prepared by: JAVIER A. PEREIRA Gatos del Monte Project Association for the Conservation and Study of Nature (ACEN) Iberá vo. B, Ciudad de Buenos Aires (1429), Argentina gatosdelmonte@acen.org.ar November
2 GATOS DEL MONTE PROJECT TEAM Lic. Javier Pereira Asociación para la Conservación y el Estudio de la Naturaleza (ACEN) Lic. Natalia Fracassi Asociación para la Conservación y el Estudio de la Naturaleza (ACEN) Instituto Nacional de Tecnología Agropecuaria (INTA) Med. Vet. Marcela Uhart Field Veterinary Program - Wildlife Conservation Society (WCS) Universidad Nacional del Centro (UNICEN) Med. Vet. Hebe Ferreyra Field Veterinary Program - Wildlife Conservation Society (WCS) Med. Vet. Carolina Marull Field Veterinary Program - Wildlife Conservation Society (WCS) DVM MPVM Pablo Beldomenico Facultad de Ciencias Veterinarias - Universidad Nacional del Litoral Wildlife Disease Group Leahurst Veterinary Field Station, University of Liverpool Web Page: 2
3 TABLE OF CONTENTS CAPTURE AND IMMOBILIZATION OF WILD CATS AT LIHUÉ CALEL NATIONAL PARK, CENTRAL ARGENTINA Javier A. Pereira, Marcela M. Uhart, Hebe del V. Ferreyra, Carolina Marull and Natalia G. Fracassi Page 4 NUMERICAL AND SPATIAL RESPONSES OF GEOFFROY S CAT (ONCIFELIS GEOFFROYI) TO A SEVERE DECLINE IN PREY ABUNDANCE IN THE MONTE DESERT, ARGENTINA Javier A. Pereira, Natalia G. Fracassi, and Marcela M. Uhart Page 10 HELMINTHS OF GEOFFROY S CAT, ONCIFELIS GEOFFROYI (CARNIVORA, FELIDAE) FROM THE MONTE DESERT, CENTRAL ARGENTINA Pablo M. Beldomenico, John M. Kinsella, Marcela M. Uhart, Gabriela L. Gutierrez, Javier A. Pereira, Hebe del V. Ferreyra, and Carolina A. Marull Page 36 HEALTH STATUS OF GEOFFROY S CAT AT LIHUE CALEL NATIONAL PARK, CENTRAL ARGENTINA Marcela M. Uhart, Javier A. Pereira, Hebe del V. Ferreyra, and Carolina Marull Page 45 3
4 CAPTURE AND IMMOBILIZATION OF WILD CATS AT LIHUÉ CALEL NATIONAL PARK, CENTRAL ARGENTINA Javier A. Pereira (1,3), Marcela M. Uhart (2,3), Hebe del V. Ferreyra (2,3), Carolina Marull (2,3), and Natalia G. Fracassi (1,3) (1) Asociación para la Conservación y el Estudio de la Naturaleza Iberá vo. "B", Buenos Aires (1429), Argentina. (2) Field Veterinary Program, Wildlife Conservation Society Estivariz 197, Puerto Madryn (9120), Chubut, Argentina (3) Gatos del Monte Project INTRODUCTION Pampas cat and Geoffroy s cat are two small wild cat species which co-exist in large areas of Argentina and Southern South America. Major ecology and biology information gaps exist for these small felines. For Pampas cat, no current information is available on its spatial ecology or habitat use because there have been no previous studies on this topic. At the same time, the interactions between this species and the Geoffroy s cat have never been evaluated. These wild cats appear to have similar conservation threats, such as habitat loss and poaching, mainly for control of predation of domestic poultry and livestock. In addition, Pampas cat and Geoffroy s cat were heavily hunted for the international fur trade until the middle of the 1980s and the high volume of this operation (at least 350,000 skins of Geoffroy s cats and 78,000 of Pampas cats between ) could have severely reduced their population numbers. At present, these species are protected in Argentina but due the scarce information available on their natural histories it is impossible to asses the impact of habitat modification on their populations or to develop scientifically sound conservation strategies to ensure their persistence in time. While the IUCN 4
5 listed these cats as low risk species at a global scale (Category 5 for Pampas cat and 4 for Geoffroy s cat), the Argentine Society for the Study of Mammals (SAREM) categorized Pampas cat and Geoffroy s cat as "vulnerable" and "potentially vulnerable" species, respectively. The SAREM criteria probably better represents the current situation of these species in Argentina, because it includes local factors and regional threats in the analysis. During March-April 2002 and May 2003, two capture stages for the project Ecology and conservation of sympatric Pampas cat and Geoffroy s cat in an endemic ecoregion of Central Argentina were conducted in order to capturing and collaring a sample of individuals of the above-mentioned species. The overall project focuses on the ecology of these sympatric felids in a protected area of central Argentina. The objectives of this project s stage were (1) to capture and radiocollar Geoffroy s cats and Pampas cats, and (2) to collect biological samples to analyze their basic physiological parameters (hematology, blood biochemistry, etc.) and to perform parasitological, genetic, and disease studies. STUDY AREA Lihué Calel National Park (37 57 S and W, 9901 hectares) is located in La Pampa province, Central Argentina. This park represents the Monte ecoregion, endemic of this country, deficiently protected (less than 2% of its 158,000 square miles) and listed as Vulnerable by WWF because of seriously damaging effects due to human activities. The protected area consists of flat desert scrub (300 m.a.s.l.) and an isolated set of bare rock hills (590 m.a.s.l.) and is surrounded by an immense plain of desert scrub divided into large cattle ranches. Vegetation is a mosaic of creosote bush or Jarilla (genus Larrea), mixed shrub patches and open areas of grasses and forbs. Twenty-eight mammal species, including four felids (Pampas cat, Geoffroy s cat, Jaguarundi and Puma) inhabit Lihué Calel. These species suffer a high hunting pressure outside the protected area, which represents one of their main 5
6 regional conservation threats. Feral cats and dogs exist near the national park boundaries and they constitute a potential health risk for the native carnivores. MATERIALS AND METHODS Small (82 x 27 x 32 cm) and large (107 x 32 x 32 cm) Tomahawk Live Traps and small (84 x 28 x 28 cm) and large (92 x 34 x 34 cm) ad hoc live traps were used to capture wild cats. All traps consisted of single door openings, which were tripped by a foot treadle. Traps were baited in their rear portions with live domestic pigeons or with a mixture of cow liver and fat with fish oil. Each trap was camouflaged using local elements as forbs, branches and thorns, mainly in the rear portion to protect the bait. Traps were generally set in groups of 2-4 along the roads, where recurrent visual sightings were done or where a lot of spoors or feces were found. Traps were visited 2 or 3 times a day to check for captures and to feed bait pigeons. After a successful capture, trap groups were moved to another place to avoid recapturing the same individual. Each captured animal was anesthetized by the intramuscular injection of Ketamine-Medetomidine and the dosages varied according to each specimens weight and general status. While under anesthesia, cats weight, sex, age (based on body condition and tooth wear) and standard body measurements (total length, tail length, left ear length, left hind foot length, neck circumference and left upper canine length) were recorded and a complete physical exam was practiced. Blood samples were collected for hematological, genetic and disease studies. Additionally, we searched for ecto-parasites and collected fecal samples for endo-parasitological studies. Vital parameters (body temperature and cardiac and respiratory rates) were monitored during cat handling. All healthy individuals were equipped with a radiocollar weighing less than 3 % of the cat s body weight. Following handling, animals were placed in a recovery plastic kennel, 6
7 monitored avery 30 minutes and released when alert and coordinated at the site of capture. Only subadult (minimum mass 2.0 kg) and adult wild cats were collared. RESULTS During the study period, 16 different Geoffroy s cats were captured, 5 in 2002 and 11 in Four (3 males and 1 female), and 10 (1 adult male, 8 adult females, and 1 subadult female) Geoffroy s cats were radiocollared in 2002 and 2003, respectively. One additional male (in 2002) and two additional females (in 2003) were also captured, examined and released without collaring. Although in 2002 only one individual was recaptured (an old, dehydrated male), five females were recaptured in Female OG04 was radiocollared and monitored during both 2002 and Identification, weight and morfometric values of these Geoffroy s cats are reported in Table 1. Table 1. Identification, weight (kg) and measurements (mm) of Geoffroy s cats captured at Lihué Calel. TL = Total length, T = Tail length, HFL = Hind foot length, EL = Ear length, UCL = Upper canine length, NC = Neck circumference ID # Sex Age 1 Weight TL T HFL EL UCL NC OG01 M A 4, ,5 11,6 190 OG02 M A 4, ,0 8,4 195 OG03 M A 3, ,1 13,4 200 OG04 F A 2, ,0 12,3 165 OG05 M A 3, ,5 13,0 185 OG06 F A 2, ,0 9,0 210 OG07 F A 2, ,0 10,1 167 OG04 F A 2, ,0 10,0 158 OG08 F A 3, ,0 9,5 182 OG09 F A 3, ,8 39,0 11,9 181 OG10 M A 3, ,5 11,1 194 OG11 F SA 2, ,3 31,8 11,3 151 OG12 F A 3, ,3 12,0 175 OG13 F A 2, ,3 45,1 11,9 186 OG14 F A 3, ,0 11, SA = Subadult, A = Adult 7
8 Studied individuals ranged in age from subadult to old adults. Except one female (OG11), which was a poor body condition, the rest of the sampled individuals were apparently in good physical condition, and no ectoparasites were found on them. All the captured cats remained relatively calm during the approach and sedation processes, and anesthetics details are given in Table 2. All cats were safely handled and no one was injured. Table 2. Details of Geoffroy s cats sedation on Lihué Calel. ID # Drug Injected Dosage ITA MTB RTC OG01 Ketamine 6,38 mg/kg. 7 min. 41 min. 240 min. Medetomidine 0,06 mg/kg. OG02 Ketamine 5,81 mg/kg. 7 min. 42 min. 120 min. Medetomidine 0,11 mg/kg. OG03 Ketamine 5,12 mg/kg. 3 min. 43 min. 120 min. Medetomidine 0,1 mg/kg. OG04 Ketamine 5,35 mg/kg. 5 min. 44 min. 150 min. Medetomidine 0,125 mg/kg. OG05 Ketamine 6,25 mg/kg. 9 min. 32 min. 120 min. Medetomidine 0,09 mg/kg. OG06 Ketamine 6,52 mg/kg. 5 min. 34 min. 85 min. Medetomidine 0,1 mg/kg. OG07 Ketamine 7,14 mg/kg. 6 min. 37 min. 94 min. Medetomidine 0,12mg/Kg. OG04 Ketamine 6,25 mg/kg. 5 min. 45 min. 63 min. Medetomidine 0,1 mg/kg. OG08 Ketamine 5,62 mg/kg. 5 min. 39 min. 37 min. Medetomidine 0,1 mg/kg. OG09 Ketamine 6,6 mg/kg. 5 min. 35 min. 99 min. Medetomidine 0,11 mg/kg. OG10 Ketamine 5,26 mg/kg. 5 min. 43 min. 83 min. Medetomidine 0,08mg/Kg. OG11 Ketamine 7,5 mg/kg. 5 min. 44 min. 51 min. Medetomidine 0,1 mg/kg. OG12 Ketamine 5,33mg/Kg. 5 min. 35 min. 85 min. Medetomidine 0,1 mg/kg. OG12 Ketamine 7,14mg/Kg. 6 min. 32 min. 97 min. (recapture) Medetomidine 0,125mg/Kg. OG13 Ketamine 6 mg/kg. 5 min. 34 min. 123 min. Medetomidine 0,1 mg/kg. OG14 Ketamine Medetomidine 8,3mg/Kg. 0,09 mg/kg. 12 min. 36 min. 97 min. A IT = Induction time: from drug injection to complete drug effect, B MT = Manipulation time: from complete drug effect until the animal was placed in the recovery kennel, C RT = Recuperation time: from the end of manipulation to the release of the animal 8
9 The lack of Pampas cat trapping success suggests that there is some inherent behavior that makes this species wary of enclosure traps and thus difficult to capture for collaring. Other capture attempts failed as well in other parts of South America. ACKNOWLEDGEMENTS The Whitley Laing Foundation and the Rufford Foundation ( Rufford Small Grant ), the Roger Williams Park Zoo ( Sophie Danforth Conservation Biology Fund ), the Cleveland Metroparks Zoo ( Scott Neotropical Fund ), and Idea Wild funded this stage of the study. We want to thank the Lihué Calel s park rangers (Raul Milne, Miguel Romero, Gisella Muller and Horacio Erasun) for their support and help during the fieldwork. To Dr. Jim Sanderson, who kindly provided valuable equipment (traps and a computer) for this study and to Drs. Andrés Novaro, Susan Walker, Marcelo Pessino, Alejandro Vila and Pablo Perovic for their help. To Dr. Claudio Chehébar (National Parks Administration) for his cooperation and support in obtaining work permits and to Anibal Parera and Mario Beade (Fundación Vida Silvestre Argentina) for the equipment (receiver and antennas) facilitation and their helpful suggestions. 9
10 Manuscript in press - Journal of Mammalogy 87(6) NUMERICAL AND SPATIAL RESPONSES OF GEOFFROY S CAT (Oncifelis geoffroyi) TO A SEVERE DECLINE IN PREY ABUNDANCE IN THE MONTE DESERT, ARGENTINA Javier A. Pereira (1,3), Natalia G. Fracassi (1,3), and Marcela M. Uhart (2,3) (1) Asociación para la Conservación y el Estudio de la Naturaleza Iberá vo. "B", Buenos Aires (1429), Argentina. (2) Field Veterinary Program, Wildlife Conservation Society Estivariz 197, Puerto Madryn (9120), Chubut, Argentina (3) Gatos del Monte Project ABSTRACT We examined the numerical and spatial responses of Geoffroy s cats (Oncifelis geoffroyi) to a strong decline in the abundance of their main prey in central Argentina between April 2002 and November The second year of the study coincided with a severe drought. European hare (Lepus europaeus) density declined from 24.6 ind/km 2 during the pre-drought period to about 2.8 ind/km 2 during drought. Small rodent biomass showed also the lowest level for the study area during the drought of (134.5 g/ha). During the pre-drought and drought periods, three males and one female, and one male and nine females Geoffroy s cats, respectively, were radiotagged and monitored. Home ranges for males of the pre-drought period averaged ha ± SD and that of the single female was 27.3 ha. During the drought period, two Geoffroy s cats abandoned the area and a female with a non-optimal body condition when initially captured was found dead few days later. Four females occupied an average home range of ha ± SD. The home-range size of the single pre-drought female increased by two times after the prey decline. No obvious change in mean daily distance traveled between both periods were observed. Geoffroy s cats predominantly used habitats of dense cover and avoided open habitats during the pre-drought period, but expanded their home range and became more habitat generalist during the drought. Four kittens were recorded during the predrought period, but recruitment apparently did not occur during the drought period. All monitored Geoffroy s cats dispersed and/or died due to starvation after the prey decline. Consequently, Geoffroy s cat density dropped from 2.9 ind/10 km 2 before 10
11 the drought to 0.3 ind/10 km 2 probably due to food scarcity. This is the first study to examine the spatial ecology and density of a small wild cat species under nutritional (energetic) stress in South America. INTRODUCTION Many predator populations are limited by food availability, and their densities fluctuate with periodic changes in prey abundance (Angerbjorn et al. 1999). In general, predator populations respond to changes in prey availability either numerically or functionally. Whereas numerical response refers to absolute changes in the number of individuals by changes in reproductive rates, survival, immigration, or emigration, functional response refers to behavioral changes, such as switching to alternative prey (Murdoch and Oaten 1975; Ward and Krebs 1985; Angerbjorn et al. 1999). Strong variation in food abundance might elicit one or both responses. Models of optimal feeding vs. territory size predict that an animal's energetic needs and the density of available food are important factors influencing home range size (McNab 1963; Mace and Harvey 1983; Schoener 1983). An increase in home-range size of felids with low prey density has been observed on temporal (i.e., Ward and Krebs 1985; Poole 1994; Norbury et al. 1998) or geographical scales (i.e., Edwards et al. 2001; Grigione et al. 2002). In general, when a prey base declines suddenly, home ranges may shift or be abandoned altogether (Norbury et al. 1998; Edwards et al. 2001). In these situations, increases in mortality and rates of emigration are also apparent (Poole 1994; Harper 2004). In arid and semi-arid regions where water is a limiting resource, drought periods can have a strong effect on the abundance and density of small rodents (i.e., Lima et al. 2003; Meserve et al. 2003) and lagomorphs (Myers and Parker 1974; Palomares et al. 2001). Despite the frequency of these changes and their importance in conservation planning, observational studies spanning them are relatively uncommon 11
12 in South America, especially with predator responses over time (but see Jaksic and Simonetti 1987 and Jaksic et al. 1997). Geoffroy s cat (Oncifelis geoffroyi) is a solitary, primarily nocturnal small felid, distributed from southern Brazil and Bolivia throughout southern Patagonia in Argentina and Chile (Ximénez 1975; Nowell and Jackson 1996). Little is known on the ecology of this species (Lucherini et al. 2004), classified as Near threatened (Nowell 2002). It has been described as an opportunistic predator (Canepuccia 1999) feeding mainly upon introduced European hares (Lepus europaeus) and small rodents (Johnson and Franklin 1991; Vuillermoz and Sapoznikow 1998; Novaro et al. 2000). In southern Chile, Johnson and Franklin (1991), in the only representative radiotelemetry study of the species, reported that Geoffroy s cats tend to use habitats with dense vegetation and probably high prey density. Most of the Geoffroy s cats range encompasses arid and semiarid environments (Ximénez 1975). In central Argentina, where the species inhabits mainly shrublands and xeric forests, a severe drought occurred in the summer of This natural disturbance provided us with the opportunity to study the effects of extreme conditions on the abundance of small- and medium-sized herbivores and its consequences for Geoffroy s cat density and spatial behavior. Specifically, we focused on variations in home range size, habitat preference, daily movements, and density of Geoffroy s cats relative to changes in prey availability. MATERIALS AND METHODS STUDY AREA. The study was conducted in Lihue Calel National Park (37 57 S and O, 9,900 ha) and surrounding lands in La Pampa province, Argentina. This area is composed of flat terrain except for a large, isolated set of bare rock hills. The vegetation is characterized by a mosaic of creosote bush flats (Larrea spp., hereafter jarilla scrubland), grasslands mainly bunch grasses (Stipa spp.), and mixed shrub 12
13 patches (e.g., Condalia microphylla, Prosopis flexuosa). There are some ephemeral ponds, but no permanent surface water exists in the study area. Five sigmodontine rodents, two hystricognath rodents (cavies) and the European hare (Lepus europaeus) form the bulk of the Geoffroy s cat's diet in the area (J. Pereira, unpublished data). The plains vizcacha (Lagostomus maximus), a large herbivorous rodent preyed upon by Geoffroy s cat (Branch 1995), disappeared from the region in The pampas fox (Pseudalopex gymnocercus), the pampas cat (Lynchailurus colocolo), the jaguarundi (Herpailurus yaguaroundi) and the puma (Puma concolor) are potential competitors of Geoffroy s cats in the area. Mean daily temperatures are <8ºC in winter and >25ºC in summer. Annual rainfall is 498 mm (SD ± 141, period ), 72% of which (range 63 82%) is concentrated within spring and summer (October-March). The amount of rain from October 2002 through March 2003, however, was markedly lower (148.7 mm) than the seasonal average, resulting in a prolonged drought until November 2003 (data from Lihue Calel weather station). PREY AVAILABILITY. European hares were counted seasonally between the fall (April-May) of 2002 and the spring (October-November) of 2003, along a fixed transect of 15.6 km traversing the main vegetation communities in the study area. Spotlighting counts were conducted by vehicle after sunset, 2-5 times per season. Because dense vegetation obstructed hares visibility, we used the Strip Transect method to estimate density (Burnham et al. 1980). Strip width was fixed as the mean diameter in which hares could be correctly detected times two. The area covered by each sampled transect was 187 ha. A rodent survey was initiated in the winter of 2003 (drought period). The density and biomass of sigmodontine rodents were determined in the jarilla scrubland, the main vegetation type in the study area. Trapping was carried out by installing two 7 x 8-grids of aluminium live traps (Sherman Traps Inc., Florida, USA) spaced 10 m apart. The grids were operated for 5 consecutive nights using rolled oats and peanuts as bait. Captured individuals were identified to species, weighed, individualized (by natural 13
14 marks, color markers or, as the last option, toe-clip), and released at the capture site. Rodent abundance was estimated based on the minimum number of individuals known to be alive (MNKA), and the density was calculated as the ratio between the abundance and the area occupied by the grid. Small rodent biomass was estimated as a product of the density and mean body mass of the individuals captured during this study. Biomass values were then compared with those obtained by S. Heinonen (National Parks Administration, unpublished report) during the winter of 1993, the only previous study conducted in the area during a non-drought period. A new rodent survey was done in winter 2004 (after the drought), following the same protocol as in GEOFFROY S CAT CAPTURES. Trapping was conducted in April 2002 (pre-drought period, trapping effort: 224 trap-nights) and May 2003 (drought period, 440 trap-nights) with Tomahawk live traps baited with domestic pigeons. Captured individuals were immobilized with ketamine and medetomidine administered intramuscularly (average dose: 6 mg/kg and 0.1 mg/kg, respectively). Geoffroy s cats were sexed, weighed, measured, and aged (based on a physical examination and tooth eruption patterns). Individuals >2.0 kg were fitted with radiocollars (Advanced Telemetry Systems, Isanti, Minnesota). During the pre-drought and drought periods, 42-g M1940 radiocollars with internal antenna and 60-g M1950 radiocollars with external antenna, respectively, were used. These radiocollars represented on average 1.1% (range ) and 2.2% (range ) of the cats body weight. Each animal was released at the capture site once it had recovered from anesthesia. The manipulation and care of animals involved during this study followed guidelines approved by the American Society of Mammalogists (Animal Care and Use Committee 1998) and by the Argentine Society for the Study of Mammals. HOME RANGE AND MOVEMENTS. The locations of Geoffroy s cats were obtained by triangulation from the ground (White and Garrott 1990), using a hand-held 5-element Yagi antenna (Wildlife Materials, Carbondale, Illinois) or an H-antenna (Telonics, Mesa, Arizona) and a portable receiver (TR-4, Telonics, Mesa, Arizona). Locations were plotted on a 1:30,000 satellite image of the study area using Universal 14
15 Transverse Mercator (UTM) coordinates. Visual sighting of radiocollared animals georeferenced using a Garmin E-Trex Legend GPS (Garmin International Inc., Olathe, Kansas) were also included in the analysis of home range sizes. Individuals were located 1 to 5 times per week, at daytime and night-time. In most (79%) instances, the distance between the observer and the monitored animals was <260 m (total range: m). Home range size was estimated using the Minimum Convex Polygon (MCP, Mohr 1947) and the Adaptive Kernel (AK, Worton 1989) methods in the CALHOME software package (Kie et al. 1996). The MCP is relatively robust with low sample sizes (Harris et al. 1990) and is the most commonly used technique for estimating the home-range size of cats. We report the 100% MCP to allow comparison with the Johnson and Franklin (1991) study. The AK are not influenced by effects of grid size and can estimate densities of any shape (Seaman and Powell 1996). We calculated the 50% AK (as an area of core utilization) and the 95% AK (as a commonly referenced contour) with a level of smoothing selected by least-squares cross-validation and a grid cell size of 30 m x 30 m. Due to the low number of evaluated individuals and the high variability in the home range sizes, no statistical analysis could be performed on the home range data. Independence of locations was assumed by taking only one location within a 24 h period interval (Swihart and Slade 1985). The minimum number of locations needed to adequately describe home-range size was estimated by plotting home-range sizes against the number of fixes to determine if this parameter reached an asymptote (Harris et al. 1990). Home range overlap were calculated by averaging percentage overlap between pairs of 100% MCP ranges. Daily movements were calculated by measuring the linear distance between consecutive 24-h radiolocations (Rabinowitz 1990; Poole 1994). This parameter was considered as an indicator of the time and effort spent searching for prey (Brand et al. 1976; Ward and Krebs 1985; Poole 1994). Differences between years were evaluated with t-test. 15
16 HABITAT USE. We defined our study area by obtaining the 100% MCP of all independent locations for all Geoffroy s cats. We developed a Geographic Information System of this area (9,572 ha) based on vegetation information obtained from 5 random transects crossing the area and from a LandSat 7 TM satellite image (bands 3, 4 and 5) from January We performed a supervised classification using the maximum likelihood decision rule (Lillesand and Kiefer 1994) and ERDAS IMAGINE 8.2 software. Locations of each Geoffroy s cat were digitized and converted as a layer using ARCVIEW 3.2/Thematic mapper. Habitat use was investigated at one scale of selection, defined as the selection of a home range within a study area (second-order selection, Johnson 1980). We considered distinct vegetation types as different habitat types (jarilla scrubland, mixed scrubland, dense grassland, xeric forest, and others). Each Geoffroy s cat location was assigned one habitat type. A chi-square goodness of fit test was used to determine if the observed frequencies of habitat use differed significantly from expected frequencies based on habitat availability (Neu et al. 1974; McClean et al. 1998). The null hypothesis tested was that usage occurs in proportion to availability considering all habitats simultaneously (Neu et al. 1974). Following a chi-square test, we used a Bonferroni correction of the z-statistic (α = 0.10) to maintain an experiment-wise error rate (Miller 1981) and to create a normal approximation of the confidence intervals to determine which habitat types were either selected, avoided, or neither. This is the most common test of habitat use (McClean et al. 1998) and has been widely used in felid studies. Notwithstanding this, we know that we violate the assumption of independence of observations within and among individuals (cf. Alldredge and Ratti 1986). However, the small sample of monitored individuals precluded their use as experimental units (Aebischer et al. 1993). Because few locations were obtained on many animals, data were pooled for all males during the predrought period and for all females (except female OG04) during the drought period in order to compare 16
17 habitat preference between both periods. Habitat preference of female OG04 was analyzed separately because we obtained a large number of locations for her. GEOFFROY S CAT DENSITY. Minimum Geoffroy s cat density was calculated seasonally between fall 2002 and spring 2003 based upon radiocollared cats in addition to the number of unmarked individuals using the 9,572-ha study area, as estimated by visual sightings (Poole 1994; Franklin et al. 1999; Palomares et al. 2001). Of the unmarked individuals, the only ones considered were those that could be positively identified as different individuals, based on sighting location, body size, color patterns, or other characteristics. As a result, density estimates were conservative and should be considered minimum values (Poole 1994). RESULTS PREY ABUNDANCE. Hare density was high between fall and spring 2002 (pre-drought), peaking at 24.6 ind/km 2 in winter. Density began to decline progressively until fall 2003, when it dropped to 2.8 ind/km 2 and remained low (<3.5 ind/km 2 ) through spring 2003 (Fig. 1). This represented a decline in hare density of >88% in 9 months. Rodent biomass estimated during the drought period was 66% lower (134.5 ± 89.2 g/ha, Fig. 2) than that estimated for a non-drought period (S. Heinonen, National Parks Administration, unpublished report). Whereas seven rodent species were captured during the non-drought period, only two species were captured during the drought. However, because of the long time period between the two rodent surveys, causes other than the drought (e.g., succession) may have been responsible for the observed species loss. On the other hand, the recovery of the rodent community after a normal rainy season was notable; the average biomass increased to ± 35.5 g/ha in winter 2004 (Fig. 2). 17
18 FIG. 1. Density (Mean ± SD) of European hares (Lepus europaeus) by season (FL = Fall, WN = Winter, SP = Spring, SM = Summer) in Lihue Calel National Park, Argentina. Hares density (Ind / km 2 ) FL02 WN02 SP02 SM03 FL03 WN03 SP03 Season FIG. 2. Biomass (Mean ± SD) of small rodents in different winters (WN) in Lihue Calel National Park, Argentina. 600 Rodent biomass (g / ha) WN93 WN03 WN04 Season Although the density of cavies (Microcavia australis and Galea musteloides) was not evaluated during the study, visual sightings recorded while travelling the study area suggested that their abundance also decreased considerably during the drought. CAT CAPTURES AND RADIOTRACKING. During the pre-drought period (2002) and the drought period (2003), four (three males and one female, all adults), and ten (one adult male, eight adult females and one subadult female) Geoffroy s cats were captured and radiocollared. One additional male (in 2002) and two additional females (in 2003) were also captured, examined and released without collaring due to poor 18
19 physical condition or equipment restrictions. Although in 2002 only one individual was recaptured (an old, dehydrated male), five females were recaptured in Female OG04 was captured and monitored during both the pre-drought and drought periods. Adult males outweighed adult females (3.96 ± 0.53 vs 2.83 ± 0.28 kg, t-test, p < 0.05) and were larger (body length; ± 39.3 vs ± 29.9 mm, t-test, p < 0.05). The four pre-drought individuals were monitored an average days (range ). The tracking period ended for the three males due to long-distance dispersal (>10 km), and for female OG04 because of transmitter failure. During the drought period, two Geoffroy s cats abandoned the area shortly after captured (the only radiotagged male dispersed 138 km) and the subadult female with a non-optimal body condition when initially captured was found dead; she had lost >15% of body weight in this time. Three females moved out of the home range that they had maintained for 39, 42, and 43 days and were found dead 1-2 days later at 11.3, 5.8 and 9.6 km, respectively, from capture points. Another three females died within their home ranges after 16, 17, and 54 days of being collared. Necropsies were conducted on five of these cats, and the deaths were attributed to starvation. The remaining female was killed by another felid (probably a jaguarundi [Herpailurus yaguaroundi] or another Geoffroy s cat, based on feces found at the death site and teeth marks found on the dead female s neck); also, she showed signs of starvation. The radio signal of female OG04 ceased after 1 month of radiocollaring for the second time and we were unable to find her thereafter. HOME RANGE AND MOVEMENTS. An asymptotic home range was not obtained for 3 females with <12 locations, all of them during the drought period, and they were not considered reliable estimates. Thus, these animals were not included in home-range analysis. 19
20 During the pre-drought period, the average home range (100% MCP, mean ± SD) of males was ± ha whereas the home-range size of female OG04 was 27.3 ha (Table 1). The areas used by males OG01 and OG02 were contiguous and non-overlapping. After the prey decline, the mean home range size of females was ± ha (Table 1). The home range size of female OG04 was two times larger than that during the pre-drought period. Three cases of home range overlap were found during the drought period between females OG07-OG09 (17.3%), OG13-OG14 (15.7%), and OG04-OG07 (1.4%). TABLE 1. Home range of Geoffroy s cats radiocollared during the pre-drought (year 2002) and drought (2003) periods in Lihue Calel National Park, Argentina. Total home ranges were estimated as the 100% minumum convex polygon (MCP) and the 95% adaptive kernel (AK) and core areas as the 50% adaptive kernel. Cat Id Period tracked No. of fixes Asymptote reached Home range size (in hectares) 100%MCP 95%AK 50%AK M OG01 Pre-drought 02Apr 18Jun 24 Yes M OG02 06Apr 09Jul 18 Yes M OG03 07Apr 19Aug 25 Yes F OG04 07Apr 12Jan 70 Yes F OG07 Drought 11May 20Jun 26 Yes F OG04 13May 13Jun 18 Yes F OG08 15May 28Jun 8 No F OG09 16May 28Jun 23 Yes F OG12 23May 17Jul 28 Yes F OG13 25May 10Jun 10 No F OG14 25May 11Jun 11 No Although 95% AK home range estimates were in average 1.5 times larger than 100% MCP estimates (Table 1), the inter-annual trend were in general similar with both methods. The average core area (50% AK, mean ± SD) of males of the pre-drought period was 70.0 ± 77.8 ha (range ), whereas those of the females of the drought period was 75.1 ± 82.9 ha (range ; Table 1). These values indicate how little of the home range (22.9 ± 3.6% and 16.4 ± 11.8% of the 95% AK during the pre-drought and the drought periods) was used intensively. No cases of core area overlap between individuals were found. 20
21 Geoffroy s cats were located on consecutive days 38 times during the pre-drought period and 52 times during the drought period (Table 2). Non-significant differences (t = -0.44, d.f. = 65, P = 0.66) were found in mean daily distance traveled between males of the pre-drought period and females of the drought period. All females with a body weight of >3 kg showed average daily movements of more than 1 km. If only these three females with body weight similar to those of the pre-drought males are considered, daily distance traveled showed again non-significant differences (t = 1.79, d.f. = 40, P = 0.08). Female OG04 exhibited greater daily movements during the drought (Table 2), but differences between years were nonsignificant (t = 1.63, d.f. = 32, P = 0.11). TABLE 2. Average daily distance travelled (in meters) by Geoffroy s cats during the pre-drought and drought periods in Lihue Calel National Park, Argentina. Pre-drought Drought n Mean ± SD Range n Mean ± SD Range Males ± Females ± Female OG ± ± HABITAT USE. The most abundant habitat type in the study area was the jarilla scrubland (59.2%), followed by dense grassland (23.8%), mixed scrubland (8.8%), xeric forest (0.7%) and others (7.7%, including rocky terrain, open grassland and bare soil). During both pre-drought and drought periods, Geoffroy s cats (except female OG04) mainly used jarilla scrubland, dense grasslands, and mixed scrubland (Table 3). However, the former habitat type was used less than expected by chance in both periods. Open habitats (grouped under "others") were used less than expected by chance during the predrought period (Table 3). Female OG04 used predominantly xeric forest during the entire study period. This habitat type was used more than expected and the jarilla scrubland was used less than expected in both pre- and drought periods (Table 4). The frequency of mixed scrubland use changed between pre- 21
22 drought and drought periods, being avoided during the former and used as expected by chance during the later (Table 4). TABLE 3. Chi-square test and Bonferroni Z-test 95% Confidence Intervals of habitat preference by Geoffroy s cats (except female OG04) during the pre-drought and drought periods in Lihue Calel, Argentina. Habitat type Availability (prop.) Observed locations (prop.) Bonferroni 95% Confidence Intervals Selection Pre-drought period Jarilla scrubland p Avoid Mixed scrubland p Grassland p Xeric forest p Other p Avoid χ² Statistic d.f. 4 P Drought period Jarilla scrubland p Avoid Mixed scrubland p Grassland p Xeric forest p Other p χ² Statistic d.f. 4 P <0.001 TABLE 4. Chi-square test and Bonferroni Z-test 95% Confidence Intervals of habitat preference by Geoffroy s cat OG04 during the pre-drought and drought periods in Lihue Calel, Argentina. Habitat type Availability (prop.) Observed locations (prop.) Bonferroni 95% Confidence Intervals Selection Pre-drought period Jarilla scrubland p Avoid Mixed scrubland p Avoid Grassland p Xeric forest p Prefer Other p χ² Statistic d.f. 4 P <0.001 Drought period Jarilla scrubland p Avoid Mixed scrubland p Grassland p Xeric forest p Prefer Other p χ² Statistic d.f. 4 P <
23 GEOFFROY S CAT DENSITY. At the beginning of the study Geoffroy s cat density was estimated to be 2.6 ind/10 km 2. Numbers remained high (>2.3 ind/10 km 2 ) until the summer 2003 and declined progressively to about 0.3 ind/10 km 2, during periods of food scarcity (Fig. 3). Despite extensive search, no Geoffroy s cat kittens were found during the drought period, while during the pre-drought period at least four kittens were recorded. Similarly, we did not detect follicles in ovaries or pregnancies in necropsied females during the drought. FIG. 3. Density of Geoffroy s cats by season (FL = Fall, WN = Winter, SP = Spring, SM = Summer) in Lihue Calel National Park, Argentina. Cat density (Ind / 10 km 2 ) FL02 WN02 SP02 SM03 FL03 WN03 SP03 Season DISCUSSION Our results indicate that before the drought, male Geoffroy s cats were sedentary, occupying home ranges up to 4 months before abandoning them. Similar behavior was reported by Johnson and Franklin (1991) in southern Chile, where male Geoffroy s cats dispersed >25 km from a home range that they had maintained for 3-5 months. The home range sizes of all collared cats in the present study (except for female OG12) were markedly smaller than those reported by Johnson and Franklin (1991) for both sexes. Body weights of males and females in the present study were on average 18 and 33% lighter than those of 23
24 males and females in Johnson and Franklin (1991) study. This could explain the differences in home range sizes between sites since in general this parameter scales allometrically with body size (McNab 1963; Harestad and Bunnell 1979; Mace and Harvey 1983; but see also Grigione et al. 2002). The estimated prey density reported by Johnson and Franklin (1994) allow us to reject the prey availability hypothesis to explain home range size differences between sites, given that actually prey density at the Chilean site (mean hare density = 86.6 inv/km 2 and mean rodent density = 62.2 ind/ha) was much greater than that at our study area. In Lihue Calel, Bonaventura et al. (1998) found a positive correlation between small rodent biomass and vegetation complexity, suggesting that vegetation plays an important role in structuring rodent communities in this area. Although quantitative data are lacking, severe drought in our study area resulted in the vegetation becoming markedly more sparse and insects less abundant (personal observations), and these changes may have been the driving force behind the decline in rodent biomass. The reduction in the herbaceous layer probably also triggered the decline of European hare density as they are herbivorous (Campos et al. 2001). A similar situation was described during drought periods in Lihue Calel for the vizcacha, a native herbivore of similar body weight (Branch et al. 1994). It is also possible that some hares were actually more vulnerable to predators due to the lack of foliage cover (cf. Brown 1988). The increase in capture success of Geoffroy s cats during the drought period may be due to felines becoming more inclined to investigate traps during periods of low food availability. Poor body condition of captured and necropsied individuals during the drought period further supports the importance of food scarcity. Bailey (1981) and Ward and Krebs (1985) reported that bobcats (Lynx rufus) and lynx (Lynx canadensis) that defended territories during periods of prey abundance became nomadic when prey declined. These authors suggest that if prey density is unpredictable or very low, it would be adaptive for these cats to become transient and search out widely separated concentrations of prey. In the present 24
25 study, during the period of prey scarcity, two Geoffroy s cats abandoned the area a few days after being radiocollared, but it is unclear if this was a behavioral response to declining prey abundance or if they were transient animals without stable territories. In one month, four female cats monitored during the drought occupied an average home range greater than those occupied by males during two to four months in the pre-drought period. Previous studies on small felids showed that males home ranges are substantially larger than those of females (i.e., Konecny 1990; Grassman 2000). For Geoffroy s cats, Johnson and Franklin (1991) reported a similar pattern, with adult males occupying home ranges more than twice as large as those of adult females. Based on these two trends, it follows that female Geoffroy s cats may have responded to declining prey abundance by expanding their home ranges, because they exceeded the home range size of males estimated for the pre-drought period. In support of this idea, female OG04 doubled the size of her home range between the pre-drought and drought periods. The movement patterns of predators have been observed to increase if prey abundance declines or prey becomes less detectable (Knowles 1985; Ward and Krebs 1985; Sunquist and Sunquist 1989). In agreement with these observations, the daily distance traveled by female OG04 increased once prey declined (non significant differences may be product of the small sample size), and those of females monitored during the drought period were greater than those of males monitored during the pre-drought period. These results suggest that Geoffroy s cats increased the time and effort spent in search for prey to fulfill their energetic needs. Despite drought-related environmental changes, little variations in habitat use were found between predrought and drought periods. However, comparisons between both periods should be considered carefully because they involve different individuals. Previous studies conducted on small felids (i.e., Dunstone et al. 2002; Harper 2004) have reported interindividual differences in habitat use. Johnson and Franklin (1991) reported that Geoffroy s cats predominantly used areas of dense cover because they provide higher prey availability and protective cover. Our results support this findings during the pre-drought period, 25
26 when Geoffroy s cats used habitats of dense cover (such as xeric forest and dense grassland) and avoided open habitats. When prey species became scarcer, Geoffroy s cats responded by expanding their home range to increase opportunities for encountering prey. As a result, habitats avoided during the pre-drought period such as mixed scrubland and open habitats (i.e., rocky terrain, open grassland) changed their frequency of use, being used in proportion to availability during the drought period. Six radiocollared Geoffroy s cats died due starvation during the drought period and these effect occurred with a hare density of <3.5 ind/km². A critical level for rodent abundance could not be determined due to a lack of rodent surveys during the decline period. As a result, Geoffroy s cat density dropped from 2.3 ind/10 km² to 0.4 ind/10 km² between the summer and winter 2003, with a mortality peak between June and July. A similarly elevated mortality rate within a 1-2 month period was reported by Edwards et al. (2001) for feral cats (Felis catus), during a period of prey scarcity in a semiarid area of Australia. Only a few individuals were present in the study area by the spring of 2003, but whether they were the surviving uncollared animals of the study population or immigrants is unknown. The absence of kitten recruitment for Geoffroy s cats during the prey decline is consistent with that observed for other felid species (Nellis et al. 1972; Brand et al. 1976; Knick 1990; Mowat 1993; Poole 1994). Lack of body fat and emaciation were common findings for all dead animals and it is well-known that undernutrition negatively affects sexual activity (i.e., Gill and Rissmann 1997; Schillo 1992; Wade et al. 1996). Body condition can also affect animal s immunity, rendering them more susceptible to opportunistic pathogens (Ullrey 1993; Lloyd 1995; Hulsewe et al. 1999). However, no evidence of clinical infectious disease was found in the necropsied cats, though parasite loads were very high in most individuals (Beldoménico et al., in press). We did not find additional evidence of disease-related mortality, such as observation of sick animals or epidemic-type simultaneous deaths. Based on the observed decrease in the abundance of preferred prey species, the exploitation of an alternative food source (prey-switching) by Geoffroy s cats might have been expected. This kind of 26
27 behavioral response during prey scarcity has been reported in other felids such as the puma (Pessino et al. 2002) and the Iberian lynx (Lynx pardinus-beltran and Delibes 1991). Johnson and Franklin (1991) and Canepuccia (1999) also reported prey-switching by Geoffroy s cat following seasonal changes in prey abundance. In Lihue Calel, potential alternative prey for Geoffroy s cat, such as armadillos (Zaedyus pichyi and Chaetophractus villosus), the mara (Dolichotis patagonum) or the elegant crested-tinamou (Eudromia elegans), were relatively uncommon (Lihue Calel National Park Conservation Value Species Register, Period ; and personal observations) and, although they were not monitored during the study period, their abundance was likely lower during the drought. Other vertebrate taxa, such as amphibians and reptiles, constitute a seasonal food resource virtually unavailable during the colder winter months. As a result, potential alternative prey may have been insufficient at a local scale to allow Geoffroy s cats to fulfill their energetic requirements, and that may have contributed to the increased mortality and emigration. Before its extinction from Lihue Calel, the vizcacha may have been important prey for Geoffroy s cats (Branch 1995). During events of decline in small rodents and hares, the vizcachas could have constituted a key resource for Geoffroy s cat survival, as they are large rodents which live in fixed communal burrow systems, thus providing a spatially predictable resource for predators (Branch 1995). On the other hand, replacement of native by introduced prey in predator diets appears to be widespread in southern South America and the European hare has become an important component of the Geoffroy's cat diet (Novaro et al. 2000). More research is required to understand the dynamics of interactions between this predator and its native and introduced prey species and how climate affects this chain of trophic interactions. ACKNOWLEDGEMENTS 27
28 We thank H. Ferreyra, M. Romero, G. Müller, F. Gallego, C. Rozzi Giménez, A. Hurtado, P. Gramuglia, D. Muñoz, V. Coronel, C. Marull, P. Teta, P. Rossio, J. de Estrada, D. Ugalde, and J. Gato for their dedicated fieldwork; R. Milne, P. Erasun, C. Toledo, A. Iriarte, G. Aprile, and D. Varela for their logistical support and field assistance; J. Sanderson, A. Novaro, S. Walker, P. Perovic, D. Villarreal, A. Parera, M. Beade, K. Schiaffino, A. Vila, L. Maffei, M. Pessino, and A. Sosa for the equipment and assistance provided; S. Heinonen for supplying her unpublished information; and C. Figueroa for the English translation. D. Villarreal, A. Noss, S. Walker, J. Sanderson, John Yunger, and anonymous reviewers provided constructive comments. This study was supported by the Asociación para la Conservación y el Estudio de la Naturaleza, the WCS Field Veterinary Program, the Cleveland Metroparks Zoo (Scott Neotropical Fund), the Roger Williams Park Zoo and Rhode Island Zoological Society (Sophie Danforth Conservation Biology Fund), the Rufford Foundation and Whitley Laing Foundation (Rufford Small Grant) and Idea Wild. LITERATURE CITED AEBISCHER, N. J., P. A. ROBERTSON, AND R. E. KENWARD Compositional analysis of habitat use from animal radio-tracking data. Ecology 74: ALLDREDGE, J. R., AND J. T. RATTI Comparison of some statistical techniques for analysis of resource selection. Journal of Wildlife Management 50: ANGERBJORN, A., M. TANNERFELDT, AND S. ERLINGE Predator-prey relationship, arctic foxes and lemmings. Journal of Animal Ecology 68: ANIMAL CARE AND USE COMMITTEE Guidelines for the capture, handling and care of mammals as approved by the American Society of Mammalogists. Journal of Mammalogy 79:
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36 Acta Parasitologica (2005) 50: HELMINTHS OF GEOFFROY S CAT, Oncifelis geoffroyi (CARNIVORA, FELIDAE) FROM THE MONTE DESERT, CENTRAL ARGENTINA Pablo M. Beldomenico (1,2), John M. Kinsella (2), Marcela M. Uhart (3,5), Gabriela L. Gutierrez (1), Javier A. Pereira (4,5), Hebe del V. Ferreyra (3,5) and Carolina A. Marull (3,5) (1) Facultad de Ciencias Veterinarias, Universidad Nacional del Litoral, RP Kreder 2805 (3080) Esperanza, Argentina. (2) HelmWest Laboratory, 2108 Hilda Avenue, Missoula, MT 59801, U.S.A. (3) Field Veterinary Program Wildlife Conservation Society, Estivariz 197, Puerto Madryn (9120), Argentina (4) Asociación para la Conservación y el Estudio de la Naturaleza, Iberá B, Buenos Aires, Argentina (5) Gatos del Monte Project - ABSTRACT Gastrointestinal parasites were collected from 7 free-ranging Geoffroy s cats (Oncifelis geoffroyi) from Lihué Calel National Park, Argentina. Also, fecal samples were analyzed from these animals and 3 other sympatric ones. The helminths were identified as Vigosospirura potekhina, Didelphonema longispiculata, Pterygodermatites cahirensis, Trichuris campanula, Ancylostoma tubaeforme, Toxocara cati, and Taenia sp. Fecal analysis revealed the presence of eggs of Capillaria sp. and an unidentified anoplocephalid tapeworm, and coccidian oocysts. The findings of V. potekhina, D. longispiculata, P. cahirensis, and T. campanula represent first records of these species in O. geoffroyi. Further, the former three had never been reported in South America. Stefañski Information on the helminths of Neotropical felids is scarce. Although 10 of the 12 existing wild felids of the Americas can be found in Argentina (Nowell and Jackson 1996), there are few reports of these carnivore parasites in that country. One of the most widespread species in southern South America is the 36
37 Geoffroy s cat, Oncifelis geoffroyi (d Orbigny et Gervais, 1844), which can be found from southern Brazil and Bolivia throughout southern Patagonia in Argentina and Chile. Although this species occurs in a wide variety of habitat types, most of this cat s range encompasses arid and semiarid environments (Gomes de Oliveira 1994, Nowell and Jackson 1996). Like most other small wild cats of the region, the biology of this species is poorly known. Its helminth fauna is one of the least explored topics. In central Argentina, the Geoffroy s cat population of Lihué Calel National Park (LCNP; S W, 9900 ha) is being studied by the Gatos del Monte project. This protected area, representing the Monte Ecoregion, is composed of flat terrain except for a large, isolated set of bare rock hills (590 m a.s.l.). The vegetation is characterized by a mosaic of creosote bush flats (Larrea sp.), grasslands (mainly bunch grasses of Stipa spp.), and mixed shrub patches (Administratión de Parques Nacionales, Buenos Aires, 1983). A severe drought occurred in 2003 in central Argentina. In some places such as the LCNP, this was the most severe drought since As a result, six radio-collared adult female Geoffroy s cats were found dead between May-June Necropsies and parasite collection were conducted. Death was tentatively attributed to starvation, which was later confirmed by histopathology. Additionally, in July 2004, an adult male cat was shot by a farmer near the park s boundaries and it was necropsied to collect parasites. A fecal sample had been previously taken from each dead cat and from three other sympatric radio-collared cats. By previously described methods (Beldomenico et al. 2003), each section of the gastrointestinal tract was searched for metazoan parasites, and fecal samples were preserved in 3.5% formalin saline until analyzed at the laboratory by a sedimentation-flotation technique. Five of each specific adult parasites or eggs were measured, and the measurements were reported as arithmetic mean and standard deviation (SD). When fewer specimens were available for measuring, the range or the unique value were reported. 37
38 Voucher specimens were deposited at the Colección de Parásitos de Vertebrados Silvestres of Universidad Nacional del Litoral, Esperanza, Argentina (Acc. no: LP00016-LP00035). Our findings are summarized in Table 1. Fig. 1. Parasites recovered from necropsied Geoffroy s cats from Lihué Calel National Park, Argentina: A cestode egg, B oral extremity of Vigisospirura potekhina, C oral extremity of Didelphonema longispiculata, D vulvar region of Trichuris campanula, E oral extremity of Pterygodermatites cahirensis, F caudal extremity of a male Pterygodermatites cahirensis. Scale bars = 25 µm 38
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