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Cattle depredation by puma (Puma concolor) and jaguar (Panthera onca) in central-western Brazil
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Cattle depredation by puma (Puma concolor) and jaguar (Panthera onca) in central-western Brazil Francesca B.L. Palmeiraa,*, Peter G. Crawshaw Jr.b, Claudio M. Haddadc, Katia Maria P.M.B. Ferrazd, Luciano M. Verdadee a
Research Department, Reserva Brasil, Av. Dr. Silva Melo, n. 520, apt. 606, Jardim Taquaral, CEP 04675-010 Sa˜o Paulo, SP, Brazil Pantanal Matogrossense National Park, IBAMA, R. Firmo de Matos, n. 479, CEP 79331-050 Corumba´, MS, Brazil c Animal Sciences Department, ‘‘Luiz de Queiroz’’ College of Agriculture (ESALQ), University of Sa˜o Paulo (USP), Av. Padua Dias, n. 11, CEP 13418-900 Piracicaba, SP, Brazil d Quantitative Methods Lab, Forest Science Department, ‘‘Luiz de Queiroz’’ College of Agriculture (ESALQ), University of Sa˜o Paulo (USP), Av. Padua Dias, n. 11, CEP 13418-900 Piracicaba, SP, Brazil e Animal Ecology Lab, Biological Science Department, ‘‘Luiz de Queiroz’’ College of Agriculture (ESALQ), University of Sa˜o Paulo (USP), Av. Padua Dias, n. 11, CEP 13418-900 Piracicaba, SP, Brazil b
A R T I C L E I N F O
A B S T R A C T
Article history:
In this study, data on cattle depredation by puma (Puma concolor) and jaguar (Panthera onca)
Received 2 November 2006
were recorded for six years (1998–2003) in a cattle ranch in central-western Brazil. Depreda-
Received in revised form
tion represented 18.9% of the overall cattle mortality, being predominant on calves. In bio-
16 September 2007
mass, kills represented 0.4% (63.8 kg/km2) of the ranch’s annual stock. In economic loss,
Accepted 16 September 2007
kills represented 0.3% of the cattle stock value. Depredation was mainly associated with
Available online 30 October 2007
cattle’s age class and location along with the time of birth of calves. The proportion of pastures next to forest with depredation (n = 33, 48.5%) was not distinguished to the proportion
Keywords:
of pastures not bordering forest with depredation (n = 35, 51.5%). However, the proportion
Cattle breeding season management
of pastures next to forest with depredation represented 54% (n = 33) of the 61 total pastures
Cattle production
that were at least partially surrounded by forest patches or riparian forests that comprised
Human–wildlife conflict
eight continuum blocks of forest fragments of different sizes in the ranch and adjacent
Landscape
areas. No kills occurred in the central portion (main house) of the farm, close to the head-
Livestock depredation
quarters where the pastures not bordering forest. The distances of the kills in relation to
Wild felids conservation
areas of native forest was 1317.48 ± 941.03 m. In order to reduce depredation, calves should be kept as far as possible from forest areas and concentrated cattle breeding and calving seasons should be encouraged. Ó 2007 Elsevier Ltd. All rights reserved.
1.
Introduction
Domestic livestock lack virtually all of their ancestor’s antipredator behaviors, and represent a relatively easily killed prey when compared to wild prey species of similar size
(Linnell et al., 1999). Recently, most predator species have attacked the domestic livestock across the world. Livestock depredation has been caused by lions (Panthera leo), cheetahs (Acinonyx jubatus), spotted hyenas (Crocuta crocuta), leopards (Panthera pardus), African wild dogs (Lycaon pictus) on ranches
* Corresponding author: Tel./fax: +55 11 56855168. E-mail addresses: [email protected] (F.B.L. Palmeira), [email protected] (P.G. Crawshaw Jr.), [email protected] (C.M. Haddad), [email protected] (K.M.P.M.B. Ferraz), [email protected] (L.M. Verdade). 0006-3207/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2007.09.015
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in Kenya (Patterson et al., 2004; Woodroffe et al., 2005; Kolowski and Holekamp, 2006), black-backed jackals (Canis mesomelas) in Tanzania (Holmern et al., 2007), tigers (Panthera tigris), Himalayan black bears (Ursus thibettanus), dholes (Cuon alpinus) in Bhutan (Wang and Mcdonald, 2006), and snow leopards (Panthera uncia) in Nepal (Oli et al., 1994) and in Ladakh, Tibet and Mongolia (Jackson and Wangchuk, 2001). Livestock depredation by jaguars (Panthera onca) and pumas (Puma concolor) has been recorded in Belize (Rabinowitz, 1986), in Venezuela (Hoogesteijn et al., 1993; Polisar et al., 2003), in Brazil (Mazzolli et al., 2002; Conforti and Azevedo, 2003; Zimmermann et al., 2005; Michalski et al., 2006; Palmeira and Barrella, 2007) and in other neotropical countries. Cattle ranching has increased deforestation disproportionately in the Brazilian Cerrado, replacing natural areas with extensive pastures. These are interspersed with remaining fragments of forest, required as legal reserves. As a consequence, pumas and jaguars have to coexist with domestic cattle. However, landowners usually use cattle depredation as a reason to justify persecution of large cats (Quigley and Crawshaw, 1992; Silva, 1994), which are threatened in several regions in the Southern and central-western areas of Brazil (Quigley et al., 1988; Fonseca et al., 1994). This control has become the main reason for the demise of large felids, more so in areas not legally protected (Jackson and Nowell, 1996; Nowell and Jackson, 1996; Linnell et al., 1999). The Cerrado was considered by Sanderson et al. (1999) as one of the priority areas for jaguar study and conservation, due to the lack of knowledge on the species’ geographic range and population status. Despite this intense conflict, very few data have shown any significant economic loss resulting from cattle depredation (Mazzolli et al., 2002; Michalski et al., 2006; Palmeira and Barrella, 2007). Quantifying cattle depredation is fundamental to allow the implementation of management actions so as to minimize losses and to guarantee the conservation of these cats. In this context, the main objectives of this study were to determine: (1) the actual proportion of cattle killed by large felids; (2) the age distribution of the cattle killed; (3) the temporal distribution of depredation to throughout the year; and (4) the spatial distribution of cattle depredation through the landscape mosaic.
2.
Methods
2.1.
Study area
This study was carried out at the Fazenda Ouro Branco, located in the municipality of Bono´polis in Northern Goia´s, in central-western Brazil (Fig. 1). Currently, soybean (Glycine max) and cattle (approximately 90,000 heads) are the main economic activities in the region. Bono´polis is located in a transitional area between the Cerrado and the Amazon Forest biomes in the Araguaia river basin (Dinerstein et al., 1995; IBGE, 2002). The climate is hot, semi-humid, with four months of the year (May–August) with a distinct dry season. The average annual precipitation of 1874 mm occurs mostly from September through April. The mean annual temperature is 25 °C and the average altitude is 290 m a.s.l. (IBGE, 2002). The vegetation mosaic is characterized by agriculture and permanent pastures, with gallery for-
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ests, dense deciduous forest and bush savanna (Miranda and Guimara˜es, 2002). The ranch is separated in six ‘‘fields’’ or management areas identified as Sede (SD), Colina Verde (CV), Liberdade (LB), Zequinha (ZQ), Xodo´ (XD) and Madeireira (MD) which each contain an average of 36.3 ± 11.6 individual pastures. Overall, cattle density in the fields was an average of 1.2 ± 0.4 individuals/ha. The predominant cattle breeds were Nelore, Fleckvieh and a mix of the two. The average annual stock of 12 538 ± 2679 cattle represents approximately 14% of the total heads in the municipality. The six fields at the ranch were designed to decentralize and facilitate the management of cattle, in terms of number of pastures, number of the animals and total area of pastures (ha) (Table 1). The number of pastures in each filed is dynamic and one pasture can be divided in two or vice versa when it is necessary to the cattle management. In general, the ranch has a total area of 20,000 ha (200 km2), approximately 80% of disturbed areas (pastures, dams and roads) and at least 20% of permanent protected areas (forest patches and riparian forests called ‘‘Cerrada˜o’’). Approximately 28% (n = 61) of the 218 total pastures in the study area were at least 25% of the perimeter surrounded by forest patches or riparian forests that comprised eight fragments of different sizes in the ranch and adjacent areas. There is not a cutoff employed in terms of the size of forest fragments. The natural non-forested areas (Cerrado) are severely treated and most natural areas were replaced to cultivate Brachiara and other exotic pastures.
2.2.
Cattle depredation
At the ranch, artificial inseminations, birth periods, stamping and vaccinations all cattle were kept in the central portion of each field. The spatial and temporal sampling effort allocated by ranch-hands for monitoring the herds at the pastures assumed a uniform sampling effort across all fields and seasons. The ranch-hands were being sampled the herds, cattle births and cattle mortality caused by depredation or other causes. Generally, ranch-hands rode horses eight hours per day during six days per week (from Monday to Saturday). The effort distributed across seasons and fields was 2.0 km/ h sampled by two ranch-hands, equivalent to 16 km covered per day or eight pastures checked per day in each field. Considering that each field has a different size (ha) the average monthly of the sampling effort was 5.8 ± 2 return visits to each pasture by each field. It should control the potential bias to differences in sampling effort allocated by ranch-hands for monitoring the herds at the pastures in each field. Bovines were grouped into five categories: calves (0–12 months old), heifers and ‘‘garrotes’’ (12–24 months old), and cows, oxen, and bulls (older than 24 months). Data on cattle depredation were recorded by ranch-hands for each field, from January 1998 to December 2003 and information on mortality due to all causes was recorded through December 2002. The cattle mortality was determined in the field by experienced ranch-hands through of peculiar characteristics of bovine carcasses (e.g., diseases, malnutrition, ingestion of toxic plants, lightning, malpractice during vaccination, diastolic parturition, snake bites and depredation by large cats). Data about precipitation (mm), cattle stock fluctuation and mone-
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Fig. 1 – Location of Fazenda Ouro Branco, Northern Goia´s, central-western Brazil, with the six fields called ‘‘Sede’’ (SD), ‘‘Colina Verde’’ (CV), ‘‘Liberdade’’ (LB), ‘‘Zequinha’’ (ZQ), ‘‘Xodo´’’ (XD) and ‘‘Madeireira’’ (MD).
Table 1 – Number of pastures, number of bovines and total area of pastures (ha) by field in March 2003 Field
Number of pastures (n)
Total area of pastures (ha)
Number of bovines (n)
CV MD LB ZQ XD SD
39 21 26 54 39 39
2458 1587 1669 3446 2420 2550
2800 1587 1650 2480 1188 1900
Total
218
14,130
11,605
36.3 ± 11.6
2355 ± 678.7
1934.1 ± 3456.6
Mean ± SD
tary values were taken from the Monthly Cattle Breeder Bulletin, Fazenda Ouro Branco, from January 1998 to December 2003. The monetary value in Brazilian currency (‘‘Reais’’) of
each bovine killed was converted into American dollars (US$ 1 = R$ 3.15, as rated in the market in July 2004) to estimate the economic loss resulting from predatory killings.
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The bottom notes records of the Bulletins were also used to estimate the biomass (kg) and economic value of each bovine category. To estimate total bovine biomass (kg/km2), the individual average body mass was multiplied by the total number of animals and divided by the area (km2), as suggested by Schaller (1983), Crawshaw and Quigley (1984) and Polisar et al. (2003). Values were expressed as mean ± SD. Statistical significance was measured at p < 0.05. Spearman (rs) non-parametric correlation was used to test the relationship between depredation (total killed) and the age (in months) of the bovines killed, monthly total depredation and monthly births of calves, monthly total depredation and monthly rainfall (mm), and monthly rainfall (mm) and monthly births of calves. The Chi-square (v2) test was used to compare the frequency of predation between sexes of the calves (Centeno, 2000; Ayres et al., 2000). The cubic model better explained (i.e., >r2,
2.3.
Predator identification
The ranch-hands were not able to routinely distinguish between jaguar and puma kills. This distinction was only possible during October 2002–May 2003 when researchers attempted to locate and examine individual carcasses resulting from depredation events. In addition to age and sex of the kill, we also looked for external or internal lesions caused by bites or claw marks, especially in the head, neck and legs, in order to identify the predator. Jaguar attacks result in dislocated or broken cervical vertebrae, either due to the impact of the animal fall on the ground or to a bite on the base of the skull or on the neck. Usually the jaguar starts eating the head, neck and pectoral area, frequently leaving the posterior parts intact (Schaller and Crawshaw, 1980; Mondolfi and Hoogesteijn, 1986; Guix, 1992; Hoogesteijn et al., 1993). The pumas, on the other hand, kill their preys by suffocation, biting their throat and also leaving marks of their claws in the shoulders and back of their preys. They start eating their prey through the ventral part, reaching
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the ribs and the muscles of the posterior legs (Pitman et al., 2002).
3.
Results
3.1.
Cattle mortality
Cattle mortality was classified into two major categories: depredation by felids and other causes. Mortality by other causes included diseases, malnutrition, ingestion of toxic plants, lightning, malpractice during vaccination, diastolic parturition, and snake bites. From January 1998 to December 2002, loss by depredation was lower than the accumulated losses by other causes (242 and 1041, respectively) (Fig. 2). From January 1998 to December 2003, total depredation was 309 heads. Average annual depredation was 51.5 ± 18.4 heads, equivalent to 0.4% of the average annual stock of the cattle in the ranch. The average annual stock of Fazenda Ouro Branco was 12 538 ± 2679 animals during the study period. This represented an average annual biomass of 15 528.3 kg/km2. The mean annual stock sold (1221 ± 875 animals) represented an average annual biomass of 1512.2 kg/km2 (approximately 10% of the annual stock). The average annual depredation resulted in an estimated biomass loss of 63.8 kg/km2, equivalent to 0.4% of the bovine average annual biomass, and 4.2% of the commercialized meat.
3.2.
Cattle depredation by age class
Calves were the most vulnerable age category to all causes of mortality (n = 722; 56.3% of all deaths) (Table 2). Depredation by pumas and jaguars had a negative correlation with age of the killed animals (p < 0.05; rs = 0.80), being more frequent in newborns (0–2 months) and practically absent in calves older than 8 months (only nine animals above 12 months old). The cubic model better explained (y = 150.0 75.92x + 14.06x2 0.8712x3; p < 0.001; R2adj ¼ 0:976; df = 7; where: y = number of killings, x = calves’ age) the relationship between depredation and age (months) of the killed bovines (Fig. 3). Calves younger than 9 months were the most frequently killed age class. Within this category, most of the biomass preyed upon was from animals 4 to 6 months old (36.2%; 10 680 kg), followed by calves younger than 3 months (28.8%; 8500 kg), and 7 to 9 months old (25%; 7380 kg) (Table 3). The mean annual depredation of calves was 51 ± 23, equivalent to 1.2% of the mean annual number of births
Fig. 2 – Cattle mortality by depredation by pumas (Puma concolor) and jaguars (Panthera onca) and by other causes, from January 1998 to December 2002.
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Table 2 – Cattle mortality by large cats’ depredation and by other causes, from January 1998 to December 2002 Bovines
Depredation (n)
Other causes (n)
Calves Heifers and ‘‘garrotes’’ Cows Oxen Bulls
239 0 3 0 0
483 195 335 13 15
722 195 338 13 15
56.3 15.2 26.4 1.0 1.1
Total
242
1041
1283
100.0
Fig. 3 – Regression equation between depredation and cattle age class (months), from January 1998 to December 2003 (y = 150.0 75.92x + 14.06x2 0.8712x3; p < 0.001; R2adj ¼ 0:976; df = 7; where: y = number of killings, x = age (months) of the killed bovines).
(4296 ± 958) during the study period. The sex ratio at birth was approximately 1:1 (50.1%:49.9%). However, males were more frequently killed by predators (n = 183; 59.2%) than females (n = 126; 40.8%) (v2 = 10.51, df = 1; p < 0.01).
3.3.
Seasonality
Fazenda Ouro Branco applies a seasonal management of the cattle, keeping bulls and cows together for only part of the year. Thus, births are concentrated during the rainy season, when food is more abundant and more nutritious. In a sixyear period, monthly rain precipitation ranged from 0 to 826 mm. June and August were characterized by a total lack of rain. The highest precipitation rate occurred between November and January. Depredation increased from October
Total cattle mortality (n)
Total cattle mortality (%)
to May, soon after the births occurred (September through December), and then decreased from June to September, after the births ceased (May–June) (Fig. 4). Depredation rates were proportional to the seasonal number of births: 23.8% (n = 6135) of the births and 28.2% (n = 87) of the kills during the dry season (May–September); and 76.2% (n = 19,644) of the births and 71.8% (n = 222) of the kills during the rainy season (October–April). Monthly killings by predators had no direct association with the births of calves (p > 0.05). The birth of calves was associated to the monthly rain precipitation (mm) (p < 0.01; rs = 0.43), from 1998 to 2003. Killings by predators were not associated to the monthly precipitation (mm) (p > 0.05), when comparing the collected data from 1998 to 2003. If data is considered separately, though, a strong association to precipitation (mm) is noticed in 1998 (p < 0.01; rs = 0.78), in 2000 (p < 0.05; rs = 0.58) and in 2002 (p < 0.05; rs = 0.61).
3.4.
Spatial distribution
Depredation occurred only in 31.2% (n = 68) of the pastures of the ranch. The proportion of pastures next to forest that had depredation (n = 33, 48.5%) was not distinguished to the proportion of pastures not bordering forest that suffered depredation (n = 35, 51.5%). However, the proportion of pastures next to forest with depredation represented 54% (n = 33) of the 61 total pastures that were at least partially surrounded by forest patches or riparian forests that comprised eight continuum blocks of forest fragments of different sizes in the ranch and adjacent areas. The number of depredation on pastures bordering forest that had depredation (n = 150, 48.5%) was also not distinguished to the number of depredation on pastures not bordering forest that suffered depredation (n = 159, 51.5%). It is of interest that only in the SD field (main house) the pastures not bordering forest and also not occurring cattle depredation. No kills occurred in the central
Table 3 – Age class (month) and estimated biomass (kg) of killed bovines by large cats, from January 1998 to December 2003 Age class (months)
Depredation (n)
Average body mass (kg)
Estimated biomass (kg)
0–3 3–6 6–9 9–12 12–24 >24
170 89 41 6 0 3
50 120 180 275 350 420
8500 10,680 7380 1650 0000 1260
Total
309
–
29,470
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uars was US$ 25,865, with an average of US$ 4310.89 per year (US$ 11.97/day). This represented an economic loss of 0.3% of the estimated stock value and 2.6% of the average annual trade, which was approximately 4.6 times less than the loss caused by other causes (US$ 19950.90/year or US$ 54.66/day).
3.6.
Fig. 4 – Average monthly frequency of depredation by pumas (Puma concolor) and jaguars (Panthera onca) and calf births, from January 1998 to December 2003. portion (main house) of the farm, close to the headquarters (Fig. 5). The distance from the center of the pasture in relation to areas of native forest was 1317.48 ± 941.03 m. There is a marginally significant Pearson correlation between total depredation on each pasture and the distance from the center of the pasture to forest cover (p < 0.05; r = 0.23).
3.5.
Economic loss
The economic value of the mean annual stock was estimated at approximately US$ 1.4 million US dollars, with an average annual trade of US$ 162,733. From 1998 to 2003, the economic loss caused by depredation by pumas and jag-
Predator identification
From October 2002 to May 2003, some of the kills (n = 69; 22.3%) were checked. The majority of the kills checked (n = 64; 92.7%) were confirmed to have been by puma tended to kill smaller calves than jaguars (Table 4). The average weight of calves lost to pumas and to jaguars was 74 kg (0–6-months old) and 144 kg (3–9-months old), respectively. The total cattle biomass damaged by pumas and jaguars during this period was 4740 kg and 720 kg, respectively.
4.
Discussion
Several researches have indicated that diseases, malnutrition and accidents are the main reasons for higher mortalities than attacks by large cats (Schaller, 1983; Quigley and Crawshaw, 1992; Hoogesteijn et al., 1993; Oli et al., 1994; Mazzolli et al., 2002). Cattle depredation by pumas and jaguars represented 18.9 % (n = 242) of the total cattle mortality at Fazenda
Fig. 5 – Spatial distribution of depredation by pumas (Puma concolor) and jaguars (Panthera onca), from January 1998 to December 2003.
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Table 4 – Cattle depredation by pumas (Puma concolor) and jaguars (Panthera onca) according to the age class (months), from October 2002 to May 2003 Age class (months)
Puma (n)
Jaguar (n)
Total (n)
0–3 3–6 6–9 9–12 >24
42 22 0 0 0
0 3 2 0 0
42 25 2 0 0
Total
64
5
69
Ouro Branco during the five-year period (1998–2002). This is higher than the depredation recorded by Polisar et al. (2003) (13% of the cattle mortality rate in a ten-year period) in the Venezuelan Llanos, and by Schaller (1983) in Brazilian Pantanal where only 10% of the cattle mortality rate during the calving season (July–October) was caused by depredation. The average annual depredation represented 0.4% of the mean annual stock at Fazenda Ouro Branco, which is similar to that from properties bordering Iguac¸u National Park, in southwestern Brazil (Conforti and Azevedo, 2003). In the Brazilian Pantanal, cattle predation by jaguar represented 2.3% of the annual cattle holdings (Zimmermann et al., 2005). In ranches adjacent to Tsavo National Park, in Kenya, 2.4% of the cattle annual stock was killed by lions and other wild animals (Patterson et al., 2004). In some regions of Qinghai and Gansu provinces in China, 0.5% of the cattle annual stock in five ranches was killed by snow leopards and wolves (Schaller et al., 1988). On the border of Annapurna Conservation Area, in Nepal, 2.6% of the total livestock was killed by snow leopards in two years (Oli et al., 1994). Generally speaking, wild felids are responsible for the loss of up to 3% of annual domestic stocks (Jackson and Nowell, 1996). The depredation by pumas and jaguars was different at Fazenda Ouro Branco. Pumas caused most of the cattle depredation. There was also a difference in the age class of animals killed by pumas and jaguars. Pumas killed younger calves (0–6 months) and jaguars killed older calves (3–9 months). There is an apparently significant difference in the food habits of pumas and jaguars, especially where they are sympatric. Jaguars tend to prey on medium- to large-sized animals, whereas pumas tend to prey on smaller animals (Scognamillo et al., 2003). In the Pantanal, predation by pumas was more frequent on calves (92%) than on cows (8%), whereas that by jaguars was predominantly on cows (57%) than on calves (32%), oxen (7%) and bulls (4%) (Crawshaw and Quigley, 1984). Similar to the pattern found at Fazenda Ouro Branco, other studies have shown that most predation by carnivores occurs on calves (Schaller, 1983; Hoogesteijn et al., 1993; Childs, 1998; Polisar et al., 2003; Michalski et al., 2006). The seasonal breeding management of the cattle results in a peak of births during the rainy season, which, in turn, leads to a peak of depredation. However, the concentrated offer of livestock prey during the rains possibly decreases overall total annual losses to carnivores. This strategy of swamping predators to reduce overall losses has been shown to increase survival rate in many animal species (e.g., wildebeests, caribou, sea-turtles, etc.) (Pianka, 1999; Caro, 2005).
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The sex ratio at birth of calves was approximately 1:1 (50.1%:49.9%), however, males were more frequently killed by predators (n = 183; 59.2%) than females (n = 126; 40.8%), possibly because males naturally move farther from their mothers than females (C.M. Haddad, pers. comm.). The occurrence of depredation was not at random and was restricted to some pastures close to the forest reserves and riparian forests in Fazenda Ouro Branco. The proportion of pastures next to forest that had depredation was not distinguished to the proportion of pastures not bordering forest that suffered depredation. However, the proportion of pastures next to forest with depredation represented 54% of the total pastures that were at least partially surrounded by forest patches or riparian. The number of depredation on pastures bordering forest that had depredation was also not distinguished to the number of depredation on pastures not bordering forest that suffered depredation. In fact no kills occurred in the central portion (main house) of the farm, close to the headquarters where the pastures not bordering forest. In the Morro do Diabo State Park, jaguars selected habitat types non-randomly in comparison to availability. They were strongly selected for primary and secondary forests. Agriculture and open water habitats were used roughly, and open pasture was strongly selected against (Cullen et al., 2005). At Fazenda Ouro Branco, the seasonal cattle breeding season tends to improve calf growth rates and decreases mortality, because grasses are more abundant and more nutritious during the rainy season. Seasonal birthing can also facilitate their protection against predators by allowing herds to concentrated spatially, away from forest patches, during this dangerous period. Instead, depredation occurs predominantly close to forest patches upon smaller and younger animals. For this reason, calves should be kept as far as possible from such areas. The birth of calves should be concentrated in the same period of the year in all farms in order to concentrate the impact caused by predators during one time per year. This strategy is just a theoretical concept called reproductive synchrony or predator satiation that can increase the survival rate of calves. The status and distribution of large cats and their wild preys should be investigated, estimating the density of large cat populations and of the availability of prey biomass. All these strategies should be tested in cattle ranches in order to evaluate their efficacy.
Acknowledgements We would like to thank the World Wildlife Fund – Brazil (WWF – Brazil), the Ford Foundation (Grant # CSR 2682002), Idea Wild (Grant # 676), Fazenda Ouro Branco, Ernesto Beilich, and Benvinda Belem Lopes for their support to this study. We also would like to thank Cristiano T. Trinca, Ana Maria T. Trinca, Marcia C. Trape´-Cardoso, Olga Z. Zumstein, Antonio Zumstein, Valdemar L. Tornisielo, Carlos A. Vetorazzi, Roberta O.A. Valente and Saulo Salgado for their help, suggestions and comments. Bruna C.R. de Carvalho, Rodrigo D. Amato, Fernando A. Haddad, Bento C.A. Neto, and the employees of Fazenda Ouro Branco provided invaluable field assistance. We are also grateful by comments from anonymous reviewers.
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R E F E R E N C E S
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Cattle depredation by puma (Puma concolor) and jaguar (Panthera onca) in central-western Brazil Francesca B.L. Palmeiraa,*, Peter G. Crawshaw Jr.b, Claudio M. Haddadc, Katia Maria P.M.B. Ferrazd, Luciano M. Verdadee a
Research Department, Reserva Brasil, Av. Dr. Silva Melo, n. 520, apt. 606, Jardim Taquaral, CEP 04675-010 Sa˜o Paulo, SP, Brazil Pantanal Matogrossense National Park, IBAMA, R. Firmo de Matos, n. 479, CEP 79331-050 Corumba´, MS, Brazil c Animal Sciences Department, ‘‘Luiz de Queiroz’’ College of Agriculture (ESALQ), University of Sa˜o Paulo (USP), Av. Padua Dias, n. 11, CEP 13418-900 Piracicaba, SP, Brazil d Quantitative Methods Lab, Forest Science Department, ‘‘Luiz de Queiroz’’ College of Agriculture (ESALQ), University of Sa˜o Paulo (USP), Av. Padua Dias, n. 11, CEP 13418-900 Piracicaba, SP, Brazil e Animal Ecology Lab, Biological Science Department, ‘‘Luiz de Queiroz’’ College of Agriculture (ESALQ), University of Sa˜o Paulo (USP), Av. Padua Dias, n. 11, CEP 13418-900 Piracicaba, SP, Brazil b
A R T I C L E I N F O
A B S T R A C T
Article history:
In this study, data on cattle depredation by puma (Puma concolor) and jaguar (Panthera onca)
Received 2 November 2006
were recorded for six years (1998–2003) in a cattle ranch in central-western Brazil. Depreda-
Received in revised form
tion represented 18.9% of the overall cattle mortality, being predominant on calves. In bio-
16 September 2007
mass, kills represented 0.4% (63.8 kg/km2) of the ranch’s annual stock. In economic loss,
Accepted 16 September 2007
kills represented 0.3% of the cattle stock value. Depredation was mainly associated with
Available online 30 October 2007
cattle’s age class and location along with the time of birth of calves. The proportion of pastures next to forest with depredation (n = 33, 48.5%) was not distinguished to the proportion
Keywords:
of pastures not bordering forest with depredation (n = 35, 51.5%). However, the proportion
Cattle breeding season management
of pastures next to forest with depredation represented 54% (n = 33) of the 61 total pastures
Cattle production
that were at least partially surrounded by forest patches or riparian forests that comprised
Human–wildlife conflict
eight continuum blocks of forest fragments of different sizes in the ranch and adjacent
Landscape
areas. No kills occurred in the central portion (main house) of the farm, close to the head-
Livestock depredation
quarters where the pastures not bordering forest. The distances of the kills in relation to
Wild felids conservation
areas of native forest was 1317.48 ± 941.03 m. In order to reduce depredation, calves should be kept as far as possible from forest areas and concentrated cattle breeding and calving seasons should be encouraged. Ó 2007 Elsevier Ltd. All rights reserved.
1.
Introduction
Domestic livestock lack virtually all of their ancestor’s antipredator behaviors, and represent a relatively easily killed prey when compared to wild prey species of similar size
(Linnell et al., 1999). Recently, most predator species have attacked the domestic livestock across the world. Livestock depredation has been caused by lions (Panthera leo), cheetahs (Acinonyx jubatus), spotted hyenas (Crocuta crocuta), leopards (Panthera pardus), African wild dogs (Lycaon pictus) on ranches
* Corresponding author: Tel./fax: +55 11 56855168. E-mail addresses: [email protected] (F.B.L. Palmeira), [email protected] (P.G. Crawshaw Jr.), [email protected] (C.M. Haddad), [email protected] (K.M.P.M.B. Ferraz), [email protected] (L.M. Verdade). 0006-3207/$ - see front matter Ó 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2007.09.015
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in Kenya (Patterson et al., 2004; Woodroffe et al., 2005; Kolowski and Holekamp, 2006), black-backed jackals (Canis mesomelas) in Tanzania (Holmern et al., 2007), tigers (Panthera tigris), Himalayan black bears (Ursus thibettanus), dholes (Cuon alpinus) in Bhutan (Wang and Mcdonald, 2006), and snow leopards (Panthera uncia) in Nepal (Oli et al., 1994) and in Ladakh, Tibet and Mongolia (Jackson and Wangchuk, 2001). Livestock depredation by jaguars (Panthera onca) and pumas (Puma concolor) has been recorded in Belize (Rabinowitz, 1986), in Venezuela (Hoogesteijn et al., 1993; Polisar et al., 2003), in Brazil (Mazzolli et al., 2002; Conforti and Azevedo, 2003; Zimmermann et al., 2005; Michalski et al., 2006; Palmeira and Barrella, 2007) and in other neotropical countries. Cattle ranching has increased deforestation disproportionately in the Brazilian Cerrado, replacing natural areas with extensive pastures. These are interspersed with remaining fragments of forest, required as legal reserves. As a consequence, pumas and jaguars have to coexist with domestic cattle. However, landowners usually use cattle depredation as a reason to justify persecution of large cats (Quigley and Crawshaw, 1992; Silva, 1994), which are threatened in several regions in the Southern and central-western areas of Brazil (Quigley et al., 1988; Fonseca et al., 1994). This control has become the main reason for the demise of large felids, more so in areas not legally protected (Jackson and Nowell, 1996; Nowell and Jackson, 1996; Linnell et al., 1999). The Cerrado was considered by Sanderson et al. (1999) as one of the priority areas for jaguar study and conservation, due to the lack of knowledge on the species’ geographic range and population status. Despite this intense conflict, very few data have shown any significant economic loss resulting from cattle depredation (Mazzolli et al., 2002; Michalski et al., 2006; Palmeira and Barrella, 2007). Quantifying cattle depredation is fundamental to allow the implementation of management actions so as to minimize losses and to guarantee the conservation of these cats. In this context, the main objectives of this study were to determine: (1) the actual proportion of cattle killed by large felids; (2) the age distribution of the cattle killed; (3) the temporal distribution of depredation to throughout the year; and (4) the spatial distribution of cattle depredation through the landscape mosaic.
2.
Methods
2.1.
Study area
This study was carried out at the Fazenda Ouro Branco, located in the municipality of Bono´polis in Northern Goia´s, in central-western Brazil (Fig. 1). Currently, soybean (Glycine max) and cattle (approximately 90,000 heads) are the main economic activities in the region. Bono´polis is located in a transitional area between the Cerrado and the Amazon Forest biomes in the Araguaia river basin (Dinerstein et al., 1995; IBGE, 2002). The climate is hot, semi-humid, with four months of the year (May–August) with a distinct dry season. The average annual precipitation of 1874 mm occurs mostly from September through April. The mean annual temperature is 25 °C and the average altitude is 290 m a.s.l. (IBGE, 2002). The vegetation mosaic is characterized by agriculture and permanent pastures, with gallery for-
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ests, dense deciduous forest and bush savanna (Miranda and Guimara˜es, 2002). The ranch is separated in six ‘‘fields’’ or management areas identified as Sede (SD), Colina Verde (CV), Liberdade (LB), Zequinha (ZQ), Xodo´ (XD) and Madeireira (MD) which each contain an average of 36.3 ± 11.6 individual pastures. Overall, cattle density in the fields was an average of 1.2 ± 0.4 individuals/ha. The predominant cattle breeds were Nelore, Fleckvieh and a mix of the two. The average annual stock of 12 538 ± 2679 cattle represents approximately 14% of the total heads in the municipality. The six fields at the ranch were designed to decentralize and facilitate the management of cattle, in terms of number of pastures, number of the animals and total area of pastures (ha) (Table 1). The number of pastures in each filed is dynamic and one pasture can be divided in two or vice versa when it is necessary to the cattle management. In general, the ranch has a total area of 20,000 ha (200 km2), approximately 80% of disturbed areas (pastures, dams and roads) and at least 20% of permanent protected areas (forest patches and riparian forests called ‘‘Cerrada˜o’’). Approximately 28% (n = 61) of the 218 total pastures in the study area were at least 25% of the perimeter surrounded by forest patches or riparian forests that comprised eight fragments of different sizes in the ranch and adjacent areas. There is not a cutoff employed in terms of the size of forest fragments. The natural non-forested areas (Cerrado) are severely treated and most natural areas were replaced to cultivate Brachiara and other exotic pastures.
2.2.
Cattle depredation
At the ranch, artificial inseminations, birth periods, stamping and vaccinations all cattle were kept in the central portion of each field. The spatial and temporal sampling effort allocated by ranch-hands for monitoring the herds at the pastures assumed a uniform sampling effort across all fields and seasons. The ranch-hands were being sampled the herds, cattle births and cattle mortality caused by depredation or other causes. Generally, ranch-hands rode horses eight hours per day during six days per week (from Monday to Saturday). The effort distributed across seasons and fields was 2.0 km/ h sampled by two ranch-hands, equivalent to 16 km covered per day or eight pastures checked per day in each field. Considering that each field has a different size (ha) the average monthly of the sampling effort was 5.8 ± 2 return visits to each pasture by each field. It should control the potential bias to differences in sampling effort allocated by ranch-hands for monitoring the herds at the pastures in each field. Bovines were grouped into five categories: calves (0–12 months old), heifers and ‘‘garrotes’’ (12–24 months old), and cows, oxen, and bulls (older than 24 months). Data on cattle depredation were recorded by ranch-hands for each field, from January 1998 to December 2003 and information on mortality due to all causes was recorded through December 2002. The cattle mortality was determined in the field by experienced ranch-hands through of peculiar characteristics of bovine carcasses (e.g., diseases, malnutrition, ingestion of toxic plants, lightning, malpractice during vaccination, diastolic parturition, snake bites and depredation by large cats). Data about precipitation (mm), cattle stock fluctuation and mone-
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Fig. 1 – Location of Fazenda Ouro Branco, Northern Goia´s, central-western Brazil, with the six fields called ‘‘Sede’’ (SD), ‘‘Colina Verde’’ (CV), ‘‘Liberdade’’ (LB), ‘‘Zequinha’’ (ZQ), ‘‘Xodo´’’ (XD) and ‘‘Madeireira’’ (MD).
Table 1 – Number of pastures, number of bovines and total area of pastures (ha) by field in March 2003 Field
Number of pastures (n)
Total area of pastures (ha)
Number of bovines (n)
CV MD LB ZQ XD SD
39 21 26 54 39 39
2458 1587 1669 3446 2420 2550
2800 1587 1650 2480 1188 1900
Total
218
14,130
11,605
36.3 ± 11.6
2355 ± 678.7
1934.1 ± 3456.6
Mean ± SD
tary values were taken from the Monthly Cattle Breeder Bulletin, Fazenda Ouro Branco, from January 1998 to December 2003. The monetary value in Brazilian currency (‘‘Reais’’) of
each bovine killed was converted into American dollars (US$ 1 = R$ 3.15, as rated in the market in July 2004) to estimate the economic loss resulting from predatory killings.
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The bottom notes records of the Bulletins were also used to estimate the biomass (kg) and economic value of each bovine category. To estimate total bovine biomass (kg/km2), the individual average body mass was multiplied by the total number of animals and divided by the area (km2), as suggested by Schaller (1983), Crawshaw and Quigley (1984) and Polisar et al. (2003). Values were expressed as mean ± SD. Statistical significance was measured at p < 0.05. Spearman (rs) non-parametric correlation was used to test the relationship between depredation (total killed) and the age (in months) of the bovines killed, monthly total depredation and monthly births of calves, monthly total depredation and monthly rainfall (mm), and monthly rainfall (mm) and monthly births of calves. The Chi-square (v2) test was used to compare the frequency of predation between sexes of the calves (Centeno, 2000; Ayres et al., 2000). The cubic model better explained (i.e., >r2,
2.3.
Predator identification
The ranch-hands were not able to routinely distinguish between jaguar and puma kills. This distinction was only possible during October 2002–May 2003 when researchers attempted to locate and examine individual carcasses resulting from depredation events. In addition to age and sex of the kill, we also looked for external or internal lesions caused by bites or claw marks, especially in the head, neck and legs, in order to identify the predator. Jaguar attacks result in dislocated or broken cervical vertebrae, either due to the impact of the animal fall on the ground or to a bite on the base of the skull or on the neck. Usually the jaguar starts eating the head, neck and pectoral area, frequently leaving the posterior parts intact (Schaller and Crawshaw, 1980; Mondolfi and Hoogesteijn, 1986; Guix, 1992; Hoogesteijn et al., 1993). The pumas, on the other hand, kill their preys by suffocation, biting their throat and also leaving marks of their claws in the shoulders and back of their preys. They start eating their prey through the ventral part, reaching
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the ribs and the muscles of the posterior legs (Pitman et al., 2002).
3.
Results
3.1.
Cattle mortality
Cattle mortality was classified into two major categories: depredation by felids and other causes. Mortality by other causes included diseases, malnutrition, ingestion of toxic plants, lightning, malpractice during vaccination, diastolic parturition, and snake bites. From January 1998 to December 2002, loss by depredation was lower than the accumulated losses by other causes (242 and 1041, respectively) (Fig. 2). From January 1998 to December 2003, total depredation was 309 heads. Average annual depredation was 51.5 ± 18.4 heads, equivalent to 0.4% of the average annual stock of the cattle in the ranch. The average annual stock of Fazenda Ouro Branco was 12 538 ± 2679 animals during the study period. This represented an average annual biomass of 15 528.3 kg/km2. The mean annual stock sold (1221 ± 875 animals) represented an average annual biomass of 1512.2 kg/km2 (approximately 10% of the annual stock). The average annual depredation resulted in an estimated biomass loss of 63.8 kg/km2, equivalent to 0.4% of the bovine average annual biomass, and 4.2% of the commercialized meat.
3.2.
Cattle depredation by age class
Calves were the most vulnerable age category to all causes of mortality (n = 722; 56.3% of all deaths) (Table 2). Depredation by pumas and jaguars had a negative correlation with age of the killed animals (p < 0.05; rs = 0.80), being more frequent in newborns (0–2 months) and practically absent in calves older than 8 months (only nine animals above 12 months old). The cubic model better explained (y = 150.0 75.92x + 14.06x2 0.8712x3; p < 0.001; R2adj ¼ 0:976; df = 7; where: y = number of killings, x = calves’ age) the relationship between depredation and age (months) of the killed bovines (Fig. 3). Calves younger than 9 months were the most frequently killed age class. Within this category, most of the biomass preyed upon was from animals 4 to 6 months old (36.2%; 10 680 kg), followed by calves younger than 3 months (28.8%; 8500 kg), and 7 to 9 months old (25%; 7380 kg) (Table 3). The mean annual depredation of calves was 51 ± 23, equivalent to 1.2% of the mean annual number of births
Fig. 2 – Cattle mortality by depredation by pumas (Puma concolor) and jaguars (Panthera onca) and by other causes, from January 1998 to December 2002.
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Table 2 – Cattle mortality by large cats’ depredation and by other causes, from January 1998 to December 2002 Bovines
Depredation (n)
Other causes (n)
Calves Heifers and ‘‘garrotes’’ Cows Oxen Bulls
239 0 3 0 0
483 195 335 13 15
722 195 338 13 15
56.3 15.2 26.4 1.0 1.1
Total
242
1041
1283
100.0
Fig. 3 – Regression equation between depredation and cattle age class (months), from January 1998 to December 2003 (y = 150.0 75.92x + 14.06x2 0.8712x3; p < 0.001; R2adj ¼ 0:976; df = 7; where: y = number of killings, x = age (months) of the killed bovines).
(4296 ± 958) during the study period. The sex ratio at birth was approximately 1:1 (50.1%:49.9%). However, males were more frequently killed by predators (n = 183; 59.2%) than females (n = 126; 40.8%) (v2 = 10.51, df = 1; p < 0.01).
3.3.
Seasonality
Fazenda Ouro Branco applies a seasonal management of the cattle, keeping bulls and cows together for only part of the year. Thus, births are concentrated during the rainy season, when food is more abundant and more nutritious. In a sixyear period, monthly rain precipitation ranged from 0 to 826 mm. June and August were characterized by a total lack of rain. The highest precipitation rate occurred between November and January. Depredation increased from October
Total cattle mortality (n)
Total cattle mortality (%)
to May, soon after the births occurred (September through December), and then decreased from June to September, after the births ceased (May–June) (Fig. 4). Depredation rates were proportional to the seasonal number of births: 23.8% (n = 6135) of the births and 28.2% (n = 87) of the kills during the dry season (May–September); and 76.2% (n = 19,644) of the births and 71.8% (n = 222) of the kills during the rainy season (October–April). Monthly killings by predators had no direct association with the births of calves (p > 0.05). The birth of calves was associated to the monthly rain precipitation (mm) (p < 0.01; rs = 0.43), from 1998 to 2003. Killings by predators were not associated to the monthly precipitation (mm) (p > 0.05), when comparing the collected data from 1998 to 2003. If data is considered separately, though, a strong association to precipitation (mm) is noticed in 1998 (p < 0.01; rs = 0.78), in 2000 (p < 0.05; rs = 0.58) and in 2002 (p < 0.05; rs = 0.61).
3.4.
Spatial distribution
Depredation occurred only in 31.2% (n = 68) of the pastures of the ranch. The proportion of pastures next to forest that had depredation (n = 33, 48.5%) was not distinguished to the proportion of pastures not bordering forest that suffered depredation (n = 35, 51.5%). However, the proportion of pastures next to forest with depredation represented 54% (n = 33) of the 61 total pastures that were at least partially surrounded by forest patches or riparian forests that comprised eight continuum blocks of forest fragments of different sizes in the ranch and adjacent areas. The number of depredation on pastures bordering forest that had depredation (n = 150, 48.5%) was also not distinguished to the number of depredation on pastures not bordering forest that suffered depredation (n = 159, 51.5%). It is of interest that only in the SD field (main house) the pastures not bordering forest and also not occurring cattle depredation. No kills occurred in the central
Table 3 – Age class (month) and estimated biomass (kg) of killed bovines by large cats, from January 1998 to December 2003 Age class (months)
Depredation (n)
Average body mass (kg)
Estimated biomass (kg)
0–3 3–6 6–9 9–12 12–24 >24
170 89 41 6 0 3
50 120 180 275 350 420
8500 10,680 7380 1650 0000 1260
Total
309
–
29,470
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uars was US$ 25,865, with an average of US$ 4310.89 per year (US$ 11.97/day). This represented an economic loss of 0.3% of the estimated stock value and 2.6% of the average annual trade, which was approximately 4.6 times less than the loss caused by other causes (US$ 19950.90/year or US$ 54.66/day).
3.6.
Fig. 4 – Average monthly frequency of depredation by pumas (Puma concolor) and jaguars (Panthera onca) and calf births, from January 1998 to December 2003. portion (main house) of the farm, close to the headquarters (Fig. 5). The distance from the center of the pasture in relation to areas of native forest was 1317.48 ± 941.03 m. There is a marginally significant Pearson correlation between total depredation on each pasture and the distance from the center of the pasture to forest cover (p < 0.05; r = 0.23).
3.5.
Economic loss
The economic value of the mean annual stock was estimated at approximately US$ 1.4 million US dollars, with an average annual trade of US$ 162,733. From 1998 to 2003, the economic loss caused by depredation by pumas and jag-
Predator identification
From October 2002 to May 2003, some of the kills (n = 69; 22.3%) were checked. The majority of the kills checked (n = 64; 92.7%) were confirmed to have been by puma tended to kill smaller calves than jaguars (Table 4). The average weight of calves lost to pumas and to jaguars was 74 kg (0–6-months old) and 144 kg (3–9-months old), respectively. The total cattle biomass damaged by pumas and jaguars during this period was 4740 kg and 720 kg, respectively.
4.
Discussion
Several researches have indicated that diseases, malnutrition and accidents are the main reasons for higher mortalities than attacks by large cats (Schaller, 1983; Quigley and Crawshaw, 1992; Hoogesteijn et al., 1993; Oli et al., 1994; Mazzolli et al., 2002). Cattle depredation by pumas and jaguars represented 18.9 % (n = 242) of the total cattle mortality at Fazenda
Fig. 5 – Spatial distribution of depredation by pumas (Puma concolor) and jaguars (Panthera onca), from January 1998 to December 2003.
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Table 4 – Cattle depredation by pumas (Puma concolor) and jaguars (Panthera onca) according to the age class (months), from October 2002 to May 2003 Age class (months)
Puma (n)
Jaguar (n)
Total (n)
0–3 3–6 6–9 9–12 >24
42 22 0 0 0
0 3 2 0 0
42 25 2 0 0
Total
64
5
69
Ouro Branco during the five-year period (1998–2002). This is higher than the depredation recorded by Polisar et al. (2003) (13% of the cattle mortality rate in a ten-year period) in the Venezuelan Llanos, and by Schaller (1983) in Brazilian Pantanal where only 10% of the cattle mortality rate during the calving season (July–October) was caused by depredation. The average annual depredation represented 0.4% of the mean annual stock at Fazenda Ouro Branco, which is similar to that from properties bordering Iguac¸u National Park, in southwestern Brazil (Conforti and Azevedo, 2003). In the Brazilian Pantanal, cattle predation by jaguar represented 2.3% of the annual cattle holdings (Zimmermann et al., 2005). In ranches adjacent to Tsavo National Park, in Kenya, 2.4% of the cattle annual stock was killed by lions and other wild animals (Patterson et al., 2004). In some regions of Qinghai and Gansu provinces in China, 0.5% of the cattle annual stock in five ranches was killed by snow leopards and wolves (Schaller et al., 1988). On the border of Annapurna Conservation Area, in Nepal, 2.6% of the total livestock was killed by snow leopards in two years (Oli et al., 1994). Generally speaking, wild felids are responsible for the loss of up to 3% of annual domestic stocks (Jackson and Nowell, 1996). The depredation by pumas and jaguars was different at Fazenda Ouro Branco. Pumas caused most of the cattle depredation. There was also a difference in the age class of animals killed by pumas and jaguars. Pumas killed younger calves (0–6 months) and jaguars killed older calves (3–9 months). There is an apparently significant difference in the food habits of pumas and jaguars, especially where they are sympatric. Jaguars tend to prey on medium- to large-sized animals, whereas pumas tend to prey on smaller animals (Scognamillo et al., 2003). In the Pantanal, predation by pumas was more frequent on calves (92%) than on cows (8%), whereas that by jaguars was predominantly on cows (57%) than on calves (32%), oxen (7%) and bulls (4%) (Crawshaw and Quigley, 1984). Similar to the pattern found at Fazenda Ouro Branco, other studies have shown that most predation by carnivores occurs on calves (Schaller, 1983; Hoogesteijn et al., 1993; Childs, 1998; Polisar et al., 2003; Michalski et al., 2006). The seasonal breeding management of the cattle results in a peak of births during the rainy season, which, in turn, leads to a peak of depredation. However, the concentrated offer of livestock prey during the rains possibly decreases overall total annual losses to carnivores. This strategy of swamping predators to reduce overall losses has been shown to increase survival rate in many animal species (e.g., wildebeests, caribou, sea-turtles, etc.) (Pianka, 1999; Caro, 2005).
1 4 1 ( 2 0 0 8 ) 1 1 8 –1 2 5
The sex ratio at birth of calves was approximately 1:1 (50.1%:49.9%), however, males were more frequently killed by predators (n = 183; 59.2%) than females (n = 126; 40.8%), possibly because males naturally move farther from their mothers than females (C.M. Haddad, pers. comm.). The occurrence of depredation was not at random and was restricted to some pastures close to the forest reserves and riparian forests in Fazenda Ouro Branco. The proportion of pastures next to forest that had depredation was not distinguished to the proportion of pastures not bordering forest that suffered depredation. However, the proportion of pastures next to forest with depredation represented 54% of the total pastures that were at least partially surrounded by forest patches or riparian. The number of depredation on pastures bordering forest that had depredation was also not distinguished to the number of depredation on pastures not bordering forest that suffered depredation. In fact no kills occurred in the central portion (main house) of the farm, close to the headquarters where the pastures not bordering forest. In the Morro do Diabo State Park, jaguars selected habitat types non-randomly in comparison to availability. They were strongly selected for primary and secondary forests. Agriculture and open water habitats were used roughly, and open pasture was strongly selected against (Cullen et al., 2005). At Fazenda Ouro Branco, the seasonal cattle breeding season tends to improve calf growth rates and decreases mortality, because grasses are more abundant and more nutritious during the rainy season. Seasonal birthing can also facilitate their protection against predators by allowing herds to concentrated spatially, away from forest patches, during this dangerous period. Instead, depredation occurs predominantly close to forest patches upon smaller and younger animals. For this reason, calves should be kept as far as possible from such areas. The birth of calves should be concentrated in the same period of the year in all farms in order to concentrate the impact caused by predators during one time per year. This strategy is just a theoretical concept called reproductive synchrony or predator satiation that can increase the survival rate of calves. The status and distribution of large cats and their wild preys should be investigated, estimating the density of large cat populations and of the availability of prey biomass. All these strategies should be tested in cattle ranches in order to evaluate their efficacy.
Acknowledgements We would like to thank the World Wildlife Fund – Brazil (WWF – Brazil), the Ford Foundation (Grant # CSR 2682002), Idea Wild (Grant # 676), Fazenda Ouro Branco, Ernesto Beilich, and Benvinda Belem Lopes for their support to this study. We also would like to thank Cristiano T. Trinca, Ana Maria T. Trinca, Marcia C. Trape´-Cardoso, Olga Z. Zumstein, Antonio Zumstein, Valdemar L. Tornisielo, Carlos A. Vetorazzi, Roberta O.A. Valente and Saulo Salgado for their help, suggestions and comments. Bruna C.R. de Carvalho, Rodrigo D. Amato, Fernando A. Haddad, Bento C.A. Neto, and the employees of Fazenda Ouro Branco provided invaluable field assistance. We are also grateful by comments from anonymous reviewers.
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