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Mountain lion depredation in southern Brazil
Marcelo Mazzollia,*, Mauricio E. Graipelb, Nigel Dunstonec
aProjeto Puma, R. J. Pio Duarte Silva, 535 Horto Florestal, 88037-000, Florianopolis-SC, BrazilbDepartmento de Ecologia e Zoologia-CCB, Universidade Federal de Santa Catarina-UFSC, 8804- 900, Florianopolis-SC, Brazil
cDepartment of Biological Sciences, University of Durham, South Road, Durham City, DH1 3LE, UK
Received 6 July 2000; received in revised form 5 July 2001; accepted 27 July 2001
Abstract
Mountain lion (Puma concolor) depredation incidents on livestock herds were recorded at 15 ranches in southern Brazil from
1993 to 1995. Maximum losses to mountain lions were 78% for goats, 84% for sheep, and 16% for cattle. Cattle mortality arisingfrom causes other than depredation assumed a greater importance in herd productivity. In contrast, attacks on sheep and goatswere more frequent than losses to other causes, but could be reduced to acceptable levels when flocks were corralled at night. Mostdepredation incidents occurred when weather and light conditions were unfavorable to human activity. We explain these patterns
and inter-ranch variation in depredation rates on the basis of a risk-avoidance strategy by the mountain lions. Stock losses can beminimized by understanding these patterns and by applying appropriate herd husbandry, thus reducing the urge to persecute thisprotected species. # 2002 Elsevier Science Ltd. All rights reserved.
Keywords: Mountain lion; Livestock depredation; Puma concolor
1. Introduction
Globally, large felid predators face three majorthreats: a diminishing prey base, direct persecution byman, and habitat modification and fragmentation(Nowell and Jackson, 1996). These three factors arelikely to operate together since they are predominantlyproducts of anthropogenic disturbance. The pre-dominance of any one factor over the other will varyaccording to local culture, economy, and land tenuresystem. In areas where ranching predominates predatorsmay suffer more from direct persecution followingattacks on livestock.Livestock loss is the greatest source of conflict
between humans and large cats, and the major causeunderlying their disappearance from considerable areasof their former range [e.g. tiger Panthera tigris frommost of China; lion Panthera leo from north Africa andsouthwest Asia; and the mountain lion or puma Pumaconcolor from most of eastern North America (Nowelland Jackson, 1996)]. Weaver et al. (1996) estimatedfrom several studies that the greater proportion (75%)
of adult mountain lion mortality resulted from conflictwith humans. In Brazil the mountain lion is currentlyconsidered endangered (Bernardes et al., 1990),although it cannot be considered so under IUCN cri-teria (IUCN, 1994; Nowell and Jackson, 1996). Theglobal range of the mountain lion is estimated to be17,120,00 km2 (Nowell and Jackson, 1996). Hence,Brazil,with a land area of 8,547,403 km2, could host halfthe global population (Mazzolli, 2000). Although thereis currently no indication that mountain lion popula-tions are declining in Brazil their lethal removal byhunting or other means is prohibited by law.Although its threatened status is controversial this
does not imply that mountain lions should not be pro-tected as they have, for example, restricted their rangemainly to mountainous areas in southern Brazil (Iher-ing, 1892; Mazzolli, 1993). While the current high rateof deforestation occurring in the lowland rainforests ofthe Amazon Basin of northern Brazil is a cause forconcern (Collins, 1990; Reading et al., 1995), and mayeventually lead to widespread fragmentation, the MoistOmbrofilous (Atlantic forest; GAPLAN, 1986; IBGE,1992) and Mixed Ombrofilous (Araucaria angustifoliapine forest, hereafter Araucaria forest) forests of easternand southern Brazil, respectively, are already fragmented
0006-3207/02/$ - see front matter # 2002 Elsevier Science Ltd. All rights reserved.
PI I : S0006-3207(01 )00178-1
Biological Conservation 105 (2002) 43–51
www.elsevier.com/locate/biocon
* Corresponding author.
E-mail address: [email protected] (M. Mazzolli).
to a large extent. About 5–12% of the Atlantic Forestremains (Brown and Brown, 1992), and of the Araucariaforest only 3% was left in 1978, of which 0.6% wasprimary forest (IBDF/FUPEF, 1978). Deforestationcontinues with 11% of the remaining Atlantic Forestareas destroyed between 1985 and 1990 (SOS MataAtlantica and INPE, 1993).In southern Brazil, livestock husbandry is commonly
practiced on a small scale with mixed species herds ofcattle, goats, sheep and swine. Farming of livestock andthe fodder crops (e.g. corn Zea mays) grown to supple-ment livestock diet have fragmented forests con-siderably. The consequences of forest fragmentationinclude the decline of prey refugia and increased conflictbetween wildlife and humans, particularly near habitatedges where wild ungulates such as deer (e.g. Mazamaspp.) and rodents such as capybara (Hydrochaerishydrochaeris) are attracted to crops and from wherethey are easily poached. Chiarello (1999) found thatsmall, isolated reserves in south-eastern Brazil lackedboth large predators and their prey base (e.g. agoutisDasyprocta leporina, pacas Agouti paca, peccariesTayassu tajacu and T. pecari, and deer).Under these circumstances, poaching of wildlife has
been shown to exacerbate the effects of forest fragmen-tation with direct effects on population genetic structureand demography (Glanz, 1991; Robinson, 1996; Turnerand Corlett, 1996; Cullen et al., 2000). In all cases, thedegree to which predator and prey are affected willdepend not only on the type of disturbance but also onhow it is implemented and whether procedures areadopted to reduce crop damage and livestock depreda-tion.Despite the fact that depredation incidents have been
widely reported and documented for mountain lions(e.g. in Chile: Yanez et al., 1986; Iriarte et al., 1991; in:Arizona, Cunningham et al., 1995), little is known ofthe factors affecting depredation success under thesecircumstances. Also within the context of predator-human interaction, few studies have dealt with the issueof large felids living entirely within productive agri-cultural and ranching systems (e.g. Schaller and Craw-shaw, 1980; Van Dyke et al., 1986; Mazzolli, 2000), andit is still unclear whether long term habitat suitabilityfor large carnivores can be maintained in such areaseven with the incorporation of alternative regimes offorest management and land use. The consideration ofthis issue is crucial to determine which developmentmodels can be recommended and implemented in thevicinity of suitable large carnivore habitat.The present study aims to contribute to the under-
standing of such predator–livestock and predator–human interactions by attempting to establish the rela-tionship between livestock losses and husbandry meth-ods in southern Brazil; to identify some of the factorsthat limit or encourage livestock attacks; to identify
local constraints to the instigation of mitigating proce-dures; and to compare the extent of mountain liondepredation with losses due to other causes. Suchinformation on predator–livestock interactions shouldultimately help to prevent or diminish livestock lossesand hence persecution of large felids. This informationis especially important in areas where these predatorsare found at low density and where there is interest inprotecting their populations and avoiding conflict withranching enterprises.
2. Study area
The study area is located in the coastal mountains ofsouthern Brazil, from approximately 49� W to 51� Wand from 26�500 S to 28�500 S (Fig. 1), and comprisestwo areas, distinguished by vegetation and landscapefeatures. The southern area is located near the SerraGeral which in southern Brazil is characterized by aseries of plateaus rifted by steep cliffs, with the eastern-most slopes covered with Atlantic forest up to 700 m,and cloud forest from 700 to 1600 m (Falkenberg andVoltolini, 1995). Extensive steppe (‘‘campos de alti-tude’’) cover the plateaus above and westward of thecliffs, where, as a consequence of anthropogenic fires setto expand livestock grazing areas, most Araucaria foresthas given way to the steppe with scattered fragments ofcloud forest and/or other shrub associations (Safford,1999).The northern part of the study area consists of a
mountain range with climate, geomorphology, andphytophysiognomy characteristic of the ‘‘Serra doMar’’, a mountain chain that harbors much of theremaining Atlantic forest in southern and southeasternBrazil. There is a predominance of Atlantic Forest andmountainous terrain with valleys, which are not soabruptly cut as the canyons in the southern area. Athigher altitudes small patches of steppe and Araucariaforest also occur.
3. Methods
Livestock depredation surveys were conducted from1993 to 1995, and earlier information regarding moun-tain lion poaching incidents are included where avail-able. Poaching records were based on analysis ofmountain lion skulls and pelts (n=25) from animalskilled whilst attacking livestock. In some cases, recordsare derived from information gathered during inter-views (n=5). In order to determine the sex ratio of thesemountain lions their skulls were collected, compared,and classified according to strong dimorphic features(Mazzolli, 1992). Individuals were classified as adult,subadult or young according to cementation of fissures
44 M. Mazzolli et al. / Biological Conservation 105 (2002) 43–51
and skull size (Mazzolli, 1992). We only attributed killsto mountain lion depredation in those cases where therancher or one of the authors had inspected the car-cass(es) and its location. Some could not be inspectedowing to difficult terrain and were included in lossesattributed to other causes. Several neighboring rancheswere also surveyed to allow consistency of depredationincidents to be checked. This allowed us to evaluate whe-ther a depredation incident on one ranch was supportedby others occurring on neighboring properties. Since wedid not wish to rely on anecdotal accounts, we alsolooked for evidence of recent kills (carcasses), and skinsand skulls of killed mountain lions kept as trophies. Noother major predator, e.g. jaguar (Panthera onca),inhabited the study area, which might otherwise haveincreased the possibility of predator misidentification.We compared depredation losses of sheep between
managed herds (i.e. confined within a corral during thenight) and free-ranging herds. We also compared themortality arising from depredation with that attribu-table to other causes, which may include theft, disease,and falls from cliffs, in all herds on an annual basis.We compared losses between mortality factors and
between managed and non-managed sheep herds by
summing the values for each factor across ranches (i.e.number of ranches involved and the value for each fac-tor on each ranch). We also compared the average lossper ranch by applying a Mann–Whitney U test. In the lat-ter comparison, only those cases where mortality occurredwere included unless otherwise mentioned. Means werecompared as percentages to allow comparison of datafrom ranches with different herd sizes at the same scale,i.e. larger herd sizes will tend to have higher losses.Livestock units were standardized by converting live-
stock head units to US Dollars/head to avoid under-estimating cattle losses. Because there were insufficientdata available on age class composition of herds, weassumed, for calculation purposes, that all animalswithin a particular herd belonged to the same weight orprice class. The following cost/head estimates whichwere based on local market value were applied: cattle$210, goat $50, sheep $50, swine $60.
4. Results
Fourteen of the 15 ranches surveyed within mountainlion range were located at altitudes above 800 m, one at
Fig. 1. The study areas are represented by county limits: Lages (Cochilha Rica-Rincao do Perigo, Bocaina do Sul-Mineiros, Capao Alto-Sto Cristo,
Capao Alto/Capao Verde), Rancho Queimado (Alto da Boa Vista), Dr. Pedrinho (Bom Sucesso and Rio do Cedro), Bom Retiro (Campo dos
Padres) and Itaiopolis (Toldo). Grey areas represent primary remnant vegetation above 800 m. Numbers indicate mountain lion locations from
Mazzolli (1993): 1, Praia Grande (Pedras Brancas); 2, Meleiro (foothill zone of Serra Geral); 3, Sao Joaquim (River Sao Mateus/Chapada Bonita/
Colegio); 4, Urupema (Morro do Capote); 5, Urubici (River Baiano); 6, Bom Retiro (Paraıso da Serra); 7, Alfredo Wagner (Caete); 8, Anitapolis
(Maracuja); 9, Sao Bonifacio (Neeketer ranch); 10, Antonio Carlos (Santa Maria/Faxinal); 11, Blumenau (Source of the Garcia River); 12, Brusque
(Limeira Alta); 13, Rio dos Cedros (Alto Palmeiras); 14, Rio Negrinho; 15, Joinville (D.Francisca Road/Waterfall 1 of River Cubatao); 16, Campo
Alegre; 17, Garuva (plains and foothill zone of Serra do Mar); 18, Itapoa (plains of Bom Futuro and Barra do Saı-Mirim); 19, Rio do Campo (Azul
River); 20, Curitibanos (das Pedras River); 21, Agua Doce; 22, Ponte Serrada (between rivers Chapeco and Chapecozinho).
M. Mazzolli et al. / Biological Conservation 105 (2002) 43–51 45
650 m. The average size of ranches was 840 ha, rangingfrom 40 to 2500 ha (see Table 1). These were categorizedinto three size classes: small (five ranches from 40 to 100ha in extent); medium (four ranches, 150–400 ha) andlarge (six ranches, 500–2500 ha). Twelve of these ran-ches reared cattle, 12 reared sheep, six reared goatand seven reared swine. Most livestock birthsoccurred during spring to protect newborn livestockfrom winter stress, when temperatures may drop to�10 �C.
Of 37 recorded depredation incidents, 15 occurred inwinter (June–September), 11 in spring (September–December), nine in autumn (March–June), and two insummer (December–March). Most attacks (92%) wererecorded at night. For those incidents when weather con-ditions were recorded (n=13), 62% were in rainy weather(including drizzle) associated or not with fog, 31% ingood weather, and 7% under fog. In addition, there wereseveral alleged incidents of mountain lion attacks duringfoggy conditions only, some of which were not verified.
Table 1
Details of ranches visited during the study, including number of head of livestock at each rancha
Ranch name Area (ha) Year Livestock Total Losses
lions
Losses
other
Managementa Obs. date
Cabanha Sto. Cristo 2000 1995 Cattle 1000 1 22 no Aug 95
Boa Vista 500 1994 Cattle 400 0 8 no Apr 95
Cerro Azul 400 1994 Cattle 19 3 4 no Apr 95
Do Aleixo 70 1994 Cattle 25 0 0 no Nov 94
Judas Tadeu 2500 1994 Cattle 600 0 4 no Apr 95
Mineiros 150 1994 Cattle 80 2 6 no Nov 94
Palmeira Velha 350 1994 Cattle 70 0 1 no Nov 94
Cabanha Sto. Cristo 2000 1994 Cattle 1000 2 12 no Aug 95
Sao Domingo 70 1994 Cattle 25 0 0 no Nov 94
Saulo Yung 2500 91-94 Cattle 840 1 20 no –
Sıtio Sao Pedro 100 1994 Cattle 86 0 11 no Aug 95
Sta. Cruz 522 1994 Cattle 150 0 10 no Nov 94
Judas Tadeu 2500 1995 Cattle 600 0 3 no Apr 95
Sıtio Ruck 40 1995 Cattle 32 0 8 no Apr 95
Sıtio Sao Pedro 100 1995 Cattle 86 0 3 no Aug 95
Estancia S.F. Rio Perdido 400 1993 Goat 80 28 0 no Apr 95
Saulo Yung 2500 1991 Goat 14 7 0 no –
Judas Tadeu 2500 1993 Goat 28 4 0 no Apr 95
Boa Vista 500 1994 Goat 11 1 0 no Apr 95
Mineiros 150 1994 Goat 28 22 0 no Nov 94
Judas Tadeu 2500 1995 Goat 14 0 2 no Apr 95
Sıtio Ruck 40 1995 Goat 62 22 2 no Apr 95
Dona Olindina 2500 1992 Sheep 25 0 0 yes Jan 93
Judas Tadeu 2500 1992 Sheep 25 13 0 no Apr 95
Do Aleixo 70 1994 Sheep 17 1 7 no Nov 94
Judas Tadeu 2500 1994 Sheep 25 8 0 no Apr 95
Mineiros 150 1994 Sheep 12 8 0 no Nov 94
Palmeira Velha 350 1994 Sheep 120 45 2 no Nov 94
Potreiro Velho 50 1994 Sheep 34 1 1 yes Aug 95
Cabanha Sto. Cristo 2000 1994 Sheep 42 12 0 no Aug 95
Sao Domingo 70 1994 Sheep 9 1 0 no Nov 94
Sta. Cruz 522 1994 Sheep 65 1 0 no Nov 94
Sta. Cruz 522 1994 Sheep 65 18 0 no Nov 94
Judas Tadeu 2500 1995 Sheep 25 0 0 no Apr 95
Cabanha Sto. Cristo 2000 1995 Sheep 60 0 5 no Aug 95
Saulo Yung 2500 1991 Sheep 20 6 0 no –
Sıtio Ruck 40 1995 Sheep 6 0 0 yes Apr 95
Sıtio Sao Pedro 100 1995 Sheep 14 2 0 yes Aug 95
Boa Vista 500 1994 Swine 8 0 1 no Apr 95
Do Aleixo 70 1994 Swine 6 0 0 no Nov 94
Judas Tadeu 2500 1994 Swine 35 3 0 no Apr 95
Mineiros 150 1994 Swine 18 0 4 no Nov 94
Potreiro Velho 50 1994 Swine 9 1 0 yes Aug 95
Sao Domingo 70 1994 Swine 7 0 1 no Nov 94
Judas Tadeu 2500 1995 Swine 35 1 0 no Apr 95
Sıtio Ruck 40 1995 Swine 39 0 0 no Apr 95
a The management column indicates whether the livestock were corralled at night or not.
46 M. Mazzolli et al. / Biological Conservation 105 (2002) 43–51
Mountain lions killed by ranchers were categorized byage and sex whenever possible. Adults were from 3 to 5years old, and all young were killed when in attendancewith the mother at sheep and goat kills (Table 2). Thesex ratio of adults and sub-adults was 1.2 M:1 F, com-prising five adult males, five adult females, five sub-adult males, three sub-adult females, and three young(<1 year old).
4.1. Cattle
Attacks on cattle occurred in only four out of 12ranches with cattle herds, although mountain lions werepresent and actively depredating other herd types in tenof these ranches. Cattle herds were not put into corralsat night. Three of the four ranches with cattle depreda-tion had losses ranging from 0.1 to 2.5% of their herdvalue, and also had depredation of other livestock types;the other ranch, which experienced a 16% loss, did notmaintain other types of livestock, and had the smallestcattle herd size of all ranches, suggesting a relationshipbetween these variables and the higher mountain lionattack rate.As a result of a generally low loss per ranch and low
number of cattle ranches affected, it can be seen fromFig. 2 that depredation represented only 0.27% (US$1,890) of the summed value of all cattle herds from allranches (US$ 699,670). In contrast, cattle losses to other
causes occurred on 10 ranches, and were 12 timesgreater than those to depredation, representing 3.37%(US$ 23,520) of the total value of all cattle herds.The difference in financial loss between the two mor-
tality factors is mainly a consequence of the largernumber of ranches with losses to other causes, which sumto a higher value. Nonetheless, on an individual basis,ranches that have suffered attacks generally lost less tomountain lions than to other causes, as percentages oftheir stock (Mann–Whitney U=10, df=12, P=0.16).
4.2. Sheep
Sheep were attacked at 10 of the 12 ranches withflocks, accounting for 32% (US$ 5,900) of the summedvalue of all flocks from all ranches (US$ 18,600) (seeFig. 2). Four ranches managed their sheep herds, corral-ling them at night. Of these, only two suffered losses todepredation, which amounted to 3 and 14%of their value.All eight ranches that left sheep to graze extensively
had depredation losses. From these, seven had losses tomountain lions ranging from 24 to 84% of their flockvalue (mean=43,%, S.D.=23). The remaining ranch,which suffered a 6% depredation loss, was the only onewith losses within the same range of values reported formanaged flocks.A large difference was found in depredation between
managed and non-managed ranches where losses
Table 2
Records of mountain lions killed at ranches from 1988 to 1995
#Colec. Procedence Sex Age Record
UFSC-0320 A. Palmeiras-Rio dos Cedros-SC Female Adult Skull
UFSC-0381 D. Francisca-Joinville-SC Female Adult Skull
UFSC-0351 Urupema-SC Female Adult Skull
UFSC-s/n Curitibanos-SC Female Adult Skull and skin
FURB-s/n A. Palmeiras-Rio dos Cedros-SC Female Adult Skull
UFSC-0352 Alto Palmeiras-Rio dos Cedros-SC Female Subadult Skull
UFSC-0605 Tainhas-Cambara do Sul-RS Female Subadult Skull
UFSC-0333 Ponte Serrada-SC Female Subadult Skin
UFSC-0397 Jaquirana-RS Female Subadult or adult Skin
UFSC-0396 Alfredo Wagner-SC Female Subadult or adult Skin
UFSC-0373 Bom Jesus-RS Female Subadult or adult Skin
None Rincao do Perigo-Lages-SC Female Unknown Interview
None Rincao do Perigo-Lages-SC Female Unkonwn Interview
UFSC-0344 Bom Jesus-RS Male Adult Skull and skin
UFSC-0387 Urubici-SC Male Adult Skull and skin
UFSC-0559 Joinville-SC Male Adult Skull
M.M. Alfredo Wagner-SC Male Adult Skull
FURB-s/n A. Palmeiras-Rio dos Cedros-SC Male Adult Skull
UFSC-0318 A. Palmeiras-Rio dos Cedros-SC Male Subadult Skull
UFSC-0557 Alfredo Wagner-SC Male Subadult Skull
UFSC-0319 Sideropolis-SC Male Subadult Skull
UFSC-0606 Sao Joaquim-SC Male Subadult Skull and skin
UFSC-0745 Rancho Queimado-SC Male Subadult Skull
UFSC-0323 A. Palmeiras-Rio dos Cedros-SC Unknown Young Skull
None Afredo Wagner-SC Male Young Interview
None Dr. Pedrinho-SC Unknown Young Interview
None Bocaina do Sul-SC Male Unknown Interview
M. Mazzolli et al. / Biological Conservation 105 (2002) 43–51 47
occurred (Mann–Whitney U=15, df=10, P=0.07).This difference is even larger when managed rancheswith no losses to depredation are included in the analy-sis (Mann–Whitney U=31, df=12, P=0.01).Mountain lions often killed several free-ranging sheep
and/or goats in a single attack, but would take only asingle animal from a corral. We witnessed two cases ofmassive depredation on free-ranging flocks, with 25goats and sheep killed in the southern area and 12 sheepnear the northern area, within a 2-week period.Among the managed sheep flocks, the highest loss
(14%) was sustained where sheep were penned farthestfrom the main house (about ca. 60 m) and very near adeep and forested canyon (about ca. 40 m). One ranch,which recorded no sheep losses, sustained high goatlosses, which probably reduced attacks on the smallsheep flock. These ranches also had dogs near thehouse, which might have helped to reduce depredation.Only four of the 12 ranches with sheep sustained los-
ses to other causes, less than half the number of ranchesaffected by depredation, accounting for 4% (US$ 744)of the summed value of all sheep flocks from all ranches(US$ 18,600) (see Fig. 2). In three of the four cases,losses to other causes were < 10% per ranch, althoughthe other ranch recorded 41% losses. The economicimpact of mountain lion depredation on sheep farmingis mostly due to the higher number of ranches sufferingattacks, although the average losses to mountain lionswere, in most cases, higher than to other causes (Mann–Whitney U=29.5, df=14, P=0.18). Differences betweenmortality factors are even larger when only those depreda-tion cases at non-managed ranches are included in theanalysis (Mann–Whitney U=25, df=10, P=0.126).
4.3. Goat
All six goat flocks were free-ranging and sustaineddepredation, ranging from 9 to 78% of their value.
Depredation losses from combined flocks was 38%(US$ 4332) of the summed value of all goat flocks fromall ranches (US$ 11,150), while that attributable toother causes accounted for <2% (US$ 205) of thisvalue (see Fig. 2). Two of the goat ranches had losses toother causes (3 and 7% of the herd), this was lower thanthat caused by depredation (range 9–78%, mean=37,S.D.=25).
4.4. Swine
Seven ranches raised swine, the total value of the herdamounted to US$ 7,320. Two ranches reported losses tomountain lions (11% loss each) amounted to only US$300. Losses of swine to other causes were reported onthree ranches, ranging from 13 to 22% (US$360) (seeFig. 2). Some of the losses attributed to other causesmay have been in fact a product of depredation.
5. Discussion
5.1. Livestock depredation
Free-ranging flocks of goats and sheep suffered morelosses to mountain lions than to causes other thandepredation. However, those ranches that corralledflocks at night were able to maintain a productive stock,providing evidence that with relatively little manage-ment effort small ranching enterprises are able to co-exist with mountain lions. Whereas all free-rangingsheep and goat flocks sustained depredation losses,managed flocks may avoid losses.There is evidence that differences found in predation
rates on managed and non-managed sheep flocks arerelated to risk-aversion by the mountain lion. Depreda-tion rates were influenced by environmental factors,which reduced visibility of prey and human activity.Mountain lion predation was modified by the proximityof human habitation and fencing in two ways. First,attacks did not occur on successive nights as was fre-quently the case with free-ranging flocks; and whenmountain lions did enter a corral, the damage inflicted,commonly killing only one or two sheep, was lower thanin the case of a single attack on free-ranging flocks.Other evidence of stress is found by the way mountain
lions handled livestock kills. The wild prey of mountainlions is usually dragged away after a kill, and coveredwith leaves following feeding (Hornocker, 1970). How-ever, when mountain lions killed many free-rangingsheep or goats during a single attack they left most ofthem uncovered where they fell. Within a corral,mountain lions were reported dragging sheep up to thefence and attempting to jump over. Where fencing wassufficiently high to prevent jumping, mountain lions atethe prey in the corral. On one occasion a mountain lions
Fig. 2. Livestock losses to mountain lions (in% of US$) from all
studied ranches (n=number of sampled ranches) to mountain lions
compared to losses due to other factors.
48 M. Mazzolli et al. / Biological Conservation 105 (2002) 43–51
was observed vomiting following such a meal pre-sumably as a result of feeding under stress (Mazzolli,pers. obs.).Cattle herds suffered more losses to diseases, falls
from cliffs, and theft, than to depredation. Similarly lowlevels of cattle depredation (0.4, 0.15 and 1.5%) byjaguar and mountain lions were also estimated at threeranches in Venezuela by Hoogesteijn et al. (1992). Cattleranchers would benefit by adopting strategies to mini-mize such losses, including fencing, adequate drugadministration, and herd security, than by attempting toeliminate mountain lions. Additional methods thatcould be adopted to deter predators include the use ofguard dogs (Andelt, 1999; Hansen and Smith, 1999),and aversion conditioning, such as electric fences anduse of nauseating substances (Gustavson et al., 1976;Andelt et al., 1999; Huygens and Hayashi 1999; Ternentand Garshelis, 1999).It is possible that some unknown number of cases of
cattle falling from cliffs could be an indirect result ofmountain lion attack. Typically, fencing was inade-quate, and pastures frequently included forested patchesthat provided suitable places of concealment for pre-dators. Adequate fencing would prevent cattle fromapproaching large forested patches and very steepslopes where large prey is known to be more vulnerableto predators (e.g. Hornocker, 1970; Schaller 1972;Courtin et al., 1980, Deutsch and Weeks, 1992; Hoo-gesteijn et al., 1992; Cunningham et al., 1995).Only one cattle herd scored a high depredation loss
(16%), and possible explanations are speculative in thiscase. This ranch had the smallest cattle herd amongranches and the only one that lacked more vulnerablealternative livestock such as goats and sheep. The size ofthe herd (e.g. Schell, 1993), learning by young mountainlions prior to independence (Leyhausen, 1979; Rabino-witz, 1986; Quigley and Crawshaw, 1992), and injuries(Rabinowitz, 1986) are all reasons that may lead a felidto depredate a herd intensively. Size and sex of themountain lion also may play its role; for example, malemountain lions which are considerably larger thanfemales (Anderson, 1983; Mazzolli, 1992; Gay and Best,1995), have been found to take larger wild prey (Jalk-otzy et al., 1992), and are more commonly involved incattle predation than females (Cunningham et al., 1995).
5.2. Local issues concerning depredation incidents
Meetings were held with a local ranching associationin the southern zone of our study area to discuss solu-tions to the widespread loss of sheep. The governmenthas subsidised sheep and goat ranching since 1990,although no consideration of the predation by moun-tain lions had been made. This resulted in a fast growthof the number of free-ranging herds, which was thenfollowed by an increase in mountain lion depredation
incidents. Depredation may also have increased indir-ectly as a result of the rigorous enforcement of envir-onmental laws, which might have lead to increasedmountain lion populations. Although illegal, ranchersstill kill mountain lions following livestock depredationincidents. They also alleged that attacks had recentlybecome more intense and they believed (incorrectly)that the environmental agency [IBAMA (Instituto Bra-sileiro de Meio Ambiente)] had released mountain lionsin the area, and was therefore to blame for predatorattacks.We observed three main reactions from the sheep and
goat ranchers who sustained the highest levels of losses:abandonment of ranching of their most vulnerablestock (sheep/goat/ pigs), but retaining cattle; attemptingto eliminate the predators or confining stock to a corralat night. The latter was the least frequently implementedoption although this had been advocated as early as1820 (Saint-Hillaire, 1978). There seems to be a reluc-tance to adopt alternative husbandry strategies, as hasbeen reported in other investigations of felid depreda-tion incidents (Oli et al., 1994; Weber and Rabinowitz,1996). The reluctance to adopt appropriate stock man-agement techniques may be reduced by appointing alocal environmental officer to implement and supervisechanges in herd management on some ranches and pos-sibly,by subsidising the cost of fencing, as a way to sti-mulate the changes. Such methods are commonly usedin Brazil to increase crop and meat production.In conclusion, our investigation suggests that the pre-
dation strategy of the mountain lion when huntingdomestic livestock in southern Brazil is predominantlycautionary. Preference is shown for attacking free-ran-ging flocks of goats and sheep and with only occasionalincidents occurring near households. In addition, pre-dation on livestock is influenced by weather condition,time of day, physical (declivity) and biotic (forest cover)characteristics of the terrain, and the predator’s physicalcondition. By attacking at night under conditions of lowvisibility (rain and fog) mountain lions not only avoidinstant retaliation from ranchers, but their prey are alsoless likely to recognise predation risk (Vasquez, 1994),thereby minimizing any prey reaction or struggle thatmay result in injury to the predator. Ranchers andwildlife managers should use this knowledge and avoidleaving domestic stock in conditions favorable toattacks, thus minimizing livestock losses, and persecu-tion of mountain lions.
Acknowledgements
We like to thank the staff of the Department of Eco-systems at IBAMA-Santa Catarina, especially to biolo-gists Americo Ribeiro Tunes and Marlise Becker fortheir continuous support. IBAMA partially funded the
M. Mazzolli et al. / Biological Conservation 105 (2002) 43–51 49
trips to check incidents, and on several occasions madevehicles available for field trips. Ana Cimardi has givenunconditional support to the surveys, and some of theearly field trips were only possible due to FATMAfunding obtained with her help. Ivo Ghizoni Junior(Parana) helped by participating on several of the mostrecent expeditions, while Marcos Da-Re participated inthe early stages. Thanks also to Kristin Nowell andPeter Jackson of the IUCN Cat Specialist Group forrecognizing Projeto Puma as a valuable initiative, andtheir willingness to provide help. Appreciation is alsodue to The UK CommonWealth Office and the BritishCouncil which provided a MSc scholarship to MarceloMazzolli.
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