Jaguar Conservation in Venezuela Against THE

Home , Genet (animal), Llanos, Regions of Venezuela, Venezuelan Llanos

Jaguar conservation in Venezuela against the backdrop of current knowledge on its biology and evolution Wlodzimierz Jedrzejewski, María Abarca, Ángel Viloria, Hugo Cerda, Daniel Lew, Howard Takiff, Édgar AbadÍa, Pablo Velozo and Krzysztof Schmidt

SUMMARY

Current knowledge is reviewed on the jaguar Panthera onca makes it a notorious livestock predator. Conflicts caused by jag- evolution, morphological, genetic and ecological variation, as uar predation on cattle are the main reason for killing them, al- a background for understanding conservation problems. Based though they are also hunted for skins, despite legal protection. on published sources and own data, the current situation of the Another threat is deforestation. Jaguar population in Venezuela jaguar in Venezuela is analyzed. Body size of today’s jaguars has decreased substantially during the last 30-40 years, and is is highly variable; the largest are found in the Brazilian Panta- currently variable: in northern Venezuela isolated and declining nal and Venezuelan Llanos (mean male body mass >100kg). The small populations survive; south of the Orinoco, a vast popula- smallest jaguars live in Central America (~56kg). Skin color- tion exists; in Los Llanos it was nearly eliminated ~40 years ago, ation pattern is highly variable and may have a geographic cor- but seems to have returned and re-colonized former territories. relation. The morphological variation correlates with its genetic The most important measure for conservation of jaguars in Vene variability, which in turn may reflect diversity of adaptations zuela is to create more protected areas, to develop ecological to different habitats and prey communities. Its diet is extreme- corridors to protect connectivity and halt deforestation of critical ly diversified. The territorial size and density of jaguars varies regions. Hunting and killing could decrease through educational amongst localities, probably reflecting variation in prey density programs, involving local communities, compensations for losses and degree of human disturbance. Adaptation to kill large prey from jaguar predation and incentivizing ecotourism.

Summary of Current Knowledge on Jaguar Biology and Evolution France, Germany, Netherlands, Great Britain, Caucasus, lower Urals, and in Evolution, morphological variation, genetics, and taxonomy of the jaguar Tajikistan (Sotnikova and Vislobokova, 1990; Koufos, 1992; Spassov, 1997; Turner and Anton, 1997; Kahlke, 2000; he jaguar Panthera Hemmer et al., 2001, 2010). Fossils of Hemmer et al., 2001, 2010; Barysh- onca evolved in Europe the European jaguar P. onca gom- nikov, 2002, 2011; Palombo and Valli, or Asia during the late Pliocene/early baszoegensis have been recorded from 2003-2004; O’Regan and Turner, 2004; Pleistocene period, at least 1.8-2.0 mil- several localities of Eurasia, including Foronova, 2005; Van Den Hoek Ostende lion years ago (Kurten, 1965, 1973; Italy, Greece, Bulgaria, Hungary, Spain, and De Vos, 2006; Petronio et al.,

KEYWORDS / Conservation / Current Distribution / Jaguar / Panthera onca / Venezuela / Received: 11/01/2011. Modified: 12/14/2011. Accepted: 12/21/2011.

Wlodzimierz Jedrzejewski. Prof. Dr. hab. in Forestry, Warsaw Agricultural University, Poland. Reseacher, Centro de Ecología, Instituto Venezolano de Investigaciones Científicas (IVIC), Venezuela, and Mammal Research Institute, Polish Academy of Sciences, 17-230 Bialowieza, Poland. e-mail: [email protected] María Abarca. B.Sc. in Biology, Universidad de Los Andes, Venezuela. M.Sc. and Ph.D. student in Ecology, IVIC, Venezuela. Ángel Viloria. Ph.D. in Biology, London University, UK. Researcher, IVIC, Venezuela. Hugo Cerda. Ph.D. in Biology, London University, UK. Professor, Universidad Simón Rodríguez, Venezuela. Daniel Lew. M.Sc. in Biodiversity, Universidad de Santiago de Compostela, Spain, Chief, Unit of Biodiversity, IVIC, Venezuela. Howard Takiff. M.D., New York University, EEUU, Researcher, IVIC, Venezuela. Édgar Abadía. Dr. in Microbiology, IVIC, Venezuela. Researcher, IVIC, Venezuela. Pablo Velozo. B.Sc. in Biology, La Universidad del Zulia, Venezuela. Professional Research Associate, IVIC, Venezuela. Krzysztof Schmidt. Ph.D. hab. in Biology, Warszaw University Deputy Director, Mammal Research Institute, Polish Academy of Sciences, Bialowieza, Poland.

954 0378-1844/11/12/954-13 $ 3.00/0 DEC 2011, VOL. 36 Nº 12 2011). Based on the oldest jaguar fossils 1) Panthera onca hernandesii (Western and Valli, 2003-2004; Kahlke et al., from North America, it colonized this Mexico) + P. o. centralis (Central 2011; Petronio et al., 2011). It is diffi- continent via the Bering Strait during America -El Salvador to Colombia) + P. cult to establish which of these herbi- the middle Pleistocene, ~850,000 years o. arizonensis (Southern Arizona to So- vores were the jaguar’s main prey, but ago (Simpson, 1941; Kurtén and Ander- nora, Mexico) + P. o. veraecrucis (Cen- the adaptations of ancient and living son, 1980; Seymour, 1989, 1993; Turner tral Texas to Southeastern Mexico) + P. jaguars to kill large prey are obvious. and Anton, 1997; Arroyo-Cabrales, o. goldmani (Yucatán Peninsula to Be- In South America, after colonizing this 2002). Very soon, also in the middle lize and Guatemala); continent in the middle Pleistocene up Pleistocene, it passed from North to until ~10,000 years ago, jaguars coex- 2) P. onca onca (Venezuela through the South America through the Panama isted with the local large herbivorous Amazon) + P. o. peruviana (Coastal (Darien) Isthmus (Webb, 1985, 2006; community composed of giant ground Peru); and Marshall and Sempere, 1991; Arroyo- sloth Megatherium, forest elephants Cu- Cabrales, 2002). Rincón and Prevosti 3) P. onca palustris (Pantanal regions of vieronius, mastodonts Stegomastodon, (2011) suggested that current dating Mato Grosso and Mato Grosso do Sur, camel-like Macrauchenia, huge arma- may result from sampling bias, as there Brazil) + P. o. paraguensis (along the dillos Glyptodon, wild horses Hippidi- have been relatively few paleontological Paraguay River into Paraguay and north- on, and Toxodons, animals similar to studies in northern South America. eastern Argentina). hippopotamus or rhinos (Lessa and Fa- They hypothesized that in general the Additionally, five ex- rina, 1996; Turner and Anton, 1997; Al- big cats came in a wave to South tinct subspecies have been proposed roy, 2001; Cione et al., 2003; Soibelzon America much earlier, as the Isthmus of (Cabrera, 1934; Simpson, 1941; Hoff- et al., 2008). Due to the extinction of Panama was already functional in Late stetter, 1952; Seymour, 1993; Hemmer South American mega-fauna 10,000- Pliocene, at least 2.7 million years ago 2001, 2010): P. o. mesembrina (Pleisto- 12,000 years ago, possibly because of (Webb, 2006; Rincón and Prevosti, cene South American jaguar), P. o. au- climate change and also human hunting 2011). However, most of jaguar fossils gusta (Pleistocene North American (Alroy, 2001), today jaguars are left from South America are dated from jaguar), P. o. gombaszoegensis (Pleis- with communities of much smaller prey. middle or late Pleistocene (Marshall tocene Euroasian jaguar), P. o. georgi- The body size of jag- and Sempere, 1991; Arroyo-Cabrales, ca (early Pleistocene European-Cauca- uars is highly variable, both at the con- 2002; Prevosti and Vizcaíno, 2006; sian jaguar, 1.8 mya), and P. o. tos- tinental and local levels, and there is Tonni et al., 2009; Ghilardi et al., cana (most archaic, early Pleistocene sexual dimorphism in body size, fe- 2011). Repeated advances of ice sheets European jaguar). males being 10-20% smaller than males. during glaciations caused the extinction The jaguar skull is ro- Females from western Brazil and from of jaguars in Europe, Asia, and North bust and massive but relatively short, Central America are the smallest, while America ~400,000 years ago. When the with wide zygomatic arches. The sagit- jaguars from Paraguay and Pantanal are climate warmed up during the Holocene tal crest is well developed to support believed to be the largest (Seymour, period, Central and North America the powerful head muscles. Condylobas- 1989; Hoogesteijn and Mondolfi, 1996). were recolonized by jaguars (Turner al length is usually 190-260mm, but in Hoogesteijn and Mondolfi (1996) ana- and Anton, 1997). The current geo- some may be longer than 275mm (Sey- lyzed body mass and skull measures of graphic range of the species covers mour, 1989; Hoogesteijn and Mondolfi, several jaguars from the Venezuelan from Mexico to northern Argentina 1996). Powerful jaws, with very long Llanos, the Brazilian Pantanal, the Am- (Sanderson et al., 2002a). and massive canines, endow it the azon basin and Central America. The Morphological studies strongest bite of all cats, capable of largest jaguars were from the Llanos placed the jaguar close to the leopard breaking the skull of a tapir or a turtle (mean body mass for males of 104.5kg Panthera pardus; however, recent mo- shell. Jaguars can kill a cow with a bite and for females of 66.9kg) and the Pan- lecular studies indicate that it is more from ear to ear, piercing through the tanal (99.5 and 76.7kg, respectively). closely related to the lion P. leo (Po- skull to the brain (Meachen-Samuels Skull measures were slightly higher for cock 1939; Johnson et al., 2006; and Van Valkenburgh, 2009). They have Pantanal males and significantly higher O’Brien et al., 2008; Werdelin et al., sharp, strong, retractile claws, which for Pantanal females. The smallest jag- 2010) or the snow-leopard P. uncia (Yu they use to grasp and hold a prey. All uars were those from Central America and Zhang, 2005). It has been also these adaptations to kill large prey are (56.1 and 41.4kg in males and females, suggested that the jaguar and the ex- better understood in the context of jag- respectively) while those from the Am- tinct American lion Panthera atrox uar evolution. They evolved as preda- azon were intermediate (83.6kg for (one of the biggest ever living cats) tors in the mega-fauna communities of males, no data for females). Based on were the two closest related species. In Pliocene/Pleistocene. In Europe, their fossil morphometrics, it has been sug- this scenario, P. o. gombaszoegensis, fossils are found together with fossils of gested that the late Pleistocene jaguars after entering North America would large herbivores: mammoths Mammuth- from North America (P. o. augusta) had evolved into the modern jaguars us trogontherii; forest elephants Ele- were 15-25% bigger than present jag- and the extinct P. atrox (Christiansen phas antiques; large bovids such as bi- uars, weighing up to 180kg. However, and Harris, 2009). Nine subspecies of son Bison schoetensacki, B. priscus, regression analyses of fossil measures, living jaguars were formerly distin- Bos sp. and musk ox Praeovibos pris- based on correlations between morpho- guished, based on morphological fea- cus; ancient wild boar Sus scrofa pris- logical measurements and body size of tures; more recently, based on a more cus; deer Cervus sp., Dama sp., Mega- living jaguars, have shown that past thorough analysis and molecular stud- loceros sp.; wooly rhinos Stephanorhi- jaguars were only slightly bigger than ies, this division has been rejected and nus sp.; horses Equus mosbachensis; present ones, weighing ~120kg (Sey- three distinct groups established (Po- ancient moose Alces latifrons; and mour, 1993). cock, 1939; Seymour, 1989; Larson, many others (Kahlke, 2000; Hemmer et The above-mentioned 1997; Wozencraft, 2005): al., 2001; Baryshnikov, 2002; Palombo morphological variation must have a

DEC 2011, VOL. 36 Nº 12 955 genetic basis, but population genetic biting the nape and dragging them species (Weckel et al., 2006; Cavalcanti studies on extant jaguars are scarce down (Seymour, 1989; Meachen-Samu- and Gese, 2009). (Eizirik et al., 2001, 2002; Ruíz-García els and Van Valkenburgh, 2009). They De Oliveira (2002) ana- et al., 2003, 2006, 2007; Haag et al., often conceal bigger kills, are capable lyzed data on jaguar diet from 23 dif- 2010). They include a large-scale of dragging heavy carcasses for long ferent studies. The mean standardized screening of genetic variation based on distances, and can consume small prey food niche breadth (Levins, 1968; Col- a relatively small sample (n= 40) of at the site of killing. well and Futuyma, 1971) of jaguar was jaguars taken from their entire range in About 85 species have 0.43, very similar to that of puma Central and South America (Eizirik et been reported to be prey of jaguars (0.45). The mean jaguar prey mass was al. 2001, 2002), and a more detailed (Seymour, 1989). The majority of stud- 31kg and that of its main prey was analysis of the Colombian population ies document that the most important 41kg, values that were twice as big as (Ruíz-García et al., 2006, 2007). Both in terms of biomass consumed are: col- for Puma concolor. However, the mean studies showed high microsatellite ge- lared peccary Tayassu tayacu, white- prey mass was highly variable between netic diversity of jaguars (H= 0.739 lipped peccary T. pecari, brocket deer different study sites and ranged from and 0.846 in the in the first second Mazama sp., white-tailed deer Odocoi- 2.4 to 62.9kg (de Oliveira, 2002). study). In the sample of 40 individuals, leus virginianus, cattle Bos Taurus, Cavalcanti and Gese Eizirik et al. (2001) identified 22 hap- paca Agouti paca, agouti Dasyprocta (2010) estimated the jaguar kill rate lotypes of mitochondrial DNA and esti- sp., armadillo Dasypus sp., capybara based on data from 10 radiotracked mated that the haplotypes originated Hydrochoerus hydrochaeris, caimans specimens in Pantanal, Brazil. One ani- some 280,000-510,000 years ago. This Caiman yacare, and coati Nasua nasua. mal killed on average a large or medi- suggests that the modern jaguar is a In some studies red howler monkeys um sized prey every 4 days, equivalent younger species than has been assumed (Alouatta seniculus) also constituted an to 84 prey individuals killed per year. based on paleontological studies. No important source of food biomass. Less These included 25 caimans, 21 cattle, signs of any past bottleneck have been important in terms of biomass but fre- 20 peccary, 8 deer, 3 giant ant eaters, detected, which might have been ex- quently killed are hares Sylvilagus and 9 other medium/large prey. With pected given both the history of their brasiliensis, opossums Didelphis mar- such high kill rates, jaguar livestock colonization of North and South Amer- supialis, medium size reptiles (especial- predation can be high in some areas, ica, and the recent human persecution ly turtles and iguanas), and large birds, depending on local jaguar densities. of the species. Two partial barriers to such as curassows Crax sp. (Rabinow- When hunting for cattle, jaguars kill gene flow were identified by Eizirik et itz and Nottingham, 1986; Emmons, mostly calves, but occasionally, some al. (2001): the Amazon River and the 1987; Chinchilla, 1997; Taber et al., individuals (probably mostly males) Darien strait. Additionally, studies by 1997; Nuñez et al., 2000; Garla et al., may kill also adult cows and even Ruíz-García et al. (2006, 2007) re- 2001; Scognamillo et al., 2003; Nowak, adult bulls. They may kill also horses, vealed a significant heterogeneity be- 2005; Weckel, et al., 2006; Cascelli de donkeys, pigs, dogs and chickens (Hoo- tween populations from two distinct re- Azevedo, 2008; Carrillo et al., 2009; gesteijn et al., 1993). In various stud- gions in Colombia, in concordance with Cavalcanti and Gese, 2009; Da Silveira ies, total reported cattle losses due to the occurrence of two subspecies dis- et al., 2010; Foster, et al., 2010). all natural mortality factors (droughts, tinguished earlier by taxonomists (P. o. Jaguars seem to be op- diseases, flooding, and predation) were onca and P. o. centralis). portunistic predators, able to hunt for a 2.1 to 4.2% of the population. About 10 variety of prey species in proportions to 30% of all natural mortalities were Jaguar ecology depending on local prey availability. caused by jaguars. Annually, jaguars However, closer studies showed that killed 0.1 to 1.1% of cattle. Calves may Jaguars inhabit a vari- jaguars actively select some species constitute 40-70% (in Mexico up to ety of environments, including various from local communities and avoid oth- 100%) of all cattle hunted by jaguars. types of tropical and subtropical for- ers. The species most often reported as Jaguars eliminated 0.8-2.9% of all ests, marshlands, mangroves, savanna- preferred and actively selected are col- calves born/year (Crawshaw and Quig- forest mosaics and shrublands; they can lared and white lipped peccaries, ley, 2002; Cascelli de Azevedo and live in lowlands and mountains, as well brocket and white tail deer, giant ant Murray, 2007a; Rosas-Rosas et al., as in coastal areas. They show clear eater Myrmecophaga tridactyla, capy- 2008). Scognamillo et al. (2002, 2003) preference for wet habitats and water- bara, and armadillo. Most avoided (tak- and Polisar et al. (2003) estimated a bodies, and are often reported to swim en less frequently than expected from similar rate of jaguar predation on cat- (Sanderson et al., 2002a). Paleo-envi- their share in the communities) are all tle at Hato Piñero. However, applying a ronmental reconstructions of habitats monkey species, tapirs Tapirus terres- kill rate from a radiotracking study by occupied by Pleistocene Eurasian jag- tris, cattle, caimans, coati, agouti, Cavalcanti and Gese (2010) to calculate uars also showed a wide habitat niche; opossums, skunks Conepatus sp., small losses in hatos (cattle ranches) with they lived in a variety of deciduous rodents, small and medium birds, and stable jaguar population leads to much woodlands, riparian forests, shrublands small reptiles (Garla et al., 2001; higher estimates. For example, such and dry meadow-steppes in both low- Nowak, 2005; Weckel et al., 2006; Cas- calculation for Hato Piñero would re- lands and mountain conditions (Hem- celli de Azevedo, 2008; Cavalcanti and sult in 17% of calves and 2.8% of the mer et al., 2001). Gese, 2009). Both diet composition and total cattle population removed by jag- Jaguars usually catch selectivity of jaguars to various prey uars per year. These figures show that their prey by stalking or ambush. In a species are highly variable at different jaguar predation can be an important fast attack, a jaguar grasps a prey with localities, and change with time in a problem for ranchers and should be se- its claws and bites the neck or head, or given locality. For example, some stud- riously considered in conservation it just hits a prey with its paws. With ies document avoidance of peccary, al- strategies. González-Fernández (1995) bigger preys they may jump over them, though generally it is a preferred prey showed that most hatos in the Venezu-

956 DEC 2011, VOL. 36 Nº 12 elan Cojedes state have problems with Quigley, 1991; Cascelli de Azevedo and iolo et al., 2008; Sollmann et al., jaguar and puma predation of livestock. Murray, 2007b; Cavalcanti and Gese, 2010). Habitat and prey productivity, Individual jaguars can show preference 2009). Jaguars mark their territories by degree of habitat alteration and intensi- for killing livestock while others will spraying urine, leaving feces, scraping ty of hunting by humans are the most do it rarely (Brock. 1963; Mondolfi and the ground and scratching trees, or likely factors responsible for this varia- Hoogesteijn, 1986; Hoogesteijn et al., with scent marking by rubbing their tion in home range size and density. 1993). It has been also shown that jag- head, neck or whole body against the uars killing cattle are often disabled ground or trees (Perry, 1970; Wemmer State of Knowledge, Current Situation individuals (injured, sick or old; Rabi- and Scow, 1977; Seymour, 1989; au- and Conservation Problems nowitz, 1986; Hoogesteijn et al., 1993, thors unpublished data). Roaring is also 2002, 2011). When these particular jag- a kind of territorial marking, playing a This section is based uars are shot or removed from an area, role in communication between males partially on a literature review and par- livestock losses cease even if there are and females as well. Male jaguars often tially on observations and data gathered other jaguars in the vicinity (Rabinow- respond to roars of other males (Em- during trips to 10 different areas, in or- itz, 1986). mons, 1987). Their roaring resembles der to prepare and conduct a project on Jaguars very rarely kill mews, grunts or short, loud cough, fast jaguar ecology and its morphological humans, although accidental attacks repeated 3-6 times. Both sexes are vo- and genetic variation in Venezuela. The may happen (Perry, 1970; Guggisberg, cal, although males vocalize more fre- visited localities were: 1) Los Llanos in 1975; Neto et al., 2011). At Mission quently. Cojedes state (Hato Piñero, El Baúl Tokuko, in the Sierra de Perijá, Zulia, Various studies docu- area), 2) Amazonas (Puerto Ayacucho, Venezuela, a Franciscan father reported mented a high variation in jaguar home Yutaje, San Juan de Manapiare), 3) Ser- that he conducted two funerals of range size across South and Central ranía de Imataca (El Palmar), 4) Delta women killed and partially eaten by America. The smallest territories were Amacuro, 5) Sucre-Monagas (Paría, jaguars, at a Yukpa and Bari communi- found in Belize and Costa Rica: 11- Guariquen, Río San Juan), 6) Río Caura ty at Saimadoyi. 12km2 for females and 33km2 for males (lower and middle Caura), 7) Sierra de Jaguars may live up to (Rabinowitz and Nottingham, 1986; Perijá (in the mountains and at a mosa- 23 years in captivity and up to 15 Carrillo et al., 2000). Small territories ic of forests and agricultural areas near years in the wild (Hoogesteijn and (36-45km2 for females and 37-67km2 Machiques), 8) forests on both sides of Mondolfi, 1992; Nowell and Jackson, for males) were also documented in Río Boconó, Trujillo state in Los An- 1996). Females become sexually mature Calakmul, Mexico (Ceballos et al., des, 9) forest west of Betijoque in west- at 2 years old, males at 3 years (Mon- 2001) and in some areas of Pantanal, ern Trujillo, and 10) Rancho Grande in dolfi and Hoogesteijn, 1986; Seymour, Brazil (Soisalo and Cavalcanti, 2006; Henri Pitter National Park, in the Cor- 1989; Hoogesteijn and Mondolfi, 1992; Cascelli de Azevedo and Murray, dillera de la Costa. During these trips Nowell and Jackson, 1996; Sunquist 2007b). The biggest average home censuses were conducted by recording and Sunquist, 2002). When in estrous, range size was also documented in the tracks on sand or mud, and camera females announce fertility by scent Pantanal: 163 and 157km2, respectively trapping (14-30 automatic cameras set marking and increased vocalization. (Crawshaw and Quigley, 1991). Radio- in the places selected as possible jaguar Gestation lasts ~3 months (Nowell and tracking of jaguars in Hato Piñero passes for periods of 2-5 weeks). Local Jackson, 1996), and 1-4 cubs are born, (Venezuela) revealed fairly large home people were interviewed about the pres- usually two (Rabinowitz and Notting- ranges of 83 and 100km2, for females ence and conservation problems of jag- ham, 1986; Hoogesteijn and Mondolfi, and males, respectively (Scognamillo et uars. Additionally, information was col- 1992; Sunquist and Sunquist, 2002). al., 2002, 2003; Polisar, et al., 2003). lected through personal contacts from The cubs are fully weaned at 3-6 Similar variation has some other parts of Venezuela. months of age (Hunt, 1967; Sunquist been observed for jaguar densities. The In Venezuela, the jag- and Sunquist, 2002). They start to fol- highest densities, 7-10 uar is commonly known as ‘el tigre’. low the mother when they are 2-5 individuals/100km2, were found for However, each aboriginal nation has months old and they stay with her up tropical rain forests in Belize, Costa usually its own name for this predator: to ~2 years old (Seymour, 1989; Hoo- Rica, Calakmul (Mexico) and a mosaic Guarani use ‘yaguá’, Piaroa ‘buo jäwi’, gesteijn and Mondolfi, 1992; Nowell of wetlands, forests and grasslands in Jodi ‘jkyo yëwi’, Bari ‘sebraaba’, Pemon and Jackson, 1996). Pantanal, Brazil. The lowest densities, ‘tümennen’ or ‘wayamoikö¡, Yekuana Jaguars are territorial 2-3 individuals/100km2, were recorded ‘mádo’, Yanomami ‘ironasi’, and Warao predators. Females maintain almost en- for dry scrublands in Caatinga and dry ‘toobe’. The names usually refer to the tirely exclusive territories from other savanna in Cerrado-Pantanal, Brazil, power or fierceness of the jaguar. females, while male territories may agricultural lands in Belize, and a mo- The jaguar population overlap widely. Male and female home saic of dry forests, wetlands and open in Venezuela is diversified regionally in ranges also overlap, with 1-3 female pastures in Hato Piñero, Venezuela terms of both morphological features territories being partially within a male (Ceballos et al., 2001; Scognamillo et (i.e. body mass and skin coloration pat- territory. Although generally males do al., 2002; Polisar et al., 2003; Scog- terns), and ecological parameters (i.e. not accompany females outside the namillo et al., 2003; Silver et al., 2004; food habits and population densities). mating period and do not take care of Harmsen, 2006; Cascelli de Azevedo As mentioned above, reports on body young, they may stay closer to females and Murray, 2007b; Salom-Pérez et al., mass of jaguars hunted during the past than expected from random distribu- 2007; Foster et al., 2010; Nájera, 2010). 50 years in different parts of Venezuela tion, and there are indications of social Densities <1.0 or as low as 0.3 indicate high morphological variation interactions between males and females individuals/100km2 were documented within an area that was believed to be throughout the year (Rabinowitz and for very dry or human altered habitats inhabited by a single subspecies (P. o. Nottingham, 1986; Crawshaw and in Brazil, Argentina and Uruguay (Pav- onca; Hoogesteijn and Mondolfi, 1996;

DEC 2011, VOL. 36 Nº 12 957 ing from ecologically different regions) can lead to reproductive barriers and cryptic population divisions, consistent with the habitat structure, climate and food distribution (Blier et al., 2001; Gershoni et al., 2009; Lane, 2009). Thus, the many haplotypes of mitochondrial DNA found in South Ameri- can jaguars by Eizirik et al. (2001, 2002) and the high genetic variation in nuclear DNA microsatellites shown by Ruíz García et al. (2006) may reflect not only neutral genetic drift, but perhaps evolutionary adaptation to diversified habitats and prey resources in South America. Jaguars that appear the same or very similar may in fact bear different sets of genes which allow them to live in different environmental conditions. If this were true, it would have significant consequences for conservation strategies and action plans for jaguars and possibly other species. To preserve the genetic diversity and adaptive potential, it would be necessary to Figure 1. Jaguar skins from Sierra de Imataca (‘serrano’ or ‘menudo’ coloration pattern, left) protect not only the jaguar as a species, and from Los Llanos (‘mariposa’ pattern, right) but also all the genetic variability in the different subpopulations inhabiting different habitats and different parts of Venezuela. Action plans and conserva- authors’ unpublished data). While jag- common in eastern and south-eastern tion strategies that ignore the possible uars in some parts of the country, such regions (Sierra de Imataca and along genetic variation and spatial genetic as Amazonas or the basin of the Caura the border with Guyana) than in the structure of the species may have a River, seem to be rather small, with the central and western parts of the coun- limited impact on species preservation. usual male weight not exceeding 80kg, try. These observations on morphologi- Originally, jaguars several reports from regions such as cal diversity and distinctiveness of jag- probably inhabited most of the territory Los Llanos describe the hunting of ex- uars in various parts of Venezuela may of what today is inland Venezuela, ex- tremely large jaguars (Hoogesteijn and have important consequences for the cluding perhaps the highest regions of Mondolfi, 1996; authors’ unpublished approach to species conservation. Re- Andes and driest areas of the country. data). According to reports of workers cent studies have shown that both mor- With increasing human population, the at Hato Piñero, one jaguar male killed phological and genetic variation in large range inhabited by jaguars decreased. there weighed 159kg and another, re- carnivores may have a spatial pattern- However, some authors have reported cently killed in the vicinity, weighed ing. The populations of some species that jaguars were widespread across 130kg. Accordingly to a worker at Hato are composed of sub-units distinguished most of Venezuela at least until 1950 Socorro, a jaguar killed on the border by differences in genetic composition (Zárraga, 1992). In response to the of Cojedes and Portuguesa states reflecting different adaptations to local fashion for furs of spotted cats in North weighed 180kg. Large jaguars are also habitats, climate, and food resources. America and Europe from the 1960s to believed to live in the area of Lake Ma- Also, gene flows are often directional, the early 1980s, intensive jaguar hunt- racaibo. Although these reports cannot with restricted or even no genetic ex- ing occurred all over South America be verified, it seems that in parts of change between nearby sub-populations, (Gieteling, 1972; Fitzgerald, 1989; Venezuela where jaguars prey on cattle, even in the absence of obvious geo- Ruíz–García et al., 2003; Payán and their body mass is generally higher. graphic barriers between them. Such Trujillo, 2006). Increased killing of jag- Also, the skin coloration pattern is di- cryptic population divisions have been uars during that period, which also hap- versified and seems to have some re- revealed for Canadian lynx (Rueness et pened within large cattle ranches, was gional variation (Figure 1 and authors’ al., 2003) and wolves (Pilot et al., probably also stimulated by parallel unpublished data). The skin coloration 2006; Musiani et al., 2007). The mech- predator control programs that were patterns are recognized in the popular anisms responsible for the differentia- promoted and introduced to hunting knowledge throughout the country. Jag- tion of populations into separate genetic management in North America, Europe uars with rosette-type spots are com- sub-groups may involve aminoacid al- and the Soviet Union in the 1950s to monly called ‘mariposa’ (butterfly), terations in nuclear and mitochondrial 1960s (Jędrzejewska et al., 1996; those with smaller spots (sometimes DNA encoding proteins responsible for Jędrzejewski et al., 1996; Fritts et al., black dots) are called ‘menudo’ in the cellular respiration, as well as a high 2003). As a result, jaguar populations plains, and ‘serrano’ in the mountains rate of adaptive mutations in mitochon- declined substantially all over Venezue- and hills in the north and east of Vene- drial DNA. The lack of compatible nu- la and became extinct in some regions zuela (Figure 1). Some jaguars are clear and mitochondrial DNA (as in the (Zárraga, 1992; Linares, 1998; Hooges- black (melanistic), and seem to be more case of two breeding individuals com- teijn et al., 2002; Ojasti and Lacabana,

958 DEC 2011, VOL. 36 Nº 12 2008). For example, about 30-40 years ago many jaguars were killed in Trujil- lo state in Los Andes and at the foothills of Sierra de Perijá, near Machiques, and today they are absent in these areas. The introduction of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) in 1975 started the grad- ual reduction of international fur trading, which reduced the markets for jaguar skins and therefore decreased jaguar hunting (Jackson, 1992; Nowell and Jackson, 1996). In Venezuela, the jaguar became a protected species in 1996; its hunting was forbidden and was de- clared a species threatened by extinction in north of the Meta-Orinoco rivers (Venezuela, 1996 a, b; Ojasti and Lacabana, 2008). More detailed studies are needed to verify the current status and recent changes in distribution of the jaguar population in Venezuela. However, some conclusions can be drawn based on published sources (Bis- bal, 1989; Hoogesteijn and Mondolfi, Figure 2. Jaguar distribution according to the Libro Rojo de la Fauna de Venezuela (Rodrí- guez and Rojas-Suárez, 2008, red color) and results of our surveys conducted with the aids of 1990, 1991; Medina Padilla et al., 1992; tracking, camera trapping, and interviews with local people. Also included is one data point Linares, 1998; Ojasti and Lacabana, in Apure from the SIMCOZ database, 2007 (www.simcoz.org.ve). Two data points in Apure, 2008), the data obtained from camera two in Amazonas, one in Guárico, and one in Zulia were provided by Ernesto Boede and Ra- trapping, tracking and interviews, and fael Hoogesteijn. jaguar specimens found in museum col- Filled rectangles: records of jaguars obtained by the authors through camera trapping or track lections (Sistema de Información de counting; empty rectangles: locations where camera-trapping and tracking were conducted but no evidence of jaguar presence was found; filled circles: jaguar presence confirmed from inter- Museos y Colecciones de Zoología, views with local people (confirmed visual observations or hunting) and information obtained SIMCOZ, www.simcoz.org.ve). Addi- from other researchers; half-filled circles: sporadic or unconfirmed jaguar observations, based on tional data on recent records of jaguar interviews with local people; empty circles: no recent records of jaguars, based on interviews populations were provided by Ernesto with local people. Boede and Rafael Hoogesteijn (Figure 2). Compared to the jaguar distribution above, the presence of jaguar popula- jedes, and Guárico (and possibly also to shown by Linares (1998), the more re- tions was confirmed in Amazonas, Río Barinas), where they have not been not- cent map presented in The Red Book of Caura, Imataca, Delta Amacuro, Río ed for the past 40-50 years (Figure 2). Venezuelan Fauna (Rodríguez and Ro- San Juan, Cojedes, and Perijá (Figure Altrichter et al. (2006) jas-Suárez, 2008) indicates a substantial 2). Based on our data, the situation of showed in Argentina that the time decline of jaguar populations through- the jaguar in Los Andes and in the elapsed since the last sighting of a jag- out the northern part of the country northern parts of the country might be uar in any locality correlates with the (Ojasti and Lacabana, 2008). According even worse than described in The Red age of local human settlements. This to this book, a contiguous jaguar range Book of Venezuelan Fauna. There have means that the growth of the human stretches from southern Apure and Bo- been only single, unconfirmed recent population in any area is unavoidably livar states south of Orinoco to Amazo- observations of jaguar tracks in Trujil- associated with jaguar decline and its nas State. In the north-eastern part of lo, in the area of Río Boconó and the eventual local extinction. Two factors Venezuela, jaguars can be found in the forests west of Betijoque, in the west- are associated with the growth of the Delta de Orinoco and along the coast in ern part of this Andean state, although human populations that have a direct Monagas and Sucre up to the Peninsula ~30 years ago jaguars were common impact on jaguar survival: habitat alter- de Paria. In the west, jaguars are pres- and often hunted there. Also, no strong ations (principally deforestation) and di- ent in Perijá up to Maracaibo Lake, and evidence was found of current jaguar rect killing (hunting and persecution). along the eastern and part of the west- presence in the eastern part of Serranía Habitat alteration and fragmentation ern slopes of Andes, with an extension de Turimiquire, in the north-east of have been postulated to be the most im- as far as Yaracuy and Falcon states. Venezuela, nor in central part of Cor- portant factor responsible for jaguar de- Small isolated populations are present dillera de la Costa (Figure 2). The pres- cline and local extinctions throughout in Serranía de Turimiquire on the bor- ence of jaguars in the area of Guatopo South and Central America (Nowell and der of Sucre and Monagas, in Falcon, N.P was confirmed (Emiliana Isasi, Jackson, 1996; Sanderson et al., 2002b). in Miranda in the Guatopo National personal communication). In contrast to Usually, deforestation is connected with Park and the area of Río El Guapo, in the northern part of the country, in Los spreading human settlements and the Cojedes, and in Falcón and Zulia (Fig- Llanos (central Venezuela) the situation development of agricultural activity. In ure 2). During the trips to the 10 dif- seems to have improved and jaguars many parts of Venezuela, there has ferent regions of Venezuela listed have returned to areas of Apure, Co- been an acceleration of deforestation

DEC 2011, VOL. 36 Nº 12 959 economic loss, even if other factors such as drought, disease, flooding and starvation cause a much higher mortality of calves and adult cattle (Hoogestei- jn et al.. 1993; Chávez and Gese, 2005; Chávez et al.. 2005; Zimmermann et al., 2005; Sarancho et al., 2006; Caval- canti, et al., 2010). The negative per- ception in the population, principally associated with livestock depredation, may lead to killing jaguars even inside national parks (Conforti and Cascelli de Azevedo, 2003; Carvalho and Pezzuti, 2010). It is very common throughout Venezuela, but especially in Los Lla- nos, that ranchers who lost even one calf from jaguar predation organize a deliberate hunt for that jaguar, following various modalities: mobilizing workers from the ranch (‘baquianos’) to track and kill the jaguar, usually with the help of dogs, hiring a specialized jaguar hunter to do the job, or trying to poison the jaguar or kill it with some kind of traps, such as self-firing guns. On the foothills of Sierra de Perijá near Figure 3. Jaguar killed in Bolivar state in 2010. Photo taken by a tourist. Machiques, ranchers reported that it usually takes them one to two weeks to kill a jaguar after they discover a first calf killed or even the first tracks of a due to cutting and burning tropical for- some areas becomes especially danger- jaguar that has settled inside their ‘fun- ests and converting them to pastures or ous. Animal population fragmentation do’ or farm. crop cultivation, which threatens large and isolation result in the cause for loss Private land owners areas of jaguar habitats. One example is of genetic diversity, due to genetic also kill jaguars simply out of fear, the Sierra de Imataca, where, according drift, and speed up population decline even though jaguars very rarely kill to reports of local conservationists, the (Jedrzejewski et al., 2008, 2009; Col- people. An encounter with a jaguar continuous forest has been shrinking at chero et al., 2010; Haag et al., 2010). makes people so scared that they try to an average of 1-2km/year. In the last 15 Small, isolated jaguar populations, such kill it, as they kill snakes, to protect years the forest border moved 30km as those in the northern part of the themselves and feel more secure. How- away from El Palmar (Padilla, 2003). country, are very vulnerable and may ever, a fairly common accident is when Another example is from the eastern soon disappear. a man riding a horse breaks his neck foothills of Los Andes, where primeval Despite the fact that due to the sudden reaction of the horse forests such as the famous Selva de the jaguar has been a protected species or the man upon encountering a jaguar. Ticoporo, have disappeared almost com- in Venezuela since 1996 and is consid- In such cases the jaguars are tracked pletely over the past 30 years. ered vulnerable by the scientific com- and killed, even if they never displayed Not only habitat altera- munity (Ojasti and Lacabana, 2008) di- any aggressiveness towards humans. tions, but also habitat fragmentation rect killing remains commonplace and Jaguars are also com- constitutes a serious threat for the fu- is another important threat to jaguar monly killed in Venezuela by ‘profes- ture of jaguars (Sanderson et al., 2002a; populations (Figure 3; compare Hooges- sional’ hunters who earn money from Ojasti and Lacabana, 2008; Rabinowitz teijn et al. 1993, 2002, 2011). Jaguars selling the skins and other parts of the and Zeller, 2010). In northern Venezue- are killed for variety of reasons. The jaguar body (Hoogesteijn et al.. 1993, la, the rapid expansion of areas with most common cause and the one that 2002; authors data; see Figure 3). As construction, industrial development has the biggest impact on jaguar popu- reported by hunters, trading with jaguar and transportation infrastructure create lation in Venezuela is persecution by skins is still very common, and in sev- barriers to animal movements, and es- ranchers and farmers due to livestock eral cities located close to large forest pecially for animal dispersal. There are depredation (Hoogesteijn, et al., 1993, expanses there are people in the busi- plans, some of which are being imple- 2002; González Fernández, 1995; ness of buying jaguar skins from hunt- mented, for building railways and high- Polisar et al.. 2003; Scognamillo et al., ers. Prices have substantially increased ways in northern and central parts of 2003). As mentioned above, jaguars recently due to a revival of the fashion Venezuela, as well as along the Orinoco may indeed become a problem for cattle for decorating houses with jaguar skins. River. Dispersal of jaguars and other breeding in some situations. However, Some 3-4 years ago the skin value was large mammals may be additionally re- the study of human-jaguar relationships less than 500BsF (80USD), and usually stricted by large open pastures, fences, in Pantanal, Brazil, documented that only 200-300BsF (30-50USD). Today a large water reservoirs, and intensively ranchers blame jaguars for many more hunter can get 3000-5000BsF (400- managed agricultural areas. A combina- losses than they actually cause; ranch- 600USD) for a skin, plus what he can tion of all above mentioned factors in ers see jaguars as the main reason for get for canine teeth (~200BsF each) or

960 DEC 2011, VOL. 36 Nº 12 for the whole skull. Jaguar fat is a de- served that groups of dogs are also of- should be enlarged and others must be sired product, as many believe it to be ten released and allowed to hunt by established to halt the rapid deforestation a remedy for arthritis. Additionally, the themselves. The large effects of the and habitat alterations. whole jaguar carcass, particularly hunting on animal populations are obvi- There is also the need to bones, can often be sold for the produc- ous by mere visual comparison of the consider creating new, large national tion of Chinese medicines. Jaguar hunt- protected areas, where hunting is absent parks in Cojedes, Portuguesa and Bari- ing is especially common in areas with or limited and animal densities are usu- nas, and it is especially important to en- high densities of them, such as in the ally high, with non-protected areas large the national parks in areas inhabited Sierra de Imataca, Río Caura and other where hunting is intense and animal by small, isolated, and genetically distinct parts of Bolívar state, but it occurs all densities are usually low. For example, subpopulations of jaguars. However, the over Venezuela, wherever they are still in Hato Piñero there is no hunting, ani- protected areas will have little effect on found. Recent killing of jaguars was in- mal densities are high, while in other jaguar conservation if they remain or be- variably documented in the ten areas of ‘hatos’ in the region of El Baul, in Co- come isolated from each other. Efficient Venezuela visited, upon interviewing jedes, there is intensive hunting and protection requires that they be joined hunters, ranchers and other local people animal densities are low. The result of into one functional geographic network, in regions where jaguars are present. a reduction of wild prey densities is an enabling animal dispersal and gene ex- Killing jaguars to sell skins and other increase in jaguar predation on cattle, change. For this reason, the design and products, and the commercial trade of which magnifies conflicts with ranchers implementation of a national system of these products was confirmed in five (Hoogesteijn et al., 2002; Polisar et al., ecological corridors, connecting protected regions (Imataca, Caura, Delta, Amazo- 2003; Loveridge et al., 2010). areas and other jaguar habitats is essen- nas, and Perijá). Although hunting is tial, with legal measures to protect eco- usually performed by specialized hunt- Proposals for Conservation and logical connectivity (Sanderson et al., ers, almost any hunter who spots a jag- Needs for Further Research 2002a, b; Dutton et al., 2008; Olmos Yat uar will shoot it, a practice known as Sing and Gonzalez-Fernandez, 2008; ‘opportunistic killing’ (Swank and Teer, Over the past 20 years, Rabinowitz and Zeller, 2010). Maintain- 1989, 1992). An exception are hunters several actions and solutions have been ing ecological connectivity and prevent- from some indigenous communities of proposed to make jaguar conservation ing deforestation along these corridors is Venezuela, who will not kill jaguars more effective in Central and South the most important goal. In order to re- because of their traditional belief that it America, including Venezuela (Quigley duce the impact of human constructed may result in something bad happening and Crawshaw, 1992; Rabinowitz, 1992; barriers, it is crucial that there be a legal to them or their family. Sport hunting Taber et al., 1997; Sanderson et al., requirement that all plans for investment of jaguars is also still practiced in Ven- 2002a; Sollmann, et al., 2008; Cavalcanti in construction or infrastructure include ezuela, and individuals are known to et al., 2010; Loveridge et al., 2010). The an ecological evaluation of their effect on regularly visit the Sierra de Imataca, creation of protected areas, such as na- animal movement and gene flow, and Río Caura and other regions to pursue tional parks, has been a basic approach in they should include proposals for mitiga- this hobby, as stated by local interview- jaguar conservation, although its efficacy tion measures, such as special animal ees. A variety of ways to hunt jaguars depends primarily on the size of the pro- passes (Jedrzejewski et al., 2009; exist (Medina, 1964; Zárraga, 1992) but tected area (Sollmann et al., 2008; Isasi- Jędrzejewski and Ławreszuk, 2009). probably it is most efficiently done with Catalá, 2010a). As mentioned above, indi- Efficient jaguar protec- dogs. Some hunters have special dogs vidual jaguars may have large territories tion needs support and involvement of trained to track jaguars, to find, follow and an area sufficient for maintaining a local rural communities (Cavalcanti et and chase them up a tree, whereupon population of jaguars must be quite ex- al., 2010; Loveridge et al., 2010), and the hunter has a relatively easy target tensive. In Venezuela, several national this entails the development of adequate to shoot. In general, hunters in Venezu- parks play an important role in protecting education programs. Active education ela usually keep many dogs that are habitats suitable for jaguars. However, would increase local knowledge of jag- used for hunting a variety of game ani- many areas that are very important for uar ecology, produce an awareness of mals, principally peccary and white- jaguar conservation are not adequately conservation problems, reduce fear of tailed deer, and also jaguars. protected: there are either no protected jaguars and, hopefuly, change the gener- Another very impor- areas, the protected areas are too small, al attitude of rural communities towards tant, although indirect, limiting factor or the category of protection is too low. It these animals. In consequence, it could for jaguars populations is the reduction seems obvious that forest reserves do not lower the rate of jaguar killing (Conforti in the densities of large and medium guarantee enough protection for areas as and Cascelli de Azevedo, 2003; Caval- size animals, the potential jaguar prey, valuable as Sierra de Imataca or Río canti et al., 2010; Isasi-Catalá, 2010b; as the result of intensive hunting (Hoo- Caura. The creation of national parks Loveridge et al., 2010). Education pro- gesteijn et al., 2002; Cascelli de Azeve- should be urgently considered for several grams targeted to local communities liv- do and Conforti, 2008; Loveridge et al., parts of Bolivar state, especially in the ing around national parks inhabited by 2010). Novack et al. (2005) have shown Sierra de Imataca and in the basins of the jaguars are especially important (Con- that hunting has had a specially high principal rivers, such as Orinoco, forti and Cascelli de Azevedo, 2003; impact on populations of peccary, the Cuchivero, Caura, Aro and Paragua. Pro- Carvalho and Pezzuti, 2010). Also, in- most important jaguar prey. In many tecting the jaguar populations in the delta creased cooperation between national rural areas of Venezuela most men are and mangrove ecosystems requires a sub- parks and local authorities, particularly hunters. Although the efficiency of stantial enlargement of the Delta de Ori- in obtaining funds for pro-conservation hunting is generally very high, in some noco and Turuepano national parks, and activities and the promotion of ecotour- parts it is increased due to the common the creation of new national parks in Del- ism, could benefit jaguar conservation. practice use of dogs. In Sierra de Ima- ta Amacuro, Monagas, and Sucre. Simi- The conflict between taca and in Los Andes the authors ob- larly, some national parks in Los Andes livestock owners and jaguar predation is

DEC 2011, VOL. 36 Nº 12 961 one of the most important problems to materials on minimizing livestock losses Yekuana association, Jose Luis Valles be solved. In order to reduce the proba- from predation should be prepared and from Mapire, Jose Antonio González, bility of jaguar attacks on cattle, special disseminated to ranchers. Helios Martínez, Avilio Martínez and management solutions and mitigation 6-Elaboration and implementation of Marcos Martínez from Villa de Rosario, measures have been proposed, including conservation programs directed at ranch- Luis and Loncho from Las Antenas in locating calving grounds far from for- ers and ranch owners that should include Perijá, Freddy, his brothers, Cacique ests, using electric fences, and mixing monetary incentives or other rewards for Francisco and the Bari community of cattle with buffalo (Hoogesteijn et al., farms where the presence of jaguars is Kriidaankaek, Bariven Bari association, 1992, 1993, 2002; Scognamillo et al., maintained. Nestor Luis Romero and Jorge Romero 2002; Polisar et al., 2003; Hoogesteijn from the farm Flor de Mayo, Jose Ra- and Hoogesteijn, 2008). It is necessary 7-Development of educational programs mon Flores, Gumercindo Medina Ramos, to prepare and disseminate educational in collaboration with the national park Bernarda Escalante, Werner Wilbert, materials with adequate instructions for services, to include school curricula and María Rincón, Daniel Muiller, Isidro ranchers. However, the most effective television programs. These efforts should Echeverría, Donald Marcano, coordina- solution would be to introduce special fi- be directed to increase awareness of na- tor of the Consejo Comunal Caserío nancial instruments, similar to those ture and jaguar conservation problems in Araguaimujo, Hugo Ramos, Gregorio functioning in the European Union and the country. Aguilera, the Araguaimujo and Mus- other countries, that include compensa- 8-Promotion of ecotourism. simurina Warao communities, Eccio tion for livestock loses and monetary re- Montilla, Julio Cesar Romero, Morocho 9-Control at locations where there is ac- wards for ranchers who demonstrate and Miguel from Nikitao, Luis Chinchil- tive trading of jaguar skins. pro-conservation management of their la, Claudio Castro, Asdrubal Garces and ranches and healthy jaguar and other the Garces and Castro families from El Acknowledgements wildlife populations on their land Raizón, Oswaldo Mendoza, Saúl Cara- (Mishra, 1997; Ciucci and Boitani, 1998; cas, and the direction and workers of Mech, 2003; Dutton et al., 2008; Zabel Collecting data for this article was possible due to financial sup- Campus Boconó de la Fundación La Salle and Holm-Muller, 2008, Cavalcanti et de Ciencias Naturales (FLASA). Finally, al., 2010). An additional approach is to port from the Instituto Venezolano de Investigaciones Científicas (IVIC), the the authors thank Ryszard Laskowski and involve owners of large cattle ranches in students from the Jagiellonian University conservation activities, with ecotourism Mammal Research Institute of the Polish of Cracow, as well as all the other peo- as an additional source of income (Hoo- Academy of Sciences and Polish Minis- ple whom were met during the trips to gesteijn et al., 1993, 2002; Hoogesteijn try of Sciences (grant NN304336339), various parts of Venezuela and who pro- and Chapman, 1997; Rosas-Rosas and Panthera Corporation (2010 Research vided information, data and knowledge Valdez, 2010). In Venezuela, programs and Conservation Grant the Liz Clai- that contributed to this article. of this nature have been recently pro- borne Art Ortenberg Jaguar Research posed for the areas of El Baúl and Hato Grant 2011). Bogumila Jedrzejewska Piñero by Olmos Yat Sing and Gonza- helped preparing grant proposals. The References authors are grateful to all researchers lez-Fernandez (2008). Alroy J (2001) A multispecies overkill simula- In order to provide an from IVIC and other institutions who, tion of the end-Pleistocene megafaunal adequate knowledge base for the imple- through various discussions, helped to mass extinction. Science 292: 1893. mentation of all the proposed conserva- better understand the problems of jaguar Altrichter M, Boaglio G, Perovic P (2006) The tion activities there should be a broad conservation in Venezuela, to Emiliana decline of jaguars Panthera onca in the research program combining a variety of Isasi for sharing her knowledge about Argentine Chaco. Oryx 40: 302-309. approaches and techniques, as has been jaguars in Guatopo N.P. and Antonio Arroyo-Cabrales J (2002) Registro fósil del proposed (Medina Padilla et al., 1992). González Fernández and his brother jaguar. In Medellin RA, Equihua CA, In summary, an ade- Pepe for introducing us to problems of Chetkiewicz CLB, Crawshaw PGJr, Rabi- nowitz A, Redford KH, Robinson JG, quate strategy and action plan for jaguar jaguar conservation in Los Llanos, to Sanderson EW, Taber A (Comps.) El Jag- conservation in Venezuela should in- Ernesto Boede and Rafael Hoogesteijn uar en el Nuevo Milenio. Fondo de Cul- clude the following points: for providing data on recent jaguar re- tura Economica/Universidad Nacional Auntonoma de Mexico/Wildlife Conserva- 1-Creation of a network of new national cords in Venezuela. Special gratitude is expressed to all collaborators, field assis- tion Society. Mexico. pp. 343-354. parks and enlargement of several exist- Baryshnikov GF (2002) Local biochronology ing parks. tants and volunteers who helped organiz- ing field expeditions and collecting data, of Middle and Late Pleistocene mammals 2-Application of laws to stop deforesta- from the Caucasus. Russ. J. Theriol. 1: and especially to Jose Gertrudes Gamar- 61-67. tion. ro, Douglas Oropeza, Maria Eugenia Baryshnikov GF (2011) Pleistocene Felidae 3-Establishment of a network of protect- Losada, Alexander Blanco, Blas Chaca- (Mammalia, Carnivora) from the Kudaro ed ecological corridors that would join ceres, Marcin Churski, Gilson Rivas, Ju- Paleolithic cave sites in the Caucasus. all jaguar populations and all protected nior Larreal, Tito Barrios, Marjory Proc. Zool. Inst. RAS 315: 197-226. habitat areas. Picott, Leonel (Toto) Lanner from Bisbal F (1989) Distribution and habitat asso- Machiques, Cristóbal Ríos Malaver, ciation of the carnivores in Venezuela. 4-Elaboration of laws to protect ecologi- Simón Nonato from the Pemon commu- Adv. Neotrop. Mammal. 2: 339-362. cal connectivity, and the requirement nity of Las Claritas, Jose Felix Raggi, Blier PU, Dufresne F, Burton RS (2001) Natu- that environmental impact assessments ral selection and the evolution of mtDNA- administrator of Yutaje, Rafael, his accompany all large scale investments. encoded peptides: evidence for interge- brothers, and the Piaroa community of nomic co-adaptation. Trends Genet. 17: 5-Institution of a compensation system Puerto Nuevo, Denis Rodríguez and 400-406 for livestock loses by ranchers from jag- Yurbi Rodríguez and the Yekuana com- Brock S (1963) The jaguar Panthera onca. J. uar depredation; in parallel, educational munity at Boca de Nichare, the Kuyujani Br. Guiana Mus. Zool. 37: 46-48.

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La conservación del jaguar en Venezuela frente al conocimiento actual sobre su biología y evolución Wlodzimierz Jedrzejewski, María Abarca, Ángel Viloria, Hugo Cerda, Daniel Lew, Howard Takiff, Édgar Abadía, Pablo Velozo y Krzysztof Schmidt RESUMEN El conocimiento actual sobre la evolución y las variaciones tante predador de ganado y los conflictos causados por ello son morfológicas, genéticas y ecológicas del jaguar Panthera onca son la principal causa de su matanza, además de caza para pieles a revisados como trasfondo para comprender problemas de conser- pesar de la protección legal. Otra amenaza es la deforestación. vación. Con base en publicaciones y datos propios, se analiza la La población de jaguares en Venezuela ha decrecido sustancial- situación actual del jaguar en Venezuela. El tamaño corporal del mente en los últimos 30-40 años y es actualmente variable: en la jaguar actual es muy variable; los más grandes se han encontrado zona norte sobreviven poblaciones aisladas pequeñas; al sur del en el Pantanal, Brasil, y Los Llanos venezolanos (masa corporal Orinoco existe una población importante; en Los Llanos casi se promedio macho >100kg). Los más pequeños viven en América eliminó hace ~40 años pero parece haber retornado y recoloni- Central (~56kg). El patrón de coloración de la piel es altamente zado antiguos territorios. La medida más importante de conser- variable y podría tener correlación geográfica. La variación mor- vación del jaguar en Venezuela es crear más áreas protegidas, fológica es confirmada por su variabilidad genética, que a su vez desarrollar corredores ecológicos para proteger la conectividad, podría reflejar la diversidad de adaptaciones a hábitats y comu- y detener la deforestación de regiones críticas. Caza y matanza nidades de presas. Su dieta es muy diversificada. El territorio y disminuirían con programas educativos involucrando comunidades la densidad de los jaguares varían entre localidades, reflejando locales, compensación por pérdidas por prelación, e incentivando posiblemente diferencias de presas y grado de disturbio antropo- el ecoturismo. génico. Su adaptación a matar presas grandes lo hacen un impor- a conservaÇÃO dO jaguar NA Venezuela PERANTE O conHEcimento atual sobre suA biologIa E evoluÇÃO Wlodzimierz Jedrzejewski, María Abarca, Ángel Viloria, Hugo Cerda, Daniel Lew, Howard Takiff, Édgar Abadía, Pablo Velozo y Krzysztof Schmidt RESUMO O conhecimento atual sobre a evolução e as variações mor- Sua adaptação a matar presas grandes faz dele um importante fológicas, genéticas e ecológicas do jaguar Panthera onca são predador de gado e os conflitos causados por isto são a princi- revisados como plano de fundo para compreender problemas de pal causa de sua matança, além de caça para peles apesar da conservação. Baseado em publicações e dados próprios, se ana- proteção legal. Outra ameaça é a deflorestação. A população de lisa a situação atual do jaguar na Venezuela. O tamanho cor- jaguares na Venezuela tem decrescido substancialmente nos úl- poral do jaguar atual é muito variável; os maiores têm sido en- timos 30-40 anos e é atualmente variável: sobrevivem na área contrados no Pantanal, Brasil, e nos Llanos venezuelanos (massa norte pequenas populações isoladas; ao sul do rio Orinoco exis- corporal média macho >100kg). Os menores vivem na América te uma população importante; nos Llanos foi quase eliminada Central (~56kg). O padrão de coloração da pele é altamente va- há ~40 anos mas parece haver retornado e recolonizado antigos riável e poderia ter correlação geográfica. A variação morfoló- territórios. A medida mais importante de conservação do jaguar gica é confirmada por sua variabilidade genética, que por sua na Venezuela é criar mais áreas protegidas, desenvolver corre- vez reflete diversidade de adaptações a hábitats e comunidades dores ecológicos para proteger a conectividade, e deter a deflo- de presas. Sua dieta é muito diversificada. O território e a den- restação de regiões críticas. A caça e a matança diminuiriam sidade dos jaguares variam entre localidades, refletindo possivel- com programas educativos envolvendo comunidades locais, com- mente diferenças de presas e grau de distúrbio antropogênico. pensação de perdas por prelação, e incentivando o ecoturismo.

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