Camera Trapping Reveals a Diverse and Unique High-Elevation Mammal Community Under Threat

Camera trapping reveals a diverse and unique high-elevation mammal community under threat

R UTHMERY P ILLCO H UARCAYA,CHRISTOPHER B EIRNE S HIRLEY J ENNIFER S ERRANO R OJAS and A NDREW W HITWORTH

Abstract The Cerros del Sira in Peru is known to hold a Introduction diverse composition of endemic birds, amphibians and plants as a result of its geographical isolation, yet its mammalian he Cerros del Sira is an isolated mountain range in community remains poorly known. There is increasing TPeru, home to a diverse and unique flora and fauna. awareness of the threats to high-elevation species but study- The summits rise from the left bank of the Ucayali River, ing them is often hindered by rugged terrain. We present the with rugged terrain that extends over five elevational zones –  first camera-trap study of the mammal community of the ( , m). Such isolation predisposes the Cerros del Cerros del Sira. We used  camera traps placed at regular Sira to host a large number of endemic species but also elevational intervals over –, m, detecting  medium- means that these species and their habitats are sensitive sized and large mammal species. Eight are listed as threatened to human-driven forest disturbance and climatic change  on the IUCN Red List, three are categorized as Data Deficient (Forero-Medina et al., ). Although climatic changes are and one is yet to be assessed. Although other authors have not of dramatic consequence for species residing in low-lying reported that the upper elevations of the Cerros del Sira are well-connected habitat, tropical species in isolated ranges, free from hunting, we found evidence of hunting activity such as the Cerros del Sira, will have no suitable habitat to occurring above , m, and inside the core protected shift to, and could be outcompeted by low-elevation species  area. In addition to this direct evidence of hunting, recent moving to higher altitudes (Tewksbury et al., ). information has identified significant amounts of canopy Marginalized and difficult-to-access areas with little loss in the northern reaches of the core zone. Despite wide- attraction for agriculture have historically been passively spread ecological degradation in the surrounding lowlands, protected from anthropogenic disturbance. These lands the high-elevation areas of the Cerros del Sira still maintain are remote, nutrient poor, and steep, and are ideal for a unique assemblage of lowland and highland tropical rain- governments to assign for protection (Nelson & Chomitz,   forest mammals. It has been assumed that the Cerros del ; Harris et al., ). However, the passive protection Sira and other similar remote locations are safe from dis- provided by such remoteness is increasingly being ques-  turbance and protected by their isolation but we suggest tioned (e.g. Poulsen et al., ). Evidence indicates that this is an increasingly dangerous assumption to make, and many remote protected regions are undergoing defaunation  secure protection strategies need to be developed. of medium-sized and large vertebrates (Fa et al., ; Galetti & Dirzo, ), and fragmented landscapes facilitate Keywords Camera traps, El Sira, habitat loss, hunting, oncilla, access for illegal activities in remote areas (Michalski & Leopardus tigrinus, Peru, spectacled bear, Tremarctos ornatus Peres, ). In the case of Sira, Novoa et al. () reported   Supplementary material for this article is available at the loss of , ha of forest to agriculture and grazing inside – https://doi.org/./S the Sira Communal Reserve during . Although surveys on birds (Terborgh & Weske, ; Mee et al., ; Corvacho et al., ; Forero-Medina et al., ), amphibians and reptiles (Duellman & Toft, RUTHMERY PILLCO HUARCAYA* (Corresponding author, orcid.org/0000-0003- ; Aichinger, ), and flowering plants (Wasshausen, 0068-3852) and ANDREW WHITWORTH† Osa Conservation, 1012 14th Street NW, Suite 625, Washington, DC 20005, USA ; Monteagudo Mendoza & Huamán Guerrero, ) E-mail [email protected] have been carried out in the Cerros del Sira, the mammalian CHRISTOPHER BEIRNE‡ Centre for Ecology and Conservation, University of Exeter, fauna remains poorly documented. Large and medium- Exeter, UK sized mammals play a fundamental role in the functioning SHIRLEY JENNIFER SERRANO ROJAS Universidad Nacional San Antonio Abad del of Amazonian ecosystems, including control of prey popu- Cusco, Cusco, Peru lations (Estes et al., ; Ripple et al., , ) and seed *Also at: Universidad Nacional San Antonio Abad del Cusco, Cusco, Peru dispersal (Miller et al., ; Fløjgaard et al., ), but their †Also at: Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, large home ranges and slow reproductive rates make them Glasgow, UK particularly sensitive to habitat fragmentation and hunting ‡Also at: Nicholas School of the Environment, Duke University, Durham, North   Carolina, USA (Fisher & Owens, ; Cardillo et al., ). Camera traps are well-suited for surveys in remote Received  April . Revision requested  June . Accepted  August . First published online  April . locations with poor local infrastructure and harsh terrain

This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, Downloadeddistribution, from https://www.cambridge.org/core and reproduction in any medium,. IP address: provided 170.106.33.19 the original work, onis 02 properly Oct 2021 cited. at 05:04:26, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/termsOryx, 2020, 54(6), 901–908 ©. https://doi.org/10.1017/S00306053180010962019 Fauna & Flora International doi:10.1017/S0030605318001096 902 R. Pillco Huarcaya et al.

FIG. 1 Location of transect used in a camera-trap survey of medium-sized and large mammals in Sira Communal Reserve, Peru, and deforestation in the region during – (Hansen et al., ).

(e.g. Jiménez et al., ; Beirne et al., ), and we present cattle farmers. Mining and oil concessions now cover almost the first camera-trap survey of the mammalian community the entire area (Finer et al., ;Novoaetal.,). of the Cerros del Sira. We used  terrestrial camera traps over  years along a previously unstudied elevational gradient, with the aim of laying a foundation for future research, Methods monitoring and conservation planning efforts in one of the last intact forest landscapes in the central region of the Camera traps (Trophy Cam, Bushnell, Overland Park, USA) Peruvian Amazon. were deployed in mid to late March and removed at the beginning of September (dry season) in  and . Cameras were placed  cm above the ground, and all low  Study area vegetation within m was cleared to standardize detection probabilities. Cameras were programmed to record a  s The Sira Communal Reserve is situated between the Ucayali video, with intervals of  s between successive triggers River to the east and the Pachitea River to the west. It is (Meek et al., ; Beirne et al., ). Of the  cameras de- the largest community reserve in Peru, forming part of ployed in ,  were set to monitor medium-sized and the Oxapampa–Asháninka–Yánesha Biosphere Reserve, large vertebrates (including the Critically Endangered Sira located in the departments of Pasco, Huánuco and Ucayali, curassow Pauxi koepckeae; Beirne et al., ) and two with an area of , ha and a buffer area of ,, ha were placed at the entry point to the ridgeline ( m eleva- (INRENA, ). The core protected area encompasses the tion) and at our principal campsite ( m), to monitor Cerros del Sira Mountains, and the surrounding buffer area hunting activity. Thirteen cameras were placed at elevation- comprises a local human population of ethnic groups of al intervals of  m between  and , m, four were Ashaninka, Asheninka, Shipibo-Conibo and Yanesha, and placed at intervals of  m between , and , m rural communities of Andean migrants (Benavides et al., and two were placed at water sources (a clay lick at , m ). Our study transect is to the east of Puerto Inca, and a stream at , m). In ,  camera traps were in the province of Huánuco (Fig. ), along a previously un- placed at elevational intervals of  m between  and studied ridgeline on the north-western border of the Sira , m to monitor the vertebrate community, and two Communal Reserve. The transect covers both lower and were placed at the same water source locations used in upper montane forests, and elfin forest towards the highest . As people were also detected by the wildlife cameras elevations. Mean annual precipitation is c. , mm in the along the ridgeline in , camera traps were not set specif- montane forest and up to , mm on the peaks (, m). ically to target hunting activity in . The camera-trap The remoteness of the area has attracted illegal extractive rate was calculated for every given species as the number industries, including coca cultivation (for cocaine produc- of videos/ camera-trap days. Non-independent events, tion), gold mining, poaching and logging. The construction defined as videos of the same species at the same location of roads since  has attracted the private sector, with tim- within  minutes of a previous detection, were excluded ber and agricultural corporations replacing many of the local from the calculation.

Oryx, 2020, 54(6), 901–908 © 2019 Fauna & Flora International doi:10.1017/S0030605318001096 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.19, on 02 Oct 2021 at 05:04:26, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605318001096 Camera trapping and mammals 903

PLATE 1 Threatened and rare mammals of the Sira Communal Reserve, Peru (Fig. ), and direct evidence of hunting activity at high elevations within the Reserve, recorded on camera traps. (a) Panthera onca and (b) Tremarctos ornatus (both captured at the same camera trap, at , m), (c) Myrmecophaga tridactyla, (d) Tapirus terrestris, (e) Dinomys branickii, (f) Leopardus tigrinus,(g)Mustela frenata, (h) Atelocynus microtis, (i) Lagothrix cana, and (j) a hunter carrying a shotgun at , m within the core area of the Reserve.

Cameras were interfered with by both people and on the study ridge (– March  and  March– wildlife: jaguars Panthera onca moved three cameras  April ). (destroying one completely), spectacled bears Tremarctos ornatus moved four cameras, and people moved two cameras into less effective positions for surveying (i.e. Results facing dense bushes or the ground). The survey team recorded all incidental audio and visual signs of medium- Overall, we detected  medium-sized and large mammal sized and large mammals while the team was present species, belonging to eight orders and  families (Plate ;

Supplementary Table ). Thirty mammal species were detected by camera traps, with , independent records over , camera-trap nights. Six species of mammals were detected via incidental records, four of which (all ar- boreal) were not detected by camera traps: the night monkey Aotus nigriceps, the tufted capuchin Sapajus macrocephalus, the burnished saki monkey Pithecia inusta and the kinkajou Potos flavus. The species accumulation curve from camera data for both  and  shows a clear plateau, indicating that the sampling effort was sufficient to characterize the community of medium-sized and large mammals (Fig. ). During the first , camera-trap days most  of the mammal community was recorded (Sobs = ), with only five more species added to the accumulation curve from an additional effort of . , camera-trap days. The distribution of species across the elevational bands indicates that diversity was highest at ,– , m, with the highest observed species richness    (Sobs = )at , m. Only five species were detected at . , m: the oncilla Leopardus tigrinus, the spectacled bear, the long-tailed weasel Mustela frenata,the FIG. 2 Species richness accumulation curve with cumulative Andean white-eared opossum Didelphis pernigra and the number of camera-trap nights, for medium-sized and large pacarana Dinomys branickii. The pacarana was captured mammals in Sira Communal Reserve (Fig. ). The grey shaded  only once during the study (at , m) and the long- area indicates the % confidence interval. tailed weasel only on two occasions, at , and , m    (Fig. ; Supplementary Tables & ). Discussion Of the  species detected, one is categorized as Endangered on the IUCN Red List (the Peruvian woolly The community of medium-sized and large mammals of the monkey Lagothrix cana), four as Vulnerable (the lowland Cerros del Sira is exceptionally diverse, with a unique tapir Tapirus terrestris, the spectacled bear, the oncilla and assemblage of species comprising typical lowland Amazonian the giant anteater Myrmecophaga tridactyla), and three as species as well as high-elevation species. Illegal hunting Near Threatened (the short-eared dog Atelocynus microtis, activity was detected at , m elevation and within the the margay Leopardus wiedii and the jaguar). Three of protected area of the Sira Communal Reserve, despite the the species detected are categorized as Data Deficient (the suggestion that the montane terrain of the Reserve probably South American red brocket deer Mazama americana, the receives little attention from hunters (Mee & Ohlson, ). agouti Dasyprocta variegata and the Amazon dwarf squirrel The species richness of medium-sized and large mam- Microsciurus flaviventer), and one is yet to be assessed mals reported here is higher than that reported from other (the burnished saki monkey). Nineteen of the species de- highland rainforest areas of Peru (Puno = , Apurimac tected are listed in the CITES Appendices (CITES, ; River = , northern Peru = , and Kosñipata = ; Pacheco Supplementary Table ). et al., , ; Jiménez et al., ; Medina et al., ). The two cameras placed specifically to monitor human Furthermore, we detected seven species additional to presence within the study site in  detected direct the  recorded at nearby lowland Panguana Biological hunting activity (i.e. people with guns or carrying dead Station (Hutterer et al., ). Four of these species have a wildlife) on seven occasions, and shotgun shells were high-elevation distribution (spectacled bear, oncilla, long- found frequently on the forest floor. According to our tailed weasel and Andean white-eared opossum); the other guides, some of the hunters observed in the videos were three are the pacarana, the short-eared dog and the agouti. cocaleros (people hired to harvest coca) and some were Of particular significance was the first detection (to our working on inventorying trees for a timber concession. knowledge) of the two largest land predators in South One of the videos shows a hunter carrying a dead razor- America at the same camera location, the spectacled bear billed curassow Mitu tuberosum. While camping at , m and the jaguar, on the highest camera, at , m. Bears we heard two gunshots. Shortly afterwards, camera-trap were captured at elevations as low as , m, and both footage confirmed the presence of hunters carrying two species were subsequently recorded at a camera station at dead woolly monkeys. , m in Soqtapata Reserve (.°S, .°W);

FIG. 3 Elevational distribution of mammal species recorded by camera traps along the survey transect in Sira Communal Reserve (Fig. ). Cameras placed on trails and at specific habitat features are represented by triangles, mean elevation of detections is represented by filled circles, and maximum and minimum elevation records are represented by vertical lines.

Rafael Pilares, pers. comm.). It has been suggested that the of the literature and museum databases (Hurtado et al., elevation range of these two species in Peru and Bolivia does ) yielded only  records from the past  years, of not overlap anywhere within a single mountain slope, which only three were since , all within the montane and overlaps only slightly at c.  m throughout the forest of the Peruvian Yungas region (Hurtado et al., Cordillera Oriental (Servheen et al., ). The closest ). Based on the current IUCN distribution map for known previous records of the bear to our study area this species, our records are c.  km west of the known are from the Pachitea Basin, determined by interviews range (Payan & Oliveira, ). conducted with the inhabitants of the Sira Communal Other large-bodied species of note detected that require Reserve, and from products derived from bears avaialable large intact habitats include the lowland tapir and the giant for sale in the local markets of Puerto Inca, El Sira anteater, suggesting a high degree of ecological integrity and Llullapichis (Figueroa, ; Hurtado et al., ). within the core Sira Reserve. The presence of many small, Compared to the range of the spectacled bear documented rare and cryptic species, including the margay, the short- on the IUCN distribution map for the species (Velez-Liendo eared dog and the pacarana, further underlines the impor- & García-Rangel, ), our photographic records together tance of the Reserve in sustaining species of conservation with those of Figueroa () and Hurtado et al. () in- significance (Bickford et al., ). The Peruvian woolly dicate an eastwards extension of c.  km. If the bear is dis- monkey is a key species for seed dispersal, supporting car- tributed throughout the high elevations of the Sira range this bon stores and maintaining ecosystem viability (Bello et al., could represent an increase in its known habitat of up to ; Estrada et al., ); it is also a favoured target for hun- , ha, enough habitat for c.  adult individuals, ters, as we discovered, and is susceptible to local extirpation based on previous density estimates from other locations (Peres, ). (Kattan et al., ; Ríos-Uzeda et al., ). As various in- In addition to the direct evidence of hunting in the dividual spectacled bears were recorded on several occasions Reserve recorded by the camera traps, informal interviews and in different locations, these are unlikely to be records of with members of local communities living within the buffer vagrant individuals. zone confirmed the disappearance of key bushmeat species Another threatened species of note is the oncilla, one of in nearby lowland areas (in particular the Peruvian woolly the smallest cats and least known Neotropical mammals monkey, the black-faced spider monkey Ateles chamek (Tortato & Oliveira, ; Hurtado et al., ). A search and the white-lipped peccary Tayassu pecari). This loss

of game vertebrates is driving hunting pressure into the to providing park rangers with the necessary facilities and upper reaches of the forest within the core reserve area. resources to implement protection of the Reserve. Subsistence hunting is conducted when the hunter has no viable alternatives, especially where other food options are Acknowledgements We thank all supporters of Exploration Sira: the  Royal Geographical Society, with the Institute of British Geographers; not readily available (Ripple et al., ). During our stay the Neville Shulman Challenge Award; Idea Wild; crowdfunder in the communities close to our study site we observed campaign supporters (http://www.crowdfunder.co.uk/exploration-sira); that they reared many animals for personal consumption the authorities of Sira Communal Reserve (SERNANP; permits (chicken, cows, ducks and pigs), and informal conversations N°004-2015-SERNANP-RCS & N°003-2016-SERNANP-RCS); confirmed that hunting is practised predominantly as a cul- Handykam; WildlifeKate; Bushnell Nature; the Scientific Exploration Society for the 2016 Cadogan Tate Explorer Award to AW; B.D. tural legacy (enjoying bushmeat, collecting trophies and a Patterson (Field Museum of Natural History, Chicago) for assistance social activity with family and friends), not for survival. in identification of Mustela frenata; and the Golondrinas community In  we witnessed illegal logging inside the core for providing us with expert local guides. area, at , m. Previously most anthropogenic impact had occurred outside the core area; however, our direct evidence, Author contributions Study design and fieldwork: RPH, AW, CB, along with satellite imagery showing canopy loss in the nor- JSR; data analysis and writing: RPH, AW, CB.  thern reaches of the core zone (Novoa et al., ), indicates Conflicts of interest None. this is a real threat to the protected core region. The National Service of Natural Protected Areas (SERNANP) is limited Ethical standards This research abided by the Oryx Code of con- in its capacity to patrol large remote forested landscapes. duct; the camera traps did not use flash, and the image of the hunter In such cases, remote sensing technologies such as camera pictured in Plate 1 was anonymized. traps and acoustic monitoring devices have been proven to be effective as complementary tools for monitoring illegal activities in protected areas (Hossain et al., ). References The remoteness and rugged terrain of the Cerros del Sira  has so far protected it from extensive human pressure within AICHINGER,M.( ) A new species of poison-dart frog (Anura: Dendrobatidae) from the Serranía de Sira, Peru. Herpetologica, the core area of the Reserve. However, evidence of defauna- , –.  tion of other remote areas (Fa et al., ; Galetti & Dirzo, BEIRNE, C., PILLCO-HUARCAYA, R., SERRANO-ROJAS, S.J. & ) suggests that we cannot rely on isolation alone to WHITWORTH,A.() Terrestrial camera traps: essential tool for protect this important conservation area. Fragmentation the detection and future monitoring of the Critically Endangered and isolation could be detrimental to the Cerros del Sira; Sira curassow Pauxi koepckeae. Endangered Species Research, , –. surrounding areas must retain sufficient integrity and con- BELLO, C., GALETTI, M., PIZO, M.A., MAGNAGO, L.F.S., ROCHA, M.F., – nectivity between key protected areas of the Oxapampa LIMA, R.A.F. et al. () Defaunation affects carbon storage in Asháninka–Yánesha Biosphere Reserve to facilitate species tropical forests. Science Advances, ,e. migration and gene flow for viable populations. BENAVIDES, M., BARCLAY,F.&SMITH, R.C. () Atlas de Although the surrounding lowlands have been subject to Comunidades Nativas de la Selva Central. Instituto del Bien Común, Lima, Peru. widespread ecological degradation, the high-elevation areas BICKFORD, D., LOHMAN, D.J., SODHI, N.S., NG, P.K.L., MEIER, R., still maintain a unique assemblage of lowland and highland WINKER, K. et al. () Cryptic species as a window on diversity tropical rainforest mammals, many of which are threatened and conservation. Trends in Ecology and Evolution, , –. or poorly known. The presence of these species indicates CARDILLO, M., MACE, G.M., JONES, K.E., BIELBY, J., BININDA-  the Reserve has a high degree of ecological integrity and re- EMONDS, O.R.P., SECHREST,W.etal.( ) Multiple causes of high extinction risk in large mammal species. Science, , –. mains one of the few intact wilderness regions (Watson et al.,   CITES ( ) Checklist of CITES species. Http://checklist.cites.org/% ). Although the species in these highlands have thus far /en [accessed  February ]. been protected from habitat loss, the increasing human CORVACHO, M.G., MACLEOD, R., BROOKS, D.M. & HENNESSEY,B. population and demand for economic development are ex- () Distinctive morphology, ecology, and first vocal descriptions erting increasing pressure on their habitats (Soh et al., of Sira curassow (Pauxi [unicornis] koepckeae): evidence for species  rank. Ornitologia Neotropical, , –. ). If such pressures are combined with climate change   DUELLMAN, W.E. & TOFT, C.A. ( ) Anurans from Serranía de Sira, induced range shifts (Forero-Medina et al., ), species Amazonian Perú: taxonomy and biogeography. Herpetologica, losses may be catastrophic. There is a need for increased , –. protection of the roadless, intact core area of the Reserve ESTES, J.A., TERBORGH, J., BRASHARES, J.S., POWER, M.E., BERGER, J., and to ensure the maintenance of connectivity to other key BOND, W.J. et al. () Trophic downgrading of planet Earth.  – protected areas (Kearney et al., ). The development of Science, , . ESTRADA, A., GARBER, P.A., RYLANDS, A.B., ROOS, C., plans in collaboration with local people and national park FERNANDEZ-DUQUE, E., DI FIORE, A. et al. () Impending authorities to help create viable sustainable livelihoods that extinction crisis of the world’s primates: why primates matter. limit impacts on biodiversity could foster this, in addition Science Advances, ,e.

FA, J.E., PERES, C.A. & MEEUWIG,J.() Bushmeat exploitation in MEE,A.&OHLSON,J.() A Preliminary Assessment of the tropical forests: an intercontinental comparison. Conservation Biological Importance of the Cerros del Sira, Department of Biology, , –. Huánuco, Peru. ProAves Perú, Piura, Peru, Instituto Nacional de FIGUEROA,J.() Ecología y conservación del oso andino Recursos Naturales, Lima, Peru, and Asociación Interétnica de (Tremarctos ornatus) en las Áreas Naturales Protegidas del Perú. Desarrollo de la Selva Peruana, Lima, Peru. PhD thesis. Universidad de Alicante, Alicante, Spain. MEE, A., OHLSON, J., STEWART, I., WILSON, M., ÖRN,P.&DIAZ FINER, M., NOVOA, S., CRUZ,C.&PEÑA,N.() Hotspot de FERREYRA,J.() The Cerros del Sira revisited: birds of Deforestación en la Selva Central (region Huánuco). MAAP: . submontane and montane forest. Cotinga, , –. Monitoring of the Andean Amazon Project. Amazon Conservation, MEEK, P.D., BALLARD, G., CLARIDGE, A., KAYS, R., MOSEBY, K., Washington, DC, USA and Conservación Amazónica, Lima, Peru. O’BRIEN,T.etal.() Recommended guiding principles for FISHER, D.O. & OWENS, I.P.F. () The comparative method reporting on camera trapping research. Biodiversity and in conservation biology. Trends in Ecology and Evolution, Conservation, , –. , –. MICHALSKI,F.&PERES, C.A. () Disturbance-mediated mammal FLØJGAARD, C., BRUUN, H.H., HANSEN, M.D.D., HEILMANN- persistence and abundance-area relationships in Amazonian forest CLAUSEN, J., SVENNING, J.-C. & EJRNAES,R.() Are ungulates in fragments. Conservation Biology, , –. forests concerns or key species for conservation and biodiversity? MILLER, B., DUGELBY, B., FOREMAN, D., MARTINEZ DEL RÍO, C., Reply to Boulanger et al. (DOI: ./gcb.). Global Change NOSS, R., PHILLIPS, M. et al. () The importance of large Biology, , –. carnivores to healthy ecosystems. Endangered Species UPDATE, FORERO-MEDINA, G., TERBORGH, J., SOCOLAR, S.J. & PIMM, S.L. , –. () Elevational ranges of birds on a tropical montane gradient lag MONTEAGUDO MENDOZA, A.L. & HUAMÁN GUERRERO,M.() behind warming temperatures. PLOS ONE, ,e. Catálogo de los árboles y afines de la Selva Central del Perú. GALETTI,M.&DIRZO,R.() Ecological and evolutionary Arnaldoa, , –. consequences of living in a defaunated world. Biological NELSON,A.&CHOMITZ, K.M. () Effectiveness of strict vs. Conservation, , –. multiple use protected areas in reducing tropical forest fires: a global HANSEN, M.C., POTAPOV, P.V., MOORE, R., HANCHER, M., analysis using matching methods. PLOS ONE, ,e. TURUBANOVA, S.A., TYUKAVINA, A. et al. () High-resolution NOVOA, S., FINER,M.&SNELGROVE,C.() Threats to Peru’sEl global maps of st-century forest cover change. Science, Sira Communal Reserve. MAAP: . Monitoring of the Andean , –. Amazon Project. Amazon Conservation, Washington, DC, USA HARRIS, J.B.C., DWI PUTRA, D., GREGORY, S.D., BROOK, B.W., and Conservación Amazónica, Lima, Peru. PRAWIRADILAGA, D.M., SODHI, N.S. et al. () Rapid PACHECO, V., SALAS, E., CAIRAMPOMA, L., NOBLECILLA, M., deforestation threatens mid-elevational endemic birds but climate QUINTANA, H., ORTIZ, F. et al. () Contribución al change is most important at higher elevations. Diversity and conocimiento de la diversidad y conservación de los mamíferos Distributions, , –. en la cuenca del río Apurímac, Perú. Revista Peruana de Biología, HOSSAIN, A.N.M., BARLOW, A., BARLOW, C.G., LYNAM, A.J., CHAKMA, , –. S. & SAVINI,T.() Assessing the efficacy of camera trapping as a PACHECO, V., MÁRQUEZ, G., SALAS,E.&CENTTY,O.() tool for increasing detection rates of wildlife crime in tropical Diversidad de mamíferos en la cuenca media del río Tambopata, protected areas. Biological Conservation, , –. Puno, Perú. Revista Peruana de Biología, , –. HURTADO, C.M., PACHECO, V., FAJARDO,Ú.&UTURUNCO,A.() PAYAN,E.&OLIVEIRA,T.() Leopardus tigrinus.InThe IUCN Red An updated analysis of the distribution of CITES-listed Peruvian List of Threatened Species : e.TA. Http://dx.doi. carnivores for conservation priorities. Mastozoología Neotropical, org/./IUCN.UK.-.RLTS.TA.en , –. [accessed  September ]. HUTTERER, R., VERHAAGH, M., DILLER,J.&PODLOUCKY,R.() PERES, C.A. () Evaluating the impact and sustainability of An inventory of mammals observed at Panguana Biological Station, subsistence hunting at multiple Amazonian forest sites. In Amazonian Peru. Ecotropica, , –. Hunting for Sustainability in Tropical Forests (eds E.L. Bennett & INRENA () DS No --AG. Crean la Reserva Comunal El J.C. Robinson), pp. –. Columbia University Press, New Sira. Http://legislacionanp.org.pe/reserva-comunal-el-sira/ York, USA. [accessed  January ]. POULSEN, J.R., KOERNER, S.E., MOORE, S., MEDJIBE, V.P., BLAKE, S., IUCN () The IUCN Red List of Threatened Species v. .. CLARK, C.J. et al. () Poaching empties critical Central Http://www.iucnredlist.org/ [accessed  March ]. African wilderness of forest elephants. Current Biology, , JIMÉNEZ, C.F., QUINTANA, H., PACHECO, V., MELTON, D., Torrealva, PR–R. J. & TELLO,G.() Camera trap survey of medium and large RÍOS-UZEDA, B., GÓMEZ,H.&WALLACE, R.B. () A preliminary mammals in a montane rainforest of northern Peru. Revista density estimate for Andean bear using camera-trapping methods. Peruana de Biología, , –. Ursus, , –. KATTAN, G., HERNÁNDEZ, O.L., GOLDSTEIN, I., ROJAS, V., MURILLO, RIPPLE, W.J., ESTES, J.A., BESCHTA, R.L., WILMERS, C.C., RITCHIE,E. O., GÓMEZ, C. et al. () Range fragmentation in the G., HEBBLEWHITE, M. et al. () Status and ecological effects of spectacled bear Tremarctos ornatus in the northern Andes. the world’s largest carnivores. Science, , . Oryx, , –. RIPPLE, W.J., ABERNETHY, K., BETTS, M.G., CHAPRON, G., DIRZO, R., KEARNEY, S.G., ADAMS, V.M., FULLER, R.A., POSSINGHAM, H.P. & GALETTI, M. et al. () Bushmeat hunting and extinction risk to WATSON, J.E.M. () Estimating the benefit of well-managed the world’s mammals. Royal Society Open Science, , . protected areas for threatened species conservation. Oryx, SERVHEEN, C., HERRERO,S.&PEYTON,B.() Spectacled bear published online  May . conservation action plan. In Status, Survey and Conservation Action MEDINA, C.E., ZEBALLOS,H.&LÓPEZ,E.() Diversidad de Plan: Bears (eds IUCN/SSC BEAR AND POLAR BEAR SPECIALIST mamiferos en los bosques montanos del valle de Kcosñipata Cusco, GROUP), pp. –. IUCN, Gland, Switzerland and Cambridge, Peru. Mastozoología Neotropical, , –. UK.

SOH, M.C.K., SODHI, N.S. & LIM, S.L.H. () High sensitivity of VELEZ-LIENDO,X.&GARCÍA-RANGEL,S.() Tremarctos montane bird communities to habitat disturbance in Peninsular ornatus.InThe IUCN Red List of Threatened Species : Malaysia. Biological Conservation, , –. e.TA. Http://dx.doi.org/./IUCN.UK.-. TERBORGH,J.&WESKE, J.S. () The role of competition in the RLTS.TA.en [accessed  January ]. distribution of Andean birds. Ecology, , –. WASSHAUSEN, D.C. () A checklist of the Acanthaceae collected in TEWKSBURY, J.J., HUEY, R.B. & DEUTSCH, C.A. () Putting the heat the “Sira mountains” of Peru. Annalen des Naturhistorischen on tropical animals. Science, , –. Museums in Wien, B, –. TORTATO, M.A. & OLIVEIRA, T.G. () Ecology of the oncilla WATSON, J.E.M., EVANS, T., VENTER, O., WILLIAMS, B., TULLOCH, A., (Leopardus tigrinus) at Serra do Tabuleiro State Park, southern STEWART, C. et al. () The exceptional value of intact forest Brazil. CAT News, , –. ecosystems. Nature Ecology & Evolution, , –.