Register      Login
Animal Production Science Animal Production Science Society
Food, fibre and pharmaceuticals from animals
RESEARCH ARTICLE

Huemul heresies: beliefs in search of supporting data. 2. Biological and ecological considerations

Werner T. Flueck A B C D E and Jo Anne M. Smith-Flueck D
+ Author Affiliations
- Author Affiliations

A National Council of Scientific and Technological Research (CONICET), Rivadavia 1917, 1033 Buenos Aires, Argentina.

B Swiss Tropical and Public Health Institute, University of Basel, Socinstrasse 57, 4051 Basel, Switzerland.

C Fundación Bariloche, C.C. 592, 8400 Bariloche, Argentina.

D Institute of Natural Resources Analysis (IARN) – Patagonia, Universidad Atlántida Argentina, C.C. 592, 8400 Bariloche, Argentina.

E Corresponding author. Email: wtf@deerlab.org

Animal Production Science 52(8) 694-706 https://doi.org/10.1071/AN11345
Submitted: 12 December 2011  Accepted: 5 June 2012   Published: 11 July 2012

Abstract

Scarce information from remnant huemul (Hippocamelus bisulcus) populations in marginal habitats can lead to erroneous interpretations of the species’ natural history, such as assumptions of being a highly inflexible species. We evaluated discrepancies between historical accounts and recent interpretations regarding biological and ecological traits to better understand factors preventing recovery of highly endangered huemul. Early information supports the theory that huemul are currently living under suboptimal conditions. We find variability to be the norm for huemul, as with other cervids, in regard to antler characteristics, group size and density, sexual segregation, and social and feeding behaviours. No evidence supports competition and disease from livestock or red deer as having caused declines or preventing recovery. Instead, livestock management, particularly presence of people and dogs, creates incompatibilities. Where red deer are sympatric with huemul, red deer are outnumbered by livestock by 2100%, and being regularly inspected at slaughter, livestock provide a good proxy for diseases afflicting red deer. Inadequate antipredator responses due to evolutionary absence of cursorial predators are unsupported as several Canis species coexisted with huemul, overlapping with dogs that arrived with Paleoindians. Three populations have increased despite high predator density. Age at maturity for huemul is 1 year, with evidence that fawns may also breed. Reported twinning needs confirmation, but occurs in congeneric taruca (H. antisensis) and other Odocoilines, and huemul frequently raise fawns successfully every year; life cycle calculations should apply these parameters. Like taruca, dominance group breeding systems have been described repeatedly. Although huemul bucks were recently claimed unique by displaying territoriality year-round, data do not support such behaviour. Two sole dispersal records (8 and 15.5 km) are unlikely to represent maximum dispersal capacity and do not support barriers assumed from few kilometres of unsuitable habitat. Huemul using 500 ha could predictably disperse up to 90 km, well within the ranges of other cervids. Mistakenly assuming barriers and underestimating reproductive capacity may distract from discovering the factors affecting recolonisations. Sustained recovery may depend on re-establishing source populations on more productive habitats, guided by zooarcheological and historical data.


References

Allen JA (1905) ‘The Princeton University expeditions to Patagonia, 1896–1899. Volume 3. Zoology. Part 1. Mammalia of Southern Patagonia.’ (E. Schweizerbart’sche Verlagshandlung: Stuttgart)

Anderson AE, Wallmo OC (1984) Odocoileus hemionus. Mammalian Species 219, 1–9.
Odocoileus hemionus.Crossref | GoogleScholarGoogle Scholar |

Anon. (1904) Excursión del gobernador del Chubut. Caras y Caretas 7, 8

Atwood TC, Gese EM, Kunkel KE (2007) Comparative patterns of predation by cougars and recolonizing wolves in Montana’s Madison range. The Journal of Wildlife Management 71, 1098–1106.
Comparative patterns of predation by cougars and recolonizing wolves in Montana’s Madison range.Crossref | GoogleScholarGoogle Scholar |

Barberena R, Méndez C, Mena F, Reyes O (2011) Endangered species, archaeology, and stable isotopes: Huemul (Hippocamelus bisulcus) isotopic ecology in Central-Western Patagonia (South America). Journal of Archaeological Science 38, 2313–2323.
Endangered species, archaeology, and stable isotopes: Huemul (Hippocamelus bisulcus) isotopic ecology in Central-Western Patagonia (South America).Crossref | GoogleScholarGoogle Scholar |

Barrio J (2010) Taruka Hippocamelus antisensis. In ‘Neotropical Cervidology’. (Eds JM Barbanti Duarte, S González) pp. 77–88. (FUNEP: Jaboticabal, Brazil)

Bauer JW, Logan KA, Sweanor LL, Boyce WM (2005) Scavenging behavior in puma. The Southwestern Naturalist 50, 466–471.
Scavenging behavior in puma.Crossref | GoogleScholarGoogle Scholar |

Belardi JB, Otero JG (1998) Anatomia economica del Huemul (Hippocamelus bisulcus): una contribucion a la interpretacion de las evidencias arqueológicas de su aprovechamiento en Patagonia. Anales del Instituto Patagonico 26, 195–207.

Benedict JB (1999) Effects of changing climate on game-animal and human use of the Colorado High Country (USA) since 1000 BC. Arctic, Antarctic, and Alpine Research 31, 1–15.
Effects of changing climate on game-animal and human use of the Colorado High Country (USA) since 1000 BC.Crossref | GoogleScholarGoogle Scholar |

Benedict JB (2005) Tundra game drives: an Arctic-Alpine comparison. Arctic, Antarctic, and Alpine Research 37, 425–434.
Tundra game drives: an Arctic-Alpine comparison.Crossref | GoogleScholarGoogle Scholar |

Beninde J (1937) ‘Zur Naturgeschichte des Rothirsches.’ (Verlag Dr. Paul Schoeps: Leipzig, Germany)

Bowman J, Jaeger JA, Fahrig L (2002) Dispersal distance of mammals is proportional to home range size. Ecology 83, 2049–2055.
Dispersal distance of mammals is proportional to home range size.Crossref | GoogleScholarGoogle Scholar |

Brandborg SM (1955) ‘Life history and management of the mountain goat in Idaho.’ (State of Idaho Department of Fish and Game: Boise, ID)

Breitenmoser U, Haller H (1993) Patterns of predation by reintroduced European lynx in the Swiss alps. The Journal of Wildlife Management 57, 135–144.
Patterns of predation by reintroduced European lynx in the Swiss alps.Crossref | GoogleScholarGoogle Scholar |

Brinkman TJ, Deperno CS, Jenks JA, Haroldson BS, Osborn RG (2005) Movement of female white-tailed deer: effects of climate and intensive row-crop agriculture. The Journal of Wildlife Management 69, 1099–1111.
Movement of female white-tailed deer: effects of climate and intensive row-crop agriculture.Crossref | GoogleScholarGoogle Scholar |

Bubenik GA, Bubenik AB (1990) ‘Horns, pronghorns, and antlers.’ (Springer-Verlag: New York)

Cabrera A, Yepes J (1940) ‘Mamíferos sudamericanos.’ (Compañia Argentina de Editores: Buenos Aires)

Caughley G (1994) Directions in conservation biology. Journal of Animal Ecology 63, 215–244.
Directions in conservation biology.Crossref | GoogleScholarGoogle Scholar |

Codron D, Clauss M (2010) Rumen physiology constrains diet niche: linking digestive physiology and food selection across wild ruminant species. Canadian Journal of Zoology 88, 1129–1138.
Rumen physiology constrains diet niche: linking digestive physiology and food selection across wild ruminant species.Crossref | GoogleScholarGoogle Scholar |

Colomés AA (1978) Biología y ecología del huemul Chileno (Hippocamelus bisulcus). Estudio de sus habitos alimentarios. PhD Dissertation, Universidad de Chile, Santiago.

Corti P (2008) Organisation sociale, dynamique de population, et conservation du cerf huemul (Hippocamelus bisulcus) dans la Patagonie du Chili. PhD Dissertation, Université de Sherbrooke, Canada.

Corti P, Wittmer HU, Festa-Bianchet M (2010) Dynamics of a small population of endangered huemul deer (Hippocamelus bisulcus) in Chilean Patagonia. Journal of Mammalogy 91, 690–697.
Dynamics of a small population of endangered huemul deer (Hippocamelus bisulcus) in Chilean Patagonia.Crossref | GoogleScholarGoogle Scholar |

Corti P, Shafer AB, Coltman DW, Festa-Bianchet M (2011) Past bottlenecks and current population fragmentation of endangered huemul deer (Hippocamelus bisulcus): implications for preservation of genetic diversity. Conservation Genetics 12, 119–128.
Past bottlenecks and current population fragmentation of endangered huemul deer (Hippocamelus bisulcus): implications for preservation of genetic diversity.Crossref | GoogleScholarGoogle Scholar |

Cruz I, Munoz AS, Caracotche M (2010) A huemul (Hippocamelus bisulcus) antler artefact in archaeological deposits of the Atlantic coast. Implications for human mobility and species distribution. Magallania 38, 287–294.

De Nigris ME (2004) Guanaco and huemul in Patagonian hunter-gatherers diet. In ‘Zooarcheology of South America. BAR International Series 1298’. (Ed. GL Mengoni Gonalons) pp. 11–21. (Archeopress: Oxford, England)

DeMiguel D, Fortelius M, Azanza B, Morales J (2008) Ancestral feeding state of ruminants reconsidered: earliest grazing adaptation claims a mixed condition for Cervidae. BMC Evolutionary Biology 8, 13
Ancestral feeding state of ruminants reconsidered: earliest grazing adaptation claims a mixed condition for Cervidae.Crossref | GoogleScholarGoogle Scholar |

DeYoung RW, Demarais S, Gee KL, Honeycutt RL, Hellickson MW, Gonzales RA (2009) Molecular evaluation of the white-tailed deer (Odocoileus virginianus) mating system. Journal of Mammalogy 90, 946–953.
Molecular evaluation of the white-tailed deer (Odocoileus virginianus) mating system.Crossref | GoogleScholarGoogle Scholar |

Díaz NI, Smith-Flueck JM (2000) ‘The Patagonian huemul. A mysterious deer on the brink of extinction.’ (Literature of Latin America: Buenos Aires)

Dolman PM, Wäber K (2008) Ecosystem and competition impacts of introduced deer. Wildlife Research 35, 202–214.
Ecosystem and competition impacts of introduced deer.Crossref | GoogleScholarGoogle Scholar |

Fernández PM (2008) Taphonomy and zooarchaeology in the Neotropics: a view from northwestern Patagonian forest and steppe. Quaternary International 180, 63–74.
Taphonomy and zooarchaeology in the Neotropics: a view from northwestern Patagonian forest and steppe.Crossref | GoogleScholarGoogle Scholar |

Fernández OA, Busso CA (1997) Arid and semi-arid rangelands: two thirds of Argentina. In ‘Proceedings from an international workshop in Iceland. Rala Report No. 200’. (Eds O Arnalds, S Archer) pp. 41–60. (Agricultural Research Institute: Reykjavik, Iceland)

Flueck WT (2001) Body reserves and pregnancy rates of introduced red deer in Patagonia (Argentina) after a period of drought. Ecología Austral 11, 11–24.

Flueck WT (2004) Observations of interactions between Puma, Puma concolor, and introduced European Red Deer, Cervus elaphus, in Patagonia. Canadian Field Naturalist 118, 132–134.

Flueck WT (2010) Exotic deer in southern Latin America: what do we know about impacts on native deer and on ecosystems? Biological Invasions 12, 1909–1922.
Exotic deer in southern Latin America: what do we know about impacts on native deer and on ecosystems?Crossref | GoogleScholarGoogle Scholar |

Flueck WT, Smith-Flueck JM (2011a) Huemul heresies: beliefs in search of supporting data. 1. Historical and zooarcheological considerations. Animal Production Science 51, cxl–cliv. [Short Communication]

Flueck WT, Smith-Flueck JM (2011b) Huemul heresies: beliefs in search of supporting data. 2. Biological and ecological considerations. Animal Production Science 51, clv–clxix. [Short Communication]

Flueck WT, Smith-Flueck JM (2011c) Huemul heresies: beliefs in search of supporting data. 3. Reproduction. Animal Production Science 51, clxx–clxxix. [Short Communication]

Flueck WT, Smith-Flueck JM (2011d) Recent advances in the nutritional ecology of the Patagonian huemul: implications for recovery. Animal Production Science 51, 311–326.

Flueck WT, Smith-Flueck JM (2012a) Diseases of red deer introduced to Patagonia and implications for native ungulates. Animal Production Science 52, 766–773.
Diseases of red deer introduced to Patagonia and implications for native ungulates.Crossref | GoogleScholarGoogle Scholar |

Flueck WT, Smith-Flueck JM (2012b) Huemul heresies: beliefs in search of supporting data. 1. Historical and zooarcheological considerations. Animal Production Science 52, 685–693.
Huemul heresies: beliefs in search of supporting data. 1. Historical and zooarcheological considerations.Crossref | GoogleScholarGoogle Scholar |

Frid A (1994) Observations on habitat use and social organization of a huemul (Hippocamelus bisulcus) coastal population in Chile. Biological Conservation 67, 13–19.
Observations on habitat use and social organization of a huemul (Hippocamelus bisulcus) coastal population in Chile.Crossref | GoogleScholarGoogle Scholar |

Frid A (1999) Huemul (Hippocamelus bisulcus) sociality at a periglacial site: sexual aggregation and habitat effects on group size. Canadian Journal of Zoology 77, 1083–1091.

Fundación RA Philippi (2009) Mamiferos de Chile. Huemul. Available at http://www.fundacionraphilippi.cl [Verified 7 June 2012]

García K, Jara A, Ortiz JC, Victoriano P (2008) Evaluation of an extinction scenario for the last population of Hippocamelus bisulcus (Molina, 1782) in Central Chile. Interciencia 33, 152–159.

Gazzolo C (2006) Botanical composition of taruka (Hippocamelus antisensis) diet during rainy season in Huascaran national park, Peru. In ‘Advances in deer biology’. (Eds L Bartos, A Dusek, R Kotrba, J Bartosova) p. 216. (Research Institute of Animal Production: Praha, Czech Republic)

Geist V (1981) Behavior: adaptive strategies in Mule deer. In ‘Mule and black-tailed deer of North America’. (Ed. OC Wallmo) pp. 157–223. (University of Nebraska Press: Lincoln, NE)

Geist V (1998) ‘Deer of the world.’ (Stackpole Books: Mechanicsburg, PA)

Giai AG (1936) Huemul, inofensivo venado de las soledades cordilleranas de la Patagonia. La Chacra (Arg.) 6, 99–101.

Gigoux EE (1929) El huemul. Revista Chilena de Historia Natural (Valparaiso, Chile) 23, 573–582.

Gill R, Saucedo C, Montero E, Millacura R (2003) Ranging behavior and habitat use of huemul (Hippocamelus bisulcus). In ‘Huemul ecology research for conservation planning’. (Darwin Initiative: Cochrane, Chile)

Gill R, Saucedo C, Aldridge D, Morgan G (2008) Ranging behavior of huemul in relation to habitat and landscape. Journal of Zoology 274, 254–260.
Ranging behavior of huemul in relation to habitat and landscape.Crossref | GoogleScholarGoogle Scholar |

Grant JW, Chapman CA, Richardson KS (1992) Defended versus undefended home range size of carnivores, ungulates and primates. Behavioral Ecology and Sociobiology 31, 149–161.
Defended versus undefended home range size of carnivores, ungulates and primates.Crossref | GoogleScholarGoogle Scholar |

Grosse A (1949) El huemul – ciervo de los Andes y emblema del escudo Chileno. The Condor 12, 10–12. [Revista Chileno Alemana]

Guérin C, Faure M (2009) The Cervidae (Mammalia, Artiodactyla) of the Upper Pleistocene/Lower Holocene deposits of the Serra da Capivara National Park Region (Piaui, Brazil). Geobios 42, 169–195.

Guineo P, Guineo Garay R, Garay G (2008) ‘Conociendo al huemul de Torres del Paine.’ (La Prensa Austral: Punta Arenas, Chile)

Hatcher JB (1903) ‘Reports of the Princeton University expeditions to Patagonia, 1896–1899. Vol. I: Narrative of the Expeditions. Geography of Southern Patagonia.’ (E. Schweizerbart’sche Verlagshandlung: Stuttgart)

Hershkovitz P (1972) The recent mammals of the neotropical region: a zoogeographic and ecological review. In ‘Evolution, mammals, and southern continents’. (Eds A Keast, FC Erk, B Glass) pp. 311–431. (State University New York Press: Albany, New York)

Hjeljord O (2001) Dispersal and migration in northern forest deer – are there unifying concepts? Alces (Thunder Bay, Ont.) 37, 353–370.

Hobbs NT, Baker DL, Gill RB (1983) Comparative nutritional ecology of montane ungulates during winter. The Journal of Wildlife Management 47, 1–16.
Comparative nutritional ecology of montane ungulates during winter.Crossref | GoogleScholarGoogle Scholar |

Housse PR (1953) ‘Animales salvajes de Chile en su clasificación moderna: su vida y sus costumbres.’ (Ediciones de la Universidad de Chile: Santiago, Chile)

Iglesias RE (1965) El huemul. La Montaña (Arg.) 7, 26–28.

Jaksic FM, Iriarte JA, Jimenez JE, Martinez DR (2002) Invaders without frontiers: cross-border invasions of exotic mammals. Biological Invasions 4, 157–173.
Invaders without frontiers: cross-border invasions of exotic mammals.Crossref | GoogleScholarGoogle Scholar |

Jepsen JU, Topping CJ (2004) Modelling roe deer (Capreolus capreolus) in a gradient of forest fragmentation: behavioural plasticity and choice of cover. Canadian Journal of Zoology 82, 1528–1541.
Modelling roe deer (Capreolus capreolus) in a gradient of forest fragmentation: behavioural plasticity and choice of cover.Crossref | GoogleScholarGoogle Scholar |

Jiménez J, Guineo G, Corti P, Smith J, Flueck W, Vila A, Gizejewski Z, Gill R, McShea B, Geist V (2008) Hippocamelus bisulcus. In ‘2008 IUCN Red List of threatened species’. (International Union for Conservation of Nature: Gland, Switzerland) Available at www.iucnredlist.org/details/10054 [verified 7 June 2012]

Jungius H (1974) Beobachtungen am Weisswedelhirsch und an anderen Cerviden in Bolivien. Zeitschrift fur Saugetierkunde 39, 373–383.

Kjellander P, Hewison AJM, Liberg O, Angibault JM, Bideau E, Cargnelutti B (2004) Experimental evidence for density-dependence of home-range size in roe deer (Capreolus capreolus L.): a comparison of two long-term studies. Oecologia 139, 478–485.
Experimental evidence for density-dependence of home-range size in roe deer (Capreolus capreolus L.): a comparison of two long-term studies.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD2MrltFKgug%3D%3D&md5=c8ab9ca9870afa16745a1853d1e58366CAS |

Knopff KH, Knopff AA, Boyce MS (2010) Scavenging makes cougars susceptible to snaring at wolf bait stations. The Journal of Wildlife Management 74, 644–653.
Scavenging makes cougars susceptible to snaring at wolf bait stations.Crossref | GoogleScholarGoogle Scholar |

Kokko H, Lopez-Sepulcre A (2006) From individual dispersal to species ranges: perspectives for a changing world. Science 313, 789–791.
From individual dispersal to species ranges: perspectives for a changing world.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28XnvVyntr8%3D&md5=b2d5f5ccfac5f75052291f4646a2be36CAS |

Krieg H (1940) ‘Als Zoologe in Steppen und Wäldern Patagoniens.’ (J.F. Lehmanns Verlag: München, Germany)

Kurtén B (1975) A new pleistocene genus of American mountain deer. Journal of Mammalogy 56, 507–508.
A new pleistocene genus of American mountain deer.Crossref | GoogleScholarGoogle Scholar |

Leiby PD, Dyer WG (1971) Cyclophyllidean tapeworms of wild carnivores. In ‘Parasitic diseases of wild mammals’. (Eds JW Davis, RC Anderson) pp. 174–234. (Iowa State University Press: Ames, IA)

Lever C (1985) ‘Naturalized mammals of the world.’ (Longman Inc.: New York)

Lord RD (2007) ‘Mammals of South America.’ (JHU Press: Baltimore, MD)

Madsen A (1948) ‘Patagonia Vieja.’ (Zagier & Urruty: Buenos Aires)

Magne de la Croix P (1937) El huemul. Caras y Caretas 40, 117

Maher CR, Lott DF (1995) Definitions of territoriality used in the study of variation in vertebrate spacing systems. Animal Behaviour 49, 1581–1597.
Definitions of territoriality used in the study of variation in vertebrate spacing systems.Crossref | GoogleScholarGoogle Scholar |

McCallum H, Dobson A (2002) Disease, habitat fragmentation and conservation. Proceedings. Biological Sciences 269, 2041–2049.
Disease, habitat fragmentation and conservation.Crossref | GoogleScholarGoogle Scholar |

Miller S, Rottman J, Raedeke K, Taber RD (1983) Endangered mammals of Chile: status and conservation. Biological Conservation 25, 335–352.
Endangered mammals of Chile: status and conservation.Crossref | GoogleScholarGoogle Scholar |

Molinari-Jobin A, Molinari P, Breitenmoser-Wursten C, Breitenmoser U (2002) Significance of lynx Lynx lynx predation for roe deer Capreolus capreolus and chamois Rupicapra rupicapra mortality in the Swiss Jura Mountains. Wildlife Biology 8, 109–115.

Murillo J, Ramb AM (1975) ‘Der Kampf der Andenhirsche.’ (K. Thienemann Verlag: Stuttgart, Germany)

Navas JR (1987) Los vertebrados exoticos introducidos en la Argentina. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia 14, 7–38.

Novaro AJ, Moraga CA, Briceno C, Funes MC, Marino A (2009) First records of culpeo (Lycalopex culpaeus) attacks and cooperative defense by guanacos (Lama guanicoe). Mammalia 73, 148–150.
First records of culpeo (Lycalopex culpaeus) attacks and cooperative defense by guanacos (Lama guanicoe).Crossref | GoogleScholarGoogle Scholar |

Nowak MC, Taylor TE, Witmer GW (2000) Prolonged scavenging by a female mountain lion in Northeastern Oregon. Northwestern Naturalist (Olympia, Wash.) 81, 63–65.
Prolonged scavenging by a female mountain lion in Northeastern Oregon.Crossref | GoogleScholarGoogle Scholar |

Nowak RM, Walker EP (1999) ‘Walker’s mammals of the world.’ (JHU Press: Baltimore, MD)

Nugent G, McShea WJ, Parkes J, Woodley S, Waithaka J, Moro J, Gutierrez R, Azorit C, Mendez Guerrero F, Flueck WT, Smith-Flueck JM (2011) Policies and management of overabundant deer (native or exotic) in protected areas. Animal Production Science 51, 384–389.

Osgood WH (1923) The Marshall Chilean Field Expedition 1922–1923. Field notes. Available at https://sites.google.com/a/fieldmuseum.org/bruce-pattersons-lab/Home/collections/osgood-files [verified 7 June 2012]

Ozaki M, Suwa G, Kaji K, Ohba T, Hosoi E, Koizumi T, Takatsuki S (2007) Correlations between feeding type and mandibular morphology in the sika deer. Journal of Zoology 272, 244–257.
Correlations between feeding type and mandibular morphology in the sika deer.Crossref | GoogleScholarGoogle Scholar |

Packard FM (1947) A study of the deer and elk herds of Rocky Mountain National Park, Colorado. Journal of Mammalogy 28, 4–12.
A study of the deer and elk herds of Rocky Mountain National Park, Colorado.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaH2s%2FlslCqtQ%3D%3D&md5=69ad3b163b0ce398c39ea8d7ab5802e9CAS |

Pérez W, Clauss M, Ungerfeld R (2008) Observations on the macroscopic anatomy of the intestinal tract and its mesenteric folds in the Pampas deer (Ozotoceros bezoarticus, Linnaeus 1758). Anatomia, Histologia, Embryologia 37, 317–321.
Observations on the macroscopic anatomy of the intestinal tract and its mesenteric folds in the Pampas deer (Ozotoceros bezoarticus, Linnaeus 1758).Crossref | GoogleScholarGoogle Scholar |

Philippi RA (1892) El guemul de Chile. Anales del Museo Nacional de Chile 2, 1–9.

Philippi RA (1895) ‘Drei Hirsche der Anden.’ (F.A. Brockhaus: Leipzig, Germany)

Pinto AA (2004) Foot-and-mouth disease in tropical wildlife. Annals of the New York Academy of Sciences 1026, 65–72.
Foot-and-mouth disease in tropical wildlife.Crossref | GoogleScholarGoogle Scholar |

Poole KG, Bachmann KD, Teske IE (2010) Mineral lick use by GPS radio-collared mountain goats in Southeastern British Columbia. Western North American Naturalist 70, 208–217.
Mineral lick use by GPS radio-collared mountain goats in Southeastern British Columbia.Crossref | GoogleScholarGoogle Scholar |

Povilitis A (1978) The Chilean Huemul Project – a case history (1975–1976). In ‘Threatened deer’. pp. 109–128. (International Union for Conservation of Nature: Morges, Switzerland)

Povilitis A (1983) The huemul in Chile: national symbol in jeopardy? Oryx 17, 34–40.
The huemul in Chile: national symbol in jeopardy?Crossref | GoogleScholarGoogle Scholar |

Povilitis A (1985) Social behavior of the huemul (Hippocamlus bisulcus) during the breeding season. Zeitschrift für Tierpsychologie 68, 261–286.
Social behavior of the huemul (Hippocamlus bisulcus) during the breeding season.Crossref | GoogleScholarGoogle Scholar |

Prichard HH (1902a) Field notes upon some of the larger mammals of Patagonia made between September 1900 and June 1901. Proceedings of the Zoological Society of London 1, 272–277.

Prichard HH (1902b) ‘Through the heart of Patagonia.’ (D. Appleton and Co.: New York)

Prichard HH (1910) ‘Hunting camps in wood and wilderness.’ (William Heinemann: London)

Putman R, Flueck WT (2011) Intraspecific variation in biology and ecology of deer: magnitude and causation. Animal Production Science 51, 277–291.
Intraspecific variation in biology and ecology of deer: magnitude and causation.Crossref | GoogleScholarGoogle Scholar |

Quay WB, Müller-Schwarze D (1970) Functional histology of integumentary glandular regions in black-tailed deer (Odocoileus hemionus columbianus). Journal of Mammalogy 51, 675–694.
Functional histology of integumentary glandular regions in black-tailed deer (Odocoileus hemionus columbianus).Crossref | GoogleScholarGoogle Scholar |

Quay WB, Müller-Schwarze D (1971) Relations of age and sex to integumentary glandular regions in Rocky Mountain mule deer (Odocoileus hemionus hemionus). Journal of Mammalogy 52, 670–685.
Relations of age and sex to integumentary glandular regions in Rocky Mountain mule deer (Odocoileus hemionus hemionus).Crossref | GoogleScholarGoogle Scholar |

Redford KH, Eisenberg JF (1992) ‘Mammals of the Neotropics: the southern cone. Volume 2. Chile, Argentina, Uruguay, Paraguay.’ (The University of Chicago Press: Chicago)

Reed DF (1981) Conflicts with civilization. In ‘Mule and black-tailed deer of North America’. (Ed. OC Wallmo) pp. 509–535. (University of Nebraska Press: Lincoln, NE)

Rottmann JS (2003) Experiencias con huemules en cautividad. In ‘4ta Reunión Chileno-Argentina sobre Estrategias de Conservación del Huemul’. (Ed. G Acosta-Jamett) pp. 40–42. (CONAF and CODEFF: Las Trancas, Chile)

Rusconi C (1936) Anomalias en las cornamentas del huemul. Anales Sociedad Cientifica Argentina 122, 288–296.

Saizar P (1987) La estacion de cría del pudu. Vida Silvestre 5, 56–58.

Saucedo C, Gill R (2004) The endangered huemul or south Andean deer Hippocamelus bisulcus. Oryx 38, 132–133.

Serrano Ferron E, Verheyden H, Hummel J, Cargnelutti B, Lourtet B, Merlet J, Gonzalez-Candela M, Angibault JM, Jonathan A, Hewison M, Clauss M (2012) Digestive plasticity as a response to woodland fragmentation in roe deer. Ecological Research 27, 77–82.
Digestive plasticity as a response to woodland fragmentation in roe deer.Crossref | GoogleScholarGoogle Scholar |

Serret A, Borghiani F (1997) Registros de avistajes y comportamiento de huemules en el seno Moyano, Parque Nacional Los Glaciares. Boletin Técnico Fundación Vida Silvestre Argentina 35, 1–24.

Sierralta D (2003) La microhistología de fecas para el estudio de dieta del huemul. In ‘Huemul ecology research for conservation planning’. (Darwin Initiative: Cochrane, Chile)

Simonetti JA (1995) Wildlife conservation outside parks is a disease-mediated task. Conservation Biology 9, 454–456.
Wildlife conservation outside parks is a disease-mediated task.Crossref | GoogleScholarGoogle Scholar |

Smith-Flueck JM (2003) The ecology of huemul (Hippocamelus bisulcus) in Andean Patagonia of Argentina and considerations about its conservation. Doctoral Dissertation, Universidad Nacional Comahue, Bariloche, Argentina.

Smith-Flueck JM, Flueck WT (2001) Natural mortality patterns in a population of southern Argentina huemul (Hippocamelus bisulcus), an endangered Andean cervid. European Journal of Wildlife Research 47, 178–188.

Smith-Flueck JM, Barrio J, Ferreyra N, Nuñez A, Tomas N, Guzman J, Flueck WT, Hinojosa A, Vidal F, Garay G, Jimenez J (2011) Advances in ecology and conservation of Hippocamelus species in South America. Animal Production Science 51, 378–383.
Advances in ecology and conservation of Hippocamelus species in South America.Crossref | GoogleScholarGoogle Scholar |

Stephens PA, Sutherland WJ (1999) Consequences of the Allee effect for behaviour, ecology and conservation. Trends in Ecology & Evolution 14, 401–405.
Consequences of the Allee effect for behaviour, ecology and conservation.Crossref | GoogleScholarGoogle Scholar |

Steward JH (1946) Handbook of South American Indians. Volume 1. The marginal tribes. (Smithsonian Institution: Washington, D.C.)

Stopher KV, Nussey DH, Clutton-Brock TH, Guinness F, Morris A, Pemberton JM (2011) The red deer rut revisited: female excursions but no evidence females move to mate with preferred males. Behavioral Ecology 22, 808–818.
The red deer rut revisited: female excursions but no evidence females move to mate with preferred males.Crossref | GoogleScholarGoogle Scholar |

Texera WA (1974) Algunos aspectos de la biología del huemul (Hippocamelus bisulcus) (Mammalia: Artiodactyla, Cervidae) en cautividad. Anales del Instituto Patagonico 5, 155–188.

Tufto J, Andersen R, Linnell J (1996) Habitat use and ecological correlates of home range size in a small cervid: the roe deer. Journal of Animal Ecology 65, 715–724.
Habitat use and ecological correlates of home range size in a small cervid: the roe deer.Crossref | GoogleScholarGoogle Scholar |

Uhart M, Chang Reisig E (2006) Ungulados exoticos: amenaza sanitaria para el huemul? In ‘5ta Reunión Binacional Argentino-Chilena sobre Estrategias de Conservación del Huemul’. pp. 25–27. (Adm. Parques Nacionales: Bariloche, Argentina)

Uribe LS (2011) Los Dominios de ‘Silencio’ en el Baker. Informativo: Coordinación Nacional Del Plan de Conservación del Huemul Chile 4, 1–22.

Urrutia AB, Ojeda AA (2008) ‘Manual de Plagas y Enfermedades del Bosque Nativo en Chile.’ (Editora e Imprenta Maval Ltda: Santiago, Chile)

Vidal F, Smith-Flueck JM, Flueck WT, Arias E (2011) The Patagonian huemul (Hippocamelus bisulcus) under captive conditions: past and current experiences. Animal Production Science 51, 340–350.
The Patagonian huemul (Hippocamelus bisulcus) under captive conditions: past and current experiences.Crossref | GoogleScholarGoogle Scholar |

Vila C, Sundqvist AK, Flagstad O, Seddon J, Bjornerfeldt S, Kojola I, Casulli A, Sand H, Wabakken P, Ellegren H (2003) Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant. Proceedings. Biological Sciences 270, 91–97.
Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant.Crossref | GoogleScholarGoogle Scholar |

Vila AR, Saucedo C, Aldridge D, Ramilo E, Corti P (2010) South Andean huemul Hippocamelus bisulcus (Molina 1782). In ‘Neotropical Cervidology’. (Eds JM Duarte, S González) pp. 89–100. (FUNEP: Jaboticabal, Brazil)

von Thüngen J, Lanari MR (2010) Profitability of sheep farming and wildlife management in Patagonia. Pastoralism 1, 274–290.

Wang J (2004) Application of the One-Migrant-per-Generation rule to conservation and management. Conservation Biology 18, 332–343.
Application of the One-Migrant-per-Generation rule to conservation and management.Crossref | GoogleScholarGoogle Scholar |

Webb SD (2000) Evolutionary history of New World Cervidae. In ‘Antelopes, deer, and relatives’. (Eds ES Vrba, GB Schaller) pp. 38–64. (Yale University Press: New York)

Wells KW, Stangl FB (2003) Superior size and antler development in populations of white-tailed deer (Odocoileus virginianus) from the North Texas rolling plains. The Texas Journal of Science 55, 337–346.

Wemmer C (1998) ‘Deer: status survey and conservation action plan.’ (IUCN/SSC Deer Specialist Group: Gland, Switzerland)

Wensing D (2005) Conservation study of the huemul (Hippocamelus bisculcus) within the Bernardo O’Higgins National Park, Chile. Thesis report NWS-I-2005–6, Utrecht University, Utrecht, The Netherlands.

Whitehead GK (1993) ‘The Whitehead encyclopedia of deer.’ (Swan Hill Press: Shrewsbury, England)

Williams D (2010) Bighorn success. New Mexico Wildlife 55, 1–15.

Willis B (1914) ‘El norte de la Patagonia.’ (Ministerio de Obras Publicas: Buenos Aires)

Wittmer HU, Corti P, Saucedo C, Galaz JL (2010) Learning to count: adapting population monitoring for endangered huemul deer (Hippocamelus bisulcus) to meet conservation objectives. Oryx 44, 516–522.
Learning to count: adapting population monitoring for endangered huemul deer (Hippocamelus bisulcus) to meet conservation objectives.Crossref | GoogleScholarGoogle Scholar |