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

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.Fil: Fluck, Werner Thomas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata; Argentina. Universidad de Basilea; Suiza. Universidad Atlantida Argentina; Argentina. Fundación Bariloche; ArgentinaFil: Smith-Flueck, Jo Anne M.. Universidad Atlantida Argentina; Argentin

A review of introduced cervids in Chile

We review the extent of exotic deer distributions in Chile, which are encountered in all provinces, including Tierra del Fuego, except for possibly Region III; many deer are contained in at least 107 enclosures. Red deer (Cervus elaphus) by far has the largest feral population of exotic cervids in southern South America, providing source animals that can easily cross the Andes between Chile and Argentina. Red deer was introduced from Europe to the central valley of Chile in 1928. Since the 1940s, feral populations have also expanded from Argentina into Chile, by way of easily accessible, low-elevation mountain passes of the Andes, accompanied by further direct shipments from Argentina. The area occupied by 1990 was estimated at 3400km2, whereas an analysis in 2003 estimated an area of 7700km2. The overall area invaded by 2003 was between 3742'S and 5455'S, and 7336'W and 6950'W (Argentina and Chile combined, although non-contiguous). Ecological impact of the red deer in Chile has been described since 1981, and red deer features in the Chilean Pest Manual. A conservative rate for the red deer expansion was estimated at 1km/year, but likely is more rapid where habitat modifications facilitate movement. The pre-Columbian northern limit of the native cervid huemul (Hippocamelus bisulcus) was 30S, and because red deer has occupied all habitat types currently used by huemul, it could thus spread 750km further north. To the south, all areas are suitable for red deer. Invasion patterns will depend on additional intentional introductions and enclosures on both sides of the Andes because of the omnipresent risk of escapes. Fallow, axis and roe deer (Dama dama, Axis axis, Capreolus capreolus, respectively) also have been introduced to Chile and occur in many enclosures. Fallow deer recently escaped on Chiló Island, became established and raised concerns because of its potential impacts on several endemic species on the island. The striking lack of information on feral deer may relate to policies and laws about firearms and restricted access to hunting areas, resulting in the apparent absence of popular hunting, which, nevertheless, could be a potential tool should the invasion continue and lead to future deer overabundance.Fil: Fluck, Werner Thomas. Universidad Atlantida Argentina; Argentina. Fundación Bariloche; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; ArgentinaFil: Smith-Flueck, Jo Anne M.. Universidad Atlantida Argentina; Argentin

Marcando con radio collar un grupo del amenazado huemul patagónico por primera vez en Argentina

En Argentina quedan 350-500 huemules en peligro de extinción. Se marcaron huemules con radio-collares durante 6 días de invierno. El encuentro de un huemul cada día promedió 93 minutos (17 grupos, 37 individuos). El período de tiempo desde la detección hasta la inmovilización promedió 46 minutos; la distancia de tiro promedió 15.7 m; y desde la inmovilización hasta recuperación promedió 45.5 minutos. Se realizaron 10 intentos de captura, de los cuales seis fueron exitosos. Comparado con la única radio-marcación anterior en Argentina, esta operación requirió 96.5% menos días-hombre/animal. De los huemules marcados, 86% presentaban patologías clínicas, posiblemente por estar restringidos a áreas marginales o sumideros, es decir, trampas ecológicas. Esta operación permitió marcar por primera vez un macho y un grupo de seis huemules en Argentina.In Argentina, 350-500 endangered huemul remain. Huemul were radio-collared in Chubut during six winter days. Encountering huemul each day averaged 93 minutes (17 groups, 37 individuals). Spotting until darting an animal averaged 46 minutes; darting distance averaged 15.7 m; and darting until recovery averaged 45.5 minutes. Ten capture attempts were made, of which six were successful. Compared to the only huemul previous radio-collared in Argentina (2016), the present operation required 96.5% less man-days/animal. Of marked huemul, 86% presented clinical pathologies, best explained by restriction to marginal if not sink areas, or ecological traps. This operation allowed the first-ever marking of huemul bucks in Argentina, and a group of six huemul.Fil: Fluck, Werner Thomas. Administración de Parques Nacionales; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad de Basilea; SuizaFil: Smith Flueck, Jo Anne M.. Universidad Nacional del Comahue; Argentin

Huemul heresies: Beliefs in search of supporting data. 1. Historical and zooarcheological considerations

Patagonian huemul (Hippocamelus bisulcus) was eliminated from many former ranges before arrival of early explorers who already acknowledged its rareness. Considering huemul analogous to ungulates in mountains back home, huemul was called mountain deer, which is repeatedly cited without validation and remains the orthodox interpretation. Yet other species considered analogous also use lowlands, flatlands, deserts and grasslands, and the only congeneric, H. antisensis, uses habitat with high affinity to Patagonian grasslands. Recent comparative analyses of the post-cranial morphology show that huemul cannot be associated with rock-climbing species, but falls within ranges of other cervids. Interpretations of past human utilisation rely on one study of economic anatomy and bone remains, frequently concluding that huemul was unimportant to hunter-gatherers. However, considering only bone fat and omitting easily removable fat is erroneous. Total energy of deer in autumnwinter partitions into 53% as fat and hunter-gatherers elsewhere focussed on deer during the peak fat cycle, using all easily removable fat (1200% more energy than in bones) and consuming fat and marrow while butchering. Natives are likely to have influenced huemul distribution and density in winter ranges due to high incentives (fat) and easiness to kill. Sparse evidence is likely the result of surveys in Patagonia having been biased towards caves, leaving out transient movements and camps. Generalist cervids passed the Panama land-bridge filter to reach South America. Considering paleoclimate, Hippocamelus dispersed east of the Andes, pushed by glaciations even to north-eastern Brazil. Hippocamelus would reach and cross Andes only after deglaciations. As a mixed feeder, huemul utilised Patagonian steppe. Hunter-gatherers arrived after the last glaciation and influenced the local distribution of Hippocamelus, especially in northern and central Chile, after early adoption of an agricultural lifestyle. Introduction of horses converted native economies through adoption of an equestrian lifestyle and arrival of millions of introduced livestock, which thus affected early writings. Only few records indicate the presence of large groups of huemul far from forests, and substantial killings. Human-caused range contractions of northern ungulates affected mainly losses at low elevations and most species persisted in the marginal periphery, including high-elevation refuges. Paleoecology, zoogeography and land-use history in southern South America indicate that mountain huemul is a secondary relict created by impacts of post-Columbian colonisation. We caution against the rigid application of modern huemul habitats in interpreting past habitat use and huemul ecology, and simply considering the few extra-Andean accounts as abnormal outliers.Fil: Fluck, Werner Thomas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata; Argentina. Universidad de Basilea; Suiza. Universidad Atlantida Argentina; Argentina. Fundación Bariloche; ArgentinaFil: Smith-Flueck, Jo Anne M.. Universidad Atlantida Argentina; Argentin

Variation in reproduction of a temperate deer, the southern pudu (Pudu puda)

Pudu (Pudu puda), occurring in the southern cone of Latin America, has been classified as vulnerable by the International Union for Conservation of Nature (IUCN), yet little is known about this animal in the wild, with most knowledge on the breeding behaviour coming from captive animals. For this second-smallest deer in the world, delayed implantation has been suggested to explain the two peaks in the annual cycle of male sexual hormones on the basis of the accepted tenet that the breeding period occurs only once a year, between March and June. However, in the present study, birth dates from fawns born at the Los Canelos semi-captive breeding centre in Chile and male courting behaviour revealed the possibility of two rutting periods: autumn and spring. To our knowledge, this is the first time that late-fall births (May through early June for 17% of fawns in the study population) have been recorded for the southern pudu; two of these four births were conceived by females in the wild. From zoo and captive-animal birth records (n = 97), only three fawns were born in the fall. For all births combined (n = 121), 77% occurred in spring. The roe deer (Capreolus capreolus) and Pere David deer (Elaphurus davidianus) have been considered the only two temperate cervids in which sexual activity is initiated by increasing daylength and which breed in early summer. Yet, the present results indicate a similar response from the southern pudu when under a wild or semi-captive environment, with breeding taking place in spring. These results suggest that this species may either have two reproductive periods per year or retains the capacity to be a breeder for a much more extended period of time than documented by earlier studies. Pudu, like other temperate deer, is responsive to photoperiod for timing its breeding period, but may further optimise its production of offspring by also responding to other environmental cues such as seasonal variation in food supply when climatic conditions are favourable.Fil: Vidal, Fernando. Universidad de La Frontera; ChileFil: Smith-Flueck, Jo Anne M.. Universidad Atlantida Argentina; ArgentinaFil: Fluck, Werner Thomas. Fundación Bariloche; Argentina. Universidad Atlantida Argentina; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; ArgentinaFil: Bartos, Ludek. No especifíca

Loss of migratory traditions makes the endangered patagonian Huemul deer a year-round refugee in its summer habitat

The huemul (Hippocamelus bisulcus) is endangered, with 1500 deer split into >100 subpopulations along 2000 km of the Andes. Currently occupied areas are claimed-erroneously-to be critical prime habitats. We analyzed historical spatiotemporal behavior since current patterns represent only a fraction of pre-Columbian ones. Given the limited knowledge, the first group (n = 6) in Argentina was radio-marked to examine spatial behavior. Historically, huemul resided year-round in winter ranges, while some migrated seasonally, some using grasslands >200 km east of their current presence, reaching the Atlantic. Moreover, huemul anatomy is adapted to open unforested habitats, also corroborated by spotless fawns. Extreme naivety towards humans resulted in early extirpation on many winter ranges—preferentially occupied by humans, resulting in refugee huemul on surrounding mountain summer ranges. Radio-marked huemul remained in small ranges with minimal altitudinal movements, as known from other subpopulations. However, these resident areas documented here are typical summer ranges as evidenced by past migrations, and current usage for livestock. The huemul is the only cervid known to use mountain summer ranges year-round in reaction to anthropogenic activities. Losing migratory traditions is a major threat, and may explain their presently prevalent skeletal diseases, reduced longevity, and lacking recolonizations for most remaining huemul subpopulations.Fil: Fluck, Werner Thomas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte; Argentina. Universidad de Basilea; Suiza. Administración de Parques Nacionales; ArgentinaFil: Smith Flueck, Jo Anne M.. Universidad Nacional del Comahue; Argentina. Parque Protegido Shoonem; Argentina. Deer Lab; ArgentinaFil: Escobar, Miguel E.. Parque Protegido Shoonem; ArgentinaFil: Zuliani, Melina Elizabeth. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte; Argentina. Fundación Bariloche; ArgentinaFil: Fuchs, Beat. Deer Lab; ArgentinaFil: Geist, Valerius. University of Calgary; CanadáFil: Heffelfinger, James R.. Arizona Game and Fish Department; Estados UnidosFil: Black de Decima, Patricia Ann. Universidad Nacional de Tucumán. Facultad de Ciencias Naturales e Instituto Miguel Lillo; ArgentinaFil: Gizejewski, Zygmunt. Polish Academy of Sciences; ArgentinaFil: Vidal, Fernando. Univerdidad Santo Tomas; Chile. Centro de Conservacion y Manejo de Vida Silvestre; ChileFil: Barrio, Javier. Centro de Ornitología y Biodiversidad; PerúFil: Molinuevo, María Silvina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Agrarias y Forestales. Departamento de Ciencias Biológicas; ArgentinaFil: Monjeau, Jorge Adrian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte; Argentina. Fundación Bariloche; ArgentinaFil: Hoby, Stefan. Berne Animal Park; SuizaFil: Jiménez, Jaime M.. University of North Texas; Estados Unido

Troubling disease syndrome in endangered live Patagonian huemul deer (Hippocamelus bisulcus) from the Protected Park Shoonem: unusually high prevalence of osteopathology

Abstract Objective The last 1500 endangered Patagonian huemul deer (Hippocamelus bisulcus) exist in > 100 groups which are not recovering. Prevalence of osteopathology in dead huemul was 57+% (Argentina), whereas similar cases in Chile were accompanied by selenium deficiency. The first clinical cases from live wild huemul confirm widespread osteopathology which explains short life spans, low recruitment, and thus absence of population recovery. Results The first-ever radio-collaring of 3 male huemul in Argentina and 3 females, plus a fresh female carcass allowed examination of 7 huemul. Of these, 86% were diseased and clinical pathophysiognomy included lameness, affected hoof, exfoliation of 2–7 incisors, other cranial osteopathologies, and muscle atrophy. The parsimonious explanation for absent population recovery is high prevalence of osteopathology as evidenced earlier in carcasses, and now by these clinical cases. Areas currently used by huemul have reduced selenium bioavailability, very deficient soil levels, and overt selenium deficiency in local livestock and plants. These areas are known to result in primary iodine deficiency which is aggravated by selenium deficiency. The nexus to nutritional ecology of huemul likely is inaccessibility to most fertile lowlands and traditional winter ranges, elimination of migratory traditions, and concomitant elimination of source populations

Diseases of red deer introduced to Patagonia and implications for native ungulates

Abstract. The red deer (Cervus elaphus) invasion in Patagonia has been continuing for nearly a century, with occurrence in all habitats between 34 S and 55 S. Their distribution, movement patterns and locally high densities raise concerns over their potential epidemiological role in maintaining disease reservoirs or transmitting diseases such as foot-and-mouth disease or tuberculosis, with potential severe health and economical impacts at the interface of humans, livestock or native wildlife. Among adult females collected by rifle and radio-collared deer that died naturally, no ectoparasites were found (n = 73). Fasciola hepatica was encountered in three surveys at prevalences ranging from 9% to 50% (n = 108). Taenia ovis krabbei was identified, and Cysticerus tenuicollis was found at a prevalence of 8% (n = 12). Ostertagia sp., Bunostomum sp. and Dictyocaulus sp. had a prevalence of 75%, 25% and 13% (n = 9), respectively. Several gastrointestinal parasites reported at low prevalence in endangered Patagonian huemul (Hippocamelus bisulcus) are common in livestock and considered commensals in domestic ruminants. Sympatry of huemul with livestock is commonplace, whereas with red deer it occurs in <2% of known populations, in which case there were 1.2 red deer, but 25.2 livestock per huemul, making livestock the determining epidemiological factor regarding disease transmission or reservoir. As red deer have been coexisting with livestock for >100 years in Argentina, both red deer and livestock play epidemiological roles for shared diseases. Research, conservation and management efforts should be directed towards livestock herd health programs or restriction of free livestock movements, particularly if diseases are shown to have an impact on recruitment of endangered natives. Livestock are routinely researched and inspected at slaughter and thus provide a proxy for diseases afflicting co-existing ungulates. Testing for antibodies to foot-and-mouth disease viral antigen was negative (n = 41). A tentative diagnosis of mycobacterial infection was based on typical visceral lesions. Antler damage occurred on 73% of shed antlers, with 36% having major breaks of tines and main beams, possibly indicating mineral imbalances. One male had both antlers, including pedicles with portions of frontal, parietal and occipital bones, broken off the skull, causing his death. The prevalence of 0.9% of campylognathia (n = 776) indicates that the disease is unlikely to be inheritable, because the founding stock of 20 animals would have had a prevalence of at least 5%. Among deer, handedness of scoliosis related significantly to the hemisphere where specimens originated (P < 0.001, n = 131). Coriolis forces are known to affect early stages of development, such as the innervation pattern of the mammalian vestibular system, or the plane of bilateral symmetry. It is, therefore, conceivable that the networks processing these environmental cues, or the mechanisms responsible for compensation, are malfunctioning and thus result in a preponderance of facial scoliosis in accordance to the earth's rotation