Reassessment of morphology and historical
distribution as factors in conservation efforts for the Endangered Patagonian
Huemul Deer Hippocamelus bisulcus (Molina 1782)
Huemul Task Force *
International
Union for Conservation of Nature (IUCN), Species Survival Commission (SSC), C\o
Chair, C.C. 592, 8400 Bariloche, Argentina,
Email: HTF@deerlab.org
Date of publication
(online): 26 November 2012
Date of publication (print): 26 November 2012
ISSN 0974-7907 (online) | 0974-7893 (print)
Editor: Norma Chapman
Manuscript details:
Ms # o3088
Received 30 January 2011
Final received 22 October 2012
Finally accepted 24 October 2012
Citation: Huemul Task Force (2012). Reassessment of morphology and historical distribution as factors
in conservation efforts for the Endangered Patagonian Huemul Deer Hippocamelus
bisulcus (Molina 1782). Journal of Threatened
Taxa 4(14): 3302–3311.
Copyright: © Huemul Task Force
2012. Creative Commons Attribution 3.0 Unported License.JoTT allows unrestricted use of this article in any medium for non-profit
purposes, reproduction and distribution by providing adequate credit to the
authors and the source of publication.
Acknowledgements: The
Huemul Task Force would like to thank the non-author group members: Paulo
Corti, Robin Gill, William McShea, and Cristian Saucedo.
* Individual authors: Eduardo G. Aisen, Fernando Vidal,
Gladys Garay, Jaime E. Jiménez, Jo Anne Smith-Flueck, Norberto Tomas, Patricia
Black de Decima, Valerius Geist, Werner Flueck, Zygmunt Gizejewski.
* Individual authors: contact details
Eduardo G. Aisen
Lab. Theriogenology, Universidad Nacional del Comahue, IDEPA
(CONICET-UNCo), c.c. 85
8303 Cinco Saltos, Rio Negro
Email: eduardoaisen@hotmail.com
Patricia Black de Decima
Instituto Miguel Lillo, Miguel Lillo 205, 4000 San Miguel de
Tucuman, Argentina
Email: patriciablack_decima@hotmail.com,
pblack@csnat.unt.edu.ar
Werner Flueck
National Council of Scientific and Technological Research
(CONICET), Buenos Aires, Swiss Tropical and Public Health Institute, University
Basel, C.C. 592, 8400 Bariloche, Argentina
Email: werner.flueck@unibas.ch
Jo Anne Smith-Flueck
Institute of Natural Resources Analysis - Patagonia,
Universidad Atlantida Argentina, C.C. 592, 8400 Bariloche, Argentina
Email: j.smith@deerlab.org (corresponding author)
Norberto Tomas
Parque Nacional Calilegua, San Lorenzo s/nº
4514 Calilegua, Jujuy, Argentina
Email: jnorbertotomas@yahoo.com.ar, ntomas@apn.gov.ar
Gladys Garay
Paraguaya 126, Barrio San Miguel, Punta Arenas, Chile
Email: gladysenviaje@yahoo.com
Jaime E. Jiménez
University of North Texas, 1155 Union Circle
Denton, Texas 76203-5017, USA
Omora
Ethnobotanical Park, Universidad de Magallanes, Puerto Williams, Chile
Email: jaime.jimenez@unt.edu
Fernando Vidal
Fundación Huilo-Huilo, Vitacura 2909 Of.
1112
Las Condes, Santiago, Chile
Email: fauna.andina@gmail.com
Valerius Geist
PO Box 1294, Station A, Port Alberni, BC, V9Y 7M2, Canada
Email: kendulf@shaw.ca
Zygmunt
Gizejewski
Polish
Academy of Sciences, Institute of Animal Reproduction and Food Research,
Pl-10-747
Olsztyn, Poland
Email: z.gizejewski@pan.olsztyn.pl
urn:lsid:zoobank.org:pub:06EA2E12-F321-40B3-BE88-86B86A2CE2BB
Abstract: To assist with conservation of
Endangered Patagonian Huemul Deer (Hippocamelus bisulcus), the Huemul Task Force (HTF)
reassessed information on appendicular morphology, paleobiogeography, and
historical distribution as potential factors in recovery efforts. Traditional
claims of being a mountain specialist of the Andes were refuted by empirical
evidence showing huemul morphology to coincide with other cervids rather than
the commonly implied homology to rock-climbing ungulates. It thus supports
historical evidence of huemul in treeless habitat and reaching the Atlantic
coast, which cannot be dismissed as past erroneous observations. Instead, pre-
and post-Columbian anthropogenic impacts resulted in huemul displacement from
productive sites and in survival mainly in remote and marginal refuge areas.
The process of range contraction was facilitated by easy hunting of huemul,
energetic incentives from seasonal fat cycles and huemul concentrations, the
change from hunting-gathering to a mobile equestrian
economy, and colonization with livestock. However, areas used presently by
huemul, as supposed mountain specialists, are also used by wild and domestic
ungulates that clearly are not considered mountain specialists, whereas the
only other Hippocamelussuccessfully uses areas homologous to tree-less Patagonia. Rigid application of
modern habitat usage to infer past habitat use and ignoring historic extra-Andean
accounts is unwarranted; these conclusions reached by the HTF indicate new
opportunities for recovery efforts by considering morphological and historical
evidence. For instance, reintroductions to other portions of the landscape used
formerly by huemul, which tend to be more productive sites than those currently
occupied by many huemul groups, would present a promising avenue.
Keywords: Adaptation,
Andes mountains, appendicular morphology,
biogeography, Hippocamelus
bisulcus, historical condition, human influence, range
contractions, skeletal ratios.
Abbreviations: HTF - Huemul Task Force; IUCN -
International Union for Conservation of Nature; SSC - Species Survival
Commission
Spanish Abstract: Con el objeto de contribuir a la
conservación del huemul Patagónico (Hippocamelus bisulcus) el grupo de trabajo, Huemul Task
Force (HTF), realizó una revisión de la información sobre la morfología
apendicular, paleobiogeografía y distribución histórica del huemul como
factores potencialmente relevantes en los esfuerzos de recuperación de la
especie. La creencia tradicional de que el huemul es unespecialista de los hábitats de montaña andinos fue refutada por la evidencia
empírica de los análisis morfológicos. La anatomía apendicular del huemul es
similar a la de otros cérvidos y difiere de las especializaciones implicadas
para la escalada en roca de otros ungulados. Por lo
tanto, se apoya la evidencia histórica del huemul en unhábitat sin árboles como la estepa Patagónica. Su presencia histórica en la
costa atlántica no puede ser considerada comoobservaciones erróneas. En su lugar, hay que entender que impactos
antropogénicos pre y post colombinos dieron como resultado el desplazamiento de
huemul desde los sitios más productivos a sitios de supervivencia, sobre todo
en las áreas de refugio remotas y marginales. El proceso de contracción de su
rango geográfico se vió facilitado por la caza fácil, por los incentivos
energéticos de los ciclos estacionales de las reservas corporales de grasa, por
las concentraciones numéricas de huemul, por el cambio de la economía
cazador-recolector a una economía ecuestre móvil, y por la colonización con
introducción de ganado doméstico. Las áreas de montaña
actualmente utilizadas por el huemul, supuesto especialista de estos hábitats,
también son ocupadas por ungulados domésticos, que claramente no son
especialistas en montaña. Además, el único otro miembro de Hippocamelusutiliza con éxito las áreas homólogos a la región. La aplicación rígida del uso del hábitat
moderno para inferir el uso del hábitat pasado, ignorando el hábitat histórico
extra-andino es injustificada. Estas conclusiones alcanzadas
por el HTF indican nuevas oportunidades para los esfuerzos de recuperación del
huemul, apoyadas por la combinación de elementos morfológicos e históricos.Por ejemplo, la reintroducción a otros sectores del paisaje utilizado
anteriormente por el huemul, que tienden a ser sitios más productivos que los
actualmente ocupados por muchos grupos de huemules, presentaría una vía
prometedora.
Introduction
In
recognition of the urgency of the crisis regarding the Endangered Patagonian
Huemul Deer Hippocamelus bisulcus, the Huemul Task Force (HTF) was
formed within the IUCN-SSC to create another tool to provide recommendations
and guidelines based on sound scientific information through which the recovery
of Huemul can be achieved. Aside
from assisting to determine the current status of Huemul and review the
existing knowledge base, the aim is to identify scientifically acceptable
methodology appropriate for the species’ recovery.
Although
Diaz (1993) showed already then how history erroneously ‘led to the assumption
that the Huemul was a deer of the mountains and that it had always inhabited
areas in proximity to rugged topography’, the importance of this fact has
remained largely unrecognized, and remnant populations in mountain sites are
commonly interpreted to be due to Huemul being particularly adapted to such
sites, which thus supposedly represent prime habitat for the species. Moreover, Huemul being a mountain deer
is often reiterated and supported by referencing key paleontological work, a studywhich, however, was erroneous (see below). Main results of these historic
influences are 3-fold, by fomenting persisting claims that: (i) Huemul are
exclusively a mountain deer, specialized to rugged terrain; (ii) that their
natural range are the Andes mountains, as evidenced by the current relict
distribution; and (iii) that the few historic accounts of extra-Andean presence
were thus erroneous, or unimportant outliers. As interpretations of the biology and
ecology of Huemul play a mayor role in conservation strategies, the HTF worked
systematically through available evidence to evaluate common depictions of this
species.
Methods
Beginning
in June of 2008, a diversity of data from various published and unpublished
sources were analyzed to address the questions if Huemul specifically is a
mountain deer, and if historic reports of extra-Andean Huemul can legitimately
and universally be dismissed as erroneous observations of past naturalists. The
HTF formulated several conclusions below.
Results
and Discussion
1. Huemul as mountain deer
Early
European explorers and naturalists described Huemul as stocky, massive and
short-legged deer of mountains, comparing them to Ibex Capra ibex and
Chamois Rupicapra rupicapra in their homeland (e.g., Krieg 1925; Kurten
1979). They assumed Huemul to be a
mountain deer, just as was the interpretation for Ibex and Chamois at that
time, ungulates which by then were mainly surviving in remote alpine areas. Similarly, early North American workers
compared Huemul to Mountain Sheep Ovis canadensis and Mountain Goats Oreamnos
americanus (e.g., Krieg 1925; Frers 1969). More recent authors, often referring to
these early writings, make similar statements. However, early writings about Huemul
generally already reported their rareness, disappearance or near extinction
(e.g., Philippi 1857; Gigoux 1929), and references to stocky and short-legged
Huemul were casual remarks made about deer found mainly in refuge areas. Moreover, behavior like the aggressive
horseshoe stance (Cowan & Geist 1961) and thick long hair (Image 1)
dissimulate stockiness by distorting body shape (reviewed in Flueck &
Smith-Flueck 2011).
A
new fossil of North American cervid Navahoceros was described by Kurten
(1975) as having ‘highly unusual adaptive characters’ among cervids,
interpreted as extreme adaptations to mountains, and resulting in its common
name ‘mountain deer’. He explicitly
compared it to alpine Chamois and Ibex, and considered Hippocamelus as
related to his fossil. Even
though his fossil species has since been shown to be a construct and declared a
‘nomen nudum’ (Morejohn & Dailey 2004), this relationship is still cited in
arguing that Huemul is a mountain deer. The only comparative osteological analysis on leg morphology of Huemul
and 12 other ungulates revealed that Huemul cannot be
associated with rock climbing species. Although intraspecific proportional leg length is influenced by
ecogeography, nutrition, physiology and factors affecting exercise, with
variances of up to 70% in better studied cervids, Huemul morphology does not
overlap with rock climbing species previously considered analogous, but falls
within the range of other cervids (Flueck & Smith-Flueck 2011).
Position
about Huemul not being a mountain deer, adopted by the HTF in 2011:
1.
Early historical descriptions of Huemul as short-legged mountain deer
comparable to Ibex, Chamois, Bighorn Sheep, or Mountain Goats were only casual
comments. The descriptions likely
resulted from thick hair coats (7–9 cm hair length) and the behavior of
using the horseshoe posture.
2.
Kurten’s technical paleontological paper (1975) established Navahocerosas a mountain deer comparable to Ibex and Chamois: Navahoceros has been
shown to be a ‘nomen nudum’ (as was concluded earlier for Kurten’s
Stilt-legged Deer, Sangamona). Kurten’s referring to Hippocamelusas related to the ancestral Navahoceros (only differing by having two
[erroneous], instead of three antler tines), has been rejected based on
revising all bones and assembled skeletons labeled as Navahoceros, which
were confirmed to be Odocoileus (Morejohn & Dailey 2004).
3.
The morphometric analysis of complete leg assemblies from Huemul,then compared to several other species, shows that Huemul completely differ
from ungulates considered rock climbers. Furthermore, intraspecific variation in proportional leg
length—largely due to ecogeographical rules and nutritional and
physiological limitations—is up to 70% and results in populations of Rangiferand even O. virginianus having much shorter legs (by 14%) than the
Huemul sample.
4.
The nutritional ecology and climatic and topographic features of localities
where Huemul currently remain indicate that leg proportions from these sites
would be at the low end of the range of variations for Huemul: irrespectively,
these proportions clearly fall within the range of other cervids (Image
2). Taruca (H. antisensis)—the
only sister species to Huemul—utilizes some forest types, but is
currently mainly found in treeless grasslands with high affinity to Patagonian
grasslands, coexisting with several camelid species.
2. Past distribution of
Huemul
The
pre-Columbian distribution of Huemul has its roots in founding stock, likely of
the Odocoileus line (Morejohn & Dailey 2004), which dispersed
through the Panama land bridge during the Great American Interchange. Having to pass this equatorial filter of
continuous savanna habitat, succeeding species were generalists and
predominantly savanna-adapted (Webb 1978). As reviewed in Flueck & Smith-Flueck (2012), Hippocameluswere established by the Pleistocene, having dispersed south on the eastern side
of the Andes through continuous savanna habitat. Several periods of glaciation kept
ancestral Hippocamelus repeatedly away from the Andes, and fossils are
even known as far northeast as 809’S & 36022’W in the
most eastern tip of Brazil, and from the plains of Argentina, Uruguay and
southeastern Brasil.
During
glacial periods, the Andes were covered with ice even to near the equator, and
a continuous sheet covered the mountains from 33–560S during
the last glacial maximum. Glaciers south of 420S dipped into the
Pacific, overlaid the Andes 1600–1800 m thick, and reached hundreds of
kilometers into eastern Patagonian plains where only treeless habitat existed,
with Patagonia-like grasslands reaching way into Brazil, and much of South
America covered by savannah and grasslands. Moreover, the sea level was about
120m lower and the Atlantic coastline located 300km or more to the east of the
present coastline in some latitudes, which greatly extended the flat
paleosteppe region eastwards (e.g., Marshall 1988; Clapperton 1993; Markgraf
& Kenny 1997).
During
glaciations, Hippocamelus thus persisted in eastern treeless lowlands
reaching the plains of Uruguay, northern Argentina and Brazil. As mixed feeders, Huemul can incorporate
notable amounts of grass in the diet. Furthermore, besides Graminae, Patagonian steppes contain large
components of shrubs, maintain important green grass production throughout
winter, and deer are known to make much use of seed heads, which further
corroborates past and even historic distributions of Huemul in non-forested
habitats. Once eastern foothill
regions became glacier-free, Huemul were able to reach Andean habitat and when deglaciation
allowed for it, eventually to cross the Andes. Faunal exchanges from the east were
foremost across low Andean passes and explain the occurrence of late
Pleistocene Huemul in Chile as far northwest as 300S by the Pacific
coast (e.g., Ochsenius 1985; Moreno et al. 1994). With the last glacial retreat, forests
spread from few western refuges, and eventually covered the southern Andes
again, reaching their current extent only 2–3000 ya.
Nomadic
hunter-gatherers arriving in the southern Andes with the last interglacial
period would have had some influence on local distribution of Huemul. However, in northern and central Chile,
human adopted sessile and agricultural lifestyles long before arrival of the Spanish,
reaching very high densities, completely changing habitat through slash and
burn, and regionally extirpating several species including Huemul and Pudu DeerPudu puda. Further south,
humans became established mainly along the Pacific coast and focused on marine
resources. Consequently, early
explorers still found coastal areas abundant with Huemul. East of the Andes, Huemul then also
existed in zones between the Andean foothills and the Patagonian mesas, still
regularly occurring in flat grasslands about 120km east of the Andes, and
already more rarely, up to another 140km further east. Several reports show this species to
have reached the Atlantic coast (e.g., MacDouall 1833; Moreno 1899).
The
Spanish arrival resulted in highly significant changes brought about by the
introduction of horses, which created an equestrian lifestyle for native people
and profoundly changed their economies. Liberated in Buenos Aires, feral horses already reached the Strait of
Magellan by 1580. Livestock also
became feral immediately and soon roamed by the millions. Darwin (1839) found that native people
knew how to use knives, forks, spoons and relished
sugar, and most of the men spoke some English and Spanish. He further noted that these natives
travelled up to 750km inland during summer to hunt in the foothills, each man
having 6–7 horses. Native
tribes dominated the region for some 300 years until displaced by wars,
followed by the colonization of Patagonia with fencing and ranching occurring
rapidly throughout the region, and with over 47% of Patagonian forests burnt
before 1914 (e.g., Willis 1914).
As
a result of the above-mentioned history, the first early writings were
posterior to significant anthropogenic changes in the distribution of Huemul, with
explorers therefore largely unaware of previous history. Their descriptions of Huemul often were
from remnant populations living in remote and inaccessible places. Subsequent naturalists found an even
more reduced distribution, but as locations coincided with the few early
accounts, it led to dogmatic descriptions. Thus, decades have gone by further ingraining the notion that Huemul are
exclusively of Andean forests and not part of lowland central Chile; specially
adapted to precipitous rocky terrain, and forest habitats of the Andes; a
Mountain Deer living above tree line; only living between 1300–1700 m or high elevation mountains; or as preferring steep, rocky,
north-facing slopes. Preference to
the high Andes, principally near the international border along the continental
divide, was considered explicitly to be due to the conditions in that area
being the most favorable to Huemul (e.g., Osgood 1943).
The
fact that native people may have influenced Huemul distribution, including
after the increase in mobility due to horses, has been discounted based on the
argument that Huemul were energetically uninteresting. However, dissectable fat of deer
contains up to 47% of total energy content, whereas marrow fat adds only 1%
more, explaining why hunter-gatherers focus on deer during the autumn/winter
peak of fat (e.g., Thomas & Toweill 1982; Lipo 2007). This has been ignored when claiming that
hunter-gatherers would not have used an animal so lean as the Huemul, this
reasoning being based only on marrow fat. In contrast, while butchering, natives of northern North America
consumed dissectable fat and transported remaining bones for marrow and tallow
extraction at camp, just as documented for Patagonian hunter-gatherers: but the
few bone remains found in old shelters only provide a partial picture. It is erroneous to ignore that
professional hunter-gatherers would certainly have taken advantage of easy
accessible fat which presents >1200% more energy content than that obtained
from bones. Hunter-gatherers,
commonly moving according to seasonal movements of prey, covering distances up
to 150km for hunting particularly in autumn and early winter, best explained
logistical mobility in low-density hunter-gatherers in northern environments. From hunting camps, groups of young men
would make roundtrips of >100km in about three days, being able to portray
detailed maps covering 240,000km2 and animal movements within. Deer being preferred, a temporary camp
would remain if there were animals within 50km. Taking this in account when considering
historic reports of winter concentrations of Huemul, foraging conditions in the
pre-Colombian era were likely even superior to historic times in terms of
significance to hunter-gatherers. Borrero (2008) acknowledged that so far surveys in Patagonia had been
biased, being focused on caves that represent permanent sites. Transient hunting camps and movements
are thus under-represented and difficult to document anyway.
Position about the past
distribution of Huemul, adopted by the HTF in 2011:
1.
Likely it was Odocoileus lucasi (viz., Navahocerus nomen nudum)
dispersing through the Panama land bridge, savanna-adapted by necessity via
that equatorial filter, and giving rise to Hippocamelus.
2.
Glaciations prevented the continuous use of Andean highlands and the Pacific
side: during glaciations, Patagonia-type habitat, and fossil Hippocamelusreached into northeastern Brazil; Patagonia was twice the current size, as the
Atlantic coast line was hundreds of kilometers further east due to much lower
sea levels.
3.
Dispersal and colonization likely occurred along the eastern fringe of the
cordillera and a coastal route (e.g., Marshall 1988). Similarly, bighorn sheep remained in
grassland and steppe areas during glaciation, then followed as glaciers
retreated; colonization is considered to have likely occurred along cordillera
going south, along glacial margins in habitats like tundra and taiga (Geist
1985).
4.
After the last glaciation, forests spread from Pacific refuges and covered the
southern Andes again, reaching their current extent only 2–3000 ya. Once ice-free, low passes allowed Huemul
to enter from the east and to populate landscapes also west of the Andes.
5.
Humans arrived with the last interglacial period, about 10-12000 ya in the
southern Andes. Pre-Columbian
hunter-gatherers likely had local impacts on Huemul distribution based on
optimal foraging among studied hunter-gatherers (including behavior of
Patagonian natives regarding Guanaco Lama guanicoe), seasonal fat cycle
in Huemul, easiness to hunt Huemul in autumn/winter. Pre-Columbian sessile natives in central
Chile exterminated local fauna including cervids like pudu and Huemul.
6.
Post-Columbian natives became equestrian, focusing on
feral domestic livestock and native ungulates. Patagonians traveled up to 750km to hunt
in eastern ecotone and foothills, burning extensive landscapes. Similarly on the Chilean side, large
numbers of livestock and equestrian people displaced Huemul such that early on
Huemul were considered rare and restricted to steep remote mountain areas
(except in southern distant Fiord areas).
7.
After 300 years of dominance, the natives on the eastern side of Andes were
subdued and a wave of fencing and ranching went through Patagonia, with heavy
impact on the few Huemul remaining on those lands.
8.
Due to pre- and post-Columbian events, the first writings were posterior to
significant anthropogenic changes in Huemul distribution, with descriptions
from remnant Huemul populations living mainly in remote and inaccessible
places. Similarly for desert
bighorn sheep, rather than becoming a relict species created by
post-pleistocene ecological changes, they have become secondary relicts with
small, isolated populations created by the impact of European settlement as
early as 1540. The overall result
was the extirpation of many populations of bighorn and the creation of smaller,
isolated herds, prone to extinction (McCutchen 1982).
9.
As a result of pre- and post-Columbian events, there are few historic documents
of Huemul still existing in extra-Andean landscapes, however:
-there were still several reports about large groups in
traditional wintering areas, i.e. groups of 100 or more
-today mainly forked antlers occur, yet there are
several reports of 4 and 5 point antlers, i.e. prior habitat sites were
superior to extant sites (the newest rediscovery of this fact: de la Croix
1937)
-besides thorough reviews by Diaz (1993, 2000) there
are several newly discovered sources including photos of hunted Huemul, with
distances from the Andes at 120km, 200km, 260km, 270km, and all the way to the
Atlantic coast (Image 3).
Note: there are several lines
of evidence that Huemul also occurred in Tierra del Fuego (see Flueck &
Smith-Flueck 2012).
Conclusions
The
few historic accounts still documenting presence of Huemul in the eastern
treeless lowlands, indicate that Huemul were well
suited to exploit those areas. This
information cannot be dismissed due to its relevance, similarly as had been
shown for Chamois and Ibex. By avoiding the application of analogies based only on the present
situation, but beginning to use comparative morphometry and the past to
understand the presence, the repeated fallacy of simply imposing the present on
the past will be omitted. The empirical comparisons showed Huemul leg morphology to fall well
within that of other cervids and can be expected to vary substantially if they
were to live in habitats formerly used. It supports the evidence that Huemul existed in treeless habitat and
colonized Andean forests and higher altitudes secondarily, and habitat breadth
of Huemul is thus more like that found in other closely related Odocoilines,
and moreover, coincides with habitat use by the only congeneric, the taruca. Although Huemul can use forests
exclusively, they can also thrive in ecotone, and (previously) in grasslands,
steppe, and deserts (like Odocoilines, Ibex, Bighorn Sheep, Red Deer Cervus
elaphus, Guanaco). Additionally, unspotted Hippocamelus fawns also point to an
origin in non-forested areas, which still presents the principal habitat use byH. antisensis. Moreover,
even small cervids thrive exclusively in treeless grasslands, like Pampas Deer
(Ozotoceros bezoarticus Perez et al., 2008) or Roe Deer (Capreolus
capreolus Abbas et al., 2012), which show extensive digestive plasticity
via behavioral and morphological adaptations.
The
seasonal fat cycle and congregations likely made Huemul a prime candidate for
hunter-gatherers, who would have therefore influenced their distribution and
density. The subsequent equestrian
mobility of natives and later colonists further displaced Huemul from
traditional valleys and winter ranges. Several other ungulates had mainly lost their low elevation habitats
from anthropogenic pressures and range contraction allowed them to persist only
in marginal peripherical habitat, concentrated at high elevations or otherwise
inaccessible sites (Channell & Lomolino 2000; Laliberte & Ripple
2004). Thus, paleoecology,
zoogeography, and history of land use in southern Latin America indicate that
Huemul exclusively remaining in high mountains are secondary relicts created by
post-Columbian anthropogenic impacts. The presence of Huemul right into historic times in grasslands and
steppe areas is likely of more importance to Huemul conservation than hitherto
acknowledged.
Although
Huemul presently still use flat areas, they did more so in the past (like
Odocoilines, Bighorn Sheep, Guanaco, but unlike Mountain Goat, Ibex). The habitat types used by Huemul in recent
times and resulting in the colloquial description as mountain deer, are not
preventing the use of these areas by species not considered mountain
specialists, like Guanaco, Wild Boar Sus scrofa, Red Deer, Fallow Deer Dama
dama, cattle (specially feral ones), goats, sheep, and horses. The habitat types used by Huemul
historically are also used by these same species above. Taruca, considered by several authors
osteologically indistinguishable from Huemul and suggested to represent two
subspecies, occur in areas used by camelids (vicuña, guanaco, llamas, alpacas),
White-Tailed Deer Odocoileus virginianus, domestic sheep, cattle,
horses, and mules. Taruca occur in
the cold desert ‘puna’, which includes large tracks of plains (altiplano):
nearly every one of its plant genera also occurs in the Patagonia steppe
(Fernández & Busso 1997). Lastly, taruca also occur in ecotone and forests (still now, but more so
in the past).
The
rigid application of modern Huemul habitat usage to infer past habitat use and
ignoring historic extra-Andean accounts as erroneous or abnormal outliers is
unwarranted. The conclusions
reached by the HTF indicate new opportunities for recovery efforts by
considering morphological and historical evidence. For instance, reintroductions to other
portions of the landscape used formerly by Huemul, which tend to be more
productive sites than those currently occupied by many Huemul groups, would
present a promising avenue (see IUCN 2012 in prep.:Guidelines for Reintroductions and other Conservation Translocations. IUCN/SSC
Reintroduction and Invasive Species Specialist Groups. IUCN, Gland,
Switzerland). Although adopting a
uniform conservation program over a large geographical area is attractive to
policy-makers and conservation planners, the large range of past geographical
and ecological sites used by Huemul indicate that conservation programs could
benefit from broadening strategies accordingly.
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