Reassessment of morphology and historical distribution as factors in conservation efforts for the Endangered Patagonian Huemul Deer Hippocamelus bisulcus (Molina 1782)

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 Hippocamelus successfully 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

Full text

Journal of Threatened Taxa | www.threatenedtaxa.org | November 2012 | 4(14): 3302–3311
JoTT COMMUNICATION 4(14): 3302–3311
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
OPEN ACCESS | FREE DOWNLOAD
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.
3302
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 Hippocamelus successfully 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.
urn:lsid:zoobank.org:pub:06EA2E12-F321-
40B3-BE88-86B86A2CE2BB
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
Abbreviations: HTF - Huemul Task Force; IUCN - International Union for Conservation
of Nature; SSC - Species Survival Commission

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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 study which, 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 Navahoceros as 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 Hippocamelus as 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

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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 Rangifer and 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),
Hippocamelus were 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.
Image 1. (a) The 7–9-cm-thick hair coat is best appreciated when being shed. Note in the upper and lower left photos that
the diameter of the upper neck is substantially less than further down in the unshed portion. (b) The aggressive stance of
huemul at close range, known as the horseshoe posture.

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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 Deer Pudu 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

Image 2. Body shapes of several extant Huemul that are not in an alert stance and with summer coat.

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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
Hippocamelus reached 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

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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

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presents the principal habitat use by H. 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
Image 3. Huemul habitat far from forests and in flat or rolling landscapes. (a) Huemul in steppe, approached by gaucho,
from Onelli 1904. (b) Huemul photographed in the 1920s by A. Grosse. (c,d) Princeton expeditions, late 1800s, hunting
huemul as far as 200km from forests, from Hatcher 1903. (e) Extant Huemul in periglacial Pacific coast, old moraines, and
flat wide valley bottoms. (note: open areas used during midday hours)
a
c
b
d
e
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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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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 un especialista 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 un hábitat sin árboles como
la estepa Patagónica. Su presencia histórica en la costa atlántica no puede ser considerada como observaciones 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 Hippocamelus utiliza 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.

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* 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