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Predicaments of endangered huemul deer, Hippocamelus bisulcus, in Argentina: A review

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A total of 350–600 huemul (Hippocamelus bisulcus) remain as fragmented groups along 1,850km of Argentine Andes. Their conservation depends on accurate knowledge of the species' requirements and the factors preventing their recovery. The Regional Delegation for Patagonian National Parks (RDP) erroneously alleged that huemul status is satisfactory, and current in situ efforts are sufficient to guarantee recovery. Therefore, conservation centers are regarded unnecessary and the associated risks too high, especially because previous attempts with manipulations have failed. No data support these claims, instead many subpopulations have disappeared recently even in national parks (NP) which hold <0.01huemul/km2. Causes preventing recovery or recolonization are unknown. Current pressures on huemul subpopulations include increased economic activities and alien species. Normal ranges for many biological parameters or population performance of huemul are unknown. Focus is on habitat studies using presence as surrogate for what should be studied on survival and reproduction. Factors important to small-sized populations or preventing recovery remain unstudied. RDPs insistence on indirect methodology prevents implementation of other potentially more promising research approaches. The lack of consensus regarding the necessity and feasibility of a conservation center prevented its establishment and related census flights in unprotected sites. RDP currently forecloses aerial census and capturing and thus prospects for a huemul conservation center, and the proposition of establishing such a center was neither discussed nor incorporated into the national recovery plan. Helicopter captures have been used successfully on deer in huemul habitat. Captures and translocation of huemul occurred since 1830; several zoos kept them successfully up to 10years, and natural tameness facilitated husbandry. Recently, Chile successfully caught and transported huemul by helicopter to stock a private center. Unknowns can be addressed easily on semicaptive deer; other questions can be studied through reintroductions, employing adaptive management. RDP places faith in NP providing viable subpopulations. However, it remains doubtful whether some 220 huemul living in >22,000km2 of parks can guarantee species survival. For Argentine cervids, absence of studies and management plans due to lack of funds is typical. Considering the actual situation and future perspectives, it appears doubtful that recovery will be achieved based on strategies similar to those employed in the past.
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Eur J Wildl Res (2006) 52: 6980
DOI 10.1007/s10344-005-0020-4
REVIEW
W. T. Flueck .J. M. Smith-Flueck
Predicaments of endangered huemul deer,
Hippocamelus
bisulcus
, in Argentina: a review
Received: 30 July 2005 / Accepted: 19 October 2005 / Published online: 6 December 2005
#Springer-Verlag 2005
Abstract A total of 350600 huemul (Hippocamelus
bisulcus) remain as fragmented groups along 1,850 km of
Argentine Andes. Their conservation depends on accurate
knowledge of the speciesrequirements and the factors
preventing their recovery. The Regional Delegation for
Patagonian National Parks (RDP) erroneously alleged that
huemul status is satisfactory, and current in situ efforts are
sufficient to guarantee recovery. Therefore, conservation
centers are regarded unnecessary and the associated risks
too high, especially because previous attempts with
manipulations have failed. No data support these claims,
instead many subpopulations have disappeared recently
even in national parks (NP) which hold <0.01 huemul/km
2
.
Causes preventing recovery or recolonization are un-
known. Current pressures on huemul subpopulations in-
clude increased economic activities and alien species.
Normal ranges for many biological parameters or popula-
tion performance of huemul are unknown. Focus is on
habitat studies using presence as surrogate for what should
be studied on survival and reproduction. Factors important
to small-sized populations or preventing recovery remain
unstudied. RDPs insistence on indirect methodology pre-
vents implementation of other potentially more promising
research approaches. The lack of consensus regarding the
necessity and feasibility of a conservation center prevented
its establishment and related census flights in unprotected
sites. RDP currently forecloses aerial census and capturing
and thus prospects for a huemul conservation center, and
the proposition of establishing such a center was neither
discussed nor incorporated into the national recovery plan.
Helicopter captures have been used successfully on deer in
huemul habitat. Captures and translocation of huemul
occurred since 1830; several zoos kept them successfully
up to 10 years, and natural tameness facilitated husbandry.
Recently, Chile successfully caught and transported
huemul by helicopter to stock a private center. Unknowns
can be addressed easily on semicaptive deer; other ques-
tions can be studied through reintroductions, employing
adaptive management. RDP places faith in NP providing
viable subpopulations. However, it remains doubtful
whether some 220 huemul living in >22,000 km
2
of
parks can guarantee species survival. For Argentine
cervids, absence of studies and management plans due to
lack of funds is typical. Considering the actual situation
and future perspectives, it appears doubtful that recovery
will be achieved based on strategies similar to those
employed in the past.
Keywords Conservation biology .Deer conservation .
Captive program .Adaptive management .Extinction
Introduction
Huemul (Hippocamelus bisulcus; Fig. 1), an endemic
cervid of Chile and Argentina, is considered at risk of
extinction due to numeric reductions by >99% and in
distribution by >50% (Díaz and Smith-Flueck 2000). The
remaining 350600 Argentine huemul are dispersed along
1,850 km of Andes in roughly 50, mostly fragmented,
subpopulations. As both countries continue to lose sub-
populations (total numbers 1,0001,500), their status is
critical (Díaz and Smith-Flueck 2000; Serret 2001).
Recovery depends on understanding factors affecting
individual subpopulations, and on well designed in and ex
situ projects. Information gaps impeding conservation can
be filled via research on semicaptive huemul, which would
also provide animals for reintroductions or restocking.
Reintroductions using adaptive management principles
(AMP) could determine factors presently preventing
recovery (Walters and Holling 1990). The fact that huemul
W. T. Flueck (*)
CONICET,
C.C. 176,
8400 Bariloche, Argentina
e-mail: deerlab@baritel.com.ar
J. M. Smith-Flueck
DeerLab,
C.C. 176,
8400 Bariloche, Argentina
remain in remote small clusters at low densities limits the
efficacy of field research (CONAF and CODEFF: Plan
para la conservacion del huemul del sur en Chile, 2001),
especially if funding for such research is low. Considering
the demands for professional standards of capture and
treatment of animals at risk of extinction (IUCN 2001a), we
analyze methodologies based on the characteristics of
huemul and its habitat.
Serret (2001) stated that huemul cannot be captured and
confined as easily as other cervids, which was also
emphasized in the Chilean recovery plan (CONAF and
CODEFF: Plan para la conservacion del huemul del sur en
Chile, 2001). According to the Argentine recovery plan, ex
situ conservation shall be evaluated when necessary, yet a
clear opposition to centers with semicaptive huemul prevails.
The dominating Argentine group in huemul conservation,
the Regional Delegation for Patagonian National Parks
(RDP), claimed in an official statement that (a) Argentine
huemul have not yet reached a critical population size
justifying implementation of ex situ programs; (b) risks,
therefore, associated with captive huemul are too high; (c)
ample areas contain huemul populations in a good
conservation state; (d) other activities have much higher
conservation value than those obtained from ex situ
programs; (e) current in situ efforts are sufficient to guarantee
long-term survival of huemul, and therefore, conservation
centers are unnecessary; and (f) previous attempts to keep
huemul in captivity have failed (Ramilo 2001). Conse-
quently, RDP prevented a feasibility study as a first step
towards establishing a conservation center.
Scientific data contradict the claim that huemuls
conservation status is satisfactory. Many subpopulations
have disappeared recently, even within national parks (NP)
(Franke 1949; Serret 2001). Low numbers and severe
population fragmentation indicate high risks of extinction
(Pimm and Bass 2002). Chilean huemul conservation is
also losing ground, even in protected areas (PA) (Povilitis
2002). We posit that (a) historic events caused fragmen-
tation and reduced subpopulations (Díaz and Smith-Flueck
2000); (b) new factors, like invading red deer (Cervus
elaphus), are now acting on remaining huemul (e.g. Flueck
et al. 2003; Smith-Flueck 2003); and (c) subpopulations are
so reduced that they are subjected to constraints of biology
of small populations (Caughley 1994). Given the situation
in Argentina, conservation benefits from semicaptive
huemul would thus be significant. As we will show,
huemul have been successfully maintained in captivity, and
the risks from manipulation are negligible. Thus, this
review may serve as a feasibility analysis for the
establishment of a conservation center as an additional
strategy for huemul recovery.
Current predicaments of Argentine huemul
Economic developments
Since 1980s, human populations in provinces with huemul
increased 57100% (Elustondo, 12/9/2003, Clarín, Argen-
tina). Increasing roads, resource use, large-scale construc-
tions and mining, some even encroaching on NP (Martin
and Chehebar 2001), limit wildlife conservation (Laliberte
and Ripple 2004; Parera 2002; Saucedo 2002). Neither
Argentina nor Chile have regulations regarding huemul
management or habitat on private land. Tourism, including
helicopter rides, has steadily increased and reached all
huemul areas (Serret 2001; Braun 2002; Moyano 2004). In
2003, recreation in Patagonian NP increased by 56%
(Administración de Parques Nacionales, Argentina 2004;
Sabatini and Iglesia 2001). Ranches are rapidly being
converted to tourist centers. Poaching still occurs in both
countries but has not yet been quantified.
Alien wildlife
Introduced red deer threaten huemul (Flueck et al. 2003;
Smith-Flueck 2003). Sometimes illegally, enterprises
import wildlife, like 26 red deer from New Zealand (NZ)
in 2002 (Sympson 2003). Mountain tahr (Hemitragus
jemlahicus), highly invasive and difficult to control, ar-
rived from NZ in 2000. Interestingly, they do not appear
in any of the governments alien species lists and are likely
to eventually roam huemul habitat. Imports of exotics are
associated with risks of introducing diseases, such as
chronic wasting disease or parasitoses. In Argentina, exotic
Taenia krabeii may now exist in a sylvatic cycle (Flueck
and Jones 2005), and several exotic nematodes are already
reported in red deer.
Huemul likely first went extinct in more favorable
habitats like ecotones and former winter ranges due to
human settlements and associated introductions of exotics.
From such source areas, red deer are invading huemul
refuges and may displace huemul (Díaz and Smith-Flueck
2000); they may cause interference competition (Stephens
et al. 2003) or may introduce diseases. Regardless, huemul
are at substantial risks from the mass effectalone of
invading red deer (Thompson et al. 2003).
Fig. 1 The Patagonian Huemul, Hippocamelus bisulcus (photo by
Smith and Flueck)
70
Limitations of protection in Argentina
While Lanin NP, since its establishment, has lost all
subpopulations, Nahuel Huapi NP lost many if not most
(Serret 1993,2001). Patagonian NP (22,618 km
2
) contain
only about 220 huemul, or <1 individual/100 km
2
(Serret
1993,2001). Huemul densities average 0.7/km
2
(Díaz and
Smith-Flueck 2000), indicating that >98% of NP are
presently void of huemul. Hence, Serret (2001) cautioned
that NP may not contain viable subpopulations: according
to Reed et al. (2003), no group qualifies as viable. Settlers
within NP raise livestock (Martin and Chehebar 2001),
utilizing up to 56% of park areas (Simberloff et al. 2003),
but commonly run several fold more livestock than
permitted (Serret et al. 1994). Although parts of NP with
huemul were declared Critical Areas, 90% of the area is
still exclusively used by cattle and only 10% by huemul.
Reducing livestock is hampered by lack of control, fencing,
and prevailing policies (Martin and Chehebar 2001;
Sabatini and Iglesia 2001), hence fueling the tragedy of
the commons. IUCNs category II for NP requires absence
of exploitation, yet extensive parts within huemul ranges
are exploited, even intangibleareas, mainly through un-
authorized livestock and hunting. Low-impact areas of NP
occur at higher elevation (=highest protection) due to
diminished human interests. In contrast, favorable habitats
in NP are privately owned, intensely utilized, characterized
by lower elevations with greater proportions of flat slopes
and wet meadows, served as winter habitat for many
species, and likely provided source populations. Low-
impact portions of NP today (Ramilo 2002) mainly
represent suboptimal conditions for huemul. Elsewhere,
species have gradually gone extinct in PA due to laissez-
faire management, whereas they recovered in areas brought
under state-of-the-art management (Saenz et al. 2001).
Thus, all factors in NP considered detrimental (APN, Proc.
First Huemul Conf., 1992) should be eliminated, including
livestock, dogs, and consumptive and nonconsumptive
uses. Once subpopulations are shown to recover, it could
be evaluated if certain land uses should be permitted again.
Whereas most PA are paper parkswith scarce or absent
implementation (Martin and Chehebar 2001; Rusch 2002),
a substantial proportion of subpopulations are outside of
PA (Serret 2001). NP contain extensive huemul habitat,
making it a prime candidate for contributing significantly
to huemul recovery.
Small population characteristics
More effort into surveying new sites might reveal some
new small groups, but the larger subpopulations are already
known. Two subpopulations contain around 100 deer each
(Serret 1993; Smith-Flueck and Flueck 2001a), another
contains about 40 deer, and the remaining quasi-extinct
subpopulations may each contain <15 individuals and can
be considered quasi-extinct (Berger 1990). Even if
estimates would be 300% higher, subpopulations still
could not be considered viable (Pimm and Bass 2002; Reed
et al. 2003). To date, no cases of dispersal nor recoloniza-
tion exist. Thus, while potential corridors can be
established, they might not benefit huemul presently.
Levels of fragmentation, reduced subpopulation sizes,
site fidelity, absence of recolonization, and continued loss
of groups indicate a tendency towards reduction in
occupied area and numbers.
State of knowledge
There are 3 doctoral thesis (1816, 1979, 2003), 2 books
(Díaz and Smith-Flueck 2000; Serret 2001), and a Cross-
Search of ISI Web-of-Knowledge and 17 external data-
bases (1945present) listed 16 entries on H. bisulcus (9
original studies). According to CONAF and CODEFF
(Plan para la conservacion del huemul del sur en Chile,
2001), Chilean data obtained during 25 years were mainly
descriptive and were neither collected systematically nor
statistically analyzed. Most information from Argentina
appears as in-house reports. RDP recorded warden
observations between 1990 and 1998 (Martin and Chehebar
2001): only 12 groups were seen. Such data lack precision
to determine absence and serve mainly to confirm presence
(Sutherland 2000). Mortalities were analyzed only once for
huemul (Smith-Flueck and Flueck 2001b). Although initial
diagnosesof huemul status were mere hypotheses (APN,
Proc. First Huemul Conf., 1992), reassertions in other
workshops (1995, 1998, 2002) resulted in dogmas without
evidence. Successful diagnoses must be based on scientific
monitoring (Sutherland 2000) which is notoriously lacking.
In essence, no information has yet been generated on
factors preventing recovery of any subpopulation.
Current management and conservation in Argentina
The past 15 years resulted in new PA, stricter laws,
awareness programs, and confirmation of huemul pres-
ence. While neglecting nonprotected subpopulations,
efforts focused on NP (local NGOs and the Wildlife
Conservation Society, WCS: huemul project 1998pres-
ent), determining habitat availability and selection and
developing predictive models of potential habitat based on
signs. RDP defined huemul research priorities for NP
(Ramilo 2003) as determining (a) habitat size and location
for rearing young, (b) habitat for summer/winter mainte-
nance, (c) unused areas (transit), (d) factors affecting
fecundity and mortality, (e) abundance trends, and (f)
interactions with red deer, livestock, puma (Puma con-
color), and tourism. However, no such research has been
conducted so far, and no projects address issues important
to small population size or with respect to factors
preventing recovery.
71
Proposed research needs
Methodological considerations
RDP does not have research programs, but determines
research needs (Ramilo 2003). For huemul, only specific
methods are allowed. Such rigidity impedes progress
(Sutinen et al. 2004). For instance, for habitat and
monitoring studies, researchers are obliged to use unified
methods to supposedly facilitate comparisons. However,
different results from applying one method would likely be
interpreted as real differences, when actually it was due to
personnel, especially if members change constantly as
through volunteering (Sutherland 2000; Johnson 2002).
Another insistency concerns applying only indirect meth-
odsinaccurately claimed to be consented by all stake-
holders in huemul conservation (Proc. 4th binational
reunion on huemul conservation). Limiting trend evalua-
tions of tiny populations to only fecal surveys is
unwarranted and ignores the enormity of this task at the
required precision. Given low density, remoteness, envi-
ronmental variability, change of sites by huemul (Ortega et
al. 2003), personnel variability to 35%, and dependence of
fecal counts on population sex and age structure (Smart et
al. 2004), to name a few, other methods are superior. We
suggest that traditional fecal surveys will not provide the
required accuracy and precision to monitor trends for most
remaining subpopulations.
Habitat studies
Current habitat selection may be different from that prior to
European colonization, due to abnormal density, sex or age
ratios; that is, subpopulations may be disorganized socio-
biologically (Bubenik 1982). Huemul now occur possibly
in marginal or sink areas (Flueck 2003). Current habitat
choice is sometimes erroneously used as a surrogate for
survival and reproductive success, which is likely no
longer true (Battin 2004). It is a fallacy to claim currently
inhabited areas as optimal areas because huemul today may
be displaced into peripheral areas by human activity, where
their habitat requirements can hardly be realized. High-
elevation Andes likely do not represent the core of huemul
distribution, rather two cores exist: one each in Chile and
Argentina at mid to low elevations. Contractions of huemul
range were thus towards peripheral habitat and driven by
anthropogenic extinction forces (Channell and Lomolino
2000). Other endangered species have likewise been driven
into refuges shown afterwards to be marginal (Craig 1994;
Caughley 1994). Nutritional constraints for current low-
density huemul appear improbable (except winter range
inaccessibility, see Flueck 2003), considering equivalent
habitats support high densities of exotic herbivores. Past
distributions predict existence of ecotypes and adaptive
feeding behavior, as corroborated by diet studies (Flueck
2003; Smith-Flueck 2003). Annual variations in habitat
variables preclude defining static requirements and explain
site use changes (Ortega et al. 2003; Osko et al. 2004).
Consequently, research within PA should be directed at
obtaining confident population parameters (reproductive
rates, recruitment, survival, mortality causes, trends) rather
than habitat studies. Studies on spatiotemporal movements
are essential to determine source-sink dynamics and
discover reasons for lack of recolonization.
Need for direct methods
Huemul captures using net gunning as best available
method in remote areas are currently not possible because
RDP considers helicopter and huemul to be incompatible.
Although aerial surveys of eight nonprotected subpopula-
tions were initially accepted, with government observers
already assigned, they were unexpectedly denied without
explanations; these would have been the first systematic
estimates in Argentina. Helicopter censuses have been
permitted with other endangered Argentine cervids like
Ozotoceros bezoarticus, for which helicopter net gunning
was proposed as the best capture method. Whereas one-
time census flights in areas outside of NP was unacceptable
to RDP, commercial use of helicopter for mining explora-
tion, tourism, as well as private use, is commonly practiced
there (Moyano 2004).
Reasons for establishing a huemul conservation
center in Argentina
IUCN (1987) emphasized that (a) vulnerability of small
populations is consistently underestimated (see Wharton
1995); (b) timely recognition is critical and depends on
valid information on population status; and (c) to reduce
extinction risk, supportive centers should be established
when wild populations still count thousands of indi-
viduals. In 2001, IUCN declared huemul to be at very
high risk of extinction.This was justified by all sub-
populations having <250 individuals and with <2,500
total mature individuals in decline (category C2a).
Furthermore, reported densities indicate that the actual
extent of occurrence is <5,000 km
2
, with severe fragmen-
tation and declining tendency, justifying even applying
category B1a. Captive huemul provide research opportu-
nities that would now be difficult or impossible with wild
populations. Accordingly, IUCN (2002) recommended
explicitly that ex situ programs foster research on
questions relevant for in situ conservation.
In Argentina, RDP currently considers a conservation
center as irrelevant (Ramilo 2001), although the available
scientific information justifies this supportive tool. Yet as
late as in 1971, the Argentine government launched the
Operativo Nacional Huemul,trying to establish a center
to avoid extinction (Díaz and Smith-Flueck 2000), though
these plans were never realized.
A centers main objectives would include conservation
biology and education, as well as fund raising. Scientific
data are fundamental for (1) diagnosing in situ problems,
(2) decision making, (3) mitigating negative impacts on
72
huemul and its habitat, and (4) evaluating conservation
actions through monitoring. While limited data exist on
basic life history strategies, data on variability of parame-
ters important to conservation efforts are completely lack-
ing (Table 1). Only twice have neonates been observed, and
11 parturition dates were based on fawns roughly aged at
16 weeks (Smith-Flueck 2003). Currently, huemul cannot
be judged regarding normal ranges of basic anatomical,
morphological, physiological, biochemical, and genetic
parameters (Díaz and Smith-Flueck 2000). Limited or no
baseline data exist to determine if individuals or popula-
tions are performing adequately, whether huemul are
highly susceptible to bovine disease, as was assumed (E.
Ramilo, cited in Wemmer 1998), and whether Cysticercus
tenuicollis causes death (Simonetti 1995). While several
unknowns can be addressed directly on semicaptive deer,
other important questions (Table 2) can be studied through
reintroductions (Riney 1967; IUCN 2001b), such as to
whether current distribution and habitat use by huemul are
artifacts of recent changes.
Due to past equivocal results and continuous loss of
opportunities, it is highly advisable to professionally
design the next center, as soon as possible, supported by
long-term financial commitments so as not to curtail
success and future attempts by others (IUCN 2002). Also,
funds for ex situ programs are frequently available for
complementary parallel in situ studies (Wharton 1995).
Establishing and running a financed center with parallel
field research in Argentina (Smith-Flueck and Flueck
2001c) was considered unnecessary (Ramilo 2001). In
contrast, Chile, with many more huemul, recognized the
need and had already proposed a center in 1989 (CONAF,
Proposal to San Diego Zoological Society). Recently, they
established a huemul center (SAG permit 1535, April
2005) with the main purpose to restock the surrounding
reserve. It is highly unlikely that Argentina will benefit, as
surplus animals will be used for reintroductions and public
education in Chile.
Modern techniques to manipulate huemul
Critical features of future studies, centers, or reintroduc-
tions should include capture, transport, handling, and
management. Guidelines recommend mechanical over
chemical restraint (Jessup et al. 1996). For flighty cervids,
modern methodsby considering behavior, biology, nu-
trition, and environmentreduce risks of exertional my-
opathy (Kreeger 1999).
Capture methods for huemul include trapping, chemical
immobilization, or net gunning. We have used clover and
panel traps, drive nets, drop and cannon nets, and drive
traps to capture cervids in USA and Patagonia (1983
1994). Baiting in Patagonia did not work during vegetative
periods due to surplus natural forage. Access in winter was
difficult, unselective traps captured omnipresent livestock,
and monitoring twice a day and readiness of capture teams
were costly. Drive netting in typical huemul habitat was
only possible by helicopter, and permanent drive traps are
costly and impractical given the low density of huemul.
Large darts are disadvantageous, but unavoidable with
drugs like xylazine/ketamine, even after concentrating by
lyophilizing. The recently introduced thiafentanil allows
the use of small darts and enables highly effective re-
versal. Thiafentanil, with faster induction time than
medetomidine/ketamine, even working in hyperexcited
animals, can therefore be recommended as one of the best
options for immobilizing huemul (Kreeger 1999). Low
densities, close approach, small targets, deer disappearing
in vegetation during induction, and required time and
personnel make darting a challenging and risky method
and should only be attempted by experienced personnel.
Table 1 Unknown or assumed traits of huemul to study in a conservation center
Trait Published parameters Source
Age at first conception As fawn; at 3 or 4 years Díaz and Smith-Flueck 2000; Serret 2001
Gestation period 200220 days Díaz and Smith-Flueck 2000
Parturition dates OctoberJanuary Diaz and Smith-Flueck 2000
Weight at birth Up to 3.5 kg; 5 kg Flueck and Smith-Flueck 2005
Weaning At 4 months Diaz and Smith-Flueck 2000
Growth pattern for both sexes ?
Morphometry by sex and age ? Reviewed in Diaz and Smith-Flueck 2000
Maximum/mean life expectancy ?
Life time reproductive success ?
Social behavior Territoriality? Serret 2001
Scent glands (anatomy, function) ? Reviewed in Diaz and Smith-Flueck 2000
Biological parameters and variability
a
?
Defecation rate (sex, age, season, feed) ?
Mineral metabolism and requirements ?
Annual biological cycle of both sexes ?
Nutritional requirements Which habitat types allow reproduction?
a
Physiology, clinical biochemistry, hematology, endocrinology, immunology, hormones, phermones, genetics, pathology, parasitology
73
Mortality rates of 1723% have occurred with cervids,
and nearly 50% were from dart trauma (Bates et al. 1985).
Capturing several huemul dispersed over a large remote
area will make darting very costly and will require much
manpower.
Helicopter captures
Helicopter captures are most appropriate where huemul are
widely dispersed and in difficult terrain. Helicopter darting
has been replaced by net gunning due to reduced trauma,
distress, morbidity, mortality, and costs (Jessup et al. 1988).
It permits (a) no drug-related side effects, (b) collection of
chemically unaffected tissues, (c) capture of whole social
groups, (d) capture in difficult areas, (e) capturing over
large areas and all day, and (f) reducing costs per animal.
As group sizes average about 23 (Díaz and Smith-Flueck
2000), capture of these can readily be accomplished. Net
gunning resulted in 6.68% mortality in stress-prone Ovis
canadensis (n=306) and 1.96% in several other ungulates
(n=2598; L.H. Carpenter 1996. Survey of post-capture
mortalities. Helicopter Wildlife Management, Utah, un-
published). Net gunning was the safest and least distressful
of five methods and 56% less risky than darting (Kock et al.
1987; Jessup et al. 1988): it was used successfully in
Patagonia (Fig. 2) to mark red deer dispersed over 130 km
2
of former huemul habitat (Flueck et al. 2005).
Huemul and helicopter
While net gunning huemul in Argentina is currently ham-
pered because of concerns related to disturbance, darting
from the ground has been used for a WCS project (Parera
2002). Aircrafts including helicopters have been employed
on cervids since the 1940s, and reactions towards noise are
well known. Cervid hearing differs completely from
humans, who hear better at <8 kHz, whereas cervids hear
better at >810 kHz (Krausman et al. 2004). Machinery
noises usually produce low frequencies: aircrafts typically
<2 kHz. Consequently, these sounds are perceived 63 times
stronger by men vs ungulates (Krausman et al. 2004). To
compare, noise at a busy Buenos Aires intersection was
101 dB (Río Negro, Argentina, 19/11/2001:11). Helicop-
ters producing 100 dB to humans represents about 80 dB to
cervids, equivalent to lorries passing people at 30 m and
60 km/h. Krausmans group concluded that low-level
flights have little consequences for ungulates, even in NP.
In contrast, Chile, where huemul helicopter census is
valued for their great utility (SAG, letter 12.3.2003),
proposed huemul helicopter captures 17 years ago, and in
April 2005, they caught and transported huemul over
1,500 km by helicopters, without mortalities (Saucedo,
Fig. 2 Net gunning of red deer in habitat used by huemul only
decades earlier
Table 2 Questions about huemul which can be studied through reintroduction or reinforcement with radio-collared animals
Population
dynamics
Rates of survival, fecundity, recruitment; causes of mortality; population trends, etc.
Dispersal
behavior
Source-sink and metapopulation dynamics; types of barrier; migrations; corridors, etc.
Physiological
ecology
How adaptable are huemul in relation to habitat types, disturbances, etc.
Behavioral
ecology
Responses to livestock, alien species, tourism, roads, etc.
Genetic
variation
How is it reflected in persistence or growth of a population?
Reproductive
biology
Life-time reproductive success of wild females; Allee effect
Habitat use Seasonal changes, year-to-year variation; in relation to density, sex, reproductive status
Predation Effect of natural and alien predators; poaching
Population
estimates
Ratio of marked to unmarked individuals; calibration with indirect methods
Natural
history
Details often reveal factors behind a decline or recovery, especially if evaluating and comparing several sites
(Sutherland 2000)
74
personal communication). Also in Chile, huemul did not
leave areas subjected to prolonged, intensive, and repeated
low-flying helicopters for logging.
Transportation, restocking, reintroduction, husbandry,
and research opportunities
Modern handling methods can minimize distress in
captured wild cervids (Jessup et al. 1996). Huemul have
been transported over large distances as exemplified by
successful translocation to NZ and Europe, and recently by
helicopter within Chile. Capturing whole social groups, as
feasible by net gunning, substantially reduces distress
during translocation and release. All huemul should be
radio-collared and closely monitored during the initial
weeks after release.
Assuring well-being of captive huemul requires natural
environments with minimal human interference. Centers
should be exclusively used for huemul, be located within
the natural habitat of the species, and provide semicaptive
environments with natural forage by having pens up to
50 ha (Smith-Flueck et al. 2004). This optimizes research
variety and quality by minimizing human contact and
providing natural stimuli. It requires double fencing 2.7 m
high, electric fencing, barbed external wire, and buried
fencing to keep out puma and dogs. Details regarding
genetic, demographic, behavioral, and disease issues and
other recommendations about facility design, husbandry,
and research topics were elaborated in Smith-Flueck et al.
(2004).
Natural tameness facilitates huemul husbandry (Franke
1949; Rottmann 2003; Smith-Flueck et al. 2004). The first
huemul received by Franke, student of Konrad Lorenz and
caretaker of a zoological station in Nahuel Huapi NP, was
tied hours in the rain with myriad dog bite wounds and
transported several hours by truck and boat. Franke
brought the soaked female inside and placed her in a box
containing hay next to the fire. During the night, she
jumped into his bed and remained there. A few days later,
she accompanied him, and his great Dane dog on a stroll
(Franke 1949). Taruca (H. antisensis) also adapted well to
mans presence. For many years in a Berlin Zoo enclosure
of 100 m
2
, they were exposed to constant visitors, yet still
reproduced regularly (Frädrich 1978). They were one of the
easiest cervids to keep at the zoo (Frädrich, personal
communication).
Even when >1,500 km from natural habitat, huemul
adapted well to feed provided at the Buenos Aires Zoo.
Several births occurred in the Dehesa Zoo, in Puerto Radal,
in southern Argentina, and possibly in the Buenos Aires
Zoo (Smith-Flueck et al. 2004). The only contemporary
person having worked with captive huemul found no
indication that captive conditions would impair huemul
husbandry (Rottmann 2003).
Concerns about manipulating huemul in the light
of history
Inappropriate captures and transports, inadequate manage-
ment after confinement, and lack of attention to diseases
were the main reasons for failed past attempts to breed
huemul in captivity. Huemul, chased by dogs for hours or
long distances until cornered, exhibited dog bites, fractured
legs, lesions, and trauma from brutalities (Díaz 2002) and
often died within hours or days after being captured
(Franke 1949). In 1932, Holmberg acclimated 16 huemul
to individual crates to be transported from the Andes to
Buenos Aires. Adverse climatic conditions resulted in deer
being taken out and hoisted into the ship against
instructions; all animals died from nervous shock
(Smith-Flueck et al. 2004). In 1937, a male arrived at
Puerto Radal with a fractured hind leg and died from
internal hemorrhages (Franke 1949). In 1941, two huemul,
lassoed for the Santiago Zoo, received fatal dog wounds
(Díaz 2002). Then in 1973, a male died from injuries and
three others died soon after arriving to Dawson Island,
having lost >50% of their weight; it was speculated that
declined physical conditions were aggravated by small pen
sizes and low variety of food (Smith-Flueck et al. 2004).
These past failures resulted in current conjectures in
Argentina that manipulating huemul is too risky (Ramilo
2001). Nonetheless, numerous successes occurred, which
surprisingly have been overlooked. Possibly the first
huemul arrived at the London Zoo (1830) and died
6 months later. Three huemul were translocated to NZ in
1870. Then, in 1881, the London Zoo obtained, from the
Paris Jardín de Aclimatación, a male who survived capture
and transport by ship to France and then to England, aside
from all terrestrial transports (Smith-Flueck et al. 2004).
The Buenos Aires Zoo captured a male and two females in
1936 and tamed them before transportation across several
hundred kilometers of dirt road and by sea to Buenos Aires.
They adapted rapidly to small pens and human presence
(Fig. 3), and only recently discovered by Díaz (2002), some
animals were still present in 1942. The zoo collector also
had maintained huemul on his property where they
reproduced successfully (Díaz and Smith-Flueck 2000).
Puerto Radal began operating in 1936 with Franke, whose
meticulous notes, compiled over several years, were
unfortunately lost, leaving only his book documenting
tameness, births, and husbandry conditions (Fig. 4; Franke
1949). It terminated because all huemul were accidentally
released during Frankes absence in 1941. Between 1942
and 1952, new individuals arrived including a female that
gave birth and a male that survived 4 years, but the program
closed in 1956 for lack of funds (Smith-Flueck et al. 2004).
In Chile, huemul were brought to Dehesa Zoo in 1979:
two males survived, three females were added in 1982, and
three birth were recorded (Smith-Flueck et al. 2004).
Huemul kept there in large pens during 10 years had no
difficulties with the diet (Iriarte, personal communication).
75
Eight huemul were reintroduced between 1977 and 1981
to Torres del Paine NP, Chile (Rau 2003). In 1989, a
Chilean proposal to capture 60 additional huemul using
helicopter (CONAF, Proposal to San Diego Zoological
Society) to stock a center for repopulating this park never
materialized. Fortunately, the population has increased
regardless (Rau 2003).
Taruca came to the Berlin Zoo in 1889, surviving for
3.5 years. In 1891, a female arrived at the Hamburg Zoo,
and in 1929, a male arrived at the Hagenbeck Zoo (Frädrich
1978). The Berlin Zoo was most successful where one
taruca pair resulted in 12 offspring born following their
1931 arrival, and a male lived 11 years before the zoo was
destroyed during WWII (Frädrich 1978). A female
survived in the Bronx Zoo from 1938 to 1943 (Frädrich
1978). A male currently in a Zoo in Lima, Peru, lives on
commercial cattle feed, alfalfa hay, and carrots.
We like to emphasize that stress is an inevitable side
effect of manipulation of wild animals. However, stress
is only of relevance if excessive (called distress), when it
negatively affects survival or fecundity (Gill et al. 2001).
Future challenges for huemul conservation in Argentina
Aerial censuses of eight nonprotected subpopulations,
which were proposed by the authors, were denied by the
responsible authorities before a formal proposal could be
submitted. Subsequently, we secured private financing to
unite nearly 70 participants for the national recovery
workshop (2001), including experts from the USA, three
representatives of WCS, and Chileans who had elaborated
their recovery plan shortly before. The meeting was spon-
sored by Bodega Norton, Turner Endangered Species Fund,
Fig. 4 Huemul in Puerto Radal,
Nahuel Huapi National Park,
19361941 (Franke 1949; photos
courtesy of H. Franke-Giron)
Fig. 3 Huemul lived in the
Buenos Aires Zoo at least from
1936 to 1942 where they
adapted well (Díaz 2002; photos
courtesy of the Editorial Atlán-
tida, Buenos Aires)
76
IUCN SSC, Earth Restoration Alliance, and Island Founda-
tion (www.iibce.edu.uy/citogenetica/deer). In our presenta-
tion, we outlined a conservation center for semicaptive
huemul, based on existing long-term financial commitments
by international NGOs, allowing application of modern
methods (Smith-Flueck and Flueck 2001c). Surveys of
eight nonprotected subpopulations again formed the basis
of the planned study that included placement of four GPS-
and two VHF-radios per site. The proposal, although a first
public presentation, was considered irrelevant for discus-
sion during the recovery plan elaboration by an orga-
nized opposition, which included representatives of WCS,
who had been provided previously with the official
position of RDP that huemul in semicaptivity are un-
necessary (Ramilo 2001). Yet the resulting plan concluded
(2001) that the actual huemul situation demands major
improvement in amount and quality of information within a
few years, to avoid risks of gradually losing many
subpopulations. Per agreement, signed by all stakeholders,
volunteers prepared the draft plan, but contrary to this
agreement, no scientific commission was established
afterwards to review drafts before delivery to government
authorities, and no consideration was given for public
review. No budget exists, external funding is assumed, and
hence, implementation is speculative. Initial recommenda-
tions towards huemul conservation (APN, Proc. First
Huemul Conf., 1992) were reiterated in subsequent
workshops (1995, 1998, 2002), but most recommendations
were not accomplished. These were therefore again in-
corporated into the recovery plan (Table 3). If the goal is
recovery, measures should be applied to most subpopula-
tions, yet two of three provinces with huemul still have not
approved the plan. Lack of funds prevented implementing
the plan thus far (IUCN SSC, quadrennial report, 2005).
Unfortunately, recommendations from the Chilean recov-
ery plan were not considered, and the need for an Argentine
center has never been evaluated. Meanwhile, Chile recently
stocked its first private center with huemul brought from
nonprotected areas and considered the situation critical
enough that its National Science Council is supporting the
cloning of huemul using red deer as surrogate mothers
(www.conicyt.cl).
Prevailing reluctance also relates to procuring founders.
RDP claimed that knowledge on Argentine huemul indicates
that efforts should be in situ; furthermore, a donating
subpopulation would first have to be found, unlike Chile,
which has a potential donor population (Ramilo 2001).
However, our outline, in accordance with IUCN policy
(2002), recommended that captured animals represent as
much genetic variability as possible. Thus, the approach was
to census eight potential donor subpopulations and evaluate
if one social group (23 animals) per site could be captured
subsequently, and together form the initial stock. Wharton
(1995) concluded that removal of few animals has virtually
no impact even for threatened populations. Insisting that
founders must come from one sole subpopulation is unwise,
considering that subpopulations had been very reduced,
isolated, and likely genetically affected. Ironically, the
assertion by RDP that no sole population is in a condition
to risk removal of several members only further emphasizes
the critical status of the Argentine huemul.
Conclusions
One challenge to establish a comprehensive conservation
center had been overcome: long-term financial commit-
ments allowing modern approaches. However, important
obstacles were the lack of consensus regarding necessity
and feasibility to use this conservation tool, and a
Table 3 ChileanArgentine workshops (1992, 1995, 1998) and the Argentine draft recovery plan of 2001: biologically and ecologically
relevant studies urged to be priorities, and analysis in subsequent meetings as to having made advances in these issues in Argentina
Issue 1992 1995 1998 2001
Urged Advances Urged Advances Urged Advances Urged
Population
Density Yes No Yes No Yes No Yes
Population tendency Yes No Yes No Yes No Yes
Sex and age structure Yes No Yes No No Yes
Group composition Yes No No No Yes
Health and nutritional status Yes No Yes No No Yes
Seasonal habitat use Yes No No No Yes
Reproduction Yes No Yes No Yes No Yes
Predation Yes No Yes No Yes No Yes
Determine factors causing declines Yes No No No Yes
Interaction with alien species Yes No Yes No No Yes
Control spread of red deer Yes No Yes No Yes No Yes
Interaction with tourism, forestry, livestock Yes No Yes No Yes No Yes
Protected areas
Provide personnel, infrastructure Yes No Yes No Yes No Yes
Prepare management plan Yes No Yes No Yes No Yes
Increase control of areas, hunting Yes No No Yes No Yes
77
controversy centered around different views of what
constitutes recovery. Judging the predicament of Argentine
huemul differs fundamentally. Are the 350600 huemul,
fragmented into several dozen subpopulations, viable and
will they remain so? Will the roughly 220 huemul living in
nearly 23,000 km
2
of NP guarantee survival of the species?
Is the claimed low level of urgency an expression of excess
confidence due to ignoring uncertainties? Reed et al.
(2003) calculated mean minimum viable population size to
be 7,316 adults in order to last 40 generations. We are not
aware of any data showing that current conditions of
huemul groups could be considered viable. Also, recovery
implies ecologically effective densities rather than only
viability (Soulé et al. 2003).
One can postulate that measures, if applied as up to now,
will also be sufficient for recoverythe current dogmatic
policy of the RDP (Ramilo 2001). Serious problems dis-
cerned for Argentine cervids include absence of studies and
management plans (Dellafiore and Maceira 1998), and con-
sidering economics and politics, chronic financial con-
straints are likely to continue. Postponing a center likely
hinders timely advances in knowledge and thus develop-
ment of efficient conservation plans. NP with similar sizes
have lost >27% of large mammals since establishment, and
small initial population size was the most critical determi-
nant of extinction (OGrady et al. 2004). Likewise, Chilean
PA have not prevented that >50% of vertebrate populations
are now endangered, and 55% of PA are too small to allow
viable large mammal populations. Huemul possibly will
follow extinction patterns of wild sheep (Berger 1990),
and without confident knowledge on factors preventing
recovery, conservation efforts might simply amount to
documenting the loss of further subpopulations.
Different views about huemul conservation are worth-
while and need to be discussed. RDP determines research
needs, authorizes projects, but effectively prevents others
from determining scientific needs and priority setting. For
instance, scientists working in NP cannot obtain permits
from RDP (Martin 2003) to collect unanticipated wildlife
remains having scientific value, on the grounds that it
impedes hypothetical future studies on such remains.
Although remains could persist for years, incidental
encounters may provide invaluable information which is
lost if not collected. Most remains, however, even bones,
disappear rapidly (Cook et al. 2004). Thus, endangered
huemul can only be studied through RDP-approved
projects, and incidental finds and observations are value-
less because they cannot be collected.
Unresolved issues were published by editors of
binational conference proceedings as consentedconclu-
sions and recommended to be incorporated in binational
treaties without reevaluations by participants. Yet, monop-
olizing important issues will likely not be beneficial to
huemul recovery. Errors translated into official policy are
cardinal under the likelihood that no funds for adequate
monitoring will be available to detect consequences in
appropriate time. Scientific input in recovery plans has
major positive effects on success (Boersma et al. 2001),
and guidelines, even federal laws (Sutinen et al. 2004), are
available on current standards for science-based conserva-
tion planning, emphasizing involvement of qualified
scientists during all phases, rigorous replicable methodol-
ogy, repetition of studies by different groups, use of best
scientific information available, acknowledgment of cri-
tiques and alternative points of view, transparency, full
public access and participation, and peer review. The
assertion that Argentine huemul status is not critical
enough to justify the implementation of modern tools
like aerial surveys or ex situ programs, that there are ample
subpopulations with good conservation status, and that
current in situ efforts are sufficient to guarantee long-term
survival (Ramilo 2001) is subject to serious doubts. This
review indicates that sufficient grounds exist to consider
the status of Argentine huemul highly precarious.
In 1936, Argentine NP created Puerto Radal station to
rear huemul for repopulating park areas already devoid of
deer. In 1971, the Operativo Nacional Huemulwas
launched to prevent huemul from extinction. Institutions
included NP Administration, National Directory for
Renewable Natural Resources, Argentine Scientific So-
ciety, Institute of Continental Ice, Buenos Aires Zoo,
National Border Police, Museum of Natural Sciences,
Nature Association, and Argentine Mountain Federation.
Objectives were to capture and breed huemul and
repopulate NP areas. For unknown reasons, this program
never materialized. The indications are that today, the
situation is much worse. Taking this into account, the
precautionary principle would not allow waiting until
more subpopuations are lost before more active strategies
are considered warranted again.
Acknowledgements We thank colleagues from the following
institutions whose insights have shaped our views: Center of
Reproduction of Endangered Species in San Diego, Wildlife
Pharmaceuticals, Turner Endangered Species Foundation, and the
Eyes of the Wild Foundation. We thank the Wildlife Conservation
Society, the Lincoln Park Zoo, the World Nature Association, but
especially G. Langes, the Club Mil Rosas and Bodega Norton for
supporting research on huemul. Bodega Norton also covered costs of
the Argentine national recovery plan workshop, permitting that all
stakeholder could unite. L.H. Carpenter kindly provided information
on capture success. N.I. Díaz, G. Bubenik, U. Kierdorf, J. Marshall,
A. Povilitis, S. Ranney, Dan Wharton, and several anonymous
reviewers provided many challenging suggestions. We like to
dedicate this work to the late Victor Arrechea, formerly super-
intendent of the Nahuel Huapi National Park, for his foresight to
organize the National Recovery workshop for huemul.
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... Esta especie se encuentra en la categoría en peligro de extinción debido a factores como la pérdida y degradación del hábitat, la caza furtiva, la interferencia con ungulados exóticos y la depredación por perros Díaz and J. Smith-Flueck 2001). Esto conlleva a que las poblaciones sean pequeñas y se encuentren fragmentadas y confinadas a sitios de mala calidad (Flueck et al., datos no publicados;Flueck and Smith-Flueck 2006;Corti et al. 2011). Nuestro modelo muestra que toda la cordillera de la Patagonia es un hábitat climáticamente apto para que la D����������� ��������� �� ��������� ������� ₂₉ especie pueda sobrevivir (Rosas et al. 2017;Quevedo et al. 2017); esto se ve reflejado en la distribución actual de esta especie. ...
Article
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Ante las previsiones sobre el cambio climático y su influencia en los rangos geográficos de las especies, es importante esclarecer qué indicadores climáticos son los que mejor explican las distribuciones actuales, como herramientas de conservación de especies amenazadas. Asimismo, es relevante conocer el porcentaje de protección que tienen los hábitats idóneos para dichas especies. En este trabajo identificamos las principales variables climáticas que explican los patrones de distribución de 16 especies de mamíferos medianos y grandes en la Patagonia mediante el modelo MaxEnt, utilizando 19 variables bioclimáticas del WorldClim, la elevación, el NDVI, el EVI y un proxy de presión antrópica (human footprint). Calculamos el porcentaje de cobertura de áreas protegidas para cada especie y encontramos que la mayor parte del hábitat ideal está fuera de ellas, y que las áreas de protección estricta (I y II) tienen más registros de presencia que las áreas de recursos manejados (III a VI). Las especies de distribución restringida son las más vulnerables a la extinción por ser menos resilientes a los cambios en la envoltura climática, sobre todo fuera de áreas protegidas. [Palabras clave: envoltura climática, modelo de distribución, Patagonia, mamíferos nativos] Potential distribution models of native mammals in Patagonia. Given the prospects of climate change and its influence on the species' geographic range, it is important to clarify which climate indicators best explain current distributions, as tools for the conservation of threatened species. It is also important to know the percentage of protection of suitable habitats for these species. In this paper we identify the main climatic variables that explain the distribution patterns of 16 species of medium and large mammals in Patagonia through the MaxEnt model, using 19 bioclimatic variables from WorldClim, elevation, NDVI and EVI and a proxy for anthropic pressure (human footprint). We calculated the percentage of coverage of protected areas for each species and found that most of the ideal habitat is outside them, and that strictly protected areas (I and II) have more records of presence than managed resource areas (III to VI). Species with restricted distributions are the most vulnerable to extinction because they are less resilient to changes in the climatic envelope, especially outside protected areas.
... Free-ranging dogs that are poorly fed are more likely to prey on and harass wildlife (Silva-Rodríguez & Sieving, 2011). Rural dogs are interference competitors with the native fox Lycalopex griseus Silva-Rodríguez, Verdugo, et al., 2010), eat the eggs of migratory birds that nest in Chile (Schüttler et al., 2009), and predate on native deer species, including pudu Pudu puda Silva-Rodríguez & Sieving, 2012) and huemul Hippocamelus bisulcus (Corti et al., 2010;Flueck & Smith-Flueck, 2006). Reports from Peru (Barrio, 2007) state that fawns of taruka, Hippocamelus antisensis, a deer also found in northern Chile, may be killed by shepherd and feral dogs. ...
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Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe), are reported to be very susceptible to predation by pack-hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack-hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies, and hunting behavior. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviors of guanacos: predator inspection of ambush predators, for example, Puma concolor, and rushing at and kicking smaller cursorial predators, for example, Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group-hunting canids east of the Andes, there were no native species creating group-hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group-hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.
... Freely ranging dogs that are poorly fed are more likely to prey on and harass wildlife (Silva-Rodríguez & Sieving 2011). Rural dogs are interference competitors with the native fox Lycalopex griseus (Silva-Rodríguez et al. 2010), eat the eggs of migratory birds that nest in Chile (Shuttler et al. 2009), and predate on native deer species, including pudu Pudu puda (Silva-Rodríguez et al. 2009;Silva-Rodríguez & Sieving 2012), and huemul Hippocamelus bisulcus (Flueck & Smith-Flueck 2006;Corti et al. 2010). Reports from Peru (Barrio 2007) state that fawns of taruka, Hippocamelus antisensis, a deer also found in northern Chile, may be killed by shepherd and feral dogs. ...
Preprint
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Artiodactyl prey species of Chile, especially guanacos ( Lama guanicoe ) are reported to be very susceptible to predation by pack hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack-hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies and hunting behaviour. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviours of guanacos: predator inspection of ambush predators, e.g. Puma concolor , and rushing at and kicking smaller cursorial predators, e.g. Lycalopex culpaeus . We propose that since the late Pleistocene extinction of hypercarnivorous group-hunting canids east of the Andes, there were no native species creating group-hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.
... Es más probable que perros que se dejan en libertad y son mal alimentados depreden o acosen a la fauna nativa (Silva-Rodríguez & Sieving 2011). Perros en áreas rurales son competidores por interferencia con el zorro nativo Lycalopex griseus , comen los huevos de aves migratorias que anidan en Chile (Schüttler et al. 2009), y depredan especies de ciervo nativo, incluyendo al pudú (Pudu puda) , Silva-Rodríguez & Sieving 2012, y al huemul (Hippocamelus bisulcus) (Flueck & Smith-Flueck 2006;Corti et al. 2010). Reportes de Perú (Barrio 2007) mencionan que cervatillos de taruka (Hippocamelus antisensis), un ciervo que también puede encontrarse en Chile septentrional, pueden ser acabados por perros, pastores y asilvestrados. ...
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Traducción al español por Benjamín Silva del manuscrito original en preprint en Authorea.
... Es más probable que perros que se dejan en libertad y son mal alimentados depreden o acosen a la fauna nativa (Silva-Rodríguez & Sieving 2011). Perros en áreas rurales son competidores por interferencia con el zorro nativo Lycalopex griseus , comen los huevos de aves migratorias que anidan en Chile (Schüttler et al. 2009), y depredan especies de ciervo nativo, incluyendo al pudú (Pudu puda) , Silva-Rodríguez & Sieving 2012, y al huemul (Hippocamelus bisulcus) (Flueck & Smith-Flueck 2006;Corti et al. 2010). Reportes de Perú (Barrio 2007) mencionan que cervatillos de taruka (Hippocamelus antisensis), un ciervo que también puede encontrarse en Chile septentrional, pueden ser acabados por perros, pastores y asilvestrados. ...
... Besides the rocky areas of the highlands of southern Andes, the Patagonian Huemul (H. bisulcus) also resides forests, shrublands, grasslands, wetlands (inland) and deserts of lowlands from Chile and Argentina (Frid, 2001;Flueck and Smith-Flueck, 2006;Black-Decima et al., 2016). Furthermore, the extinct species Hippocamelus sulcatus was never found at Andean deposits, and besides the Brazilian record of Vidal (1946), it is restricted to lowlands of Buenos Aires and Corrientes provinces in Argentina (Alcaraz, 2010;Alcaraz and Francia, 2013;Chimento et al., 2019), and possibly also at Córdoba Province (see Castellanos, 1944). ...
Article
The tragic fire at the National Museum (Brazil) in September 2018 caused invaluable losses to South American natural and cultural heritage. However, previous visits to the paleontological collection generated photographic and descriptive records of mammalian fossil specimens from the Pleistocene of Brazil. Thus, it was possible to redescribe and revise the taxonomy of Cervidae fossils from two important paleontological sites in the Brazilian Intertropical Region (João Cativo and Lage Grande). These fossils include antlers and lower teeth, and they were described and taxonomically identified by morphological comparisons with all extant and extinct South American deers. A new diversity of Cervidae was recognized, differently from that proposed in the literature, which recognized the presence of Hippocamelus sulcatus, Mazama americana, and Ozotoceros bezoarticus. Instead, the Cervidae fossils from João Cativo and Lage Grande sites belong to the extinct deer Morenelaphus. Although some specimens could not be identified at a specific level (Cervidae indet.), a comparative analysis of the estimated body mass pointed out the most massive deer in South America (including extinct and extant) and recognized Morenelaphus as the second heaviest. The record of these large deers with huge antlers suggests the presence of natural grasslands areas and milder climatic conditions for Late Pleistocene in the Brazilian Intertropical Region. Climatic and environmental changes may have driven these large deers to extinction in the Pleistocene/Holocene transition. This study reveals an unprecedented Cervidae diversity for the Brazilian Intertropical Region, emphasizes the relevance of dental features, together with antlers morphology, in taxonomic studies of Cervidae, and also highlights the importance of South American natural heritage in the retrieving of knowledge and new scientific discoveries through collection specimens.
... Freely ranging dogs that are poorly fed are more likely to prey on and harass wildlife (Silva-Rodríguez & Sieving 2011). Rural dogs are interference competitors with the native fox Lycalopex griseus (Silva-Rodríguez et al. 2010), eat the eggs of migratory birds that nest in Chile (Shuttler et al. 2009), and predate on native deer species, including pudu Pudu puda (Silva-Rodríguez et al. 2009;Silva-Rodríguez & Sieving 2012), and huemul Hippocamelus bisulcus (Flueck & Smith-Flueck 2006;Corti et al. 2010). Reports from Peru (Barrio 2007) state that fawns of taruka, Hippocamelus antisensis, a deer also found in northern Chile, may be killed by shepherd and feral dogs. ...
Preprint
Full-text available
Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe) are reported to be very susceptible to predation by pack hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack-hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies and hunting behaviour. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviours of guanacos: predator inspection of ambush predators, e.g. Puma concolor, and rushing at and kicking smaller cursorial predators, e.g. Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group-hunting canids east of the Andes, there were no native species creating group-hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.
... Their population trend is References: F= female, M= male, Bp= sequence base pairs. # Accession numbers of sequences retrieved by BLAST alignment decreasing as a consequence of habitat loss, threats related to human activity (mainly for the presence of dogs Canis lupus familiaris), and competition with introduced exotic ungulates, such as red deer and livestock (Flueck and Smith-Flueck 2006; Duarte and González 2010; IUCN 2020). Therefore, these species (and specially the huemul), are in numerical regression and require the establishment of habitat corridors and the enlargement of protected areas to increase effective population size (Corti et al. 2011). ...
Article
Sarcocystis spp. are intracellular protozoan parasites with heteroxenous life cycles. This study described Sarcocystis spp. infection in adult South American native deer huemul (Hippocamelus bisulcus) and pudu (Pudu puda). Heart, diaphragm, tongue, and skeletal muscle samples were collected from 5 huemuls and 2 pudus, found dead in National Parks. Direct microscopic examination, transmission electron microscopy, PCR, and sequencing were performed. Sarcocystis spp. microscopic thin-walled cysts were identified in 3 huemuls and 1 pudu. Several cysts from 1 huemul and 1 pudu were observed by TEM; ultrastructure was similar to previously reported as cyst wall type 17 and types 2 and 8, respectively. Fragments of the 18S rRNA and cytochrome c oxidase subunit I (cox1) genes were amplified and sequenced from 3 individual cysts from 2 huemuls and 2 cysts from the pudu. The sequences from huemuls showed a high identity among them (> 99%) at both amplified targets. The highest identities were > 99.7% at 18S rRNA and 93% at cox1 with S. tarandivulpes sequences. The 18S rRNA gene sequences from pudus showed an identity > 99.7% with Sarcocystis sp., S. taeniata, and S. linearis sequences, while the cox1 sequences were different, one showing 99.42% identity with S. venatoria and the other 98.22% with S. linearis. A single species, similar to S. tarandivulpes, was identified in all huemul samples while 2 molecularly different Sarcocystis spp. were found in 1 pudu with high similarities to either S. venatoria or to S. linearis, S. taeniata-like, and S. morae. Based on the cox1 sequence identities, at least the Sarcocystis sp. in huemuls might represent a new species, primarily occurring in this host. Additional sarcocyst isolates from both hosts need to be examined molecularly in order to firmly establish whether these species are indeed native to huemuls and/or pudus or are derived from introduced deer species.
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The huemul (Hippocamelus bisulcus) is endangered, with 1500 deer split into >100 subpopulations along 2000 km of 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.
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Early naturalists already considered huemul rare, refuged and a stocky, short‐legged mountain deer, 163 years before declared endangered (1972). Anatomically, huemul do not overlap with rock‐climbers previously considered analogous, as corroborated in this paper by including additional huemul cases. Assertions that population declines are caused principally via livestock infections remain unfounded. Instead, osteopathology in multiple populations across 1,000 km, affecting 57% among dead and 86% among live specimens, may relate to micronutrient deficiencies. Historically classified a mountain deer, widespread osteopathology, micronutrient deficiencies and lack of recovery qualify huemul as a refugee species. Recovery strategies thus must include repopulating historical distribution sites.
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The red deer (Cervus elaphus), among the world's 14 most invasive exotic mammals, has recently arrived in Patagonia. Forty-seven deer were captured, marked with radio collars, and monitored in order to determine survival rates and identify causes of death. Net gunning from a helicopter allowed captures to be evenly distributed through the study area and in a timely manner. The absence of capture related mortality in our study agrees with previously reported low rates for this method. Animals were monitored for periods ranging from 5 to 2611 days. Ten animals provided censored values due to radio failures or being shot: they survived on average 811 days (SE = 221) after capture. Ten animals experienced natural deaths with an average time alive after capture of 413 days (SE = 106): eight were killed by puma (Puma concolor). The remaining 27 animals were alive for an average of 974 days (SE = 52). Twenty-one percent of the animals died naturally during 38.549 deer-days. Using the Kaplan-Meier procedure, the annual survival rates for the years 2001-2003 were 0.89, 0.91 and 0.92 respectively (n = 47); the average annual survival rate being 91%. Although predation was the most important cause of adult mortality, all mortality sources combined (puma predation, legal and illegal hunting, disease, emigration) have not maintained deer population density low enough to prevent food limitation from occurring in the recent past.
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The huemul (Hippocamelus bisulcus), an endangered deer living in the Andes of southern South America, numbers 600-700 in Argentina, from at least 63 fragmented subpopulations. Very limited information exists on most populations, but there are indications that the distribution and population sizes continue to diminish, possibly through reduced reproductive rates, elevated mortality rates or both. This is the first analysis of the population ecology of a subpopulation in Argentina, made possible through the only existing compilation of numerous remains of dead huemul (n = 19). Surveys were conducted between 1993 and 1999 to locate and collect remains, which were found concentrated at lower elevations. The sex ratio of adults and subadults (n = 12) was 1:1, but could not be determined for fawns. Clear signs of predation was found in 37% of all deer: 50% among calves, 50% among subadults and 31% among adults. These signs occurred in 29% of adult females and 50% of adult males. Deaths occurred during winter as judged by the state of the male skulls, which indicated the phase of the antler cycle. Femur fat content of 3 deer averaged 93%. Predation signs frequently were clearly from puma (Puma concolor), which is the only natural predator of huemul. The high predation rate, high marrow fat content and no indications of deaths resulting from disease corroborate the pristine state of the of mature lenga (Nothofagus pumilio) forest. The area with a concentration of huemul carcasses appears to be the huemul's major winter habitat.
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Textbooks on the principles of conservation biology abound. Yet, how does one put this theoretical knowledge into practice? The aim of The Conservation Handbook is to provide clear guidance on the implementation of conservation techniques. The wide range of methods described include those for ecological research, monitoring, planning, education, habitat management and combining conservation with development. Nineteen case studies illustrate how the methods have been applied. The book will be of interest to conservation biology students and practicing conservationists worldwide. For each copy of the book sold, another copy will be sent free to a practicing conservationist outside Western Europe, North America, Australia, New Zealand and Japan. Foreword by E. O. Wilson. Concise, practical guide packed full of ideas, methods and advice. Provides solutions for the main conservation problems most commonly encountered. 18 global case studies illustrate the application of techniques. The Conservation Handbook Donations Project this book is being sent free to those practising conservationists outside Western Europe, North America, Australia, New Zealand and Japan who are otherwise unlikely to obtain a copy. These copies are provided at cost price by Blackwell Science, the publisher, and paid for with the author''s royalties. Each book sold means another one will be donated.
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