 Hippocamelus
bisulcus
Patagonian huemul, South Andean huemul |
Taxonomy | Description
| Reproduction | Ecology
| Behavior | Distribution
| Conservation | Remarks
| Literature |
| Taxonomy
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Hippocamelus bisulcus [Molina, 1782].
-
Citation: Sagg. Stor. Nat. Chile, p. 320.
-
Type locality: Chilean Andes, Colchagua Prov.
The taxonomic record (above) is taken from Wilson and Reeder (1993). No
subspecies are recognized for the Patagonian huemul, although regional
differences have been noted by various authors (e.g., Frid, 1994).
Although closely related to the
taruca or North Andean huemul
(H. antisensis), the two
are consistently recognized as separate species (Geist, 1998). Invalid
synonyms for Hippocamelus bisulcus include andicus,
cerasina, chilensis, dubius, equinus,
huemel, and leucotis (Wilson and Reeder, 1993).
General Characteristics
The Patagonian huemul is the largest herbivore of the South Andean region
(MacNamara, 1990). It is a stout deer, similar in form to many
mountain-dwelling members of the
Caprinae (Redford and Eisenberg,
1992). Males are larger and heavier than females, notably around their
neck (Redford and Eisenberg, 1992; Diaz, 2000). Reported body weights
for this species fall primarily between 70 and 100 kg, and head and body
length is approximately 150 cm. This species is larger and darker in
coloration than its northern counterpart, the
taruca
(H. antisensis) (MacNamara,
1990; Geist, 1998).
Reported measurements for Patagonian huemul (Hippocamelus
bisulcus) |
| Source |
Adult Weight |
Head & Body Length |
Shoulder Height
|
Tail Length |
| Diaz, 2000 |
- |
163 cm 
151 cm  |
90 cm
81 cm |
13 cm |
| MacNamara, 1990 |
45-65 kg |
150-170 cm |
91 cm |
11.5-13 cm |
Nowak, 1991
for Hippocamelus |
45-65 kg |
140-165 cm |
77.5-90 cm |
11.5-3 cm [sic] |
| Redford and Eisenberg, 1992 |
70.0 kg |
149.7 cm |
80.0-90.0 cm |
12-13 cm |
| Smith-Flueck, 2000 |
100 kg |
140-175 cm
140-157 cm |
79-100 cm
78-84 cm |
- |
| Whitehead, 1993 |
70-100 kg |
- |
80-90 cm |
- |
The dense, coarse pelage of H. bisulcus is a uniform dark brown, varying
in summer from dark rust to deep coffee in color (MacNamara, 1990; Redford
and Eisenberg, 1992; Geist, 1998; Diaz, 2000). The winter coat is paler,
with a yellow or grey tint (Redford and Eisenberg, 1992; Smith-Flueck, 2000).
The undersides, especially the anal and inguinal regions, are whitish,
contrasting sharply with the dark upper coat (MacNamara, 1990; Diaz, 2000).
There is no lightly-colored rump or throat patch as seen in
H. antisensis, but a darker
spot is often present on the hindquarters (MacNamara, 1990; Geist, 1998).
The tail is short with a white underside (Diaz, 2000)
The coat is comprised of two layers: a long, greasy, bristly layer of guard
hairs which covers a dense, woolly undercoat (MacNamara, 1990; Nowak, 1991).
The brittle guard hairs are hollow which, combined with the undercoat,
provides insulation to -50oC (MacNamara, 1990; Diaz, 2000).
The pelage in winter is comprised of guard hairs 5-7 cm long, which are shed
in large clumps during the spring and replaced with a summer coat with hairs
3-4 cm in length (Diaz, 2000; Smith-Flueck, 2000). In all seasons,
the hairs are slightly crinkled, growing the longest on the underside, tail,
and legs (Nowak, 1991; Smith-Flueck, 2000).
The build of the Patagonian huemul is distinctly stocky, a result of an arched
back and relatively short legs (Nowak, 1991; Smith-Flueck, 2000). The
resulting crouching position is typical of many mountain-dwelling ungulates,
as are the hard, pointed, sharply-edged hooves (Geist, 1998; Smith-Flueck,
2000). Tarsal glands are present on the rear legs, but metatarsal glands
are absent (Geist, 1998).
The ears are long and narrow (described by MacNamara (1990) as "donkey ears");
they measure around 17 cm in length, and are covered on their inner surface
with white hair (Diaz, 2000; Smith-Flueck, 2000). The only facial marking
is a dark forked blaze on ridge of the nose, which runs from the dark rhinarium
and splits in between the eyes to form 'eyebrows' (Smith-Flueck, 2000).
While this blaze is usually attributed to males alone, both sexes possessed
such markings in a coastal Chilean population studied by Frid (1994). The
eyes are surrounded by a vague area of lighter colored hair, which accentuates
the large preorbital glands (Geist, 1998; Diaz, 2000). Varying authors
report either a presence (MacNamara, 1990; Nowak, 1991) or absence (Smith-Flueck,
2000) of upper canine teeth.
Only males grow antlers; these are usually simple in form and typically branch
just once, although antlers with up to five tines have been recorded (Nowak,
1991; Redford and Eisenberg, 1992; Smith-Flueck, 2000). In a typical forked
antler, the rear tine is the longer, typically measuring 30-35 cm in length
from the burr (where the antler joins the skull) to the tip of the tine (Nowak,
1991; Redford and Eisenberg, 1992, Diaz, 2000). In young males the
first set of antlers appears around eighteen months of age, although they
may appear as early as five months of age (Redford and Eisenberg, 1992;
Smith-Flueck, 2000). Males cast off their antlers near the end of the austral
winter, from mid July to early September (Smith-Flueck, 2000), although Geist
(1998) states that they are dropped in October-November. New antlers begin
growing soon after old ones are shed, and the full size is reached (and
velvet shed) after around five months of growth (Smith-Flueck, 2000).
Ontogeny and Reproduction
The breeding season of H. bisulcus is poorly defined and may differ
widely between populations, but is typically at some time between mid-February
and May (Smith-Flueck, 2000). The Patagonian huemul appears to have
a dominance group mating system, wherein one dominant male will mate with
most of the females in a group (Smith-Flueck, 2000). Although a male
(buck) will mate with many females (does) as they become receptive, the
association with a single doe may last for as long as 8-9 weeks, including
a long period of time after mating (Smith-Flueck, 2000).
The gestation period has never been measured directly, but estimates range
from 6-8 months - reported numbers include 180-210 days (Redford and Eisenberg,
1992), 200-220 days (Smith-Flueck, 2000) and 240 days (Geist, 1998).
Females almost always give birth to a single young, usually in the
months of November and December, a time when the local vegetation is highly
nutritious (Frid, 1994; Smith-Flueck, 2000). Fawns are born with a
thick, woolly coat which is not spotted (unlike many deer species), but rather
a solid dark coffee-brown color (MacNamara, 1990; Redford and Eisenberg,
1992; Smith-Flueck, 2000). A single newborn huemul examined by Smith-Flueck
(2000) weighed 2035 g, with a total body length of 61.5 cm. Milk teeth
show at birth (Smith-Flueck, 2000).
Prior to giving birth, females seek seclusion; born in an isolated area (often
along high bluffs), the newborn fawn will spend most of their time bedded
down and separated from their mother, with the resting site being changed
frequently to avoid predators. Eventually, the fawn will begin to follow
its mother, although at a stomp of the dam's feet (indicating danger) the
fawn immediately drops to the ground and freezes. Weaning occurs at 4-5 months
of age (Smith-Flueck, 2000).
Despite the fact that females may reach sexual maturity as early as 6 months
of age, the fecundity of H. bisulcus is low, with fewer than 50% of
females breeding on an annual basis. Smith-Flueck (2000) suggests that
this may be due to a long interbirth interval, resulting in females only
breeding every other year. The maximum observed life span for this
species is around 14 years of age (Smith-Flueck, 2000).
Ecology
H. bisulcus inhabits rugged and diverse terrain in the southern Andes
(MacNamara, 1990; Smith-Flueck, 2000). While most populations are presently
restricted to high, relatively inaccessible regions between 900 and 1,700
meters above sea level, huemuls in coastal Chile have been observed at very
low altitudes, even on the beach; i.e., at sea level (Nowak, 1991; Frid,
1994; Smith-Flueck, 2000). Snow cover is a limiting factor in habitat
choice - this species is not found in areas with over 30 cm of snow
(Smith-Flueck, 2000).
Huemuls are frequently observed on north or north-western facing slopes,
often with a pitch between 30 and 40 degrees (MacNamara, 1990; Redford and
Eisenberg, 1992; Smith-Flueck, 2000). In Chubut, Argentina, however,
observations of this species have been made on slopes generally less than
25 degrees (Smith-Flueck, 2000). Patagonian huemul are usually found
at or below the tree line, principally inhabiting brushy areas with dense
shrub cover (65% of observations in Argentina) and temperate forests (20%
of observations in Argentina) (MacNamara, 1990; Nowak, 1991; Redford and
Eisenberg, 1992; Smith-Flueck, 2000). These subalpine forests are dominated
by false-beeches Nothofagus sp. (including N. antarctica, N.
betuloides, and N. pumilio; Fagaceae), with shrub cover including
Discaria sp. (Rhamnaceae), Gaultheria sp.(Ericaceae),
Embothrium coccineum (Proteaceae), Escallonia rubra
(Escalloniaeae), and Mulinum spinosum (Umbelliferae) (MacNamara, 1990;
Galende et al., 2005).
Seasonal movements are well-documented in this species; huemuls inhabit high
altitudes during summer, with frequent usage of pasture and brush above the
treeline (80%) or in forests (12.5%) (Nowak, 1991; Smith-Flueck, 2000).
Females especially prefer high cliffs and ridges (likely for the protection
they offer from terrestrial predators), while males are found almost exclusively
on flatter meadows (Frid, 1994; Smith-Flueck, 2000). From these high
elevations, huemuls move downslope in autumn in order to winter in valleys
and denser forests (Nowak, 1991; Redford and Eisenberg, 1992); seasonal movements
on the magnitude of two or three kilometers were documented in one animal
(Smith-Flueck, 2000).
Patagonian huemuls are either solitary or found in small groups; these are
generally composed of 2-5 individuals, although aggregations of up to ten
individuals have been observed at low altitudes (Redford and Eisenberg, 1992;
Smith-Flueck, 2000). The structure of these groups is highly flexible,
and a single individual may associate in several 'groups' during the course
of a single day (Smith-Flueck, 2000). Adults of both sexes are often seen
alone, while the most commonly observed grouping is a mother-offspring pair
(29%). Pairs of an adult male and female, either alone or accompanied
by juveniles, are also seen regularly, while groups of adult males are infrequent
and all-female groups never observed. Adult males tolerate the presence of
subordinate males, and it has thus been inferred that territoriality is absent
in this species (Smith-Flueck, 2000).
Local population density estimates range from 0.02 to 5.66 animals per square
kilometer (Smith-Flueck, 2000), although Redford and Eisenberg (1992) report
only the lower end of this range (0.02 to 1.2 per square kilometer).
Annual home range size in Tamango Nature Reserve, Chile was estimated
to be 300 hectares (Smith-Flueck, 2000), a figure which compares well to
the 340 hectares reported in Redford and Eisenberg (1992). During the
breeding season (between March and May), huemuls confine their movements
to ranges of 36 to 73 hectares, although this is likely a gross
underestimate of the area used throughout the year (Smith-Flueck, 2000).
On a daily basis, individuals may travel 3-3.9 kilometers as they search
for food and shelter (Smith-Flueck, 2000).
Predators of the huemul include the puma (Puma concolor) and domestic
dogs (Canis lupus familiarus), while the culpeo fox (Pseudoalopex
[Dusicyon] culpaeus), wild cats (Oncifelis guigna and O.
geoffroyi), and raptors, especially condors, may be a threat to young
(Geist, 1998; Smith-Flueck, 2000). When threatened by predators, H.
bisulcus attempts avoidance by first hiding, but if discovered at close
range will flee rapidly (Geist, 1998). This species has low endurance
when running, and will escape persistent pursuers by seeking refuge on cliffs
or by swimming into lakes - this species swims with ease (Geist, 1998; Diaz,
2000; Smith-Flueck, 2000).
H. bisulcus is a browser, feeding almost exclusively on tender vegetation
and only rarely on grasses or lichens (Geist, 1998; Smith-Flueck, 2000).
Several studies on food consumption have been conducted, which show
a strong trend with some regional variation. In addition to the species
listed below, captive animals have shown a preference for willow (Salix
sp.) (Smith-Flueck, 2000).
Galende et al. (2005), observing huemul in Nahuel Huapi National Park,
identified 32 plant species consumed by H. bisulcus. Shrubs and trees
comprised the greatest part of the diet (58.6% and 19.5% of fragments identified
in feces, respectively), while forbs and grasses were occasionally consumed,
each contributing to just over 9% of the diet. The shrub Ribes
magellanicum (Grossulariaceae) and the tree Nothofagus pumilio
(Fagaceae) were the principal components of the diet during the summer (28.5%
and 33.4%, respectively), while in winter (when these plants drop their leaves)
the shrubs Maytenus sp. (Celastraceae) and Gaultheria sp.
(Ericaceae) were used heavily (50.8% and 28.7%, respectively).
Smith-Flueck (2000) reports 48 plant species which have been identified in
the diet, comprised of 27-46% forbs and 31-72% woody plants when averaged
across the year. Maytenus were a very important food item, making
up as much as 60% of the annual diet in Argentina. Other frequently used
species include Alstroemeria (9%), Chlorea viridiflora
(Orchidaceae; 8%), Geranium (5%), and Schinus (5%).
The population living along the Chilean coast observed by Frid (1994) had
a more restricted diet, with only 11 species of plant being eaten. The perennial
forb Gunnera magellanica (Gunneraceae) was heavily preferred, with
consumption of this plant accounting for as much as 82.5% of observed feeding
time in grasslands and bluffs. Feeding in moorland habitat was concentrated
on two species: the tree Nothofagus antarctica (Fagaceae; 84.7% of
observation time) and evergreen shrub Embothrium coccineum (Proteacea;
15.3% of observation time).
Behavior
H. bisulcus is primarily diurnal in its activity patterns, but may
be active at night as well (Redford and Eisenberg, 1992). They have keen
senses of sight, hearing, and smell, but (in undisturbed areas) are remarkably
tolerant of humans and can be approached within a few meters (Diaz, 2000;
Smith-Flueck, 2000). In heavily disturbed regions, huemul are much more flighty
due to harassment by humans and especially dogs; in such areas, a huemul
may take flight if approached within 300 meters (Smith-Flueck, 2000).
The courtship behavior of the Patagonian huemul is similar to other
New World deer, and has
been described in detail by Smith-Flueck (2000). As the breeding season
approaches, males thrash their antlers against vegetation, helping to shed
the velvet from the fully formed antlers and advertising their presence (they
may fray the bark off of tree trunks from just above the ground to 113 cm
high). This behavior continues through the courtship period, and is
coupled with scent marking using the facial glands (Nowak, 1991). A
male huemul will follow a potentially receptive female, sniffing her anogenital
region and employing flehmen (lip-curl) after smelling her urine in order
to determine whether or not the female is in estrus. After determining
the reproductive status of a female, the male will approach receptive mates
with the head stretched low, performing a high amount of "foreplay" and
non-copulatory mounting before achieving intromission. Dominant males
may chase off subordinates if they approach within 300 meters of an estrus
female (Geist, 1998). Otherwise, the low population densities at which
huemuls live reduces amount of conflict between males, and may explain the
reduced antlers of this species as well as the low frequency of dominance
displays (Smith-Flueck, 2000).
Smith-Flueck (2000) describes this species' repertoire of vocalizations,
many of which are audible to humans at a distance of 30 meters or more.
During the breeding season, males either a grunt or produce a series
of soft whines while pursuing females; a doe, in return, will produce a snort
when she is the object of a male's attention. Agonistic encounters
between males are accompanied by a rapid series of sharp notes, lasting 3-6
seconds and descending in volume throughout the series; Smith-Flueck (2000)
suggests that older males may produce deeper noises, thereby advertising
the male's status. The alarm call of the Patagonian huemul is a snort,
which is usually accompanied by the stamping of the front legs. Fawns
bleat in alarm; these cries usually bring the mother immediately to
the fawn.
Genetics
The karyotype of the Patagonian huemul is 2n = 70 (Spotorno et al.,
1987 in Redford and Eisenberg, 1992).
Distribution
Prior to the arrival of Europeans in South America, the Patagonian huemul
was abundant in southern Chile and Argentina between 34° and 53°
S latitude, being found primarily along the spine of the Andes, but also
on steppes east of the mountains (Redford and Eisenberg, 1992). As
the map below indicates, the present distribution is significantly reduced,
with the majority of subpopulations being restricted to the high Andes.
Redford and Eisenberg (1992) report that H. bisulcus is presently
found in the Chilean provinces of Aisén, Magallanes, Nuble, and
continental Chiloé. The Argentinian populations are restricted
to provinces of Neuquén, Rio Negro, Chubut and Santa Cruz..
Countries: Argentina, Chile (IUCN, 2006).
Range Map (Redrawn from Diaz, 2000)
Conservation Status
H. bisulcus is classified as endangered (C2a) by the IUCN (2006),
and is listed on Appendix I of CITES (2007) along with the congeneric
taruca
(H. antisensis). The
total population of this species is estimated to be between 1,000 and 1,500
individuals, of which 350-600 inhabit Argentina (Flueck and Smith-Flueck,
2006) - both Redford and Eisenberg (1992) and Smith-Flueck (2000) suggest
that current numbers may be less than 1% of this species' pre-European abundance.
Coupled with the reduction in numbers, a reduction in range of over
50% has been noted [Weber and Gonzalez (2003) state that this species inhabits
only 30% of its former range], with the result that wild huemul populations
are now highly fragmented (Smith-Flueck, 2000).
Threats to huemuls are both ongoing and numerous. Humans affect huemul
directly through hunting (poaching) and by encroaching on this species' habitat
(IUCN, 2006). In the Nevados area of Chile (the site of the most northerly
population of huemul), land settlement, construction of pipelines, and logging
(along with the construction of logging roads) have all served to reduce
native habitats and increase the presence of humans in the region (Povilitis,
1998). An influx of tourists has also occured, although the impacts
of tourism have yet to be determined (Flueck and Smith-Flueck, 2006).
However, perhaps the largest threat comes from domestic stock and
introduced species (Redford and Eisenberg, 1992; IUCN, 2006). Harassment
and depredation by domestic dogs, susceptibility to diseases and parasites
of introduced species (especially domestic cattle and sheep), and competition
from introduced species (e.g., red deer) are all major factors in the continued
decline of this species (Povilitis, 1998; Flueck and Smith-Flueck, 2006;
IUCN, 2006)
This species is actively protected in Chile, and a conservation agreement
between Argentina and Chile is in effect (Weber and Gonzalez, 2003).
However, Flueck and Smith-Flueck (2006) expressed concern regarding
the actual conservation status in Argentina; these authors advocate increased
conservation measures be taken if this species is to continue to survive.
In 2007, the Huemul Task Force was created under the IUCN Species Survival
Commission to provide recommendations - based on sound scientific information
- through which the recovery of huemul can be achieved.
Remarks
H. bisulcus is the national animal of Chile (Weber and Gonzalez, 2003),
and is the only large herbivore to inhabit the subantarctic false-beech habitat
of Patagonia (Smith-Flueck, 2000).
The common name huemul (pronounced hw~y~mul) is of Araucanian
origin, and is said to derive from the verb "huemin", meaning "to follow";
this is apparently in reference to a custom of this species of walking in
single file (MacNamara, 1990; Smith-Flueck, 2000). The name is also
occasionally spelled as guemal. In the past, this species was popularly
known as the "Chilean huemul". However, since many populations inhabit
Argentina, the use of this name has been replaced by the more appropriate
name "Patagonian huemul" (Smith-Flueck, 2000). An increase in
the use of the local name "taruca" for
H. antisensis
has meant that H. bisulcus is now sometimes referred to
simply as the "huemul".
The genus name Hippocamelus is a combination of the Greek words
hippos (a horse) and kamelos (a camel) - while huemuls are
not closely related to either, they were first described (from hearsay
information) as a type of wild ass (Diaz, 2000). The species name
bisulcus translates from Latin as "having two furrows", or "cloven";
which Diaz (2000) suggests refers to original classification of this deer
as a cloven-hoofed horse.
-
Local names
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Huemul [from Araucanian Indians] (MacNamara, 1990; Smith-Flueck, 2000)
-
-
French
-
Huémul des Andes méridionales (IUCN, 2006)
-
Huemul du Chili (MacNamara, 1990)
-
-
German
-
Südlicher Andenhirsch (MacNamara, 1990)
-
-
Spanish
-
Huemul Patagónico (Redford and Eisenberg, 1992)
Ciervo Andino meridional (IUCN, 2006)
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