Alpine ibex
Male
Female
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Family: Bovidae
Subfamily: Caprinae
Tribe: Caprini
Genus: Capra
Species:
C. ibex
Binomial name
Capra ibex
Distribution of the Alpine ibex (population in Bulgaria not included)

The Alpine ibex (Capra ibex), also known as the steinbock, is a species of goat that lives in the Alps of Europe. It is one of ten species in the genus Capra and its closest living relative is the Iberian ibex. The Alpine ibex is a sexually dimorphic species: males are larger and carry longer horns than females. Its coat colour is brownish grey. Alpine ibexes tend to live in steep, rough terrain and open alpine meadows. They can be found at elevations as high as 3,300 m (10,800 ft) and their sharp hooves allow them to scale their mountainous habitat.

Alpine ibexes primarily feed on grass and are active throughout the year. They are also social, although adult males and females segregate for most of the year, coming together only to mate. During the breeding season, males fight for access to females using their long horns. Ibexes have few predators but do succumb to various parasites and diseases.

After being extirpated from most areas by the 19th century, the Alpine ibex was successfully reintroduced to parts of its historical range. All individuals living today descend from the stock in Gran Paradiso National Park in Italy. The species is currently listed as of least concern by the IUCN, but went through a population bottleneck of fewer than 100 individuals during its near-extinction event. This has led to very low genetic diversity across populations.

Taxonomy

The Alpine ibex was first described by Carl Linnaeus in 1758. It is classified in the genus Capra with nine other species of goats.[2] Capra is Latin for "she-goat"[3] while the species name "ibex" is a Latin name translated as "chamois" and is possibly derived from an earlier Alpine language.[4]

Fossils of the genus Tossunnoria are found in late Miocene deposits in China and they appear to have been transitional between goats and their ancestors.[5] The genus Capra may have originated in Central Asia and colonised Europe, the Caucasus, and East Africa from the Pliocene and into the Pleistocene. Mitochondrial and Y chromosome evidence show hybridisation of species in this lineage.[6] Fossils of the Alpine ibex date to the late Pleistocene (during the last glacial period) in France and Italy. It and the Iberian ibex (C. pyrenaica) probably evolved from the extinct Pleistocene species Capra camburgensis, whose fossils have been found in Germany. Alpine ibex appear to have been larger during the Pleistocene than modern day.[5]

The Nubian (C. nubiana), walia (C. walie), and Siberian ibex (C. sibirica) were considered to be subspecies of the Alpine ibex in the 20th century, giving populations in the Alps the trinomial of C. i. ibex.[7] 2006 genetic evidence has supported them being separate species.[6]

The following cladogram of seven Capra species is based on 2022 mitochondrial evidence:[8]

Capra

Siberian ibex

Nubian ibex

Markhor

Domestic goat

Bezoar ibex

Alpine ibex

Iberian ibex

Appearance

Male ibex skull

Alpine ibexes are sexually dimorphic.[2] Males grow to a height of 90 to 101 cm (35 to 40 in) at the withers, with a body length of 149–171 cm (59–67 in) and weigh 67–117 kg (148–258 lb). Females are much smaller and have a shoulder height of 73–84 cm (29–33 in), a body length of 121–141 cm (48–56 in), and weigh 17–32 kg (37–71 lb).[5]

The Alpine ibex is a stocky animal, with a tough neck and robust legs with short metapodials. Compared with most other wild goats, the species has a wide, shortened snout. Adaptations for climbing include sharp, highly separated hooves and a rubbery callus under the front feet.[2][5] Both male and female Alpine ibexes have large, backwards-curving horns with an elliptical cross-section and a trilateral-shaped core. The front surface of the horns have transverse ridges marking an otherwise flat surface. At 69–98 cm (27–39 in), the horns of males are substantially longer than those of females, which reach only 18–35 cm (7.1–13.8 in) in length.[5]

The species has brownish-grey hair over most of the body, lighter on the belly, with darker markings on the chin and throat. The hair on the chest region is nearly black while stripes exist along the dorsal (back) surface. It is duller coloured than other members of its genus. As with other goats, only males have a beard. Ibexes moult in spring, when their thick winter coat, consisting of woolly underfur, is replaced with a short, thin summer coat. Their winter coat grows back again in the fall. As a member of Capra, the Alpine ibex has glands near the eyes, and on the groin and feet, but not on the face.[5]

Distribution and habitat

Ibex walking along its mountain habitat

The Alpine ibex is native to the Alps, its range including France, Switzerland, Liechtenstein, Italy, Germany and Austria.[9] Fossils of the species have been found as far south as Greece, where they went extinct due to hunting over 7,500 years ago.[10] Between the 16th and 18th centuries, the species disappeared from much of its range due to hunting, leaving only one population surviving in and around Gran Paradiso by the 19th century. They have since been reintroduced into parts of their former range,[9] as well as new areas such as Slovenia[11] and Bulgaria.[9]

An excellent climber, the Alpine ibex occupies steep, rough terrain at elevations of 1,800 to 3,300 m (5,900 to 10,800 ft). They prefer to live an open areas,[5] but when there is little snow, adult males may gather in larch and mixed larch-spruce woodland depending on population density.[12] Outside the breeding season, the sexes live in separate habitats.[12][13] Females are more likely to be found on steep slopes while males prefer more level ground. Males inhabit lowland meadows during the spring, when fresh grass appears.[5] They then climb to alpine meadows during the summer.[12] Both males and females move to steep, rocky slopes when winter arrives, to avoid dense buildups of snow.[14] They prefer slopes of 30–45° and take refuge in small caves and overhangs.[15]

Behaviour and ecology

Alpine ibexes are strictly herbivorous, with the majority of their diet consisting of grass. Grass is preferred every season; during the summer, ibex supplement their diet with herbs, while during fall and winter they include dwarf shrubs and conifer shoots.[16] Grass genera that are most commonly eaten are Agrostis, Avena, Calamagrostis, Festuca, Phleum, Poa, Sesleria, and Trisetum.[5] The amount of time spent feeding during the day is similar between the sexes during the spring, while in summer, females feed more than males, particularly those that are lactating.[17] Hotter temperatures give large adult males heat stress, thus reducing their feeding time, but they may avoid this problem by feeding at night.[18]

In Gran Paradiso, home ranges can reach over 700 ha (1,700 acres), while in reintroduced populations they may approach 3,000 ha (7,400 acres).[5][19] Home range size depends on the availability of resources and the time of year. Home ranges tend to be largest during summer and autumn, smallest in winter, and intermediate in spring. Female home ranges are usually smaller than those of males.[5][14][19] Ibexes do not hibernate during the winter; they take shelter on cold winter nights and bask in the morning. They also reduce their heart rate and metabolism.[20] The Alpine ibex may compete with the chamois and red deer for resources.[2] The presence of those species may force the ibex to occupy higher elevations.[21]

The climbing ability of the Alpine ibex is such that it has been observed scaling the 57° slopes of the Cingino Dam in Piedmont, Italy, where they lick the artificial salts. Only females and young (kids) will climb the steep dam due to their lighter weight and shorter legs. Kids have been recorded to reach heights of 49 m (161 ft), ascending in a zig-zag path while descending in straight lines.[22]

Social behaviour

Male ibexes fighting

Although the Alpine ibex is a social species, they tend to live in separate groups based on sex and age.[5] For most of the year, adult males group separately from females, and older males live separately from younger males.[23] Female groups consist of 5–10 members, while male groups have 2–16 members but can have as many as 50 or more.[2][24] Dependent kids live with their mothers in female groups. Segregation between the sexes is a gradual process and males younger than nine years old may still associate with female groups.[25] Adult males are more likely to be found alone than females, particularly older males.[24] Social spacing tends to be looser in the summer, when there is more room to feed. Ibexes have stable social connections, and consistently regroup with the same individuals when ecological conditions force them back together.[26] Female groups tend to be more stable than male groups.[2][25]

Adult males and females gather during the breeding season between December and January. By April and May, the adults separate again.[5] A dominance hierarchy exists among males based on size, age, and horn length.[27] Hierarchies are established outside the breeding season, allowing males to focus more on mating and less on fighting. Males use their horns for combat, and will attack the sides or clash head-to-head, the latter often involves them standing bipedally and clashing downward.[28]

Alpine ibexes communicate mainly through whistles, described as "short and sharp". They primarily serve as alarm calls and may occur singularly or in succession with short gaps. Females and their young communicate by bleating.[2]

Reproduction and growth

Mother with young

The mating season begins in December, and typically lasts around six weeks. During this time, male herds break up into smaller groups that search for females. The rut takes place in two phases. In the first phase, the males interact with the females as a group. In the second phase, one male separates from his group to follow an individual female in oestrus.[5] Dominant, older males (9–12 year olds) will follow a female and guard her from rivals, while more subordinate, younger males (2–6 year olds) will try to sneak past the tending male when he is distracted. If a female flees, both dominant and subordinate males will try to follow her. During courtship, the male stretches the neck, flicks the tongue, curls the upper lip, urinates and sniffs the female.[29] After copulation, a male rejoins his group and restarts the first phase of the rut.[5] Enviromental conditions can affect courtship in the species: for example, snow can limit the males' ability to follow and mate with females.[30]

A female is in oestrus for around 20 days, while gestation averages around five months, and typically results in the birth of one but sometimes two kids.[31] Females give birth away from their social groups and on rocky slopes which are relatively safe from predators.[32] After a few days, the kids can already move on their own. Mothers and kids gather into nursery groups where young are nursed for up to five months.[2] Nursery groups can also include non-lactating females.[17] Alpine ibexes reach sexual maturity at 18 months, but females continue to grow until they are around five or six years old, and males nine to eleven years old.[5]

The horns grow throughout life. Young are born without horns; they are visible as tiny tips at one month and reach 20–25 mm (0.8–1.0 in) in their second month.[2] In males, the horns grow at about 8 cm (3.1 in) per year for the first five-and-a-half years, eventually slowing to half that rate once the animal reaches 10 years of age.[5] The slowing of horn growth coincides with aging in males.[33] The age of an ibex can be determined by annual growth rings in the horns, which stop growing in winter.[34][35][2]

Mortality and health

Males live for around 16 years while females live around 20 years. The species has a particularly high adult survival rate compared to other herbivores around its size.[36][37] In one study, all kids reached two years of age and the majority of adults lived to 13 years, though most 13-year-old males did not reach the age of 15.[36] Alpine ibexes have a low rate of predation.[5] Their mountain habitat keeps them safe from predators like wolves, though golden eagles may prey on young.[2] In Gran Paradiso, sources of mortality are old age, lack of food, and disease. They are also killed by avalanches.[5]

The species may suffer necrosis and fibrosis caused by the bacteria Brucella melitensis,[38] and foot rot caused by Dichelobacter nodosus.[39] Infections from Mycoplasma conjunctivae damage the eye via keratoconjunctivitis and can lead to mortality rates of up to 30%.[40] Ibexes can host gastrointestinal parasites, such as coccidia, strongyles,[41] Teladorsagia circumcincta and Marshallagi amarshalli[42] as well as lungworms, mainly Muellerius capillaris.[43] Several individuals were recorded to have died from heart diseases including arteriosclerosis, cardiac fibrosis, sarcosporidiosis, and valvular heart disease.[44]

Conservation

Biologist transporting an ibex for reintroduction

During the Middle Ages, the Alpine ibex ranged throughout the Alpine region of Europe.[9] Starting in the early 16th century, the overall population declined due almost entirely to hunting by humans, particularly with the introduction of firearms.[45] By the 19th century, the species survived only in and around the Gran Paradiso of northwest Italy and on the Italian-French border, with a population of about 100 individuals.[5][45][9] Hunting of the ibex was banned in 1821 by the government of Piedmont, and Gran Paradiso was declared a royal hunting reserve in 1854 by Victor Emmanuel II.[5][2] In 1920, his grandson Victor Emmanuel III of Italy donated the land to the state of Italy and it was established as a national park.[46] The ibex population reached 4,000 by 1933 but subsequent mismanagement by the Fascist government caused it to drop to around 400 by 1945.[47][48] Their protection improved after the war, and there were 4,000 in the park in 2005.[46] In the late 20th century, the Gran Paradiso population was used for reintroductions into other parts of Italy.[45]

Starting in 1902, several ibexes from the Gran Paradiso were taken into captive facilities in Switzerland for selective breeding and reintroduction into the wild. Until 1948, translocated founders were captive-bred. Afterwards, there were reintroductions of wild-born specimens from established populations in Piz Albris, Le Pleureur and Augstmatthorn. These gave rise to the populations in France and Austria. Ibexes also recolonised areas on their own.[5] The total Alpine ibex population reached 3,020 in 1914, 20,000 in 1991 and 55,297 in 2015, and by 1975, the species occupied much of its medieval range.[5][9][45] In the 1890s, ibexes were introduced to Slovenia, despite the lack of evidence of their presence there following the last glacial period.[11] In 1980, ibexes were translocated to Bulgaria.[9]

Between 2015 and 2017, there were around 9,000 ibexes in 30 colonies in France, over 17,800 individuals and 30 colonies in Switzerland, over 16,400 ibexes in 67 colonies in Italy, around 9,000 in 27 colonies in Austria, around 500 in five colonies in Germany and almost 280 ibexes and four colonies in Slovenia.[9] As of 2020, the Alpine ibex is considered to be of Least Concern by the IUCN with a stable population trend. It was given a recovery score of 79%, making it "moderately depleted". While the species would likely have gone extinct without conservation efforts in the 19th and 20th centuries, as of 2021 it has a low conservation dependence. The IUCN assesses that without current protections, the population decline of the species would be minimal. Some countries allow limited hunting.[1]

Having gone through a genetic bottleneck, the ibex population has low genetic diversity and is at risk of inbreeding depression.[1][49] A 2020 analysis found that highly deleterious mutations were lost in these new populations but they also gained mildly deleterious ones.[50] The genetic purity of the species may also be threatened by hybridisation with domestic goats, which have been allowed to roam in their habitat.[51] The genetic bottleneck of populations may increase vulnerability to infectious diseases as their immune system has low major histocompatibility complex diversity.[52] In the Bornes Massif region of the French Alps, management actions, including a test-and-cull program (a method to control outbreaks), effectively reduced Brucella infection prevalence in adult females from 51% in 2013 to 21% in 2018, with active infections also declining significantly.[53]

Cultural significance

The Alpine ibex is called the steinbock, from the Old High German steinboc, literally "stone buck".[54][55] Several European names for the animal developed from this, including the French bouquetin, and Italian stambecco.[55] The Alpine ibex is one of many animals depicted in the art of the Late Pleistocene-era Magdalenian culture in Western Europe.[56] Ibexes were used by local people for traditional medicine.[45] The horn material was used to counter cramps, poisoning and hysteria, while the blood was thought to prevent stones from developing in the bladder.[57] Its value as a source of medicine is what led to its near extinction.[45] Since its recovery, the Alpine ibex has been seen as a resilient symbol of the mountain range. The species is featured on the coat of arms of the Swiss canton of Grisons.[58]

References

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  32. ↑ Grignoli, S.; Rossi, I.; Bertolotto, E.; Bassano, B.; Apollonio, M. (2007). "Influence of the kid on space use and habitat selection of female Alpine ibex". The Journal of Wildlife Management. 71 (3): 713–719. Bibcode:2007JWMan..71..713G. doi:10.2193/2005-675. JSTOR 4495243. S2CID 84661674.
  33. ↑ von Hardenberg, A.; Bassano, B.; Zumel Arranz, M. del P.; Bogliani, G. (2004). "Horn growth but not asymmetry heralds the onset of senescence in male Alpine ibex (Capra ibex)". Journal of Zoology. 263 (4): 425–432. doi:10.1017/S0952836904005485.
  34. ↑ Michallet, J; Loison, A; Gaillard, J; Gauthier, D (1994). "Valeur de critères biométriques externes pour la détermination de l'âge du bouquetin des Alpes (Capra ibex ibex): rôle du sexe et de l'habitat". Gibier Faune Sauvage (in French). 11: 99–118. S2CID 132957886.
  35. ↑ Ratti, P; K. H (1977). "Untersuchungen zur Altersschätzung und Altersbestimmung beim Alpensteinbock (Capra ibex ibex) im Kanton Graubünden". Zeitschrift für Jagdwissenschaft (in German). 23 (4): 188–213. doi:10.1007/BF01905758. S2CID 20248387.
  36. 1 2 ToÏgo, C.; Gaillard, J.-M.; Festa-Bianchett, M.; Largo, E.; Michallet, J.; Maillard, D. (2007). "Sex- and age-specific survival of the highly dimorphic Alpine ibex: evidence for a conservative life-history tactic". Journal of Animal Ecology. 76 (4): 679–686. doi:10.1111/j.1365-2656.2007.01254.x. PMID 17584373.
  37. ↑ Girard, I; Toigo, C; Gaillard, J-M; Gauthier, D; Martinot, J-P (1999). "Patron de survie chez le bouquetin des Alpes (Capra ibex ibex) dans le parc national de la Vanoise". Revue d'Écologie (in French). 54 (3): 235–251. doi:10.3406/revec.1999.2294. S2CID 193212971.
  38. ↑ Ferrogilo, E.; Tolari, F.; Bassano, B. (1998). "Isolation of Brucella melitensis from alpine ibex". Journal of Wildlife Diseases. 34 (2): 400–402. doi:10.7589/0090-3558-34.2.400. PMID 9577795. S2CID 43739687.
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