Alpine ibex: Difference between revisions

Alpine ibex
	Male
	Female
Conservation status
	; Least Concern (IUCN 3.1)
Scientific 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; Linnaeus, 1758
	Distribution of the Alpine ibex (population in Bulgaria not included)

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The Alpine ibex (Capra ibex), also known as the steinbock, is a European species of goat that lives in the Alps. 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 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 the steep slopes and cliffs of their mountainous habitat.

Alpine ibexes primarily feed on grass and are active throughout the year. Although they are social animals, adult males and females segregate for most of the year, coming together only to mate. During the breeding season, males use their long horns to fight for access to females. Ibexes have few predators but may succumb to parasites and diseases.

By the 19th century, the Alpine ibex had been extirpated from most of its range and it went through a population bottleneck of fewer than 100 individuals during its near-extinction event, leading to very low genetic diversity across populations. The species has been successfully reintroduced to parts of its historical range. All individuals living today descend from the stock in Gran Paradiso National Park, Italy. As of 2020^[update], the IUCN lists the species as being of least concern.

Taxonomy

Carl Linnaeus first described the Alpine ibex in 1758. It is classified in the genus Capra with nine other species of goat.^[2] Capra is Latin for 'she-goat'^[3] while the species name ibex is translated from Latin 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; these fossils appear to have been transitional between goats and their ancestors.^[5] The genus Capra may have originated in Central Asia and spread to 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 dating from the last glacial period during the late Pleistocene have been found in France and Italy. The Alpine Ibex and the Iberian ibex (C. pyrenaica) probably evolved from the extinct Pleistocene species Capra camburgensis, whose fossils have been found in Germany. The Alpine ibex appears to have been larger during the Pleistocene than in the modern day.^[5]

In the 20th century, the Nubian (C. nubiana), walia (C. walie), and Siberian ibex (C. sibirica) were considered to be subspecies of the Alpine ibex; populations in the Alps were given the trinomial of C. i. ibex.^[7] Genetic evidence from 2006 has supported the status of these Ibexes as 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

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. Transverse ridges on the front surface of the horns mark 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 its body but lighter in colour on the belly with dark markings on the chin and throat. The hair on the chest region is nearly black and there are stripes along the dorsal (back) surface. The Alpine ibex 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 in the autumn. As in other members of Capra, the Alpine ibex has glands near the eyes, groin, and feet but none on the face.^[5]

Distribution and habitat

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

The Alpine Ibex is an excellent climber; it occupies steep, rough terrain at elevations of 1,800 to 3,300 m (5,900 to 10,800 ft). It prefers to live in open areas,^[5] but when there is little snow and depending on population density, adult males may gather in larch and mixed larch-spruce woodland.^[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] and climb to alpine meadows during the summer.^[12] In early winter, both males and females move to steep, rocky slopes to avoid dense buildups of snow.^[14] Alpine ibexes prefer slopes of 30–45°, and take refuge in small caves and overhangs.^[15]

Behaviour and ecology

Alpine ibexes on Augstmatthorn in the Bernese Highlands, Switzerland

The Alpine ibex is strictly herbivorous; its diet consists mostly of grass, which is preferred all year; during the summer, ibexes supplement their diet with herbs, while during autumn and winter they also eat dwarf shrubs and conifer shoots.^[16] The most-commonly eaten grass genera are Agrostis, Avena, Calamagrostis, Festuca, Phleum, Poa, Sesleria, and Trisetum.^[5] In the spring, animals of both sexes spend about the same amount of time feeding during the day, while in summer, females, particularly those that are lactating, eat more than males.^[17] High temperatures cause heat stress in large adult males, reducing their feeding time, but they may avoid this problem by feeding at night.^[18]

In Gran Paradiso, home ranges of the Alpine Ibex can exceed 700 ha (1,700 acres) and in reintroduced populations, home ranges 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. Females' 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 mornings. They also reduce their heart rate and metabolism.^[20]

The Alpine ibex may compete for resources with chamois and red deer;^[2] the presence of these species may force the ibex to occupy higher elevations.^[21] The Alpine ibex's climbing ability is such that it has been observed scaling the 57-degree slopes of the Cingino Dam in Piedmont, Italy, where it licks salts. Only females and kids, which are lighter and have shorter legs than adult males, will climb the steep dam. Kids have been observed at 49 m (161 ft), ascending in a zig-zag path while descending in straight paths.^[22]

Social behaviour

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

Adult males and females gather together in December and January, the breeding season, then separate again in April and May.^[5] Among males, a dominance hierarchy based on size, age, and horn length exists.^[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; they bash rivals' sides or clash head-to-head often by rearing then clashing downwards.^[28]

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

Reproduction and growth

The mating season begins in December and typically lasts for around six weeks. During this time, male herds break up into smaller groups and search for females. The rut takes place in two phases; in the first phase, males interact with females as a group and in the second phase, one male separates from his group to follow a female in oestrus.^[5] Dominant males between nine and twelve years old follow a female and guard her from rivals while subordinate, younger males between two and six years old try to sneak past the tending male when he is distracted. If the 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, the male rejoins his group and restarts the first phase of the rut.^[5] Environmental conditions can affect courtship in the species; for example, snow can limit the males' ability to follow females and mate with them.^[30]

The female is in oestrus for around 20 days and gestation averages around five months, and typically results in the birth of one or sometimes two kids.^[31] Females give birth away from their social groups on rocky slopes that are relatively safe from predators.^[32] After a few days, the kids can 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 in 18 months but continue to grow until females are five to six and males are nine to eleven years old.^[5]

The horns grow throughout life. Young are born without horns, which become visible as tiny tips at one month and reach 20–25 mm (0.8–1.0 in) in the 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, slowing to half that rate once the animal reaches 10 years of age.^[5] The slowing of horn growth in males coincides with aging.^[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

Male Alpine Ibexes live for around 16 years while females live for around 20 years. The species has a high adult survival rate compared with other herbivores around its size.^[36]^[37] In one study, all kids reached two years of age and the majority of adults lived for 13 years, although 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, causes of death are old age, lack of food, and disease. They are also killed by avalanches.^[5]

Alpine ibexes 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 death 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 have died from heart diseases, including arteriosclerosis, cardiac fibrosis, sarcosporidiosis, and valvular heart disease.^[44]

Conservation

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, especially with the introduction of firearms.^[45] By the 19th century, only around 100 individuals remained in and around Gran Paradiso in north-west Italy and on the Italian-French border.^[5]^[45]^[9] In 1821, the Government of Piedmont banned hunting of the Alpine ibex and in 1854, Victor Emmanuel II declared Gran Paradiso a royal hunting reserve.^[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] By 1933, the Alpine ibex population reached 4,000 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 by 2005, there were 4,000 in the national park.^[46] In the late 20th century, the Gran Paradiso population was used for reintroductions into other parts of Italy.^[45]

Starting in 1902, several Alpine ibexes from Gran Paradiso were taken into captive facilities in Switzerland for selective breeding and reintroduction into the wild. Until 1948, translocated founder animals 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. Alpine ibexes also recolonised areas on their own.^[5] The 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 IUCN considers the Alpine ibex to be of Least Concern 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. According to the IUCN, 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 Alpine ibex population has low genetic diversity and is at risk of inbreeding depression.^[1]^[49] A 2020 analysis found highly deleterious mutations were lost in these new populations but they had also gained mildly deleterious ones.^[50] The genetic purity of the species may be threatened by hybridisation with domestic goats, which have been allowed to roam in the Alpine Ibex's habitat.^[51] The genetic bottleneck of populations may increase vulnerability to infectious diseases because 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 to control outbreaks, effectively reduced Brucella infection prevalence in adult females from 51% in 2013 to 21% in 2018, and active infections also significantly declined.^[53]

Cultural significance

The Alpine ibex is called the steinbock, which originated from the Old High German word steinboc, literally "stone buck".^[54]^[55] Several European names for the animal developed from this, including the French bouquetin and the 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] Local people used Ibexes 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] The species' value as a source of medicine 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 depicted on the coat of arms of the Swiss canton of Grisons.^[58]

References

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^ Girard, I.; Toïgo, 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.
^ 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.
^ Moore-Jones, G.; Dürr, S.; Willisch, C.; Ryser-Degiorgis, M.-P. (2021). "Occurrence of footrot in free-ranging alpine ibex (Capra ibex) colonies in Switzerland". Journal of Wildlife Diseases. 57 (2): 327–337. doi:10.7589/JWD-D-20-00050. PMID 33822150. S2CID 232357703.
^ Giacometti, M.; Janovsky, M.; Belloy, L.; Frey, J. (2002). "Infectious keratoconjunctivitis of ibex, chamois and other Caprinae". Revue Scientifique et Technique (International Office of Epizootics). 21 (2): 335–345. doi:10.20506/rst.21.2.1338. PMID 11974619.
^ Carcereri, A.; Stancampiano, L.; Marchiori, E.; Sturaro, E.; Ramanzin, M.; Cassini, R. (2021). "Factors influencing gastrointestinal parasites in a colony of Alpine ibex (Capra ibex) interacting with domestic ruminants". Hystrix, the Italian Journal of Mammalogy. 32 (1): 95–101. doi:10.4404/hystrix-00393-2020.
^ Lanfranchi, P.; Manfredi, M. T.; Zaffaroni, E.; Fraquelli, C.; Ratti, P.; Giacometti, M. (1995). "Eine dreijährige Untersuchung der Labmagen-Helminthenfauna beim Alpensteinbock (Capra i. ibex) der Kolonie Albris, Graubünden, Schweiz". Zeitschrift für Jagdwissenschaft (in German). 41: 24–35. doi:10.1007/BF02241870. S2CID 36051526.
^ Cassini, R.; Párraga, M. A.; Signorini, M.; Frangipane di Regalbono, A.; Sturaro, E.; Rossi, L.; Ramanzin, M. (2015). "Lungworms in Alpine ibex (Capra ibex) in the eastern Alps, Italy: An ecological approach". Veterinary Parasitology. 214 (1–2): 132–138. doi:10.1016/j.vetpar.2015.09.026. PMID 26476915.
^ Guarda, F.; Bollo, E.; Bassono, B.; Peracino, V.; Giraldo, A. (2001). "Patologia Cardiaca Del Camoscio (Rupicapra Rupicapra) E Dello Stambecco (Capra Ibex) Nel Parco Nazionale Gran Paradiso" (PDF). Large Animals Review (in Italian). 7 (2): 47–52. Archived (PDF) from the original on 21 December 2023. Retrieved 21 December 2023.
^ ^a ^b ^c ^d ^e ^f Stüwe, M.; Nievergelt, B. (1991). "Recovery of Alpine ibex from near extinction: the result of effective protection, captive breeding, and reintroductions". Applied Animal Behaviour Science. 29 (1–4): 379–387. doi:10.1016/0168-1591(91)90262-V.
^ ^a ^b Riley, L.; Riley, W. (2005). Nature's Strongholds: The World's Great Wildlife Reserves. Princeton University Press. pp. 390. ISBN 0-691-12219-9.
^ Graf von Hardenberg, W. (2021). A Monastery for the Ibex: Conservation, State, and Conflict on the Gran Paradiso 1919–1949. University of Pittsburgh Press. p. 32. ISBN 978-0822946359.
^ Graf von Hardenberg, W. (2010). "Act Locally, Think Naturally: A Brief History of Access Rights and Environmental Conflicts in Fascist Italy". In Armiero, M.; Hall, M. (eds.). Nature and History in Modern Italy. Ohio University Press. p. 149. ISBN 9780821443477.
^ Biebach, I.; Keller, L. F. (2009). "A strong genetic footprint of the re-introduction history of Alpine ibex (Capra ibex ibex)". Molecular Ecology. 18 (24): 5046–5058. Bibcode:2009MolEc..18.5046B. doi:10.1111/j.1365-294X.2009.04420.x. PMID 19912536. S2CID 36215646.
^ Grossen, C.; Guillaume, F.; Keller, L. F.; Croll, D. (2020). "Purging of highly deleterious mutations through severe bottlenecks in Alpine ibex". Nature Communications. 11 (1): 1001. Bibcode:2020NatCo..11.1001G. doi:10.1038/s41467-020-14803-1. PMC 7035315. PMID 32081890.
^ Giacometti, M.; Roganti, R.; De Tann, D.; Stahlberger-Saitbekova, N.; Obexer-Ruff, G. (2004). "Alpine ibex Capra ibex ibex x domestic goat C. aegagrus domestica hybrids in a restricted area of southern Switzerland". Wildlife Biology. 10 (2): 137–143. doi:10.2981/wlb.2004.018. S2CID 55900555.
^ Brambilla, A.; Keller, L.; Bassuno, B.; Grossen, C. (2017). "Heterozygosity–fitness correlation at the major histocompatibility complex despite low variation in Alpine ibex (Capra ibex)". Evolutionary Applications. 11 (5): 631–644. doi:10.1111/eva.12575. PMC 5979623. PMID 29875807.
^ Calenge, C.; Lambert, S.; Petit, E.; Thébault, A.; Gilot-Fromont, E.; Toïgo, C.; Rossi, S. (2021). "Estimating disease prevalence and temporal dynamics using biased capture serological data in a wildlife reservoir: The example of brucellosis in Alpine ibex (Capra ibex)". Preventive Veterinary Medicine. 187: 105239. doi:10.1016/j.prevetmed.2020.105239. PMID 33373957. S2CID 229721738.
^ "steinbock". Merriam-Webster. Archived from the original on 8 January 2024. Retrieved 7 January 2024.
^ ^a ^b García-González, R.; Herrero, J.; Nores, C. (2021). "The names of southwestern European goats: is Iberian ibex the best common name for Capra pyrenaica?". Animal Biodiversity and Conservation. 44 (1): 1–16. doi:10.32800/abc.2021.44.0001. hdl:10261/264864. S2CID 229546091.
^ Castelli, A. (2010). "Ibex Images from the Magdalenian Culture" (PDF). Archived (PDF) from the original on 20 December 2023. Retrieved 19 December 2023.
^ Thorington, J. M. (1944). "The ibex and chamois in ancient medicine". Bulletin of the History of Medicine. 15 (1): 65–78. JSTOR 44442799.
^ Schirpke, U.; Meisch, C.; Tappeiner, U. (2018). "Symbolic species as a cultural ecosystem service in the European Alps: insights and open issues". Landscape Ecology. 33 (5): 711–730. Bibcode:2018LaEco..33..711S. doi:10.1007/s10980-018-0628-x.

External links

Alpine Ibex European Specialist Group (GSE-AIESG)

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[15] Wiersema, G. (1984). "Seasonal use and quality assessment of ibex habitat". Acta Zoologica Fennica. 172: 89–90.

[16] Klansek, E.; Vavra, I.; Onderscheka, K. (1995). "Die Äsungszusammensetzung des Alpensteinwildes (Capra i. ibex L.) in Abhängigkeit von Jahreszeit, Alter und Äsungsangebot in Graubünden". Zeitschrift für Jagdwissenschaft (in German). 41 (3): 171–181. doi:10.1007/BF02239946. S2CID 41249118.

[Neuhaus-17] Neuhaus, P.; Ruckstuhl, K. E. (2002). "Foraging behaviour in Alpine ibex (Capra ibex): consequences of reproductive status, body size, age and sex". Ethology Ecology & Evolution. 14 (4): 373–381. Bibcode:2002EtEcE..14..373N. doi:10.1080/08927014.2002.9522738. S2CID 84995631.

[18] Aublet, J.-F.; Festa-Bianchet, M.; Bergero, D.; Bassano, B. (2009). "Temperature constraints on foraging behavior of male Alpine ibex (Capra ibex) in summer". Oecologia. 159 (1): 237–247. Bibcode:2009Oecol.159..237A. doi:10.1007/s00442-008-1198-4. PMID 18987895. S2CID 1755798.

[Parrini2003-19] Parrini, F.; Grignolio, S.; Luccarini, S.; Bassano, B.; Apollonio, M. (2003). "Spatial behaviour of adult male Alpine ibex Capra ibex ibex in the Gran Paradiso National Park, Italy". Acta Theriologica. 48 (3): 411–423. doi:10.1007/BF03194179. S2CID 6211702.

[20] Signer, C.; Ruf, T.; Arnold, W. (2011). "Hypometabolism and basking: the strategies of Alpine ibex to endure harsh over-wintering conditions". Functional Ecology. 25 (3): 537–547. Bibcode:2011FuEco..25..537S. doi:10.1111/j.1365-2435.2010.01806.x.

[21] Herfindal, I.; Anderwald, P.; Filli, F.; Andri, S. C.; Rempfler, T. (2019). "Climate, competition and weather conditions drive vertical displacement and habitat use of an alpine ungulate in a highly topographic landscape". Landscape Ecology. 34 (11): 2523–2539. Bibcode:2019LaEco..34.2523H. doi:10.1007/s10980-019-00902-y. S2CID 202570193.

[Biancardi2017-22] Biancardi, C. M.; Minetti, A. E. (2017). "Gradient limits and safety factor of Alpine ibex (Capra ibex) locomotion". Hystrix, the Italian Journal of Mammalogy. 28 (1): 56–60. doi:10.4404/hystrix-28.1-11504.

[23] Bon, R.; Rideau, C. S.; Villaret, J.-C.; Joachim, J. (2001). "Segregation is not only a matter of sex in Alpine ibex, Capra ibex ibex". Animal Behaviour. 62 (3): 495–504. doi:10.1006/anbe.2001.1776. S2CID 53205174.

[Villaret-24] Villaret, J.-C.; Bon, R. (1998). "Sociality and relationships in Alpine ibex (Capra ibex)". Revue d'Écologie. 53 (2): 153–170. doi:10.3406/revec.1998.2251. S2CID 58921073. Archived from the original on 7 November 2023. Retrieved 7 November 2023.

[Villaret1995-25] Villaret, J. C.; Bon, R. (1995). "Social and spatial segregation in Alpine ibex (Capra ibex) in Bargy, French Alps". Ethology. 101 (4): 291–300. Bibcode:1995Ethol.101..291V. doi:10.1111/j.1439-0310.1995.tb00366.x.

[26] Brambilla, A.; von Hardenberg, A.; Canedoli, C.; Brivio, F.; Sueur, C.; Stanley, C. R. (2022). "Long term analysis of social structure: evidence of age-based consistent associations in male Alpine ibex". Oikos. 2022 (8): e09511. Bibcode:2022Oikos2022E9511B. doi:10.1111/oik.09511. hdl:10281/402427.

[27] Bergeron, P.; Grignolio, S.; Apollonio, M.; Shipley, B.; Festa-Bianchet, M. (2010). "Secondary sexual characters signal fighting ability and determine social rank in Alpine ibex (Capra ibex)". Behavioral Ecology and Sociobiology. 64 (8): 1299–1307. Bibcode:2010BEcoS..64.1299B. doi:10.1007/s00265-010-0944-x. S2CID 36262460.

[28] Willisch, C. S.; Neuhaus, P. (2010). "Social dominance and conflict reduction in rutting male Alpine ibex, Capra ibex". Behavioral Ecology. 21 (2): 372–380. doi:10.1093/beheco/arp200.

[29] Willisch, C. S.; Neuhaus, P. (2009). "Alternative mating tactics and their impact on survival in adult male Alpine ibex (Capra ibex ibex)". Journal of Mammalogy. 90 (6): 1421–1430. doi:10.1644/08-MAMM-A-316R1.1. S2CID 86267022.

[30] Rossi, I.; Grignolio, S.; Bassano, B.; Apollonius, M. (2003). "Strategie riproduttive dello Stambecco (Capra ibex ibex) nel Parco Nazionale del Gran Paradiso". Hystrix, the Italian Journal of Mammalogy (in Italian). 14 (ATIt Congress Supplement) (Special Issue: IV ATIt Congress). doi:10.4404/hystrix-14.0-4292 (inactive 2 November 2024).{{cite journal}}: CS1 maint: DOI inactive as of November 2024 (link)

[Stuwe1987-31] Stüwe, M.; Grodinsky, C. (1987). "Reproductive biology of captive Alpine ibex (Capra i. ibex)". Zoo Biology. 6 (4): 331–339. doi:10.1002/zoo.1430060407.

[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.

[Toigo2007-36] 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. Bibcode:2007JAnEc..76..679T. doi:10.1111/j.1365-2656.2007.01254.x. PMID 17584373.

[37] Girard, I.; Toïgo, 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.

[39] Moore-Jones, G.; Dürr, S.; Willisch, C.; Ryser-Degiorgis, M.-P. (2021). "Occurrence of footrot in free-ranging alpine ibex (Capra ibex) colonies in Switzerland". Journal of Wildlife Diseases. 57 (2): 327–337. doi:10.7589/JWD-D-20-00050. PMID 33822150. S2CID 232357703.

[40] Giacometti, M.; Janovsky, M.; Belloy, L.; Frey, J. (2002). "Infectious keratoconjunctivitis of ibex, chamois and other Caprinae". Revue Scientifique et Technique (International Office of Epizootics). 21 (2): 335–345. doi:10.20506/rst.21.2.1338. PMID 11974619.

[41] Carcereri, A.; Stancampiano, L.; Marchiori, E.; Sturaro, E.; Ramanzin, M.; Cassini, R. (2021). "Factors influencing gastrointestinal parasites in a colony of Alpine ibex (Capra ibex) interacting with domestic ruminants". Hystrix, the Italian Journal of Mammalogy. 32 (1): 95–101. doi:10.4404/hystrix-00393-2020.

[42] Lanfranchi, P.; Manfredi, M. T.; Zaffaroni, E.; Fraquelli, C.; Ratti, P.; Giacometti, M. (1995). "Eine dreijährige Untersuchung der Labmagen-Helminthenfauna beim Alpensteinbock (Capra i. ibex) der Kolonie Albris, Graubünden, Schweiz". Zeitschrift für Jagdwissenschaft (in German). 41: 24–35. doi:10.1007/BF02241870. S2CID 36051526.

[43] Cassini, R.; Párraga, M. A.; Signorini, M.; Frangipane di Regalbono, A.; Sturaro, E.; Rossi, L.; Ramanzin, M. (2015). "Lungworms in Alpine ibex (Capra ibex) in the eastern Alps, Italy: An ecological approach". Veterinary Parasitology. 214 (1–2): 132–138. doi:10.1016/j.vetpar.2015.09.026. PMID 26476915.

[44] Guarda, F.; Bollo, E.; Bassono, B.; Peracino, V.; Giraldo, A. (2001). "Patologia Cardiaca Del Camoscio (Rupicapra Rupicapra) E Dello Stambecco (Capra Ibex) Nel Parco Nazionale Gran Paradiso" (PDF). Large Animals Review (in Italian). 7 (2): 47–52. Archived (PDF) from the original on 21 December 2023. Retrieved 21 December 2023.

[recovery-45] ^ ^a ^b ^c ^d ^e ^f Stüwe, M.; Nievergelt, B. (1991). "Recovery of Alpine ibex from near extinction: the result of effective protection, captive breeding, and reintroductions". Applied Animal Behaviour Science. 29 (1–4): 379–387. doi:10.1016/0168-1591(91)90262-V.

[Riley-46] Riley, L.; Riley, W. (2005). Nature's Strongholds: The World's Great Wildlife Reserves. Princeton University Press. pp. 390. ISBN 0-691-12219-9.

[Wilko_Graf_von_Hardenberg_2021-47] Graf von Hardenberg, W. (2021). A Monastery for the Ibex: Conservation, State, and Conflict on the Gran Paradiso 1919–1949. University of Pittsburgh Press. p. 32. ISBN 978-0822946359.

[48] Graf von Hardenberg, W. (2010). "Act Locally, Think Naturally: A Brief History of Access Rights and Environmental Conflicts in Fascist Italy". In Armiero, M.; Hall, M. (eds.). Nature and History in Modern Italy. Ohio University Press. p. 149. ISBN 9780821443477.

[Biebach,_I_2009-49] Biebach, I.; Keller, L. F. (2009). "A strong genetic footprint of the re-introduction history of Alpine ibex (Capra ibex ibex)". Molecular Ecology. 18 (24): 5046–5058. Bibcode:2009MolEc..18.5046B. doi:10.1111/j.1365-294X.2009.04420.x. PMID 19912536. S2CID 36215646.

[Grossen2020-50] Grossen, C.; Guillaume, F.; Keller, L. F.; Croll, D. (2020). "Purging of highly deleterious mutations through severe bottlenecks in Alpine ibex". Nature Communications. 11 (1): 1001. Bibcode:2020NatCo..11.1001G. doi:10.1038/s41467-020-14803-1. PMC 7035315. PMID 32081890.

[51] Giacometti, M.; Roganti, R.; De Tann, D.; Stahlberger-Saitbekova, N.; Obexer-Ruff, G. (2004). "Alpine ibex Capra ibex ibex x domestic goat C. aegagrus domestica hybrids in a restricted area of southern Switzerland". Wildlife Biology. 10 (2): 137–143. doi:10.2981/wlb.2004.018. S2CID 55900555.

[52] Brambilla, A.; Keller, L.; Bassuno, B.; Grossen, C. (2017). "Heterozygosity–fitness correlation at the major histocompatibility complex despite low variation in Alpine ibex (Capra ibex)". Evolutionary Applications. 11 (5): 631–644. doi:10.1111/eva.12575. PMC 5979623. PMID 29875807.

[53] Calenge, C.; Lambert, S.; Petit, E.; Thébault, A.; Gilot-Fromont, E.; Toïgo, C.; Rossi, S. (2021). "Estimating disease prevalence and temporal dynamics using biased capture serological data in a wildlife reservoir: The example of brucellosis in Alpine ibex (Capra ibex)". Preventive Veterinary Medicine. 187: 105239. doi:10.1016/j.prevetmed.2020.105239. PMID 33373957. S2CID 229721738.

[54] "steinbock". Merriam-Webster. Archived from the original on 8 January 2024. Retrieved 7 January 2024.

[García-González-55] García-González, R.; Herrero, J.; Nores, C. (2021). "The names of southwestern European goats: is Iberian ibex the best common name for Capra pyrenaica?". Animal Biodiversity and Conservation. 44 (1): 1–16. doi:10.32800/abc.2021.44.0001. hdl:10261/264864. S2CID 229546091.

[56] Castelli, A. (2010). "Ibex Images from the Magdalenian Culture" (PDF). Archived (PDF) from the original on 20 December 2023. Retrieved 19 December 2023.

[57] Thorington, J. M. (1944). "The ibex and chamois in ancient medicine". Bulletin of the History of Medicine. 15 (1): 65–78. JSTOR 44442799.

[58] Schirpke, U.; Meisch, C.; Tappeiner, U. (2018). "Symbolic species as a cultural ecosystem service in the European Alps: insights and open issues". Landscape Ecology. 33 (5): 711–730. Bibcode:2018LaEco..33..711S. doi:10.1007/s10980-018-0628-x.

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-The '''Alpine ibex''' ('''''Capra ibex'''''), also known as the '''steinbock''', is a European species of [[Capra (genus)|goat]] that lives in the [[Alps]]. It is one of ten species in the genus ''Capra'' and its closest living relative is the [[Iberian ibex]]. The Alpine ibex is a [[Sexual dimorphism|sexually dimorphic]] species; males are larger and carry longer horns than females. Its [[animal coat|coat]] 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 {{convert|3300|m|abbr=on}} and their sharp hooves allow them to scale their mountainous habitat.
+The '''Alpine ibex''' ('''''Capra ibex'''''), also known as the '''steinbock''', is a European species of [[Capra (genus)|goat]] that lives in the [[Alps]]. It is one of ten species in the genus ''Capra'' and its closest living relative is the [[Iberian ibex]]. The Alpine ibex is a [[Sexual dimorphism|sexually dimorphic]] species; males are larger and carry longer horns than females. Its [[animal coat|coat]] 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 {{convert|3300|m|abbr=on}} and their sharp hooves allow them to scale the steep slopes and cliffs of their mountainous habitat.
 Alpine ibexes primarily feed on grass and are active throughout the year. Although they are social animals, adult males and females segregate for most of the year, coming together only to mate. During the breeding season, males use their long horns to fight for access to females. Ibexes have few predators but may succumb to parasites and diseases.
-By the 19th century, the Alpine Ibex had been [[Local extinction|extirpated]] from most of its range and it went through a [[population bottleneck]] of fewer than 100 individuals during its near-extinction event, leading to very low genetic diversity across populations. The species has been successfully reintroduced to parts of its historical range. All individuals living today descend from the stock in [[Gran Paradiso National Park]], Italy. {{As of| 2020}}, the [[IUCN]] lists the species as being of least concern.
+By the 19th century, the Alpine ibex had been [[Local extinction|extirpated]] from most of its range and it went through a [[population bottleneck]] of fewer than 100 individuals during its near-extinction event, leading to very low genetic diversity across populations. The species has been successfully reintroduced to parts of its historical range. All individuals living today descend from the stock in [[Gran Paradiso National Park]], Italy. {{As of| 2020}}, the [[IUCN]] lists the species as being of least concern.
 ==Taxonomy==
-[[Carl Linnaeus]] first described the Alpine ibex in 1758. It is classified in the [[genus]] ''[[Capra (genus)|Capra]]'' with nine other species of goat.<ref name=Brambilla2022/> ''Capra'' is Latin for "she-goat"<ref>{{cite web |title=Capri |website=[[Online Etymology Dictionary]] |access-date=11 December 2023 |url=https://www.etymonline.com/word/capri |archive-date=5 May 2022 |archive-url=https://web.archive.org/web/20220505025521/https://www.etymonline.com/word/Capri |url-status=live }}</ref> while the species name "ibex" is translated from Latin as "[[chamois]]" and is possibly derived from an earlier Alpine language.<ref>{{cite web |title=ibex |website=[[Online Etymology Dictionary]] |access-date=11 December 2023 |url=https://www.etymonline.com/word/ibex#etymonline_v_1464 |archive-date=11 December 2023 |archive-url=https://web.archive.org/web/20231211153217/https://www.etymonline.com/word/ibex#etymonline_v_1464 |url-status=live }}</ref>
+[[Carl Linnaeus]] first described the Alpine ibex in 1758. It is classified in the [[genus]] ''[[Capra (genus)|Capra]]'' with nine other species of goat.<ref name=Brambilla2022/> {{lang|la|Capra}} is Latin for {{gloss|she-goat}}<ref>{{cite web |title=Capri |website=[[Online Etymology Dictionary]] |access-date=11 December 2023 |url=https://www.etymonline.com/word/capri |archive-date=5 May 2022 |archive-url=https://web.archive.org/web/20220505025521/https://www.etymonline.com/word/Capri |url-status=live }}</ref> while the species name {{lang|la|ibex}} is translated from Latin as {{gloss|[[chamois]]}} and is possibly derived from an earlier Alpine language.<ref>{{cite web |title=ibex |website=[[Online Etymology Dictionary]] |access-date=11 December 2023 |url=https://www.etymonline.com/word/ibex#etymonline_v_1464 |archive-date=11 December 2023 |archive-url=https://web.archive.org/web/20231211153217/https://www.etymonline.com/word/ibex#etymonline_v_1464 |url-status=live }}</ref>
 Fossils of the genus ''[[Tossunnoria]]'' are found in late [[Miocene]] deposits in China; these fossils appear to have been [[transitional fossil |transitional]] between goats and their ancestors.<ref name=Parrini2009/> The genus ''Capra'' may have originated in Central Asia and spread to Europe, the Caucasus, and East Africa from the [[Pliocene]] and into the [[Pleistocene]]. [[Mitochondrial DNA |Mitochondrial]] and [[Y chromosome]] evidence show [[Hybrid (biology) |hybridisation]] of species in this lineage.<ref name=Pidancier2006>{{cite journal |last1=Pidancier |first1=N. |last2=Jordan |first2=S. |last3=Luikart |first3=G. |last4=Taberlet |first4=P. |year=2006 |title=Evolutionary history of the genus ''Capra'' (Mammalia, Artiodactyla): Discordance between mitochondrial DNA and Y-chromosome phylogenies |journal=Molecular Phylogenetics and Evolution |volume=40 |issue=3 |pages=739–749 |doi=10.1016/j.ympev.2006.04.002 |pmid=16757184 |bibcode=2006MolPE..40..739P |s2cid=12743878}}</ref> Fossils of the Alpine ibex dating from the [[last glacial period]] during the late Pleistocene have been found in France and Italy. The Alpine Ibex and the [[Iberian ibex]] (''C. pyrenaica'') probably evolved from the extinct Pleistocene species ''[[Capra camburgensis]]'', whose fossils have been found in Germany. The Alpine ibex appears to have been larger during the Pleistocene than in the modern day.<ref name=Parrini2009/>
@@ Line 64: / Line 63: @@
 The Alpine ibex is a stocky animal with a tough neck and robust legs with short [[metapodial]]s. 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.<ref name=Brambilla2022/><ref name=Parrini2009/> Both male and female Alpine ibexes have large, backwards-curving horns with an elliptical cross-section and a trilateral-shaped core. Transverse ridges on the front surface of the horns mark an otherwise flat surface. At {{convert|69|–|98|cm|abbr=on}}, the horns of males are substantially longer than those of females, which reach only {{convert|18|–|35|cm|abbr=on}} in length.<ref name=Parrini2009>{{cite journal |last1=Parrini|first1=F. III|last2=Cain |first2=J. W.|last3=Krausman|first3=P. R. |year=2009 |title=''Capra ibex'' (Artiodactyla: Bovidae) |journal=Mammalian Species | issue = 830 | pages = 1–12 | doi = 10.1644/830.1| doi-access = free }}</ref>
-The species has brownish-grey hair over most of its body; this hair's colour is lighter on the belly, and there are dark markings on the chin and throat. The hair on the chest region is nearly black and there are stripes along the dorsal (back) surface. The Alpine ibex 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  in the autumn. As in other members of ''Capra'', the Alpine ibex has glands near the eyes, groin and feet but there are none on the face.<ref name=Parrini2009/>
+The species has brownish-grey hair over most of its body but lighter in colour on the belly with dark markings on the chin and throat. The hair on the chest region is nearly black and there are stripes along the dorsal (back) surface. The Alpine ibex 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  in the autumn. As in other members of ''Capra'', the Alpine ibex has glands near the eyes, groin, and feet but none on the face.<ref name=Parrini2009/>
 ==Distribution and habitat==
@@ Line 71: / Line 70: @@
 The Alpine ibex is native to the [[Alps]] of central Europe; its range includes France, Switzerland, Liechtenstein, Italy, Germany, and Austria.<ref name=Bramnilla2020/> Fossils of the species have been found as far south as Greece, where it became [[local extinction |locally extinct]] over 7,500 years ago due to human predation.<ref>{{Cite journal |doi=10.1007/s13364-012-0094-9 |title=Past and present distribution of the genus ''Capra'' in Greece |date=2013 |last1=Geskos |first1=A. |journal=Acta Theriologica |volume=58 |pages=1–11 |s2cid=256122729 }}</ref> Between the 16th and 18th centuries, the species disappeared from much of its range due to hunting, leaving by the 19th century one surviving population in and around [[Gran Paradiso]], Italy. The species has since been reintroduced into parts of its former range,<ref name=Bramnilla2020>{{cite journal |last1=Bramnilla |first1=A. |last2=von Hadenberg |first2=A. |last3=Nelli |first3=L. |last4=Bassono |first4=B. |year=2020 |title=Distribution, status, and recent population dynamics of Alpine ibex ''Capra ibex'' in Europe |journal=Mammal Review |volume=40 |issue=3 |pages=267–277 |doi=10.1111/mam.12194 |s2cid=218784224 |url=https://www.zora.uzh.ch/id/eprint/187150/8/MAMMAL-18-54.R5_Proof_fl.pdf |access-date=2023-12-01 |archive-date=2023-12-03 |archive-url=https://web.archive.org/web/20231203020732/https://www.zora.uzh.ch/id/eprint/187150/8/MAMMAL-18-54.R5_Proof_fl.pdf |url-status=live}}</ref> as well as new areas such as Slovenia<ref name=management>{{cite book |last1=Adamič |first1=M. |last2=Jerina |first2=K. |year=2010 |contribution=Ungulates and their management in Slovenia |title=European Ungulates and Their Management in the 21st Century |publisher=Cambridge University Press |editor1=Apollonio, M. |editor2=Andersen, R. |editor3=Putman, R. |pages=514 |isbn=9780521760614}}</ref> and Bulgaria.<ref name=Bramnilla2020/>
-The Alpine Ibex is an excellent climber; it occupies steep, rough terrain at elevations of {{convert |1800 |to |3300 |m |abbr=on}}. It prefers to live an open areas<ref name=Parrini2009/> but when there is little snow, and depending on population density, adult males may gather in [[larch]] and mixed larch-[[spruce]] woodland.<ref name=Grignolio2003/> Outside the breeding season, the sexes live in separate habitats.<ref name=Grignolio2003/><ref>{{cite journal |last1 =Toïgo |first1=C. |last2=Gaillard |first2=J. M. |last3=Michallet |first3=J. |year=1997 |title=Adult survival pattern of the sexually dimorphic Alpine ibex (''Capra ibex ibex'') |journal=Canadian Journal of Zoology |volume=75 |pages=75–79 |doi=10.1139/z97-009}}</ref> 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,<ref name=Parrini2009/> and climb to alpine meadows during the summer.<ref name=Grignolio2003>{{cite journal |last1=Grignolio |first1=S. |last2=Parrini |first2=F. |last3=Bassano |first3=B. |last4=Luccarini |first4=S. |last5=Apollonio |first5=M. |year=2003 |title=Habitat selection in adult males of Alpine ibex, ''Capra ibex ibex'' |journal=Folia Zoologica |volume=52 |issue=2 |pages=113–120 |url=http://www.ivb.cz/folia/52/2/113-120.pdf |access-date=2014-12-18 |archive-date=2013-05-22 |archive-url=https://web.archive.org/web/20130522213611/http://www.ivb.cz/folia/52/2/113-120.pdf |url-status=dead}}</ref> In early winter, both males and females move to steep, rocky slopes to avoid dense buildups of snow.<ref name=Grignolio2004>{{cite journal |last1=Grignolio | first1=S. |last2=Rossi |first2=I. |last3=Bassano |first3=B. |last4=Parrini |first4=F. |last5=Apollonio |first5=M. |year=2004 |title=Seasonal variations of spatial behaviour in female Alpine ibex (''Capra ibex ibex'') in relation to climatic conditions and age |journal=Ethology Ecology and Evolution |volume=16 |issue=3 |pages=255–264 |doi=10.1080/08927014.2004.9522636 | bibcode=2004EtEcE..16..255G |s2cid=85380031}}</ref> Alpine ibexes prefer slopes of 30–45°, and take refuge in small caves and overhangs.<ref>{{cite journal |last=Wiersema |first=G. |year=1984 |title=Seasonal use and quality assessment of ibex habitat |journal=Acta Zoologica Fennica |volume=172 |pages=89–90}}</ref>
+The Alpine Ibex is an excellent climber; it occupies steep, rough terrain at elevations of {{convert |1800 |to |3300 |m |abbr=on}}. It prefers to live in open areas,<ref name=Parrini2009/> but when there is little snow and depending on population density, adult males may gather in [[larch]] and mixed larch-[[spruce]] woodland.<ref name=Grignolio2003/> Outside the breeding season, the sexes live in separate habitats.<ref name=Grignolio2003/><ref>{{cite journal |last1 =Toïgo |first1=C. |last2=Gaillard |first2=J. M. |last3=Michallet |first3=J. |year=1997 |title=Adult survival pattern of the sexually dimorphic Alpine ibex (''Capra ibex ibex'') |journal=Canadian Journal of Zoology |volume=75 |pages=75–79 |doi=10.1139/z97-009}}</ref> 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,<ref name=Parrini2009/> and climb to alpine meadows during the summer.<ref name=Grignolio2003>{{cite journal |last1=Grignolio |first1=S. |last2=Parrini |first2=F. |last3=Bassano |first3=B. |last4=Luccarini |first4=S. |last5=Apollonio |first5=M. |year=2003 |title=Habitat selection in adult males of Alpine ibex, ''Capra ibex ibex'' |journal=Folia Zoologica |volume=52 |issue=2 |pages=113–120 |url=http://www.ivb.cz/folia/52/2/113-120.pdf |access-date=2014-12-18 |archive-date=2013-05-22 |archive-url=https://web.archive.org/web/20130522213611/http://www.ivb.cz/folia/52/2/113-120.pdf |url-status=dead}}</ref> In early winter, both males and females move to steep, rocky slopes to avoid dense buildups of snow.<ref name=Grignolio2004>{{cite journal |last1=Grignolio | first1=S. |last2=Rossi |first2=I. |last3=Bassano |first3=B. |last4=Parrini |first4=F. |last5=Apollonio |first5=M. |year=2004 |title=Seasonal variations of spatial behaviour in female Alpine ibex (''Capra ibex ibex'') in relation to climatic conditions and age |journal=Ethology Ecology and Evolution |volume=16 |issue=3 |pages=255–264 |doi=10.1080/08927014.2004.9522636 | bibcode=2004EtEcE..16..255G |s2cid=85380031}}</ref> Alpine ibexes prefer slopes of 30–45°, and take refuge in small caves and overhangs.<ref>{{cite journal |last=Wiersema |first=G. |year=1984 |title=Seasonal use and quality assessment of ibex habitat |journal=Acta Zoologica Fennica |volume=172 |pages=89–90}}</ref>
 ==Behaviour and ecology==
@@ Line 80: / Line 79: @@
 In Gran Paradiso, [[home range]]s of the Alpine Ibex can exceed {{convert|700|ha|abbr=on}} and in reintroduced populations, home ranges may approach {{convert|3000|ha|abbr=on}}.<ref name=Parrini2009/><ref name=Parrini2003/> 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. Females' home ranges are usually smaller than those of males.<ref name=Parrini2009/><ref name=Grignolio2004/><ref name=Parrini2003>{{cite journal |last1=Parrini |first1=F. |last2=Grignolio |first2=S. |last3=Luccarini |first3=S. |last4=Bassano |first4=B. |last5=Apollonio |first5=M. | year=2003 |title=Spatial behaviour of adult male Alpine ibex ''Capra ibex ibex'' in the Gran Paradiso National Park, Italy |journal=Acta Theriologica |volume=48 |issue=3 |pages=411–423 |doi=10.1007/BF03194179 | s2cid=6211702}}</ref> Ibexes do not [[hibernation |hibernate]] during the winter; they take shelter on cold winter nights and [[sunning (behaviour) |bask]] in the mornings. They also reduce their heart rate and metabolism.<ref>{{cite journal |last1=Signer |first1=C. |last2=Ruf |first2=T. |last3=Arnold |first3=W. |year=2011 |title=Hypometabolism and basking: the strategies of Alpine ibex to endure harsh over-wintering conditions |journal=Functional Ecology |volume=25 |issue=3 |pages=537–547 |doi=10.1111/j.1365-2435.2010.01806.x |bibcode=2011FuEco..25..537S}}</ref>
-The Alpine ibex may compete for resources with [[chamois]] and [[red deer]];<ref name=Brambilla2022/> the presence of these species may force the ibex to occupy high elevations.<ref>{{cite journal |last1=Herfindal |first1=I. |last2=Anderwald |first2=P. |last3=Filli |first3=F. |last4=Andri |first4=S. C. |last5=Rempfler |first5=T. |year=2019 |title=Climate, competition and weather conditions drive vertical displacement and habitat use of an alpine ungulate in a highly topographic landscape |journal=Landscape Ecology |volume=34 |issue=11 |pages=2523–2539 |doi=10.1007/s10980-019-00902-y |bibcode=2019LaEco..34.2523H |s2cid=202570193}}</ref> The Alpine ibex's climbing ability is such that it has been observed scaling the 57-degree slopes of the [[Cingino Dam]] in [[Piedmont]], Italy, where it [[salt lick |licks salt]]s. Only females and kids, which are lighter and have shorter legs than adult males, will climb the steep dam. Kids have been observed at  {{convert |49 |m |abbr=on}}, ascending in a zig-zag path while descending in straight paths.<ref name=Biancardi2017>{{cite journal |last1=Biancardi |first1=C. M. |last2=Minetti |first2=A. E. |year=2017 |title=Gradient limits and safety factor of Alpine ibex (''Capra ibex'') locomotion |journal=Hystrix, the Italian Journal of Mammalogy |volume=28 |issue=1 |pages=56–60 |doi=10.4404/hystrix-28.1-11504}}</ref>
+The Alpine ibex may compete for resources with [[chamois]] and [[red deer]];<ref name=Brambilla2022/> the presence of these species may force the ibex to occupy higher elevations.<ref>{{cite journal |last1=Herfindal |first1=I. |last2=Anderwald |first2=P. |last3=Filli |first3=F. |last4=Andri |first4=S. C. |last5=Rempfler |first5=T. |year=2019 |title=Climate, competition and weather conditions drive vertical displacement and habitat use of an alpine ungulate in a highly topographic landscape |journal=Landscape Ecology |volume=34 |issue=11 |pages=2523–2539 |doi=10.1007/s10980-019-00902-y |bibcode=2019LaEco..34.2523H |s2cid=202570193}}</ref> The Alpine ibex's climbing ability is such that it has been observed scaling the 57-degree slopes of the [[Cingino Dam]] in [[Piedmont]], Italy, where it [[salt lick |licks salt]]s. Only females and kids, which are lighter and have shorter legs than adult males, will climb the steep dam. Kids have been observed at  {{convert |49 |m |abbr=on}}, ascending in a zig-zag path while descending in straight paths.<ref name=Biancardi2017>{{cite journal |last1=Biancardi |first1=C. M. |last2=Minetti |first2=A. E. |year=2017 |title=Gradient limits and safety factor of Alpine ibex (''Capra ibex'') locomotion |journal=Hystrix, the Italian Journal of Mammalogy |volume=28 |issue=1 |pages=56–60 |doi=10.4404/hystrix-28.1-11504}}</ref>
 ===Social behaviour===
-[[File:LOTTA TRA STAMBECCHI 1.jpg|thumb|Male ibexes fighting|alt=In a rocky landscape with some snow, two goat-like creatures with large, backward-facing horns face each other. The animal on the right has raised its forelegs.]]
+[[File:Two male Alpine ibex (Capra ibex) fighting.jpg|thumb|Male ibexes fighting|alt=In a rocky landscape with some snow, two goat-like creatures with large, backward-facing horns face each other. The animal on the right has raised its forelegs.]]
 The Alpine ibex is a [[Sociality|social]] species but it tends to live in groups that are based on sex and age.<ref name=Parrini2009/> For most of the year, adult males group separately from females, and older males live separately from young males.<ref>{{cite journal|last1=Bon|first1=R.|last2=Rideau|first2=C. S.|last3=Villaret|first3=J.-C.|last4=Joachim|first4=J.|year=2001|title=Segregation is not only a matter of sex in Alpine ibex, ''Capra ibex ibex''|journal=Animal Behaviour|volume=62|issue=3|pages=495–504|doi=10.1006/anbe.2001.1776|s2cid=53205174}}</ref> Female groups consist of 5–10 members and male groups usually have 2–16 members but sometimes have more than 50.<ref name=Brambilla2022/><ref name=Villaret/> Dependent kids live with their mothers in female groups. Segregation between the sexes is a gradual process; males younger than nine years may still associate with female groups.<ref name=Villaret1995>{{cite journal|author1=Villaret, J. C.|author2=Bon, R. |year=1995|title=Social and spatial segregation in Alpine ibex (''Capra ibex'') in Bargy, French Alps|journal=Ethology|volume=101|issue=4|pages=291–300|doi=10.1111/j.1439-0310.1995.tb00366.x|bibcode=1995Ethol.101..291V }}</ref>  Adult males, particularly older males, are more likely to be found alone than females.<ref name=Villaret>{{cite journal|last1=Villaret|first1=J.-C.|last2=Bon|first2=R.|year=1998|title=Sociality and relationships in Alpine ibex (''Capra ibex'')|journal=Revue d'Écologie|volume=53|issue=2|pages=153–170|doi=10.3406/revec.1998.2251|s2cid=58921073|url=https://hal.science/hal-03528980/|access-date=2023-11-07|archive-date=2023-11-07|archive-url=https://web.archive.org/web/20231107154232/https://hal.science/hal-03528980/|url-status=live}}</ref> Social spacing tends to be looser in the summer, when there is more room to feed. Ibexes have stable social connections; they consistently regroup with the same individuals when ecological conditions force them together.<ref>{{cite journal|last1=Brambilla|first1=A.|last2=von Hardenberg|first2=A.|last3=Canedoli|first3=C.|last4=Brivio|first4=F.|last5=Sueur|first5=C. |last6=Stanley|first6=C. R.|year=2022|title=Long term analysis of social structure: evidence of age-based consistent associations in male Alpine ibex|journal=Oikos|volume=2022|issue=8|page=e09511|doi=10.1111/oik.09511|bibcode=2022Oikos2022E9511B |doi-access=free|hdl=10281/402427|hdl-access=free}}</ref> Female groups tend to be more stable than male groups.<ref name=Brambilla2022/><ref name=Villaret1995/>
-In the breeding season, which occurs in December and January, adult males and females gather together, and separate again in April and May.<ref name=Parrini2009/> Among males, a [[dominance hierarchy]] based on size, age, and horn length exists.<ref>{{cite journal|last1=Bergeron|first1=P.|last2=Grignolio|first2=S.|last3=Apollonio|first3=M.|last4=Shipley|first4=B. |last5=Festa-Bianchet|first5=M.|year=2010|title=Secondary sexual characters signal fighting ability and determine social rank in Alpine ibex (''Capra ibex'')|journal=
+Adult males and females gather together in December and January, the breeding season, then separate again in April and May.<ref name=Parrini2009/> Among males, a [[dominance hierarchy]] based on size, age, and horn length exists.<ref>{{cite journal|last1=Bergeron|first1=P.|last2=Grignolio|first2=S.|last3=Apollonio|first3=M.|last4=Shipley|first4=B. |last5=Festa-Bianchet|first5=M.|year=2010|title=Secondary sexual characters signal fighting ability and determine social rank in Alpine ibex (''Capra ibex'')|journal=
-Behavioral Ecology and Sociobiology|volume=64|issue=8|pages=1299–1307|doi=10.1007/s00265-010-0944-x|s2cid=36262460}}</ref> Hierarchies are established outside the breeding season, allowing males to focus more on mating and less on fighting. Males use their horns for combat; they will bash the sides of rivals or clash head-to-head, the latter often involves them standing bipedally and clashing downwards.<ref>{{cite journal|last1=Willisch|first1=C. S.|last2=Neuhaus|first2=P.|year=2010|title=Social dominance and conflict reduction in rutting male Alpine ibex, ''Capra ibex''|journal=Behavioral Ecology|volume=21|issue=2|pages=372–380|doi=10.1093/beheco/arp200|doi-access=free}}</ref>
+Behavioral Ecology and Sociobiology|volume=64|issue=8|pages=1299–1307|doi=10.1007/s00265-010-0944-x|bibcode=2010BEcoS..64.1299B |s2cid=36262460}}</ref> Hierarchies are established outside the breeding season, allowing males to focus more on mating and less on fighting. Males use their horns for combat; they bash rivals' sides or clash head-to-head often by rearing then clashing downwards.<ref>{{cite journal|last1=Willisch|first1=C. S.|last2=Neuhaus|first2=P.|year=2010|title=Social dominance and conflict reduction in rutting male Alpine ibex, ''Capra ibex''|journal=Behavioral Ecology|volume=21|issue=2|pages=372–380|doi=10.1093/beheco/arp200|doi-access=free}}</ref>
 Alpine ibexes communicate mainly through short, sharp whistles that serve mostly as alarm calls and may occur singularly or in succession with short gaps. Females and their young communicate by bleating.<ref name=Brambilla2022/>
@@ Line 95: / Line 94: @@
 [[File:Alpine ibex Cima di Terrarossa 7.jpg|thumb|Mother with young|alt=Two goat-like animals with horns stand on a rocky slope with grass. The animal on the left is smaller and younger than the one on the right. The sky behind the slope is blue.]]
-The mating season begins in December and typically lasts for around six weeks. During this time, male herds break up into smaller groups and search for females. The [[rut (mammalian reproduction)|rut]] takes place in two phases; in the first phase, males interact with females as a group and in the second phase, one male separates from his group to follow a female in [[oestrus]].<ref name=Parrini2009/> Dominant males between nine and twelve years old follow a female and guard her from rivals while subordinate, younger males between two and six years old try to sneak past the tending male when he is distracted. If the female flees, both dominant and subordinate males will try to follow her. During courtship, the male stretches the neck, flicks the tongue, [[flehmen response|curls the upper lip]], urinates, and sniffs the female.<ref>{{cite journal |last1=Willisch |first1=C. S. |last2=Neuhaus |first2=P. |year=2009 |title=Alternative mating tactics and their impact on survival in adult male Alpine ibex (''Capra ibex ibex'') |journal=Journal of Mammalogy |volume=90 |issue=6 |pages=1421–1430 |doi=10.1644/08-MAMM-A-316R1.1 |s2cid=86267022 |doi-access=free}}</ref> After [[copulation (zoology) |copulation]], the male rejoins his group and restarts the first phase of the rut.<ref name=Parrini2009/> Environmental conditions can affect courtship in the species; for example, snow can limit the males' ability to follow females and mate with them.<ref>{{cite journal |last1=Rossi |first1=I. |last2=Grignolio |first2=S. |last3=Bassano |first3=B. |last4=Apollonius |first4=M. |year=2003 |title=Strategie riproduttive dello Stambecco (Capra ibex ibex) nel Parco Nazionale del Gran Paradiso |journal=Hystrix, the Italian Journal of Mammalogy |volume=14 (ATIt Congress Supplement) |issue=Special Issue: IV ATIt Congress |doi=10.4404/hystrix-14.0-4292 |doi-broken-date=17 March 2024 |language=it}}</ref>
+The mating season begins in December and typically lasts for around six weeks. During this time, male herds break up into smaller groups and search for females. The [[rut (mammalian reproduction)|rut]] takes place in two phases; in the first phase, males interact with females as a group and in the second phase, one male separates from his group to follow a female in [[oestrus]].<ref name=Parrini2009/> Dominant males between nine and twelve years old follow a female and guard her from rivals while subordinate, younger males between two and six years old try to sneak past the tending male when he is distracted. If the female flees, both dominant and subordinate males will try to follow her. During courtship, the male stretches the neck, flicks the tongue, [[flehmen response|curls the upper lip]], urinates, and sniffs the female.<ref>{{cite journal |last1=Willisch |first1=C. S. |last2=Neuhaus |first2=P. |year=2009 |title=Alternative mating tactics and their impact on survival in adult male Alpine ibex (''Capra ibex ibex'') |journal=Journal of Mammalogy |volume=90 |issue=6 |pages=1421–1430 |doi=10.1644/08-MAMM-A-316R1.1 |s2cid=86267022 |doi-access=free}}</ref> After [[copulation (zoology) |copulation]], the male rejoins his group and restarts the first phase of the rut.<ref name=Parrini2009/> Environmental conditions can affect courtship in the species; for example, snow can limit the males' ability to follow females and mate with them.<ref>{{cite journal |last1=Rossi |first1=I. |last2=Grignolio |first2=S. |last3=Bassano |first3=B. |last4=Apollonius |first4=M. |year=2003 |title=Strategie riproduttive dello Stambecco (Capra ibex ibex) nel Parco Nazionale del Gran Paradiso |journal=Hystrix, the Italian Journal of Mammalogy |volume=14 (ATIt Congress Supplement) |issue=Special Issue: IV ATIt Congress |doi=10.4404/hystrix-14.0-4292 |doi-broken-date=2 November 2024 |language=it}}</ref>
-The female is in oestrus for around 20 days and gestation averages around five months, and typically results in the birth of one or sometimes two kids.<ref name=Stuwe1987>{{cite journal |last1=Stüwe |first1=M. |last2=Grodinsky |first2=C. |year=1987 |title=Reproductive biology of captive Alpine ibex (''Capra i. ibex'') |journal=Zoo Biology |volume=6 |issue=4 |pages=331–339 |doi=10.1002/zoo.1430060407}}</ref> Females give birth away from their social groups on rocky slopes that are relatively safe from predators.<ref>{{cite journal |last1=Grignoli |first1=S. |last2=Rossi |first2=I. |last3=Bertolotto |first3=E. |last4=Bassano |first4=B. |last5=Apollonio |first5=M. |year=2007 |title=Influence of the kid on space use and habitat selection of female Alpine ibex |journal=The Journal of Wildlife Management |volume=71 |issue=3 |pages=713–719 |doi=10.2193/2005-675 |jstor=4495243 |bibcode=2007JWMan..71..713G |s2cid=84661674}}</ref> After a few days, the kids can move on their own. Mothers and kids gather into [[Crèche (zoology)|nursery]] groups, where young are nursed for up to five months.<ref name=Brambilla2022/> Nursery groups can also include non-lactating females.<ref name=Neuhaus/> Alpine ibexes reach sexual maturity at 18 months but females continue to grow until they are around five or six years old, and males are nine to eleven years old.<ref name=Parrini2009/>
+The female is in oestrus for around 20 days and gestation averages around five months, and typically results in the birth of one or sometimes two kids.<ref name=Stuwe1987>{{cite journal |last1=Stüwe |first1=M. |last2=Grodinsky |first2=C. |year=1987 |title=Reproductive biology of captive Alpine ibex (''Capra i. ibex'') |journal=Zoo Biology |volume=6 |issue=4 |pages=331–339 |doi=10.1002/zoo.1430060407}}</ref> Females give birth away from their social groups on rocky slopes that are relatively safe from predators.<ref>{{cite journal |last1=Grignoli |first1=S. |last2=Rossi |first2=I. |last3=Bertolotto |first3=E. |last4=Bassano |first4=B. |last5=Apollonio |first5=M. |year=2007 |title=Influence of the kid on space use and habitat selection of female Alpine ibex |journal=The Journal of Wildlife Management |volume=71 |issue=3 |pages=713–719 |doi=10.2193/2005-675 |jstor=4495243 |bibcode=2007JWMan..71..713G |s2cid=84661674}}</ref> After a few days, the kids can move on their own. Mothers and kids gather into [[Crèche (zoology)|nursery]] groups, where young are nursed for up to five months.<ref name=Brambilla2022/> Nursery groups can also include non-lactating females.<ref name=Neuhaus/> Alpine ibexes reach sexual maturity in 18 months but continue to grow until females are five to six and males are nine to eleven years old.<ref name=Parrini2009/>
 The horns grow throughout life. Young are born without horns, which become visible as tiny tips at one month and reach {{convert|20|–|25|mm|1|abbr=on}} in the second month.<ref name=Brambilla2022>{{cite book |last1=Brambilla |first1=A. |last2=Bassano |first2=B. |last3=Biebach |first3=I. |last4= Bollmann |first4=K. |last5=Keller |first5=L. |last6=Toïgo |first6=C. |last7=von Hardenberg |first7=A. |contribution=Alpine Ibex ''Capra ibex'' Linnaeus, 1758 |title=Terrestrial Cetartiodactyla – Handbook of the Mammals of Europe |editor1=Corlatti, L. |editor2=Zachos, F. E. |year=2022 |publisher=Springer |pages=383–408 |doi=10.1007/978-3-030-24475-0_32 |isbn=978-3-030-24474-3 |issn=2730-7387}}</ref> In males, the horns grow at about {{convert|8|cm|abbr=on}} per year for the first five-and-a-half years, slowing to half that rate once the animal reaches 10 years of age.<ref name=Parrini2009/> The slowing of horn growth in males coincides with [[senescence|aging]].<ref>{{cite journal |last1=von Hardenberg |first1=A. |last2=Bassano |first2=B. |last3= Zumel Arranz |first3=M. del P. |last4=Bogliani |first4=G. |year=2004 |title=Horn growth but not asymmetry heralds the onset of senescence in male Alpine ibex (''Capra ibex'') |journal=Journal of Zoology |volume=263 |issue=4 |pages=425–432 |doi=10.1017/S0952836904005485}}</ref> The age of an ibex can be determined by annual growth rings in the horns, which stop growing in winter.<ref>{{cite journal |last1=Michallet |first1=J. |last2=Loison |first2=A. |last3=Gaillard |first3=J. |last4=Gauthier |first4=D. |year=1994 |title=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 |journal=Gibier Faune Sauvage |volume=11 |pages=99–118 |s2cid=132957886 |language=fr}}</ref><ref>{{cite journal |last1=Ratti |first1=P. |last2=K. H. |year=1977 |title=Untersuchungen zur Altersschätzung und Altersbestimmung beim Alpensteinbock (Capra ibex ibex) im Kanton Graubünden |journal=Zeitschrift für Jagdwissenschaft |volume=23 |issue=4 |pages=188–213 |doi=10.1007/BF01905758 |s2cid=20248387 |language=de}}</ref><ref name=Brambilla2022/>
@@ Line 104: / Line 103: @@
 Male Alpine Ibexes live for around 16 years while females live for around 20 years. The species has a high adult survival rate compared with other herbivores around its size.<ref name=Toigo2007/><ref>{{cite journal |last1=Girard |first1=I. |last2=Toïgo |first2=C. |last3=Gaillard |first3=J.-M. |last4=Gauthier |first4=D. |last5=Martinot |first5=J.-P. |year=1999 |title=Patron de survie chez le bouquetin des Alpes (''Capra ibex ibex'') dans le parc national de la Vanoise |journal=Revue d'Écologie |volume=54 |issue=3 |pages=235–251 |doi=10.3406/revec.1999.2294 |s2cid=193212971 |language=fr}}</ref> In one study, all kids reached two years of age and the majority of adults lived for 13 years, although most 13-year-old males did not reach the age of 15.<ref name=Toigo2007>{{cite journal |last1=Toïgo |first1=C. |last2=Gaillard |first2=J.-M. |last3=Festa-Bianchett |first3=M. |last4=Largo |first4=E. |last5=Michallet |first5=J. |last6=Maillard |first6=D. |year=2007 |title=Sex- and age-specific survival of the highly dimorphic Alpine ibex: evidence for a conservative life-history tactic |journal=Journal of Animal Ecology |volume=76 |issue=4 |pages=679–686 |doi=10.1111/j.1365-2656.2007.01254.x |pmid=17584373|bibcode=2007JAnEc..76..679T }}</ref> Alpine ibexes have a low rate of predation;<ref name=Parrini2009/> their mountain habitat keeps them safe from predators like [[wolves]], though [[golden eagle]]s may prey on young.<ref name=Brambilla2022/> In Gran Paradiso, causes of death are old age, lack of food, and disease. They are also killed by avalanches.<ref name=Parrini2009/>
-Alpine ibexes may suffer [[necrosis]] and [[fibrosis]] caused by the bacteria ''[[Brucella melitensis]]'',<ref>{{cite journal |last1=Ferrogilo |first1=E. |last2=Tolari |first2=F. |last3=Bassano |first3=B. |year=1998 |title=Isolation of ''Brucella melitensis'' from alpine ibex |journal=Journal of Wildlife Diseases |volume=34 |issue=2 |pages=400–402 |doi=10.7589/0090-3558-34.2.400 |pmid=9577795 |s2cid=43739687 }}</ref> and [[foot rot]] caused by ''[[Dichelobacter nodosus]]''.<ref>{{cite journal |last1=Moore-Jones |first1=G. |last2=Dürr |first2=S. |last3=Willisch |first3=C. |last4=Ryser-Degiorgis |first4=M.-P. |year=2021 |title=Occurrence of footrot in free-ranging alpine ibex (''Capra ibex'') colonies in Switzerland |journal=Journal of Wildlife Diseases |volume=57 |issue=2 |pages=327–337 |doi=10.7589/JWD-D-20-00050 |pmid=33822150 |s2cid=232357703}}</ref> Infections from ''[[Mycoplasma |Mycoplasma conjunctivae]]'' damage the eye via [[keratoconjunctivitis]] and can lead to death rates of up to 30%.<ref>{{cite journal |last1=Giacometti |first1=M. |last2=Janovsky |first2=M. |last3=Belloy |first3=L. |last4=Frey |first4=J. |year=2002 |title=Infectious keratoconjunctivitis of ibex, chamois and other Caprinae |journal=Revue Scientifique et Technique (International Office of Epizootics) |volume=21 |issue=2 |pages=335–345 |doi=10.20506/rst.21.2.1338 |pmid=11974619}}</ref> Ibexes can host gastrointestinal parasite]s such as [[coccidia]], [[Strongylidae |strongyles]],<ref>{{cite journal |last1=Carcereri |first1=A. |last2=Stancampiano |first2=L. |last3=Marchiori |first3=E. |last4=Sturaro |first4=E. |last5=Ramanzin |first5=M. |last6=Cassini |first6=R. |year=2021 |title=Factors influencing gastrointestinal parasites in a colony of Alpine ibex (''Capra ibex'') interacting with domestic ruminants |journal=Hystrix, the Italian Journal of Mammalogy |volume=32 |issue=1 |pages=95–101 |doi=10.4404/hystrix-00393-2020}}</ref> ''[[Teladorsagia circumcincta]]'', and ''[[Marshallagi amarshalli]]''<ref>{{cite journal |last1=Lanfranchi |first1=P. |last2=Manfredi |first2=M. T. |last3=Zaffaroni |first3=E. |last4=Fraquelli |first4=C. |last5=Ratti |first5=P. |last6=Giacometti |first6=M. |year=1995 |title=Eine dreijährige Untersuchung der Labmagen-Helminthenfauna beim Alpensteinbock (Capra i. ibex) der Kolonie Albris, Graubünden, Schweiz |journal=Zeitschrift für Jagdwissenschaft |volume=41 |pages=24–35 |doi=10.1007/BF02241870 |s2cid=36051526 |language=de}}</ref> as well as  [[lungworms]], mainly ''[[Muellerius capillaris]]''.<ref>{{cite journal |last1=Cassini |first1=R. |last2=Párraga |first2=M. A. |last3=Signorini |first3=M. |last4=Frangipane di Regalbono |first4=A. |last5=Sturaro |first5=E. |last6=Rossi |first6=L. |last7=Ramanzin |first7=M. |year=2015 |title=Lungworms in Alpine ibex (''Capra ibex'') in the eastern Alps, Italy: An ecological approach |journal=Veterinary Parasitology |volume=214 |issue=1–2 |pages=132–138 |doi=10.1016/j.vetpar.2015.09.026 |pmid=26476915}}</ref> Several individuals have died from heart diseases, including [[arteriosclerosis]], [[cardiac fibrosis]], [[sarcosporidiosis]], and [[valvular heart disease]].<ref>{{cite journal |last1=Guarda |first1=F. |last2=Bollo |first2=E. |last3=Bassono |first3=B. |last4=Peracino |first4=V. |last5=Giraldo |first5=A. |year=2001 |title=Patologia Cardiaca Del Camoscio (Rupicapra Rupicapra) E Dello Stambecco (Capra Ibex) Nel Parco Nazionale Gran Paradiso |journal=Large Animals Review |volume=7 |issue=2 |pages=47–52 |url=https://vetjournal.it/images/archive/pdf_riviste/2632.pdf |language=it |access-date=2023-12-21 |archive-date=2023-12-21 |archive-url=https://web.archive.org/web/20231221202157/https://vetjournal.it/images/archive/pdf_riviste/2632.pdf |url-status=live}}</ref>
+Alpine ibexes may suffer [[necrosis]] and [[fibrosis]] caused by the bacteria ''[[Brucella melitensis]]'',<ref>{{cite journal |last1=Ferrogilo |first1=E. |last2=Tolari |first2=F. |last3=Bassano |first3=B. |year=1998 |title=Isolation of ''Brucella melitensis'' from alpine ibex |journal=Journal of Wildlife Diseases |volume=34 |issue=2 |pages=400–402 |doi=10.7589/0090-3558-34.2.400 |pmid=9577795 |s2cid=43739687 }}</ref> and [[foot rot]] caused by ''[[Dichelobacter nodosus]]''.<ref>{{cite journal |last1=Moore-Jones |first1=G. |last2=Dürr |first2=S. |last3=Willisch |first3=C. |last4=Ryser-Degiorgis |first4=M.-P. |year=2021 |title=Occurrence of footrot in free-ranging alpine ibex (''Capra ibex'') colonies in Switzerland |journal=Journal of Wildlife Diseases |volume=57 |issue=2 |pages=327–337 |doi=10.7589/JWD-D-20-00050 |pmid=33822150 |s2cid=232357703}}</ref> Infections from ''[[Mycoplasma |Mycoplasma conjunctivae]]'' damage the eye via [[keratoconjunctivitis]] and can lead to death rates of up to 30%.<ref>{{cite journal |last1=Giacometti |first1=M. |last2=Janovsky |first2=M. |last3=Belloy |first3=L. |last4=Frey |first4=J. |year=2002 |title=Infectious keratoconjunctivitis of ibex, chamois and other Caprinae |journal=Revue Scientifique et Technique (International Office of Epizootics) |volume=21 |issue=2 |pages=335–345 |doi=10.20506/rst.21.2.1338 |pmid=11974619}}</ref> Ibexes can host gastrointestinal parasites such as [[coccidia]], [[Strongylidae |strongyles]],<ref>{{cite journal |last1=Carcereri |first1=A. |last2=Stancampiano |first2=L. |last3=Marchiori |first3=E. |last4=Sturaro |first4=E. |last5=Ramanzin |first5=M. |last6=Cassini |first6=R. |year=2021 |title=Factors influencing gastrointestinal parasites in a colony of Alpine ibex (''Capra ibex'') interacting with domestic ruminants |journal=Hystrix, the Italian Journal of Mammalogy |volume=32 |issue=1 |pages=95–101 |doi=10.4404/hystrix-00393-2020}}</ref> ''[[Teladorsagia circumcincta]]'', and ''[[Marshallagi amarshalli]]''<ref>{{cite journal |last1=Lanfranchi |first1=P. |last2=Manfredi |first2=M. T. |last3=Zaffaroni |first3=E. |last4=Fraquelli |first4=C. |last5=Ratti |first5=P. |last6=Giacometti |first6=M. |year=1995 |title=Eine dreijährige Untersuchung der Labmagen-Helminthenfauna beim Alpensteinbock (Capra i. ibex) der Kolonie Albris, Graubünden, Schweiz |journal=Zeitschrift für Jagdwissenschaft |volume=41 |pages=24–35 |doi=10.1007/BF02241870 |s2cid=36051526 |language=de}}</ref> as well as  [[lungworms]], mainly ''[[Muellerius capillaris]]''.<ref>{{cite journal |last1=Cassini |first1=R. |last2=Párraga |first2=M. A. |last3=Signorini |first3=M. |last4=Frangipane di Regalbono |first4=A. |last5=Sturaro |first5=E. |last6=Rossi |first6=L. |last7=Ramanzin |first7=M. |year=2015 |title=Lungworms in Alpine ibex (''Capra ibex'') in the eastern Alps, Italy: An ecological approach |journal=Veterinary Parasitology |volume=214 |issue=1–2 |pages=132–138 |doi=10.1016/j.vetpar.2015.09.026 |pmid=26476915}}</ref> Several individuals have died from heart diseases, including [[arteriosclerosis]], [[cardiac fibrosis]], [[sarcosporidiosis]], and [[valvular heart disease]].<ref>{{cite journal |last1=Guarda |first1=F. |last2=Bollo |first2=E. |last3=Bassono |first3=B. |last4=Peracino |first4=V. |last5=Giraldo |first5=A. |year=2001 |title=Patologia Cardiaca Del Camoscio (Rupicapra Rupicapra) E Dello Stambecco (Capra Ibex) Nel Parco Nazionale Gran Paradiso |journal=Large Animals Review |volume=7 |issue=2 |pages=47–52 |url=https://vetjournal.it/images/archive/pdf_riviste/2632.pdf |language=it |access-date=2023-12-21 |archive-date=2023-12-21 |archive-url=https://web.archive.org/web/20231221202157/https://vetjournal.it/images/archive/pdf_riviste/2632.pdf |url-status=live}}</ref>
 ==Conservation==