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{{Short description|Species of mammal}}
{{Redirect|Steinbock|the African antelope sometimes known as "steinbuck" or "steinbok"|Steenbok|other uses|Steinbock (disambiguation)}}
{{speciesbox
| name = Alpine ibex
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name=iucn>{{cite iucn |title=''Capra ibex'' |author=Toïgo, C. |author2=Brambilla, A. |author3=Grignolio, S. |author4=Pedrotti, L. |date=2020 |page=e.T42397A161916377 |doi=10.2305/IUCN.UK.2020-2.RLTS.T42397A161916377.en |access-date=19 November 2021}}</ref>
| image = Alpensteinbock (Capra ibex) Zoo Salzburg 2014 h.jpg
| image_caption = Male
| image2 = Bouquetin01.jpg
| image2_caption = Female
| range_map=Leefgebied_alpensteenbok.JPG
| range_map_caption=Range map in the Alps
| taxon = Capra ibex
| authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]
}}

The '''Alpine ibex''' ('''''Capra ibex'''''), also known as the '''steinbock''', '''bouquetin''', or simply '''ibex''', is a species of [[Capra (genus)|wild goat]] that lives in the [[mountain]]s of the [[Europe]]an [[Alps]]. It is a [[Sexual dimorphism|sexually dimorphic]] species: males are larger and carry longer, curved horns than females. Its coat colour is typically brownish grey. Alpine ibex tend to live in steep, rough terrain near the [[snow line]]. They are also social, although adult males and females segregate for most of the year, coming together only to mate. Four distinct groups exist; adult male groups, female-offspring groups, groups of young individuals, and mixed-sex groups.

During the breeding season, males fight for access to females, and use their long horns in [[agonistic behaviour]]. After being [[Local extinction|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 [[Aosta Valley]] and [[Piedmont]] ([[Italy]]), a national park created to help the ibex thrive. The ibex is the emblem of both the Gran Paradiso National Park, and the contiguous [[Vanoise National Park]] across the French border. 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 and phylogeny==
The Alpine ibex was first described by [[Carl Linnaeus]] in 1758. It is classified in the [[genus]] ''[[Capra (genus)|Capra]]'' (Latin for "goat") with at least seven other species of wild goats. Both ''Capra'' and ''[[Ovis]]'' (sheep) descended from a [[goral]]-like animal from the [[Miocene]] and early [[Pliocene]], whose fossils are found in Kenya, China, and Slovenia. The genus ''[[Tossunnoria]]'' appears in China during the late Miocene and appears to have been intermediate between gorals and goats. Fossils of Alpine ibex date back to the late [[Pleistocene]], when the [[Spanish ibex]] and it probably evolved from the extinct Pleistocene species ''[[Capra camburgensis]]''.<ref name=Parrini2009/> The [[Nubian ibex|Nubian]] (''C. nubiana''), [[Walia ibex|walia]] (''C. walie''), and [[Siberian ibex]]es (''C. sibirica'') were previously considered to be subspecies of the Alpine ibex, giving populations in the Alps the [[Trinomial nomenclature|trinomial]] of ''C. i. ibex''.<ref>{{cite book|author =Shackleton, D. W.|year=1997|title=Wild Sheep and Goats and Their Relatives: Status Survey and Action Plan for Caprinae|publisher=International Union for Conservation of Nature and Natural Resources. Species Survival Commission. Caprinae Specialist Group|page=12|isbn=2831703530}}</ref>

==Appearance==
[[File:Steinwild Augstmatthorn.OGG|thumb|left|thumbtime=215|Video of a herd of Alpine ibex on [[Augstmatthorn]] in the [[Berner Oberland|Bernese Highlands]], Switzerland]]
Compared with other members of its genus, the Alpine ibex has a short, broad head and a duller coat. It has brownish-grey hair over most of the body, a pale abdomen, and slightly darker markings on the chin and throat and in a stripe along the back. They [[moult]] twice a year, firstly in April or May, and then again in September, when they replace the short summer coat with thicker hair and a woolly undercoat.<ref name=Parrini2009>{{cite journal |author1=Parrini, F. |author2=Cain III, J. W. |author3=Krausman, 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>

Males commonly grow to a height of {{convert|90|to|101|cm|abbr=on}} at the [[withers]], with a body length of {{convert|149|to|171|cm|abbr=on}} and weigh from {{convert|67|to|117|kg|abbr=on}}. Females are noticeably smaller, with a shoulder height of {{convert|73|to|84|cm|abbr=on}}, a body length of {{convert|121|to|141|cm|abbr=on}}, and a weight of {{convert|17|to|32|kg|abbr=on}}. Both male and female Alpine ibex have large, backwards-curving horns with numerous ridges along their length. At {{convert|69|to|98|cm|abbr=on}}, those of the males are substantially larger than those of females, which reach only {{convert|18|to|35|cm|abbr=on}} in length.<ref name=Parrini2009/>

==Distribution and ecology==
[[File:AlpineIbex.jpg|thumb|Ibex standing on cliff in winter]]
During the [[Last Glacial Period]] the species range was reaching as far south as [[Greece]], where it disappeared in the [[Holocene]], after 5,600 BC due to hunting, with the last known remains being found on mountains surrounding [[Thessaly]].<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=Alkiviadis |journal=Acta Theriologica |volume=58 |pages=1–11 |s2cid=256122729 }}</ref> The Alpine ibex was, at one point, restricted only to the Gran Paradiso National Park in northern [[Italy]], and the contiguous [[Maurienne]] Valley in the [[French Alps]]<ref>{{Cite web |url=http://www.parcnational-vanoise.fr/fr/documentation-en-ligne/doc_download/1983-dynamique-des-populations-a-extension-geographique-du-bouquetin-des-alpes-dans-le-parc-de-la-vanoise.html |title=Dynamique des populations et expansion geographique du bouquetin des alpes (''Capra ibex ibex'') dans le parc national de la vanoise |access-date=2013-09-07 |archive-url=https://web.archive.org/web/20131228151739/http://www.parcnational-vanoise.fr/fr/documentation-en-ligne/doc_download/1983-dynamique-des-populations-a-extension-geographique-du-bouquetin-des-alpes-dans-le-parc-de-la-vanoise.html |author=Girard, Irène|year=2000|publisher=PhD thesis, Université Savoie Mont Blanc|archive-date=2013-12-28 |url-status=dead }}</ref> but it was [[Reintroduction|reintroduced]] to most of the [[Alps]]. Reintroductions started in 1906 into Switzerland. Alpine ibex are now found in most or all of the Italian and French alpine ranges, southern [[Germany]], [[List of mammals of Switzerland#History|Switzerland]]<ref name=Staffelbach>{{in lang|fr}} Heinz Staffelbach, ''Manuel des Alpes suisses. Flore, faune, roches et météorologie'', Rossolis, 2009 ({{ISBN|978-2-940365-30-2}}). Also available in German: Heinz Staffelbach, ''Handbuch Schweizer Alpen. Pflanzen, Tiere, Gesteine und Wetter'', Haupt Verlag, 2008 ({{ISBN|978-3-258-07638-6}}).</ref> and [[Austria]]. It has also been [[Introduced species|introduced]] to [[Bulgaria]] and [[Slovenia]].<ref name=iucn/>

An excellent climber, its preferred [[habitat]] is the rocky region along the [[snow line]] above [[alpine forest]]s, where it occupies steep, rough terrain at elevations of {{convert|1800|to|3300|m|abbr=on}}.<ref name=Parrini2003>{{cite journal | author = Parrini, F. | 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 |display-authors=etal}}</ref> Alpine ibex are typically absent from woodland areas,<ref name=Parrini2009/> although adult males in densely populated areas may stay in [[larch]] and mixed larch-spruce woodland if no snow has fallen.<ref name=Grignolio2003/> Males spend the winter in [[coniferous]] forests.<ref name=Parrini2009/> For most of the year, males and females occupy different habitats.<ref>{{cite journal | author = ToÏgo C. | author2 = Gaillard J. M. | author3 = Michallet 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 rely on steep terrain more so than males.<ref name=Francisci1985>Francisci, F., S. Focardi, and L. Boitani. (1985) "Male and female Alpine ibex: phenology of space use and herd size", pp. 124–133. in ''The biology and management of mountain ungulates''. Lovari, S. Croom Helm. London, United Kingdom. {{ISBN|978-94-015-7346-7}}</ref> Males use lowland meadows during the spring, which is when snow melts and green grass appears.<ref name=Francisci1985/> They then climb to alpine meadows during the summer.<ref name=Grignolio2003>{{cite journal|author1=Grignolio, S.|author2=Parrini, F.|author3=Bassano, B.|author4=Luccarini, S.|author5=Apollonio, M.|year=2003|title=Habitat selection in adult males of Alpine ibex, ''Capra ibex ibex''|journal=Folia Zoologica|volume=52|issue=2|pages=113–20|url=http://www.ivb.cz/folia/52/2/113-120.pdf|access-date=2012-12-26|archive-url=https://web.archive.org/web/20130522213611/http://www.ivb.cz/folia/52/2/113-120.pdf|archive-date=2013-05-22|url-status=dead}}</ref> When winter arrives, both sexes move to steep, rocky slopes that amass little snow.<ref name=Grignolio2004>{{cite journal | author = Grignolio, S. | 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| s2cid = 85380031 |display-authors=etal}}</ref> They prefer slopes of 30–45° and use small caves and overhangs for shelter.<ref>{{cite journal|author =Wiersema, G.|year=1984|title=Seasonal use and quality assessment of ibex habitat|journal=Acta Zoologica Fennica|volume=172|pages=89–90}}</ref> Home ranges are highly variable, depending on the availability of resources, and vary in size throughout the year. Figures from {{convert|180|to|2800|ha|sqmi|abbr=on}} have been recorded.<ref name=Parrini2009/><ref name=Grignolio2004/> Home ranges tend to be largest during summer and autumn, smallest in winter, and intermediate in spring.<ref name=Parrini2009/> Female home ranges are usually smaller than those of males. Alpine ibex appear to have a low rate of predation and in Gran Paradiso typically die of age, starvation, or disease.<ref name=Parrini2009/>

===Foraging===
[[File:Stambecchi sulla diga del Cingino.jpg|thumb|Ibex on the wall of a dam]]
Alpine ibex are strictly [[herbivorous]], with over half of their diet consisting of grasses, and the remainder being a mixture of mosses, flowers, leaves, and twigs.<ref name=Parrini2009/> If leaves and shoots are out of reach, they often stand on their rear legs to reach this food. Grass genera that are the most commonly eaten are ''[[Agrostis]]'', ''[[Avena]]'', ''[[Calamagrostis]]'', ''[[Festuca]]'', ''[[Phleum]]'', ''[[Poa]]'', ''[[Sesleria]]'', and ''[[Trisetum]]''.<ref name=Parrini2009/> The climbing ability of the Alpine ibex is such that it has been observed standing on the sheer face of the [[Cingino Dam]] in [[Piedmont]], [[Italy]], where it licks the stonework to obtain mineral salts.<ref>{{cite news |url=https://www.theguardian.com/environment/2010/nov/03/goats-italy-dam-precipitous-heights |title=The goats with a head for heights |first=Terry |last=Nutkins |date=3 November 2010 |work=Guardian.co.uk |publisher=Guardian News and Media Limited |access-date=4 November 2010}}</ref>

==Life history==
Although the Alpine ibex is a social species, they segregate sexually and spatially depending on the season.<ref name=Francisci1985/> Four types of groups exist: Adult male groups, female-offspring groups, groups of young individuals 2–3 years old, and mixed-sex groups.<ref name=Parrini2009/><ref name=Villaret1995/> Young groups are numerous at the beginning of summer, but are expelled by females at the end of their gestation period. Female and offspring groups occur year-round, at least in an area of the French Alps.<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}}</ref> Mixed-sex groups of adult males and females occur during breeding, which lasts from December to January. By April and May, the adults separate.<ref name=Villaret1995/> The largest aggregations of either sex occur during June and July. Gatherings of males begin to decrease during October and November, and are lowest from the rut from December to March.<ref name=Villaret1995/> The males then leave their separate wintering areas and gather again.<ref>{{cite journal|author1=Parrini, F. |author2=Grignolio, S. |author3=Luccarini, S. |author4=Bassano, B. |author5=Apollonio, 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=411–23|doi=10.1007/BF03194179|s2cid=6211702 }}</ref>

[[File:Bouquetins combat.jpg|thumb|right|Male ibex locking horns]]
A linear dominance hierarchy exists among males. In small populations, which are more cohesive, males know their place in the hierarchy based on memories of past encounters,<ref name=Parrini2009/> while in mobile and large groups, where encounters with strangers are common, rank is based on horn size.<ref>Schaller, G. B. (1977) ''Mountain monarchs: wild sheep and goats of the Himalaya''. University of Chicago Press. Chicago, Illinois.</ref> Antagonistic behavior in males can come in the form of "direct" or "indirect" aggression. With direct aggression, one male bumps another with his horns or places himself in front of his opponent. He stands on his hind legs and comes down on his opponent with his horns. This may signal that he is ready to clash or may be attempting a real clash.<ref name=Parrini2009/> Indirect aggression is mostly intimidation displays.<ref name=Parrini2009/>
[[File:Alpine ibex Cima di Terrarossa 7.jpg|left|thumb|Mother with young]]

===Reproduction and growth===
The breeding season starts 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 male groups interact with the females that are all in [[oestrous]]. The higher the male's rank, the closer he can get to a female.<ref name=Parrini2009/> Males perform [[courtship display]]s. In the second phase of the [[rut (mammalian reproduction)|rut]], one male separates from his group to follow an individual female. He displays to her and guards her from other males. Before [[copulation (zoology)|copulation]], the female moves her tail and courtship becomes more intense. They copulate and then he rejoins his group and reverts to the first phase.<ref name=Parrini2009/> Gestation lasts around 167 days, and results in the birth of one or two kids, with twins making up about 20% of births.<ref name=Stuwe1987>{{cite journal |author1=Stüwe, M. |author2=Grodinsky, 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>

Alpine ibex reach sexual maturity at 18 months, but females do not reach their maximum body size for five to six years, and males not for 9–11 years. The horns grow throughout life, growing most rapidly during the second year of life, and thereafter by about {{convert|8|cm|abbr=on}} a year, eventually slowing to half that rate once the animal reaches 10 years of age. Alpine ibex live for up to 19 years in the wild.<ref name=Toigo2007>{{cite journal | author = ToÏgo, C. | 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 |display-authors=etal}}</ref>

===Genome===
A [[genome|genomic]] analysis of the unusually well-characterized bottlenecks of Alpine ibex populations, that were once nearly extinct, was carried out.<ref name="Grossen2020">{{Cite journal |pmc=7035315 |date=2020 |last1=Grossen |first1=C. |last2=Guillaume |first2=F. |last3=Keller |first3=L. F. |last4=Croll |first4=D. |title=Purging of highly deleterious mutations through severe bottlenecks in Alpine ibex |journal=Nature Communications |volume=11 |issue=1 |page=1001 |doi=10.1038/s41467-020-14803-1 |pmid=32081890 }}</ref> This analysis provided genomic evidence of purging of highly deleterious [[mutation]]s within these populations, but also evidence of concurrent accumulation of mildly deleterious mutations.<ref name = Grossen2020/>

==Conservation status==
[[File:Jeune bouquetin sur un rocher.jpg|thumb|right|Young Alpine ibex on a cliff]]
The Alpine ibex historically ranged through France, Italy, Switzerland, Liechtenstein, Bavaria, Austria and Slovenia. Starting in the early 16th century and with firearms becoming common, the overall population declined due to [[overexploitation]] and [[poaching]]. The ibex became extirpated in Switzerland and Germany by the 18th century, and was extirpated in Austria and northeastern Italy by the 19th century. They remained only in and around the adjacent [[Gran Paradiso]] and [[Vanoise Massif|Vanoise]] Massifs, then both part of the [[Kingdom of Sardinia (1720–1861)|Kingdom of Sardinia]]. Located in the western Italian Alps and the Maurienne valley in the north eastern French alps, bordering the Vanoise and Gran Paradiso Massif, the park was declared a royal hunting reserve in 1854 with the name of "Royal hunting reserve of Gran Paradiso" by [[Victor Emmanuel II]], the first king of a united Italy.<ref name="recovery">{{cite journal|author1=Stüwe, M.|author2=Nievergelt, B.|year=1991|title=Recovery of Alpine ibex from near extinction: the result of effective protection, captive breeding, and reintroductions|journal=Applied Animal Behaviour Science |volume=29|issue=1–4|pages=379–87|doi=10.1016/0168-1591(91)90262-V}}</ref>

Ibex were protected from poaching and their numbers increased, reaching 3,020 in 1914. The ibex enjoyed further protection when Gran Paradiso was made into a national park in 1922. Animals from this population naturally dispersed into surrounding regions. However, reintroductions have been the predominant source of new populations. Today, the total population of Alpine ibex is over 30,000 and is considered to be of [[Least Concern]] by the [[IUCN]]. However, Alpine ibex have low genetic diversity putting them at risk of [[inbreeding depression]].<ref name=iucn/><ref name="Biebach, I 2009">{{cite journal|author1=Biebach, I. |author2=Keller, L. F. |year=2009 |title=A strong genetic footprint of the re-introduction history of Alpine ibex (''Capra ibex ibex'')|journal=Molecular Ecology|volume=18|issue=24|pages=5046–58|doi=10.1111/j.1365-294X.2009.04420.x |pmid=19912536|s2cid=36215646}}</ref>

==References==
{{Reflist}}

==External links==

* [http://gse.pngp.it/ Alpine Ibex European Specialists Group]
* [http://www.ultimateungulate.com/Artiodactyla/Capra_ibex.html Ultimate Ungulate factsheet]
* [http://animal.discovery.com/guides/mammals/habitat/alpine/ibex.html Animal Planet Mammal Guide]

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[[Category:Capra (genus)]]
[[Category:Mammals of Europe]]
[[Category:Mammals described in 1758]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Articles containing video clips]]

Alpine ibex - Revision history

Ajay Kumar: Created a new article