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'{{Short description|Genus of antelope}} {{About|the genus of animal}} {{Redirect|Gnu|the free software collection|GNU|the free software project|GNU Project|other uses}} {{Redirect|Wilde beast|the roller coaster|Wilde Beast}} {{Good article}} {{Use dmy dates|date=June 2021}} {{Use British English|date=June 2014}} {{Automatic taxobox | name = Wildebeest | fossil_range = {{Fossil range|2.5|0}}Early [[Pleistocene]] – present | image = Blue Wildebeest, Ngorongoro.jpg | image_caption = [[Blue wildebeest]], ''C. taurinus''<br/>[[Ngorongoro Crater]], [[Tanzania]] | image2 = Black_wildebeest,_or_white-tailed_gnu,_Connochaetes_gnou_at_Krugersdorp_Game_Reserve,_Gauteng,_South_Africa_(30570451563).jpg | image2_caption = [[Black wildebeest]], ''C. gnou''<br/>[[Krugersdorp Game Reserve]], [[South Africa]] | taxon = Connochaetes | authority = [[Hinrich Lichtenstein]], 1812 | range_map = Connochaetes map.png | range_map_caption = Range map: <br/>Black wildebeest shown yellow<br/>Blue wildebeest shown blue<br/>Overlapping range shown brown | subdivision_ranks = Species | subdivision = ''Connochaetes gnou'' – [[black wildebeest]]<br/>''Connochaetes taurinus'' – [[blue wildebeest]] }} '''Wildebeest''' ({{IPAc-en|ˈ|w|ɪ|l|d|ɪ|b|iː|s|t}} {{respell|WIL|dib|eest}},<ref name="df">{{cite web|url=http://dictionary.reference.com/browse/wildebeest |title=Wildebeest &#124; Define Wildebeest at Dictionary.com |publisher=Dictionary.reference.com |access-date=25 March 2014}}</ref><ref>{{cite web|url=http://dictionary.cambridge.org/dictionary/british/wildebeest |title=wildebeest noun – definition in the British English Dictionary & Thesaurus – Cambridge Dictionaries Online |publisher=Dictionary.cambridge.org |date=18 February 2014 |access-date=25 March 2014}}</ref><ref name="tfd">{{cite web|url=http://www.thefreedictionary.com/wildebeest |title=wildebeest – definition of wildebeest by the Free Online Dictionary, Thesaurus and Encyclopedia |publisher=Thefreedictionary.com |access-date=25 March 2014}}</ref> {{IPAc-en|ˈ|v|ɪ|l|-}} {{respell|VIL|-}},<ref name="tfd"/> {{IPAc-en|-|d|ə|b|-}}), also called '''gnu''' ({{IPAc-en|nj|uː}} {{respell|NEW}} or {{IPAc-en|n|uː}} {{respell|NOO}}),<ref name="df2">{{cite web|url=http://dictionary.reference.com/browse/gnu |title=Gnu &#124; Define Gnu at Dictionary.com |publisher=Dictionary.reference.com |date=12 April 1997 |access-date=25 March 2014}}</ref><ref>{{cite web|url=http://dictionary.cambridge.org/dictionary/british/gnu |title=gnu noun – definition in the British English Dictionary & Thesaurus – Cambridge Dictionaries Online |publisher=Dictionary.cambridge.org |date=18 February 2014 |access-date=25 March 2014}}</ref><ref name="tfd2">{{cite web|url=http://www.thefreedictionary.com/gnu |title=gnu – definition of gnu by the Free Online Dictionary, Thesaurus and Encyclopedia |publisher=Thefreedictionary.com |access-date=25 March 2014}}</ref> are [[antelope]]s of the genus '''''Connochaetes''''' and native to Eastern and Southern Africa. They belong to the family [[Bovidae]], which includes true antelopes, cattle, goats, sheep, and other even-toed horned [[ungulate]]s. There are two species of wildebeest: the [[black wildebeest]] or white-tailed gnu (''C. gnou''), and the [[blue wildebeest]] or brindled gnu (''C. taurinus''). Fossil records suggest these two species diverged about one million years ago, resulting in a northern and a southern species. The blue wildebeest remained in its original range and changed very little from the ancestral species, while the black wildebeest changed more as adaptation to its open grassland habitat in the south. The most obvious way of telling the two species apart are the differences in their colouring and in the way their horns are oriented.<ref>{{Cite web|title=Wildebeests in Africa! Visit Africa|url=http://visitafrica.site/wildebeests-in-africa.html|access-date=2020-07-16|website=visitafrica.site|language=en-GB}}</ref> In [[East Africa]], the blue wildebeest is the most abundant [[Game (hunting)|big-game species]]; some populations perform an annual migration to new grazing grounds, but the black wildebeest is merely nomadic. Breeding in both takes place over a short period of time at the end of the rainy season and the calves are soon active and are able to move with the herd, a fact necessary for their survival. Nevertheless, some fall prey to large [[carnivore]]s, especially the [[spotted hyena]]. Wildebeest often graze in mixed herds with [[zebra]], which gives heightened awareness of potential [[Predation|predators]]. They are also alert to the warning signals emitted by other animals such as [[baboon]]s. Wildebeest are a tourist attraction but compete with domesticated livestock for pasture and are sometimes blamed by farmers for transferring diseases and parasites to their cattle. Illegal hunting does take place but the population trend is fairly stable and, with some in national parks or on private land. The [[International Union for Conservation of Nature]] lists both kinds of Wildebeest as [[least-concern species]]. ==Etymology== {{lang|nl|Wildebeest}} is [[Dutch language|Dutch]] for "wild beast" or "wild cattle" in [[Afrikaans]] (''[[:wikt:bees#Afrikaans|bees]]'' "[[cattle]]"), while ''Connochaetes'' derives from the [[Ancient Greek]] words {{lang|grc|κόννος}}, {{transl|grc|kónnos}}, "[[beard]]",<ref>{{LSJ|ko/nnos|κόννος|ref}}.</ref> and [[:wikt:χαίτη|χαίτη]], {{transl|grc|khaítē}}, "flowing hair", "[[lion#mane|mane]]".<ref>{{LSJ|xai/th|χαίτη|shortref}}.</ref><ref name=Benirschke>{{cite web |url=http://placentation.ucsd.edu/gnu.html |title=Wildebeest, Gnu |author=Benirschke, Kurt |year=2002 |work=Comparative Placentation |access-date=25 March 2014}}</ref> Some sources claim the name "gnu" originates from the [[Khoekhoe]] name for these animals, ''t'gnu''{{which lang|date=January 2022}}.<ref name=mw>{{cite dictionary|title=Gnu|url=http://www.merriam-webster.com/dictionary/gnu|dictionary=[[Merriam-Webster]] |access-date=14 January 2014}}</ref> Others contend the name and its pronunciation in English go back to the word ''!nu:''{{which lang|date=January 2022}} used for the [[black wildebeest]] by the [[San people]].<ref name=nu!>{{cite web|title=Whence the Wildebeest: An Implausibility of Gnus|url=http://veneryterms.blogspot.com/2014/04/whence-wildebeest-implausibility-of-gnus.html|author=Hurst, Richard E. |access-date=7 May 2014|date=2014-04-27}}</ref> ==Classification== ===Taxonomy and evolution=== The wildebeest, genus ''Connochaetes'', is placed under the family Bovidae and subfamily [[Alcelaphinae]], where its closest relatives are the hartebeest (''[[Alcelaphus]]'' spp.), the [[hirola]] (''Beatragus hunteri''), and species in the genus ''[[Damaliscus]]'', such as the [[topi]], the [[Common tsessebe|tsessebe]], the [[blesbok]] and the [[bontebok]].<ref name=msw3/> The name ''Connochaetes'' was given by German [[zoology|zoologist]] [[Hinrich Lichtenstein]] in 1812.<ref name=msw3/><ref name=groves>{{cite book|last=Groves|first=C.|title=Ungulate Taxonomy |year=2011 |publisher=Johns Hopkins University Press|location=Baltimore, Maryland|isbn=978-1-4214-0093-8|author2=Grubb, P. }}</ref> [[Dutch people|Dutch]] settlers first discovered wildebeest in about 1700, on their way to the interior of South Africa. Due to their resemblance to wild cattle, these people called them "wild ox" or "wildebeest". The blue wildebeest was first known to westerners in the northern part of South Africa a century later, in the 1800s.<ref name=talbot>{{cite book|last1=Talbot |first1=L. M. |last2=Talbot |first2= M. H. |title=Wildlife Monographs:The Wildebeest in Western Masailand, East Africa|year=1963|publisher=National Academies |pages=20–31}}</ref> In the early 20th century, one species of the wildebeest, ''C. albojubatus'', was identified in eastern Africa. In 1914, two separate races of the wildebeest were introduced,hbhdghsdfsdgfygfoygorgiruuhfasdhfguiguefuifdgdfgdugdsuhdgeoougugdfsgsagsuaghoiuaogiudgiuodfo namely ''Gorgon a. albojubatus'' ("Athi white-bearded wildebeest") and ''G. a. mearnsi'' ("Loita white-bearded wildebeest"). However, in 1939, the two were once again merged into a single race, ''Connochaetes taurinus albojubatus''. In the mid-20th century, two separate forms were recognised, ''Gorgon taurinus hecki'' and ''G. t. albojubatus''.<ref name=talbot/> Finally, two distinct types of wildebeest – the blue and black wildebeest – were identified. The blue wildebeest was at first placed under a separate genus, ''Gorgon'',<ref name=nowak/><ref name=corbet>{{cite journal|last1=Corbet|first1=S. W.|last2=Robinson |first2= T. J.|title=Genetic divergence in South African Wildebeest: comparative cytogenetics and analysis of mitochondrial DNA|journal=The Journal of Heredity|date=November–December 1991 |volume=82 |issue=6 |pages=447–52 |pmid=1795096 |doi=10.1093/oxfordjournals.jhered.a111126}}</ref> while the black wildebeest belonged to the genus ''Connochaetes''. Today, they are united in the single genus ''Connochaetes'', with the black wildebeest being named (''C. gnou'') and the blue wildebeest (''C. taurinus'').<ref name=msw3>{{MSW3|id=14200508|page=676}}</ref> According to a [[mitochondrial DNA]] analysis, the black wildebeest are estimated to have diverged from the main lineage during the [[Middle Pleistocene]] and became a distinct species around a million years ago.<ref name=bassi>{{cite book |last=Bassi |first=J. |title=Pilot in the Wild: Flights of Conservation and Survival |year=2013 |publisher=Jacana Media |isbn=978-1-4314-0871-9 |pages=116–118}}</ref> A divergence rate around 2% has been calculated.<ref name=corbet/> The split does not seem to have been driven by [[Competition (biology)|competition]] for resources, but instead because each species adopted a different [[ecological niche]] and occupied a different [[trophic level]].<ref name=Codron>{{cite journal |author1=Codron, Daryl |author2=Brink, James S. |year=2007 |title=Trophic ecology of two savanna grazers, blue wildebeest ''Connochaetes taurinus'' and black wildebeest ''Connochaetes gnou'' |journal=European Journal of Wildlife Research |volume=53 |issue=2 |pages=90–99 |doi=10.1007/s10344-006-0070-2 |s2cid=26717246 |url=http://www.zora.uzh.ch/id/eprint/25325/2/D_Codron.pdf }}</ref>{{failed verification|date=December 2021}} Blue wildebeest fossils dating back some 2.5 million years ago are common and widespread. They have been found in the fossil-bearing caves at the [[Cradle of Humankind]] north of [[Johannesburg]]. Elsewhere in South Africa, they are plentiful at such sites as [[Saldanha Bay|Elandsfontein]], [[Cornelia, Free State|Cornelia]], and [[Florisbad]].<ref name=hb>{{cite book|last=B.|first=Hilton-Barber|title=Field Guide to the Cradle of Humankind : Sterkfontein, Swartkrans, Kromdraai & Environs World Heritage Site|year=2004 |publisher=Struik|location=Cape Town|isbn=978-177-0070-653 |pages=162–163 |edition=2nd |author2=Berger, L. R. }}</ref> The earliest fossils of the black wildebeest were found in sedimentary rock in Cornelia in the [[Free State (province)|Free State]] and dated back about 800,000 years.<ref name=Codron/> Today, five subspecies of the blue wildebeest are recognised, while the black wildebeest has no named subspecies.<ref name=ackermann>{{cite journal|last=Ackermann|first=Rebecca|author2=James S. Brink |author3=Savvas Vrahimis |author4=Bonita de Klerk |title=Hybrid Wildebeest (Artiodactyla: Bovidae) Provide Further Evidence For Shared Signatures of Admixture in Mammalian Crania|journal=South African Journal of Science|year=2010 |volume=106 |issue=11/12 |pages=90–94 |doi=10.4102/sajs.v106i11/12.423|doi-access=free }}</ref><ref>{{Cite encyclopedia|url=https://www.britannica.com/animal/gnu|title=gnu {{!}} mammal|encyclopedia=Encyclopedia Britannica|access-date=2017-08-23|language=en}}</ref> ===Genetics and hybrids=== The diploid number of [[chromosome]]s in the wildebeest is 58.<ref name=skinner>{{cite book|last=Skinner|first=J. D.|title=The Mammals of the Southern African Subregion|year=2005|publisher=Cambridge University Press|location=Cambridge|isbn=978-0-521-84418-5|pages=645–8|edition=3rd|author2=Chimimba, C. T. }}</ref> Chromosomes were studied in a male and a female wildebeest. In the female, all except a pair of very large [[submetacentric]] chromosomes were found to be [[acrocentric]]. [[Metaphase]]s were studied in the male's chromosomes, and very large submetacentric chromosomes were found there, as well, similar to those in the female both in size and morphology. Other chromosomes were acrocentric. The [[X chromosome]] is a large acrocentric and the [[Y chromosome]] a minute one.<ref name=groves/><ref name=wallace>{{cite journal|last=Wallace|first=C.|title=Chromosome analysis in the Kruger National Park: The chromosomes of the blue wildebeest ''Connochaetes taurinus''|journal=Koedoe|year=1978|volume=21|issue=1|pages=195–6|doi=10.4102/koedoe.v21i1.974|doi-access=free}}</ref> The two species of the wildebeest are known to [[Hybrid (biology)|hybridise]]. Male black wildebeest have been reported to mate with female blue wildebeest and vice versa.<ref name=hybrid1>{{cite journal|last=Grobler|first=J. P.|author2=Rushworth, I. |author3=Brink, J. S. |author4=Bloomer, P. |author5=Kotze, A. |author6=Reilly, B. |author7= Vrahimis, S. |title=Management of hybridization in an endemic species: decision making in the face of imperfect information in the case of the black wildebeest—''Connochaetes gnou''|journal=European Journal of Wildlife Research|date=5 August 2011|volume=57|issue=5|pages=997–1006|doi=10.1007/s10344-011-0567-1|issn=1439-0574|hdl=2263/19462|s2cid=23964988|hdl-access=free}}</ref> The differences in social behaviour and habitats have historically prevented interspecific hybridisation between the species, but hybridisation may occur when they are both confined within the same area. The resulting offspring are usually fertile. A study of these hybrid animals at [[Spioenkop Dam Nature Reserve]] in South Africa revealed that many had disadvantageous abnormalities relating to their teeth, horns, and the [[wormian bones]] in the skull.<ref name=hybrid2>{{cite journal|last=Ackermann|first=R. R.|author2=Brink, J. S. |author3=Vrahimis, S. |author4= De Klerk, B. |title=Hybrid wildebeest (Artiodactyla: Bovidae) provide further evidence for shared signatures of admixture in mammalian crania|journal=South African Journal of Science|date=29 October 2010 |volume=106 |issue=11/12 |pages=1–4 |doi=10.4102/sajs.v106i11/12.423|doi-access=free }}</ref> Another study reported an increase in the size of the hybrid as compared to either of its parents. In some animals, the [[tympanic part of the temporal bone]] is highly deformed, and in others, the [[Radius (bone)|radius]] and [[ulna]] are fused.<ref name=hybrid3>{{cite journal|last=De Klerk|first=B.|title=An osteological documentation of hybrid wildebeest and its bearing on black wildebeest (''Connochaetes gnou'') evolution (Doctoral dissertation)|year=2008}}</ref> ===Characteristics of the species=== {{Multiple image | direction = horizontal | align = top | header = Wildebeest | width1 = 220 | image1 = Blue Wildebeest Mikumi clipped.jpg | caption1 = Blue wildebeest | width2 = 220 | image2 = Black Wildebeest.jpg | caption2 = Black wildebeest }} Both species of wildebeest are even-toed, horned, greyish-brown ungulates resembling cattle. Males are larger than females and both have heavy forequarters compared to their hindquarters. They have broad muzzles, Roman noses, and shaggy manes and tails.<ref name=ulfstrand /> The most striking morphological differences between the black and blue wildebeest are the orientation and curvature of their horns and the colour of their coats. The blue wildebeest is the bigger of the two species. In males, blue wildebeest stand {{convert|150|cm|abbr=on}} tall at the shoulder and weigh around {{convert|250|kg|abbr=on}}, while the black wildebeest stands {{convert|111|–|120|cm|abbr=on}} tall<ref>{{cite web|last=Lundrigan|first=Barbara|title=Connochaetes gnou |url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Connochaetes_gnou.html |work=Animal Diversity Web|access-date=25 March 2014}}</ref> and weighs about {{convert|180|kg|abbr=on}}. In females, blue wildebeest have a shoulder height of {{convert|135|cm|abbr=on}} and weigh {{convert|180|kg|abbr=on}} while black wildebeest females stand {{convert|108|cm|abbr=on}} at the shoulder and weigh {{convert|155|kg|abbr=on}}. The horns of blue wildebeest protrude to the side, then curve downwards before curving up back towards the skull, while the horns of the black wildebeest curve forward then downward before curving upwards at the tips. Blue wildebeest tend to be a dark grey colour with stripes, but may have a bluish sheen. The black wildebeest has brown-coloured hair, with a mane that ranges in colour from cream to black, and a cream-coloured tail. The blue wildebeest lives in a wide variety of habitats, including woodlands and grasslands, while the black wildebeest tends to reside exclusively in open grassland areas.<ref name=ackermann /> In some areas, the blue wildebeest migrates over long distances in the winter, whereas the black wildebeest does not.<ref name=hoffman>{{cite journal|last1=Hoffman|first1=Louw|title=Effect of Season and Gender on the Physical and Chemical Composition of Black Wildebeest (''Connochaetus gnou'') Meat|journal=South African Journal of Wildlife Research|year=2009|volume=39|issue=2 |pages=170–174 |doi=10.3957/056.039.0208 |last2=Schalkwyk |first2=Sunet van |last3=Muller|first3=Nina|s2cid=83957360}}</ref> The milk of the female black wildebeest contains a higher protein, lower fat, and lower lactose content than the milk of the blue wildebeest.<ref name=Osthoff>{{cite journal|last=Osthoff|first=G.|author2=A. Hugo |author3=M. de Wit |title=Comparison of the Milk Composition of Free-ranging Blesbok, Black Wildebeest and Blue Wildebeest of the Subfamily Alcelaphinae (family: Bovidae)|journal=Comparative Biochemistry and Physiology B|year=2009|volume=154|issue=1|pages=48–54|doi=10.1016/j.cbpb.2009.04.015|pmid=19426824}}</ref> Wildebeest can live more than 40 years, though their average lifespan is around 20 years.<ref name="National geographic">{{cite web|title= Wildebeest |url= http://animals.nationalgeographic.com/animals/mammals/wildebeest/ | publisher=[[National Geographic Society|National Geographic]] |access-date=25 March 2014|date= 2010-11-11 }}</ref> ==Distribution and habitat== Wildebeest inhabit the plains and open woodlands of parts of Africa south of the Sahara. The black wildebeest is native to the southernmost parts of the continent.<ref>{{Cite news|url=http://www.nationalgeographic.com/animals/mammals/b/blue-wildebeest/|title=Wildebeest {{!}} National Geographic|date=2010-11-11|access-date=2017-08-23}}</ref> Its historical range included South Africa, Eswatini, and Lesotho, but in the latter two countries, it was hunted to extinction in the 19th century. It has now been reintroduced to them and also introduced to Namibia, where it has become well established.<ref name=iucngnou/> It inhabits open plains, grasslands, and [[Karoo|Karoo shrublands]] in both steep mountainous regions and lower undulating hills at altitudes varying between {{convert|1350|and|2150|m|ft|abbr=on}}.<ref name=iucntaur/> In the past, it inhabited the [[highveld]] [[temperate grassland]]s during the dry winter season and the arid Karoo region during the rains. However, as a result of widespread hunting, the black wildebeest no longer occupies its historical range or makes migrations, and is now largely limited to game farms and protected reserves.<ref name=estes1991>{{cite book|last=Estes|first=R. D.|title=The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates|year=2004|publisher=University of California Press|isbn=978-052-0080-850|page=[https://archive.org/details/isbn_0520080858/page/133 133]|url=https://archive.org/details/isbn_0520080858/page/133}}</ref> The blue wildebeest is native to eastern and southern Africa. Its range includes [[Kenya]], [[Tanzania]], [[Botswana]], [[Zambia]], [[Zimbabwe]], [[Mozambique]], [[South Africa]], [[Swaziland]] and [[Angola]].<ref>{{Cite web|url=http://www.maasaimara.com/entries/great-wildebeest-migration-maasai-mara|title=Great Wildebeest Migration {{!}} Maasai Mara|website=maasaimara.com|access-date=2017-08-23}}</ref> It is no longer found in [[Malawi]] but has been successfully reintroduced into [[Namibia]]. Blue wildebeest are mainly found in short grass plains bordering bush-covered acacia [[savanna]]s, thriving in areas that are neither too wet nor too dry. They can be found in habitats that vary from overgrazed areas with dense bush to open [[woodland]] floodplains.<ref name=iucntaur/> In East Africa, the blue wildebeest is the most abundant big [[Game (food)|game]] species, both in population and biomass.<ref name=ulfstrand>{{cite book|last=Ulfstrand|first=Staffan|title=Savannah Lives: Animal Life and Human Evolution in Africa |url=https://archive.org/details/savannahlivesani00ulfs|url-access=registration|year=2002 |publisher=Oxford University Press|isbn=978-0-19-850925-7}}</ref> It is a notable feature of the [[Serengeti National Park]] in Tanzania, the [[Maasai Mara|Maasai Mara National Reserve]] in Kenya and the [[Liuwa Plain National Park]] in Zambia.<ref name="National geographic"/> ===Migration=== {{See also|Serengeti#Great migration}} {{Multiple image | perrow = 2 | direction = horizontal | align = top | header = Migration in wildebeest | total_width = 520 | image1 = Wbeest Mara.jpg | caption1 = Wildebeest herding and following zebra in the Serengeti National Park | image2 = Wildebeest-during-Great-Migration.JPG | caption2 = Wildebeest in Masai Mara during the [[Serengeti#Great migration|Great Migration]] | image3 = Wildebeest_Jumping_Into_the_Mara_River.jpg | caption3 = Wildebeest leaping into the Mara River [[Serengeti#Great migration|Great Migration]] | alt1 = }} Not all wildebeest are migratory. Black wildebeest herds are often nomadic or may have a regular home range of {{convert|1|km2|sqmi|lk=in|abbr=on}}. Bulls may occupy territories, usually about {{convert|100|to|400|m|-2|abbr=on}} apart, but this spacing varies according to the quality of the habitat. In favourable conditions, they may be as close as {{convert|9|m|ft|abbr=on}}, or they may be as far apart as {{convert|1600|m|ft|abbr=on}} in poor habitat.<ref name=nowak>{{cite book|last=Nowak|first=R. M.|title=Walker's Mammals of the World |url=https://archive.org/details/walkersmammalsof0002nowa|url-access=registration|year=1999 |publisher=Johns Hopkins University Press |isbn=978-0-8018-5789-8 |pages=[https://archive.org/details/walkersmammalsof0002nowa/page/1184 1184–6] |edition=6th}}</ref> Female herds have home ranges of about {{convert|250|acre|ha sqmi|}} in size. Herds of nonterritorial bachelor males roam at will and do not seem to have any restrictions on where they wander.<ref name=uu>{{cite web|last=Huffman |first=B. |title=''Connochaetes gnou'': White-tailed gnu, Black wildebeest |url=http://www.ultimateungulate.com/Artiodactyla/Connochaetes_gnou.html |publisher=Ultimate Ungulate |access-date=19 January 2014}}</ref> Blue wildebeest have both migratory and sedentary populations. In the [[Ngorongoro]], most animals are sedentary and males maintain a network of territories throughout the year, though breeding is seasonal in nature. Females and young form groups of about 10 individuals or join together in larger aggregations, and nonterritorial males form bachelor groups.<ref name=Leuthold>{{cite book |author=Leuthold, Walter |title=African Ungulates |year=1977 |chapter=The influence of environmental factors on the spatial and social organization |journal=Zoophysiology |volume=8 |pages=227–235 |doi=10.1007/978-3-642-81073-2_18 |isbn=978-3-642-81075-6 }}</ref> In the [[Serengeti]] and [[Tarangire Ecosystem|Tarangire]] ecosystems, populations are mostly migratory, with herds consisting of both sexes frequently moving, but resident subpopulations also exist.<ref name=":0">{{Cite journal|last1=Morrison|first1=Thomas A.|last2=Link|first2=William A.|last3=Newmark|first3=William D.|last4=Foley|first4=Charles A. H.|last5=Bolger|first5=Douglas T.|date=2016-05-01|title=Tarangire revisited: Consequences of declining connectivity in a tropical ungulate population|journal=Biological Conservation|volume=197|pages=53–60|doi=10.1016/j.biocon.2016.02.034|url=http://eprints.gla.ac.uk/117078/1/117078.pdf}}</ref> During the rutting season, the males may form temporary territories for a few hours or a day or so, and attempt to gather together a few females with which to mate, but soon they have to move on, often moving ahead to set up another temporary territory.<ref name=Leuthold/> In the [[Maasai Mara]] game reserve, a non-migratory population of blue wildebeest had dwindled from about 119,000 animals in 1977 to about 22,000 in 1997. The reason for the decline is thought to be the increasing competition between cattle and wildebeest for a diminishing area of grazing land as a result of changes in agricultural practices, and possibly fluctuations in rainfall.<ref>{{cite journal |author1=Ottichiloa, Wilber K. |author2=de Leeuwa, Jan |author3=Prins, Herbert H. T. |year=2001 |title=Population trends of resident wildebeest [''Connochaetes taurinus hecki'' (Neumann)] and factors influencing them in the Masai Mara ecosystem, Kenya |journal=Biological Conservation |volume=97 |issue=3 |pages=271–282 |doi=10.1016/S0006-3207(00)00090-2 }}</ref> Each year, some East African populations of blue wildebeest have a long-distance migration, seemingly timed to coincide with the annual pattern of rainfall and grass growth.<ref>{{Cite journal|last1=Bond|first1=Monica L.|last2=Bradley|first2=Curtis M.|last3=Kiffner|first3=Christian|last4=Morrison|first4=Thomas A.|last5=Lee|first5=Derek E.|date=2017-05-26|title=A multi-method approach to delineate and validate migratory corridors|journal=Landscape Ecology|volume=32|issue=8|language=en|pages=1705–1721|doi=10.1007/s10980-017-0537-4|s2cid=24743662|issn=0921-2973|url=http://eprints.gla.ac.uk/143652/1/143652.pdf}}</ref> The timing of their migrations in both the rainy and dry seasons can vary considerably (by months) from year to year. At the end of the [[wet season]] (May or June in East Africa), wildebeest migrate to dry-season areas in response to a lack of surface (drinking) water. When the rainy season begins again (months later), animals quickly move back to their wet-season ranges. Factors suspected to affect migration include food abundance, surface water availability, predators, and phosphorus content in grasses. Phosphorus is a crucial element for all life forms, particularly for lactating female bovids. As a result, during the rainy season, wildebeest select grazing areas that contain particularly high phosphorus levels.<ref name=ulfstrand /> One study found, in addition to phosphorus, wildebeest select ranges containing grass with relatively high nitrogen content.<ref name=benshahar>{{cite journal|last=Ben-Shahar|first=Raphael|author2=Coe, Malcolm J. |title=The relationships between soil factors, grass nutrients, and the foraging behaviour of wildebeest and zebra |journal=Oecologia |year=1992 |volume=90 |issue=3 |pages=422–428 |doi=10.1007/BF00317701|pmid=28313531|bibcode=1992Oecol..90..422B|s2cid=25556587}}</ref> Aerial photography has revealed that a level of organisation occurs in the movement of the herd that cannot be apparent to each individual animal; for example, the migratory herd exhibits a wavy front, and this suggests that some degree of local decision-making is taking place.<ref>{{cite journal |author1=Gueron, Shay |author2=Levin, Simon A. |year=1993 |title=Self-organization of front patterns in large wildebeest herds |journal=Journal of Theoretical Biology |volume=165 |issue=4 |pages=541–552 |doi=10.1006/jtbi.1993.1206 |bibcode=1993JThBi.165..541G }}</ref> Numerous documentaries feature wildebeest crossing rivers, with many being eaten by [[Nile crocodile|crocodile]]s or drowning in the attempt. While having the appearance of a frenzy, recent research has shown a herd of wildebeest possesses what is known as a "[[swarm intelligence]]", whereby the animals systematically explore and overcome the obstacle as one.<ref>{{cite book |author1=Corne, David W. |author2=Reynolds, Alan |author3=Bonabeau, Eric |year=2012 |title=Swarm intelligence |journal=Handbook of Natural Computing |pages=1599–1622 |doi=10.1007/978-3-540-92910-9_48 |isbn=978-3-540-92909-3 }}</ref> ==Ecology== ===Predators=== Major predators that feed on wildebeest include the [[lion]], [[hyena]], [[African wild dog]], [[cheetah]], [[leopard]], and [[Nile crocodile|crocodile]], which seem to favour the wildebeest over other prey.<ref name="National geographic"/> Wildebeest, however, are very strong, and can inflict considerable injury even to a lion. Wildebeest have a maximum running speed of around {{convert|80|km/h|abbr=on}}.<ref>{{cite web| last = PBS| title = Animal Guide: Blue Wildebeest| work = Nature| access-date =8 January 2013| url = https://www.pbs.org/wnet/nature/animal-guides/animal-guide-blue-wildebeest/3255/}}</ref><ref>{{cite book| publisher = Cambridge University Press| isbn = 9780521576734| last = McGowan| first = Christopher| title = A Practical Guide to Vertebrate Mechanics| date = 28 February 1999|page=162}}</ref> The primary defensive tactic is herding, where the young animals are protected by the older, larger ones, while the herd runs as a group. Typically, the predators attempt to isolate a young or ill animal and attack without having to worry about the herd. Wildebeest have developed additional sophisticated cooperative behaviours, such as animals taking turns sleeping while others stand guard against a night attack by invading predators. Wildebeest migrations are closely followed by [[vulture]]s, as wildebeest carcasses are an important source of food for these scavengers. The vultures consume about 70% of the wildebeest carcasses available. Decreases in the number of migrating wildebeest have also had a negative effect on the vultures.<ref name=Virani>{{cite journal|last=Virani|first=Munir Z.|author2=Kendall, Corinne |author3=Njoroge, Peter |author4=Thomsett, Simon |title=Major declines in the abundance of vultures and other scavenging raptors in and around the Masai Mara ecosystem, Kenya|journal=Biological Conservation |year=2011 |volume=144 |pages=746–752 |doi=10.1016/j.biocon.2010.10.024 |issue=2}}</ref> In the Serengeti ecosystem, Tanzania, wildebeest may help facilitate the migration of other, smaller-bodied grazers, such as [[Thomson's gazelle]]s (''Eudorcas thomsonii''), which eat the new-growth grasses stimulated by wildebeest foraging.<ref>{{cite journal |author=McNaughton, S. J. |year=1979 |title=Grazing as an optimization process: grass-ungulate relationships in the Serengeti |journal=The American Naturalist |volume=113 |issue=5 |pages=691–703 |jstor=2459961 |doi=10.1086/283426|s2cid=85338998 }}</ref> ===Interactions with nonpredators=== [[Zebra]]s and wildebeest group together in open savannah environments with high chances of predation. This grouping strategy reduces predation risk because larger groups decrease each individual's chance of being hunted, and predators are more easily seen in open areas.<ref name=Thaker>{{cite journal|last=Thaker|first=Maria|author2=Abi T. Vanak |author3=Cailey R. Owen |author4=Monika B. Ogden |author5=Rob Slotow |title=Group Dynamics of Zebra and Wildebeest in a Woodland Savanna: Effects of Predation Risk and Habitat Density|journal=PLOS ONE|year=2010|volume=5|issue=9|doi=10.1371/journal.pone.0012758|editor1-last=Getz|editor1-first=Wayne M.|pages=e12758|pmid=20862216|pmc=2942830|bibcode=2010PLoSO...512758T|doi-access=free}}</ref> The seasonal presence of thousands of migratory wildebeests reduces local lion predation on giraffe calves, resulting in greater survival of giraffes.<ref>{{Cite journal|last1=Lee|first1=Derek E.|last2=Kissui|first2=Bernard M.|last3=Kiwango|first3=Yustina A.|last4=Bond|first4=Monica L.|date=2016-10-01|title=Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes|journal=Ecology and Evolution|volume=6|issue=23|language=en|pages=8402–8411|doi=10.1002/ece3.2561|pmid=28031792|pmc=5167056|issn=2045-7758}}</ref> Wildebeest can also listen in on the alarm calls of other species, and by doing so, can reduce their risk of predation. One study showed, along with other ungulates, wildebeests responded more strongly to the baboon alarm calls compared to the baboon contest calls, though both types of calls had similar patterns, amplitudes, and durations. The alarm calls were a response of the baboons to lions, and the contest calls were recorded when a dispute between two males occurred.<ref name=Kitchen>{{cite journal|last=Kitchen|first=Dawn M.|author2=Thore J. Berman |author3=Dorothy L. Cheney |author4=James R. Nicholson |author5=Robert M. Seyfarth |title=Comparing Responses of Four Ungulate Species to Playbacks of Baboon Alarm Calls|journal=Animal Cognition|year=2010|volume=13|issue=6|pages=861–870|doi=10.1007/s10071-010-0334-9|pmid=20607576|s2cid=5736705}}</ref> Wildebeest compete with domesticated livestock for pasture and are sometimes blamed by farmers for transferring diseases and parasites to their cattle.<ref>{{Cite web|title=Wildebeests in Africa! Visit Africa|url=https://visitafrica.site/wildebeests-in-africa.html|access-date=2021-02-25|website=visitafrica.site|language=en-GB}}</ref> ===Breeding and reproduction=== [[File:Drink From Mom Wildebeest.ogv|thumb|Video of wildebeest feeding its calf]] Wildebeest do not form permanent pair bonds and during the mating season, or [[rut (mammalian reproduction)|rut]], the males establish temporary territories and try to attract females into them. These small territories are about {{convert|3000|m2|sqft acre|||}}, with up to 300 territories per {{convert|1|km2|sqmi acre|||}}. The males defend these territories from other males while competing for females that are coming into oestrus. The males use grunts and distinctive behaviour to entice females into their territories. Wildebeest usually breed at the end of the rainy season when the animals are well fed and at their peak of fitness.<ref name=ulfstrand /> This usually occurs between May and July, and birthing usually takes place between January and March, at the start of the wet season. Wildebeest females breed seasonally and ovulate spontaneously.<ref name=Clay>{{cite journal|last=Clay|first=A. Moss|author2=Estes, R. D.|author3=Thompson, K. V.|author4=Wildt, D. E.|author5=Monfort, S. L.|title=Endocrine Patterns of the Estrous Cycle and Pregnancy of Wildebeest in the Serengeti Ecosystem|journal=General and Comparative Endocrinology |year=2010 |volume=166 |issue=2 |pages=365–371|doi=10.1016/j.ygcen.2009.12.005|pmid=20036667}}</ref> The [[estrous cycle]] is about 23 days and the gestation period lasts 250 to 260 days. The calves weigh about {{convert|21|kg|0|abbr=on}} at birth<ref name=Benirschke/> and scramble to their feet within minutes, being able to move with the herd soon afterwards,<ref name="National geographic"/> a fact on which their survival relies.<ref>{{cite web |url=https://wildlifetv.wordpress.com/2013/05/11/how-do-baby-wildebeest-survive/ |title=How do baby Wildebeest survive? |date=11 May 2013 |publisher=Wildlife TV |access-date=7 October 2017}}</ref> The main predator of the calves is the [[spotted hyena]]. The calving peak period lasts for 2–3 weeks, and in small subpopulations and isolated groups, mortality of calves may be as high as 50%. However, in larger aggregations, or small groups living near large herds, mortality rates may be under 20%.<ref>{{cite journal |author=Estes, Richard D. |year=1976 |title=The significance of breeding synchrony in the wildebeest |journal=African Journal of Ecology |volume=14 |issue=2 |pages=135–152 |doi=10.1111/j.1365-2028.1976.tb00158.x }}</ref> Groups of wildebeest females and young live in the small areas established by the male. When groups of wildebeest join together, the female to male ratio is higher because the females choose to move to the areas held by a smaller number of males.<ref name=Thaker /> This female-dominated sex ratio may be due to illegal hunting and human disturbance, with higher male mortality having been attributed to hunting.<ref name=Nidablema>{{cite journal |last=Ndibalema |first=Vedasto G.|title=A comparison of sex ratio, birth periods and calf survival among Serengeti wildebeest sub-populations, Tanzania |journal=African Journal of Ecology |year=2009 |volume=47 |issue=4 |pages=574–582 |doi=10.1111/j.1365-2028.2008.00994.x}}</ref> ==Threats and conservation== The black wildebeest has been classified as a least-concern species by the International Union for Conservation of Nature in the ''[[IUCN Red List]]''. The populations of this species are on an increase. Now, more than 18,000 individuals are believed to remain, 7,000 of which are in Namibia, outside their natural range, and where it is farmed. Around 80% of the wildebeest occurs in private areas, while the other 20% is confined in protected areas. Its introduction into Namibia has been a success and numbers have increased substantially there from 150 in 1982 to 7,000 in 1992.<ref name=iucngnou>{{cite iucn |author=Vrahimis, S. |author2=Grobler, P. |author3=Brink, J. |author4=Viljoen, P. |author5=Schulze, E. |date=2017 |title=''Connochaetes gnou'' |volume=2017 |page=e.T5228A50184962 |doi=10.2305/IUCN.UK.2017-2.RLTS.T5228A50184962.en |access-date=12 November 2021}}</ref> [[File:Furs by Justin Hodge, Cape Town (08).jpg|thumb|Bag made with wildebeest skin]] The blue wildebeest has also been rated as of least concern. The population trend is stable, and their numbers are estimated to be around 1,500,000 – mainly due to the increase of the populations in [[Serengeti National Park]] (Tanzania) to 1,300,000.{{citation needed|date=July 2020}} However, the numbers of one of the subspecies, the eastern white-bearded wildebeest (''C. t. albojubatus'') have seen a steep decline.<ref name=":0" /> Population density ranges from 0.15/km². in [[Hwange National Park|Hwange]] and [[Etosha National Park]]s to 35/km² in [[Ngorongoro Conservation Area]] and Serengeti National Park.<ref name=iucntaur>{{cite iucn |author=IUCN SSC Antelope Specialist Group |date=2016 |title=''Connochaetes taurinus'' |volume=2016 |page=e.T5229A163322525 |doi=10.2305/IUCN.UK.2016-2.RLTS.T5229A163322525.en |access-date=12 November 2021}}</ref> Overland migration as a biological process requires large, connected landscapes, which are increasingly difficult to maintain, particularly over the long term, when human demands on the landscape compete. The most acute threat comes from migration barriers, such as fences and roads. In one of the more striking examples of the consequences of fence-building on terrestrial migrations, Botswanan authorities placed thousands of kilometres of fences across the Kalahari that prevented wildebeests from reaching watering holes and grazing grounds, resulting in the deaths of tens of thousands of individuals, reducing the wildebeest population to less than 10% of its previous size.<ref name=Williamson>{{cite journal|last=Williamson|first=D.T.|author2=B. Mbano |title=Wildebeest Mortality During 1983 at Lake Xau, Botswana|journal=African Journal of Ecology|year=1988|volume=26|issue=4|pages=341–344|doi=10.1111/j.1365-2028.1988.tb00987.x}}</ref> Illegal hunting is a major conservation concern in many areas, along with natural threats posed by main predators (which include lions, leopards, African hunting dogs, cheetahs and hyenas). Where the black and blue wildebeest share a common range, the two can hybridise, and this is regarded as a potential threat to the black wildebeest.<ref name=hybrid1/> ==Uses and interaction with humans== [[File:Taxidermied Lion and Blue Wildebeest, Namibia.jpg|thumb|Taxidermied [[Lion]] and [[Blue Wildebeest]], [[Namibia]]]] Wildebeest provide several useful animal products. The hide makes good-quality leather and the flesh is coarse, dry and rather hard.<ref name=nowak/> Wildebeest are killed for food, especially to make [[biltong]] in Southern Africa. This dried game meat is a delicacy and an important food item in Africa.<ref name=ulfstrand /> The meat of females is more tender than that of males, and is the most tender during the autumn season.<ref name=hoffman /> Wildebeest are a regular target for illegal meat hunters because their numbers make them easy to find. Cooks preparing the wildebeest carcass usually cut it into 11 pieces. The estimated price for wildebeest meat was about US$0.47 per {{convert|1|kg||||}} around 2008.<ref name=Nidabelema1>{{cite journal|last=Ndibalema|first=Vedasto G.|author2=Songorwa, Alexander N. |title=Illegal meat hunting in Serengeti: dynamics in consumption and preferences|journal=African Journal of Ecology|year=2008|volume=46|issue=3|pages=311–319|doi=10.1111/j.1365-2028.2007.00836.x|doi-access=free}}</ref> The silky, flowing tail of the black wildebeest is used to make [[fly-whisk]]s or ''chowries''.<ref name=nowak/> Wildebeest benefit the ecosystem by increasing soil fertility with their excreta. They are economically important for human beings, as they are a major tourist attraction. They also provide important products, such as leather, to humans.<ref name=adw>{{cite web|last=Geraci |first=G. |title=''Connochaetes taurinus'' : blue wildebeest |url=http://animaldiversity.ummz.umich.edu/accounts/Connochaetes_taurinus/ |work=University of Michigan Museum of Zoology|publisher=Animal Diversity Web|access-date=22 January 2014}}</ref> Wildebeest, however, can also have a negative impact on humans. Wild individuals can be competitors of commercial livestock, and can transmit diseases and cause epidemics among animals, particularly domestic cattle. They can also spread [[Ticks of domestic animals|ticks]], [[lungworm]]s, [[tapeworm]]s, flies, and [[Paramphistomum|paramphistome fluke]]s.<ref name=talbot/> === Cultural depictions === The wildebeest is depicted on the [[coat of arms]] of the [[Natal Province|Province of Natal]] and later [[KwaZulu-Natal]] in South Africa. Over the years, the South African authorities have issued several stamps displaying the animal and the South African Mint has struck a two-cent piece with a prancing black wildebeest.<ref name=mammal>{{cite journal|last=von Richter|first=W.|title=Connochaetes gnou|journal=[[Mammalian Species]]|year=1974|issue=50|pages=1–6}}</ref> Movies and television shows also feature wildebeests, including ''[[Khumba]]'' (Mama V), ''[[The Wild]]'' (Kazar and his minions), ''[[All Hail King Julien]]'' (Vigman Wildebeest), ''[[Phineas and Ferb]]'' (Newton the Gnu), ''[[The Great Space Coaster]]'' (newscaster Gary Gnu), and ''[[The Lion King]]'' (the wildebeest stampede that resulted in Mufasa's death). [[Michael Flanders]] wrote a humorous song called "[[The Gnu]]", which was very popular when he performed it, with [[Donald Swann]], in a revue called ''[[At the Drop of a Hat]]'', which opened in London on 31 December 1956. In the 1970s, [[United Kingdom|British]] [[tea]] brand [[Typhoo]] ran a series of [[Television]] advertisements featuring an animated anthropomorphic Gnu character. [[File:Heckert_GNU_white.svg|thumb|The wildebeest is the mascot of the open source software project [[GNU]].]] The wildebeest is the mascot of the [[GNU Project]]<ref>{{cite web|url=https://www.gnu.org/graphics/agnuhead.html|title=A GNU Head|access-date=15 July 2017|date=29 March 2015|publisher=[[GNU Project]]}}</ref> and [[GNU]] operating system. In the ''Llama Llama'' picture-book series by [[Anna Dewdney]], an anthropomorphised wildebeest named Nelly Gnu is the main character, Llama Llama's best friend, and is also featured in a title of her own, ''Nelly Gnu and Daddy Too''.<ref name="nellygnu">{{Cite book|title=Amazon - ''Nelly Gnu and Daddy Too''|isbn = 978-0670012275|last1 = Dewdney|first1 = Anna|year = 2014}}</ref> ==See also== * [[Keystone species]] ==References== {{Reflist}} {{Wiktionary|wildebeest|gnu}} {{Commons category|Connochaetes|''Connochaetes''}} {{Artiodactyla|R.2}} {{Taxonbar|from=Q7609}} [[Category:Alcelaphinae]] [[Category:Mammals of Africa]] [[Category:Extant Pleistocene first appearances]] [[Category:Herbivorous mammals]] [[Category:Articles containing video clips]]'