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The bluebuck (
Afrikaans Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans gr ...
: bloubok ) or blue antelope (''Hippotragus leucophaeus'') is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
of
antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mamm ...
that lived in South Africa until around 1800. It was smaller than the other two species in its genus ''
Hippotragus ''Hippotragus'' is a genus of antelopes which includes two living and one recently extinct species, as well as some fossil relatives. The name comes from Greek ἵππος (''híppos''), "horse", and τράγος (''trágos''), "he-goat". Foss ...
,'' the roan antelope and
sable antelope The sable antelope (''Hippotragus niger'') is an antelope which inhabits wooded savanna in East and Southern Africa, from the south of Kenya to South Africa, with a separate population in Angola. Taxonomy The sable antelope shares the genus ''Hi ...
. The bluebuck was sometimes considered a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of the roan antelope, but a genetic study has confirmed it as a distinct species. The largest mounted bluebuck specimen is tall at the
withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle a ...
. Its horns measure along the curve. The
coat A coat typically is an outer garment for the upper body as worn by either gender for warmth or fashion. Coats typically have long sleeves and are open down the front and closing by means of buttons, zippers, hook-and-loop fasteners, toggles, ...
was a uniform bluish-grey, with a pale whitish belly. The forehead was brown, darker than the face. Its mane was not as developed as in the roan and sable antelopes; its ears were shorter and blunter, not tipped with black; and it had a darker tail tuft and smaller teeth. It also lacked the contrasting black and white patterns seen on the heads of its relatives. The bluebuck was a
grazer Grazer may refer to: *grazer, an animal that grazes *Grazer, a native or inhabitant of Graz *GRAZER, the shoegaze band GRAZER Astronomy * Earth Grazer, Earth-grazing fireball that enters the Earth's atmosphere and leaves again * Mercury grazer, ...
, and may have calved where rainfall, and thus the availability of grasses, would peak. The bluebuck was confined to the southwestern
Cape A cape is a clothing accessory or a sleeveless outer garment which drapes the wearer's back, arms, and chest, and connects at the neck. History Capes were common in medieval Europe, especially when combined with a hood in the chaperon. Th ...
when encountered by Europeans, but fossil evidence and
rock paintings In archaeology, rock art is human-made markings placed on natural surfaces, typically vertical stone surfaces. A high proportion of surviving historic and prehistoric rock art is found in caves or partly enclosed rock shelters; this type also m ...
show that it originally had a larger distribution. Europeans encountered the bluebuck in the 17th century, but it was already uncommon by then, perhaps due to its preferred grassland habitat having been reduced to a range, mainly along the southern coast of South Africa. Sea level changes during the early
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
may also have contributed to its decline by disrupting the population. The first published mention of the bluebuck is from 1681, and few descriptions of the animal were written while it existed. The few 18th-century illustrations appear to have been based on stuffed specimens. Hunted by European settlers, the bluebuck became extinct around 1800; it was the first large African
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
to face extinction in historical times, followed by the quagga in 1883. Only four mounted skins remain, in museums in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration w ...
,
Stockholm Stockholm () is the capital and largest city of Sweden as well as the largest urban area in Scandinavia. Approximately 980,000 people live in the municipality, with 1.6 million in the urban area, and 2.4 million in the metropo ...
,
Vienna en, Viennese , iso_code = AT-9 , registration_plate = W , postal_code_type = Postal code , postal_code = , timezone = CET , utc_offset = +1 , timezone_DST ...
, and
Paris Paris () is the capital and most populous city of France, with an estimated population of 2,165,423 residents in 2019 in an area of more than 105 km² (41 sq mi), making it the 30th most densely populated city in the world in 2020. Si ...
, along with horns and possible bones in various museums.


Taxonomy

According to German zoologist Erna Mohr's 1967 book about the bluebuck, the 1719 account of the
Cape of Good Hope The Cape of Good Hope ( af, Kaap die Goeie Hoop ) ;''Kaap'' in isolation: pt, Cabo da Boa Esperança is a rocky headland on the Atlantic coast of the Cape Peninsula in South Africa. A common misconception is that the Cape of Good Hope is ...
published by the traveler Peter Kolbe appears to be the first publication containing a mention of the species. Kolbe also included an illustration, which Mohr believed was based on memory and notes. In 1975, Husson and Holthuis examined the original Dutch version of Kolbe's book and concluded that the illustration did not depict a bluebuck, but rather a
greater kudu The greater kudu (''Tragelaphus strepsiceros'') is a woodland antelope found throughout eastern and southern Africa. Despite occupying such widespread territory, they are sparsely populated in most areas due to declining habitat, deforestation, ...
(''Tragelaphus strepsiceros''), and that the error was due to a mistranslation into German. The first published illustration of the bluebuck is therefore instead a depiction of a horn from 1764. It has also been pointed out that the animal had already been mentioned (as "blaue Böcke") on a list of South African mammals in 1681. The Welsh naturalist
Thomas Pennant Thomas Pennant (14 June OS 172616 December 1798) was a Welsh naturalist, traveller, writer and antiquarian. He was born and lived his whole life at his family estate, Downing Hall near Whitford, Flintshire, in Wales. As a naturalist he had ...
made the next published illustration, and included an account of the antelope, calling it "blue goat", in his 1771 ''Synopsis of Quadrupeds'', based on a skin from the Cape of Good Hope purchased from Amsterdam. In 1778, a drawing by the Swiss-Dutch natural philosopher
Jean-Nicolas-Sébastien Allamand Jean-Nicolas-Sébastien Allamand ( nl, Johannes Nicolaas Sebastiaan Allamand) (18 October 1716, or according to others, 18 September 1713 – 2 March 1787) was a Swiss-Dutch natural philosopher. Life He was ...
was included in
Comte de Buffon Georges-Louis Leclerc, Comte de Buffon (; 7 September 1707 – 16 April 1788) was a French naturalist, mathematician, cosmologist, and encyclopédiste. His works influenced the next two generations of naturalists, including two prominent F ...
's ''
Histoire Naturelle The ''Histoire Naturelle, générale et particulière, avec la description du Cabinet du Roi'' (; en, Natural History, General and Particular, with a Description of the King's Cabinet, italic=yes) is an encyclopaedic collection of 36 large (qu ...
''; he called the antelope ''tzeiran'', the Siberian name for the goitered gazelle (''Gazella subgutturosa''). The illustration is widely believed to be based on the specimen in Leiden. This drawing is the first published illustration that shows the entire animal. Another record of the bluebuck appears in the travel memoirs of French explorer
François Levaillant François Levaillant (born Vaillant, later in life as Le Vaillant, ''"The Valiant"'') (6 August 1753 – 22 November 1824) was a French author, explorer, naturalist, zoological collector, and noted ornithologist. He described many new species of ...
, published in the 1780s, describing his quest to discover the land to the east of the Cape of Good Hope, "
Hottentots Holland The Hottentots Holland Mountains are part of the Cape Fold Belt in the Western Cape, South Africa. The mountain range forms a barrier between the Cape Town metropolitan area and the southern Overberg coast. The range is primarily composed of T ...
". The German zoologist Martin Lichtenstein wrote about the bluebuck in 1812, but the species was mentioned less frequently in subsequent literature. In 1776, the German zoologist
Peter Simon Pallas Peter Simon Pallas FRS FRSE (22 September 1741 – 8 September 1811) was a Prussian zoologist and botanist who worked in Russia between 1767 and 1810. Life and work Peter Simon Pallas was born in Berlin, the son of Professor of Surgery ...
formally described the bluebuck as ''Antilope leucophaeus''. In 1853, the Dutch zoologist
Coenraad Jacob Temminck Coenraad Jacob Temminck (; 31 March 1778 – 30 January 1858) was a Dutch aristocrat, zoologist and museum director. Biography Coenraad Jacob Temminck was born on 31 March 1778 in Amsterdam in the Dutch Republic. From his father, Jacob Temmi ...
stated that the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
was an adult male skin now in the
Naturalis Biodiversity Center Naturalis Biodiversity Center ( nl, Nederlands Centrum voor Biodiversiteit Naturalis) is a national museum of natural history and a research center on biodiversity in Leiden, Netherlands. It was named the European Museum of the Year 2021. ...
in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration w ...
(formerly
Rijksmuseum van Natuurlijke Historie The Rijksmuseum van Natuurlijke Historie (National Museum of Natural History) was a museum on the Rapenburg in Leiden, the Netherlands. It was founded in 1820 by Royal Decree from a merger of several existing collections. This happened on the initi ...
), collected in
Swellendam Swellendam is the fifth oldest town in South Africa (after Cape Town, Stellenbosch, Simon's Town, and Paarl), a town with 17,537 inhabitants situated in the Western Cape province. The town has over 50 provincial heritage sites, most of them ...
and present in
Haarlem Haarlem (; predecessor of ''Harlem'' in English) is a city and municipality in the Netherlands. It is the capital of the province of North Holland. Haarlem is situated at the northern edge of the Randstad, one of the most populated metropoli ...
before 1776. It has been questioned whether this was actually the type specimen, but in 1969, the Dutch zoologists Antonius M. Husson and
Lipke Holthuis Lipke Bijdeley Holthuis (21 April 1921 – 7 March 2008) was a Dutch carcinologist, considered one of the "undisputed greats" of carcinology, and "the greatest carcinologist of our time". Holthuis was born in Probolinggo, East Java and obtained ...
selected it as the
lectotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
of a syntype series, as Pallas may have based his description on multiple specimens. In 1846, the Swedish zoologist
Carl Jakob Sundevall Carl Jakob Sundevall (22 October 1801, Högestad – 2 February 1875) was a Swedish zoologist. Sundevall studied at Lund University, where he became a Ph.D. in 1823. After traveling to East Asia, he studied medicine, graduating as Doctor of Me ...
moved the bluebuck and its closest relatives to the genus ''
Hippotragus ''Hippotragus'' is a genus of antelopes which includes two living and one recently extinct species, as well as some fossil relatives. The name comes from Greek ἵππος (''híppos''), "horse", and τράγος (''trágos''), "he-goat". Foss ...
''; he had originally named this genus for the roan antelope (''H. equinus'') in 1845. This revision was commonly accepted by other writers, such as the British zoologists
Philip Sclater Philip Lutley Sclater (4 November 1829 – 27 June 1913) was an England, English lawyer and zoologist. In zoology, he was an expert ornithologist, and identified the main zoogeographic regions of the world. He was Secretary of the Zoological ...
and
Oldfield Thomas Michael Rogers Oldfield Thomas (21 February 1858 – 16 June 1929) was a British zoologist. Career Thomas worked at the Natural History Museum on mammals, describing about 2,000 new species and subspecies for the first time. He was appo ...
, who restricted the genus ''Antilope'' to the
blackbuck The blackbuck (''Antilope cervicapra''), also known as the Indian antelope, is an antelope native to India and Nepal. It inhabits grassy plains and lightly forested areas with perennial water sources. It stands up to high at the shoulder. Ma ...
(''A. cervicapra'') in 1899. In 1914, the name ''Hippotragus'' was submitted for
conservation Conservation is the preservation or efficient use of resources, or the conservation of various quantities under physical laws. Conservation may also refer to: Environment and natural resources * Nature conservation, the protection and manageme ...
(so older, unused genus names could be suppressed) to the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
(ICZN) with the bluebuck as the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
. However, the original 1845 naming of the genus with the roan antelope as a single species was overlooked and later suppressed by the ICZN, leading to some taxonomic confusion. In 2001, the British ecologist Peter J. Grubb proposed that the ICZN should rescind its suppression of the 1845 naming and make the roan antelope the type species of ''Hippotragus'', since too little is known about the bluebuck for it to be a reliable type species. This was accepted by the commission in 2003. The common names "bluebuck" and "blue antelope" are English for the original
Afrikaans Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans gr ...
name "blaubok" . The name is a
compound Compound may refer to: Architecture and built environments * Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall ** Compound (fortification), a version of the above fortified with defensive struc ...
of ''blauw'' and ''bok'' ("male antelope" or "male goat"). Variants of this name include "blaawwbok" and "blawebock". The generic name ''Hippotragus'' is
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
for "horse-goat", while the specific name ''leucophaeus'' is a fusion of two Greek words: ''leukos'' ("white") and ''phaios'' ("brilliant").


Preserved specimens

Four mounted skins of the bluebuck remain: the adult male in Leiden, a young male at the Zoological Museum of Stockholm, an adult female in the Vienna Museum of Natural History, and an adult male in the Museum of Natural History in Paris. A mounted skin was housed in the Zoological Museum in Uppsala until the 19th century, but now only the horns remain. There are also records of a skin in Haarlem, but its current whereabouts are unknown. Several of these skins have been identified in various 18th-century illustrations. Skeletal remains have been found in both archaeological and palaeontological contexts. In 2021, the German geneticist Elisabeth Hempel and colleagues examined sixteen bluebuck specimens to resolve their identities, and found that only four of them were bluebucks. The skins in Stockholm and Vienna were confirmed as belonging to bluebucks, as were skull fragments in Leiden that may belong to the lectotype specimen, and the horns in Uppsala. Four assigned skulls (those in Glasgow, Leiden, Paris and Berlin) were shown to belong to either sable or roan antelopes, as were two pairs of horns (in Cape Town and St. Andrews). As a result, the bluebuck is rarer in museum collections than previously though, and no complete skulls are known of the species. The researchers pointed out that there are four more potential specimens that could be confirmed through testing; two skulls in Berlin, a pair of horns in London, and either a skull or pair of horns in Brussels.


Evolution

Based on studies of morphology, the bluebuck has historically been classified as either a distinct species or as a
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of the roan antelope. After its extinction, some 19th-century naturalists began to doubt its validity as a species, with some believing the museum specimens to be small or immature roan antelopes, and both species were lumped together under the name ''A.leucophaeus'' by the English zoologist
George Robert Gray George Robert Gray FRS (8 July 1808 – 6 May 1872) was an English zoologist and author, and head of the ornithological section of the British Museum, now the Natural History Museum, in London for forty-one years. He was the younger brother ...
in 1821. The Austrian zoologist Franz Friedrich Kohl pointed out the distinct features of the bluebuck in 1866, followed by Sclater and Thomas, who rejected the synonymy in 1899. In 1974, the American biologist Richard G. Klein showed (based on fossils) that the bluebuck and roan antelope occurred sympatrically on the coastal plain of the southwestern Cape from Oakhurst to Uniondale during the early
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
, supporting their status as separate species. In 1996, an analysis of
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
extracted from the bluebuck specimen in Vienna by South African biologist Terence J. Robinson's and colleagues showed that it was outside the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
containing the roan and sable antelopes. The study therefore concluded that the bluebuck is a distinct species, and not merely a subspecies of the roan antelope as was supposed. In 2017, a reconstruction of the entire bluebuck
mitogenome Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
by Portuguese biologists Gonçalo Espregueira Themudo and Paula F. Campos, based on bone powder extracted from the horns in Uppsala, contradicted the 1996 results. This study instead placed the bluebuck as a
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to the sable antelope, with the roan antelope as an outgroup. The bluebuck and sable antelope diverged from each other 2.8 million years ago, while the roan antelope diverged from both of them 4.17 million years ago. Africa was going through climatic oscillations between 3.5 and 2 million years ago, and during a colder period, the ancestors of the sable antelope and bluebuck may have been separated, and the population in southern Africa eventually became a new species. The
cladograms A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below shows the placement of the bluebuck according to the 1996 and 2017 DNA studies: Robinson and colleagues, 1996: Themudo and Campos, 2017: Based on their larger sample of specimens, Hempel and colleagues also found the bluebuck genetically closest to the sable antelope in 2021. This was confirmed by a 2022 study by Hempel and colleagues, which managed to sample DNA from a fossil bluebuck specimen for the first time, at 9,800–9,300 years old the oldest paleogenome from Africa.


Description

The adult male bluebuck in Leiden is tall at the
withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle a ...
, and is possibly the largest known specimen. According to Sclater and Thomas, the tallest specimen is the one in Paris, a male that stands at the shoulder; the specimen in Vienna, on the other hand, is the shortest, a tall female. The bluebuck was notably smaller than the roan and sable antelopes, and therefore the smallest member of its genus. The
coat A coat typically is an outer garment for the upper body as worn by either gender for warmth or fashion. Coats typically have long sleeves and are open down the front and closing by means of buttons, zippers, hook-and-loop fasteners, toggles, ...
was a uniform bluish-grey, with a pale whitish belly, which was not contrasted on the flanks. The limbs had a faint dark line along their front surface. The forehead was brown, darker than the face, and the upper lip and patch in front of the eyes were lighter than the body. The neck-mane was directed forwards and not as developed as in the roan and sable antelopes, and the throat-mane was almost absent. Other differences between the bluebuck and its extant relatives included its shorter and blunter ears not tipped with black, a darker tail tuft (though little darker than its general colour), and smaller teeth. The bluebuck also lacked the contrasting black and white patterns seen on the heads of its relatives. As the old skins are presumably faded, it may be difficult to reconstruct the original colour of the bluebuck from them. Pennant observed that the eyes had white patches below them and the underbelly was white; the coat was a "fine blue" in living specimens, while it changed to "bluish grey, with a mixture of white" in dead animals. He also suggested that the length of the bluebuck's hair and the morphology of its horns formed a link between antelopes and
goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...
. He went on to describe the ears as pointed and over long and the tail as long, terminating in a long tuft. The horns of the bluebuck were significantly shorter and thinner than those of the roan antelope, but perhaps proportionally longer. The horns of the Leiden specimen measure along the curve. Pennant gave the horn length as . He added that the horns, sharp and curving backward, consist of twenty rings. Text Archive The horns of the bluebuck appear to have hollow pedicles (bony structures from which the horns emerge).


Behaviour and ecology

The bluebuck, as Klein puts it, became extinct before "qualified scientists could make observations on live specimens". According to historical accounts, the bluebuck formed groups of up to 20 individuals. Similarities to the roan and the sable antelopes in terms of dental morphology make it highly probable that the bluebuck was predominantly a selective
grazer Grazer may refer to: *grazer, an animal that grazes *Grazer, a native or inhabitant of Graz *GRAZER, the shoegaze band GRAZER Astronomy * Earth Grazer, Earth-grazing fireball that enters the Earth's atmosphere and leaves again * Mercury grazer, ...
, and fed mainly on grasses. The row of
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s was longer than in others of the genus, implying the presence of
dicot The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, ...
s in the diet. A 2013 study by the Australian palaeontologist J. Tyler Faith and colleagues noted the scarcity of morphological evidence to show that the bluebuck could have survived the summers in the western margin of the Cape Floristic Region (CFR), when the grasses are neither palatable nor nutritious. This might have induced a west-to-east migration, because the eastern margin receives rainfall throughout the year while precipitation in the western margin is limited to winter. An 18th-century account suggests that females might have left their newborn calves in isolation and returned regularly to suckle them until the calves were old enough to join herds, which is similar to the behaviour of roan and sable antelopes. Akin to other grazing antelopes, the bluebuck probably calved mainly where rainfall, and thus the availability of grasses, peaked. Such locations could be the western margin of the CFR during winter and the eastern margin of the CFR during summer. Faith and colleagues found that the occurrence of juveniles in bluebuck fossils decreases linearly from the west to the east, indicating that most births took place in the western CFR; due to the preference for rainfall, it may be further assumed that most births occurred during winter, when the western CFR receives most of its rainfall. The annual west-to-east migration would have followed in summer, consistent with the greater number of older juveniles in the east that would have joined herds. Juvenile fossils also occur in other places across the range, but appear to be concentrated in the western CFR.


Distribution and habitat

Endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
to South Africa, the bluebuck was confined to the southwestern
Cape A cape is a clothing accessory or a sleeveless outer garment which drapes the wearer's back, arms, and chest, and connects at the neck. History Capes were common in medieval Europe, especially when combined with a hood in the chaperon. Th ...
. A 2003 study estimated the expanse of the historic range of the bluebuck at , mainly along the southern coast of South Africa; fossils, however, have been discovered in a broader area that includes the southern and western CFR and even the highlands of
Lesotho Lesotho ( ), officially the Kingdom of Lesotho, is a country landlocked as an enclave in South Africa. It is situated in the Maloti Mountains and contains the highest mountains in Southern Africa. It has an area of over and has a population ...
. Historical records give a rough estimate of its range. On 20January 1774, Swedish naturalist
Carl Peter Thunberg Carl Peter Thunberg, also known as Karl Peter von Thunberg, Carl Pehr Thunberg, or Carl Per Thunberg (11 November 1743 – 8 August 1828), was a Swedish naturalist and an "apostle" of Carl Linnaeus. After studying under Linnaeus at Uppsala U ...
recorded a sighting in Tigerhoek,
Mpumalanga Mpumalanga () is a province of South Africa. The name means "East", or literally "The Place Where the Sun Rises" in the Swazi, Xhosa, Ndebele and Zulu languages. Mpumalanga lies in eastern South Africa, bordering Eswatini and Mozambique. ...
. In March or April 1783, Levalliant claimed to have witnessed two specimens in Soetemelksvlei,
Western Cape The Western Cape is a province of South Africa, situated on the south-western coast of the country. It is the fourth largest of the nine provinces with an area of , and the third most populous, with an estimated 7 million inhabitants in 202 ...
. Based on these notes, a 2009 study by the South African zoologist Graham I. H. Kerley and colleagues estimated the range of the bluebuck to be limited within a triangular area in the Western Cape, bounded by Caledon to the west, Swellendam to the northeast and
Bredasdorp Bredasdorp is a town in the Southern Overberg region of the Western Cape, South Africa, and the main economic and service hub of that region. It lies on the northern edge of the Agulhas Plain, about south-east of Cape Town and north of Cape Agu ...
to the south.
Rock paintings In archaeology, rock art is human-made markings placed on natural surfaces, typically vertical stone surfaces. A high proportion of surviving historic and prehistoric rock art is found in caves or partly enclosed rock shelters; this type also m ...
in the Caledon river valley of the Free State province in eastern South Africa have been identified as bluebucks, which also confirms the once wider distribution of the species. In 1974, Klein studied the fossils of ''Hippotragus'' species in South Africa. Most of these were found to represent the bluebuck and the roan antelope. The fossil record suggested that the bluebuck occurred in large numbers during the last glacial period (nearly 0.1million years ago), and was more common than sympatric antelopes. The bluebuck could adapt to more open habitats than could the roan antelope, a notable point of difference between these species. Fossils of the bluebuck have been found in the Klaises River and the
Nelson Bay Cave Nelson Bay Cave also known as Wagenaar's Cave is a Stone Age archaeological site located in the Robberg Nature Reserve on the Robberg Peninsula and facing Nelson's Bay near Plettenberg Bay in South Africa, and showing evidence of human occupat ...
(near Plettenberg Bay) and Swartklip (to the west of the
Hottentots Holland mountains The Hottentots Holland Mountains are part of the Cape Fold Belt in the Western Cape, South Africa. The mountain range forms a barrier between the Cape Town metropolitan area and the southern Overberg coast. The range is primarily composed of Ta ...
). Faith and colleagues noted that the western and southern CFR were separated by
biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
barriers, such as the
Cape Fold Belt The Cape Fold Belt is a fold and thrust belt of late Paleozoic age, which affected the sequence of sedimentary rock layers of the Cape Supergroup in the southwestern corner of South Africa. It was originally continuous with the Ventana Mount ...
and
afromontane The Afromontane regions are subregions of the Afrotropical realm, one of the Earth's eight biogeographic realms, covering the plant and animal species found in the mountains of Africa and the southern Arabian Peninsula. The Afromontane regions of ...
forests. A 2011 study suggested that low sea levels facilitated migrations for large mammals; therefore the rise in sea levels with the beginning of the Holocene would have led to fragmented bluebuck populations and distanced many populations from the western coast (fossils dating to this period are scarce in the western coast but have been recorded from the southern coast). Thus, a mass extinction could have taken place, leaving behind mainly the populations that remained in the resource-rich western CFR. The causes of the drastic decline in bluebuck populations just before the 15th and 16th centuries have not been investigated; competition with livestock and habitat deterioration could have been major factors in its depletion. Faith and colleagues further suggested that the bluebuck, being a grazer, probably favoured grassland habitats. This hypothesis is supported by fossil evidencebluebuck fossils appear in significant numbers along with those of grassland antelopes. Kerley and colleagues suggested that the bluebuck frequented grasslands and shunned wooded areas and thickets. In a 1976 study of fossils in the
Southern Cape The Western Cape is a province of South Africa, situated on the south-western coast of the country. It is the fourth largest of the nine provinces with an area of , and the third most populous, with an estimated 7 million inhabitants in 2020 ...
, Klein observed that the bluebuck's habitat preferences were similar to those of the African buffalo (''Syncerus caffer'') and the
reedbuck Reedbuck is a common name for Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it ...
s (''Redunca'').


Relationship with humans


Extinction

Due to the small range of the bluebuck at the time of European settlement of the Cape region in the 17th and 18th centuries compared to the much wider area evidenced by fossil remains, it is thought the species was already in decline before this time. The bluebuck was the sole species of ''Hippotragus'' in the region until 70,00035,000 years ago, but the roan antelope appears to have become predominant about 11,000 years ago. This might have coincided with grasslands being replaced by, for example, brush and forest, thereby reducing what is presumed to be the preferred habitat of the bluebuck, the grasslands. The results of the sea level changes in the early Holocene may also have played a role in the decline of the species, and left only the southern population to survive into historical times. Hempel and colleagues found a low level of genetic diversity between the four confirmed bluebuck specimens, which confirms its population size was low by the time of European colonisation. The bluebuck was hunted to extinction by European settlers; in 1774 Thunberg noted that it was becoming increasingly rare. The German biologist
Hinrich Lichtenstein Martin Hinrich Carl Lichtenstein (10 January 1780 – 2 September 1857) was a German physician, explorer, botanist and zoologist. Biography Born in Hamburg, Lichtenstein was the son of Anton August Heinrich Lichtenstein. He studied medicine ...
claimed that the last bluebuck had been shot in 1799 or 1800. The bluebuck is the first historically recorded large African mammal to become extinct, followed by the quagga (''Equus quagga quagga''), which died out in 1883. Around the time of its extinction, the bluebuck occurred in what would be known as the
Overberg __NOTOC__ Overberg is a region in South Africa to the east of Cape Town beyond the Hottentots-Holland mountains. It lies along the Western Cape Province's south coast between the Cape Peninsula and the region known as the Garden Route in the ...
region (Western Cape), probably concentrated in Swellendam. In 1990, the South African zoologist Brian D. Colohan argued that an 1853 eyewitness report of a "bastard gemsbok" seen near
Bethlehem, Free State Bethlehem is a town in the eastern Free State province of South Africa that is situated on the Liebenbergs river (also called Liebenbergs Vlei) along a fertile valley just north of the Rooiberg Mountains on the N5 road. It is the fastest growin ...
, actually referred to a bluebuck, 50 years after the last individuals in Swellendam were shot. The
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biolo ...
accepts Lichtenstein's dates of extinction. The related roan and sable antelopes have also disappeared from much of their former range.


Cultural significance

The bluebuck rock paintings from the Caledon river valley have been attributed to
Bushmen The San peoples (also Saan), or Bushmen, are members of various Khoe languages, Khoe, Tuu languages, Tuu, or Kxʼa languages, Kxʼa-speaking indigenous hunter-gatherer cultures that are the Indigenous peoples of Africa, first cultures of Sout ...
. They show six antelopes facing a man, and were supposedly inspired by
shamanic Shamanism is a religious practice that involves a practitioner (shaman) interacting with what they believe to be a spirit world through altered states of consciousness, such as trance. The goal of this is usually to direct spirits or spiri ...
trance Trance is a state of semi-consciousness in which a person is not self-aware and is either altogether unresponsive to external stimuli (but nevertheless capable of pursuing and realizing an aim) or is selectively responsive in following the dir ...
; they may depict a Bushman visiting the spirit-world through a tunnel. The Bushmen possibly believed that the bluebuck had a supernatural potency, like other animals in their environment. The animals in the paintings are similar in proportion to the reedbuck, but the large ears, horns, and lack of a mane rule out species other than the bluebuck. A South African fable, ''The Story of the Hare'', mentions a bluebuck (referred to as ''inputi'') that, among other animals, is appointed to guard a
kraal Kraal (also spelled ''craal'' or ''kraul'') is an Afrikaans and Dutch word, also used in South African English, for an enclosure for cattle or other livestock, located within a Southern African settlement or village surrounded by a fence of t ...
. The bluebuck is also mentioned in French novelist
Jules Verne Jules Gabriel Verne (;''Longman Pronunciation Dictionary''. ; 8 February 1828 – 24 March 1905) was a French novelist, poet, and playwright. His collaboration with the publisher Pierre-Jules Hetzel led to the creation of the '' Voyages extra ...
's ''
Five Weeks in a Balloon ''Five Weeks in a Balloon, or, A Journey of Discovery by Three Englishmen in Africa'' (french: Cinq semaines en ballon) is an adventure novel by Jules Verne, published in 1863. It is the first novel in which he perfected the "ingredients" of hi ...
'' (1863); the animal is described as a "superb animal of a pale-bluish colour shading upon the gray, but with the belly and the insides of the legs as white as the driven snow".


References


Citations


Sources

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External links


Interactive 3D scan of the Vienna specimen at Sketchfab
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Blue Antelope Correspondences
{{Authority control Grazing antelopes Mammals of South Africa Endemic fauna of South Africa Extinct mammals of Africa Mammal extinctions since 1500 Species made extinct by human activities Mammals described in 1776 Taxa named by Peter Simon Pallas Species endangered by habitat loss