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Falkland Islands wolf

Falkland Islands wolf[1]
Illustration by John Gerrard Keulemans (1842–1912)

Extinct  (1876)  (IUCN 3.1)[2]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Tribe: Canini
Genus: Dusicyon
Species: D. australis
Binomial name
Dusicyon australis
(Kerr, 1792)
Location of the Falkland Islands

The Falkland Islands wolf (Dusicyon australis), also known as the warrah and occasionally as the Falkland Islands dog, Falkland Islands fox, or Antarctic wolf, was the only native land mammal of the Falkland Islands. This endemic canid became extinct in 1876, the first known canid to have become extinct in historical times. It was the only modern species in the genus Dusicyon.

Traditionally it had been supposed that the most closely related genus was Lycalopex, including the culpeo, which has been introduced to the Falkland Islands in modern times. However, in 2009, a cladistic analysis of DNA identified the Falkland Island wolf's closest living relative as the maned wolf (Chrysocyon brachyurus)—an unusually long-legged, fox-like South American canid, from which it separated about 6.7 million years ago.[3]

The Falkland Islands wolf existed on both West and East Falkland, but Charles Darwin was uncertain if they were differentiated varieties.[4] Its fur had a tawny colour and the tip of the tail was white. Its diet is unknown, but, due to the absence of native rodents on the Falklands, probably consisted of ground-nesting birds such as geese and penguins, grubs and insects, as well as seashore scavenging.[5] It has sometimes been said that it may have lived in burrows.[6]

Contents

  • History 1
    • Darwin's description 1.1
  • Biogeography and evolution 2
    • Darwin's comments 2.1
    • Related species 2.2
    • Biogeographical isolation on the Falklands 2.3
  • Commemorations 3
  • References 4
    • Footnotes 4.1
  • External links 5
  • External links 6

History

Illustration of "Dusicyon australis" from Zoology of the Voyage of H.M.S. Beagle

The first recorded sighting was by Capt. John Strong in 1690.[7] Captain Strong took one of the animals on his ship, but during the voyage back to Europe the creature became frightened by the firing of the ship's cannon and jumped overboard.[7][8] Louis Antoine de Bougainville, who established the first settlement in the Falkland Islands termed it a loup-renard ("fox-wolf"). The name "warrah" is a corruption of the term aguará (meaning "fox" in Guaraní, a Native American language), because of its similarity to the maned wolf ("aguará guazú".) The warrah's Latin name translates into "foolish dog of the south", because of its apparent lack of fear of people.

Engraving of "Antarctic wolf", The Illustrated London News (1873).

When Charles Darwin visited the islands in 1833 he found the species present in both West and East Falkland and tame. However, at the time of his visit the animal was already very rare on East Falkland, and even on West Falkland its numbers were declining rapidly. By 1865, it was no longer found on the eastern part of East Falkland. He predicted that the animal would join the dodo among the extinct within "a very few years." It was hunted for its valuable fur, and the settlers, regarding the wolf as a threat to their sheep, poisoned it.[9] However, the belief that Falkland Islands wolf was a threat to sheep was probably due to the sheep mistaking the Falkland Islands wolves for dogs (especially at night), and, in terror, the sheep ran into bogs and swamps, where they became lost. [10] There were no forests for the animal to hide in, and it had no fear of man; it was possible to lure the animal with a chunk of meat held in one hand, and kill it with a knife or stick held in the other. However, it would defend itself occasionally if it needed to, as Admiral George Grey noted when they landed on West Falkland at Port Edgar on December 17, 1836:

I landed in the creek and had hardly put a foot on shore, when one of the foxes of the country was chased by Pilot. I ran up as they were fighting and came to the poor dog's assistance who had nearly met his match, and a rifle ball soon settled the business, but the Pilot had received a terrible bite in the leg.

A live wolf was taken to London Zoo, England in 1868.[11] Another "Antarctic wolf" arrived in 1870. Neither animal survived long. Only a dozen or so museum specimens exist today.[3]

In 1880, after the animal became extinct, Thomas Huxley classified it as related to the coyote. In 1914, Oldfield Thomas moved it to the genus Dusicyon, with the culpeo and other South American foxes. (These other canids have since been removed to Lycalopex.)

Darwin's description

Darwin writing about his 1834 visit to the Falklands in his Journal and Remarks (The Voyage of the Beagle) has the following to say of Canis antarcticus:

Biogeography and evolution

Darwin's comments

When organising his notes on the last stage of the Beagle expedition, Darwin wrote of his growing suspicions that the differences between the various Galápagos Islands mockingbirds and tortoises, as well as the possible dissimilarity of West Falkland and East Falkland Islands wolves, were but variants that differed depending on which island they came from:

The term "would" was added after the words had been written, suggesting a cautious qualification from his initial bold statement.[13] He later wrote that such facts "seemed to me to throw some light on the origin of species".[14]

Related species

A recent DNA analysis,[15] and a study of comparative brain anatomy [16] suggest that the closest living relative of the Falkland Islands wolf is the South American maned wolf.[3] Their most recent common ancestor was estimated to have lived some 6 million years ago and was close to the most recent common ancestor of all South American canids, Eucyon or a close relative. It would seem that the lineages of the maned wolf and the Falkland Islands wolf separated in North America; canids did not appear in South America until roughly 3 million years ago in a paleozoogeographical event called the Great American Biotic Interchange, in which the continents of North and South America were newly connected by the formation of the Isthmus of Panama. However, no fossil from North America can be assigned to the Falkland Islands wolf or its immediate ancestors.[3]

Dusicyon avus, known from fossils from southern South America as recent as 1600 years ago, was a close mainland relative of the Falkland Islands wolf.[17]

Biogeographical isolation on the Falklands

The route by which the Falkland Islands wolf was established in the islands was until recently a puzzle as the islands have never been connected to the mainland[18] and there are no other native land mammals.[19] No other oceanic island as remote as the Falklands has a native canid; the island fox of California, and Darwin's fox of Chile both inhabit islands much closer to a continent.

Berta and other authors suggest that it was unlikely that the wolf's ancestors could have survived the last Ice Age on the Falklands and they must therefore have arrived later, within the last ten thousand years, crossing a wide expanse of the South Atlantic.[20] Its close relative, Dusicyon avus, did survive in South America until a few thousand years ago, but swimming such a distance or even drifting on a floating log would appear effectively impossible for the wolf.[21] Some speculated that the species had originally arrived with natives visiting the islands after the settlement of the Americas, having been kept by them in a semi-domesticated state.[22] There is however no definite evidence that humans ever visited the Falklands before the Age of Discovery (the islands are not visible from the mainland, and traveling to the Falklands in open canoes would have been very difficult and extremely dangerous). Nor is there any evidence that the ancestors of the wolf were ever domesticated.

DNA of the extinct mainland relative, D. avus, analyzed in 2013 suggests that its genetic history diverged from the Falkland Islands wolf only some 16,000 years ago, during the last glacial phase.[21] This is strong evidence that the ancestors of the wolf were isolated on the islands only since the last glacial maximum. A 2009 analysis of mitochondrial DNA from five museum specimens of the Falkland Islands wolf indicated that they had multiple mitochondrial haplotypes whose most recent common ancestor lived about 330,000 years ago,[15] giving some idea of the genetic diversity of the founding population.

An Ice Age land bridge or ice connection between the Falkland Islands and South America, enabling the species' ancestors to traverse the gap, has long been suggested. There was never a true land bridge between the islands and South America, but submarine terraces have been found on the Argentine coastal shelf, formed by low sea-stands during the last glacial phase.[21] This suggests that there was a shallow strait as narrow as 20 km, which may have frozen completely at times.[21] It seems likely that the founding population of the wolf crossed on this ice bridge during the last Ice Age.[21][15] This would explain the absence of other mainland mammals on the islands; an ice crossing is much more difficult than crossing a land bridge.[21]

Commemorations

Locations:

Other:

  • The fifty pence piece features the warrah
  • The Warrah, a Falkland Islands' conservation magazine [2]
  • "Warrah Knitwear", a company formerly based in Fox Bay.
  • "Warrah Design", a company based in Fox Bay
  • The warrah was featured on a 2009 Falkland Islands commemorative stamp issued for the Charles Darwin bicentennial.
  • "The Warrah", an episode from Radio 4, From Fact to Fiction.[23]
  • The warrah also features on a Falkland Islands Football League pin badge.

References

  • History of a Voyage to the Malouin Islands (1764), Dom Pernetty.
  • Voyage Round the World, Louis Antoine de Bougainville
  • "Antarctic Wolf" (p. 441) in Harmsworth Natural History, 1910, R. Lydekker, Sir H. Johnston and Prof J. R. Ainsworth-Davis

Footnotes

  1. ^ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–628.  
  2. ^ IUCN SSC Canid Specialist Group (2008). Dusicyon australis. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 5 January 2008.
  3. ^ a b c d "New Clues To Extinct Falklands Wolf Mystery". EurekAlert. Science Daily. 2009-11-03. Retrieved 2011-09-03. 
  4. ^ a b  
  5. ^ Glover Morrill Allen (June 1942). Extinct and vanishing mammals of the Western Hemisphere: with the marine species of all the oceans. Cooper Square Publishers.  
  6. ^ Massam, Steve. "The Evans Warrah, The Identification". Falkland Islands Museum and National Trust. Retrieved 2011-09-04. 
  7. ^ a b "The First Sighting". Falkland Islands Museum and National Trust. Retrieved 2011-09-08. 
  8. ^ Rosamond Wolff Purcell (20 May 1999). Swift as a shadow: extinct and endangered animals. Houghton Mifflin Co.  
  9. ^ "Dusicyon australis". International Union for Conservation of Nature. 4 September 2011. Retrieved 2011-09-04. 
  10. ^ http://www.falklands-museum.com/demise-of-the-warrah/190-renshaws-comment-on-extinction
  11. ^ "A Falklands Wolf in London". Falkland Islands Museum and National Trust. Retrieved 2011-09-08. 
  12. ^ Charles Darwin (2000). Richard Keynes, ed. Charles Darwin's zoology notes & specimen lists from H.M.S. Beagle. Cambridge University Press.  
  13. ^  
  14. ^  
  15. ^ a b c Slater, G. J.; Thalmann, O.; Leonard, J. A.; Schweizer, R. M.; Koepfli, K.-P.; Pollinger, J. P.; Rawlence, N. J.; Austin, J. J.; Cooper, A.; Wayne, R. K. (2009-11-03). "Evolutionary history of the Falklands wolf" (PDF). Current Biology 19 (20): R937–R938.  
  16. ^ Lyras, G. A.; Van Der Geer, A. A. E. (2003-08-04). "External brain anatomy in relation to the phylogeny of Caninae (Carnivora: Canidae)" (PDF). Zoological Journal of the Linnean Society 138 (4): 505–522.  
  17. ^ Francisco Prevosti, Fernando Santiago, Luciano Prates, Mónica Salemme, and Fabiana Martin (2010), (Carnivora, Canidae) during the late Holocene"Dusicyon avus"Constraining the time of extinction of the South American fox (PDF), Geophysical Research Abstracts 12 (EGU2010-577-1)  EGU General Assembly 2010 
  18. ^ Cione, A. L.; Tonni, E. P.; Soibelzon, L. (2003). "The Broken Zig-Zag: Late Cenozoic large mammal and tortoise extinction in South America" (PDF). Rev. Mus. Argentino Cienc. Nat., n.s. 5 (1): 1–19.   see figure 3
  19. ^ But even though the study has gone some way to clarifying the evolutionary origins of the Falklands wolf, the species still represents a great bio-geographical conundrum because the Falklands have never been connected to the South American mainland and no other land mammals – not even small rodents – have managed to live there. Connor, Steve (2009-11-03). "How scientists cracked puzzle of the Falklands wolf". The Independent (Mercopress). Retrieved 2011-09-01. 
  20. ^ Berta, A. (1987). Origin, diversification, and zoogeography of the South American Canidae. Fieldiana: Zoology, 39, 455–471
  21. ^ a b c d e f Cooper, A.; Mena, F.; Austin, J. J.; Soubrier, J.; Prevosti, F.; Prates, L.; Trejo, V. (2013). "The origin of the enigmatic Falkland Islands wolf". Nature Communications 4: 1552.  
  22. ^ Clutton-Brock, J.; Corbet, G.G.; Hills, M. (1976). "A review of the family Canidae, with a classification by numerical methods". Bull. Brit. Mus. Nat. Hist. 29 (3): 117–199. Retrieved 2013-03-10. 
  23. ^ http://www.bbc.co.uk/programmes/b01c6kr3

External links

  • ("Falkland Islands")  
  • The warrah

External links

  • Falklands Conservation
  • Site, includes three pictures of the Warrah
  • Darwin's Wolf Mystery Solved
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