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New World vulture

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New World vultures
Temporal range: PaleoceneHolocene, 64.5–0 Ma
Large black bird with red, unfeathered head, perched on a rock and sitting looking to right of cameraman
Turkey vulture
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Accipitriformes
Family: Cathartidae
Lafresnaye, 1839
Genera

Cathartes
Coragyps
Gymnogyps
Sarcoramphus
Vultur

Approximate Cathartidae range map
  Summer-only range of turkey vulture 
  At least one species present year-round
Synonyms
  • Vulturidae Illiger, 1811

Cathartidae, known commonly as New World vultures or condors, are a family of birds of prey consisting of seven extant species in five genera. It includes five extant vultures and two extant condors found in the Americas. They are known as "New World" vultures to distinguish them from Old World vultures, with which the Cathartidae does not form a single clade despite the two being similar in appearance and behavior as a result of convergent evolution.

Like other vultures, New World vultures are scavengers, having evolved to feed off of the carcasses of dead animals without any notable ill effects. Some species of New World vulture have a good sense of smell, whereas Old World vultures find carcasses exclusively by sight. Other adaptations shared by both Old and New World vultures include a bald head, devoid of feathers to prevent rotting matter from accumulating while feeding, and an extremely disease-resistant digestive system to protect against scavenging-related germs.

Taxonomy and systematics

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Sketch of head showing an unobstructed view through the nostrils
A pervious nostril is typical of the family

The family Cathartidae was introduced (as the subfamily Cathartinae) by the French ornithologist Frédéric de Lafresnaye in 1839.[1][2] The New World vultures comprise seven species in five genera, being Coragyps, Cathartes, Gymnogyps, Sarcoramphus, and Vultur. Of these, only Cathartes is not monotypic.[3] The family's scientific name, Cathartidae, comes from cathartes, Greek for "purifier".[4] Although New World vultures and Old World vultures are not very closely related, they share many resemblances because of convergent evolution.[5] Phylogenetic analyses including all Cathartidae species found two primary clades. The first consists of black vultures (Coragyps atratus) together with the three Cathartes species (lesser yellow-headed vultures (C. burrovianus), greater yellow-headed vultures (C. melambrotus), and turkey vultures (C. aura)), while the second consists of king vultures (Sarcoramphus papa), California condors (Gymnogyps californianus) and Andean condors (Vultur gryphus).[6]

New World vultures were traditionally placed in a family of their own in the Falconiformes.[7] However, in the late 20th century some ornithologists argued that they are more closely related to storks on the basis of karyotype,[8] morphological,[9] and behavioral[10] data. Thus some authorities placed them in the Ciconiiformes with storks and herons; Sibley and Monroe (1990) even considered them a subfamily of the storks. This was criticized,[11][12] and an early DNA sequence study[13] was based on erroneous data and subsequently retracted.[14][15][16] There was then an attempt to raise the New World vultures to the rank of an independent order, Cathartiformes, not closely associated with either the birds of prey or the storks and herons.[17]

Recent multi-locus DNA studies on the evolutionary relationships between bird groups[18][19] indicate that New World vultures are related to the other birds of prey, excluding the Falconidae. This analysis argues that New World vultures should either be a part of a new order Accipitriformes[19] or part of an order (Cathartiformes) closely related to, but distinct from, other birds of prey.[18] New World vultures are a sister group to Accipitriformes,[18] a group consisting of Accipitridae, the osprey and secretarybird.[20] Both groups are basal members of the recently recognized clade Afroaves.[18]

Cathartidae Extant species
Genus Common and binomial names Image Range Population estimate Conservation status
Coragyps Le Maout, 1853 Black vulture
Coragyps atratus
South America and north to US Least concern
Cathartes Illiger, 1811 Turkey vulture
Cathartes aura
Throughout the Americas to southern Canada Least concern
Lesser yellow-headed vulture
Cathartes burrovianus
South America and north to Mexico 500,000-4,999,999[21] mature individuals Least concern
Greater yellow-headed vulture
Cathartes melambrotus
Amazon Basin of tropical South America Least concern
Gymnogyps Lesson, 1842 California condor
Gymnogyps californianus
California and parts of northern Arizona, formerly widespread throughout the mountain ranges of Western North America 201,[22] of which approximately 93 mature individuals Critically endangered
Vultur Linnaeus, 1758 Andean condor
Vultur gryphus
Andes[23] Maximum estimate: 10,000[24] Vulnerable
Sarcoramphus Duméril, 1805 King vulture
Sarcoramphus papa
Southern Mexico to northern Argentina 10,000-100,000[25] Least concern

Extinct species and fossils

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The fossil history of the Cathartidae is complex, and many taxa that may possibly have been New World vultures have at some stage been treated as early representatives of the family.[26] There is no unequivocal European record from the Neogene.

Fossil of the extinct Breagyps clarki

It is clear that the Cathartidae had a much higher diversity in the Plio-Pleistocene, rivalling the current diversity of Old World vultures and their relatives in shapes, sizes, and ecological niches. Extinct taxa include:

  • Diatropornis ("European vulture") Late Eocene/Early Oligocene – ?Middle Oligocene of France[27]
  • Phasmagyps Chadronian of Colorado[27][28]
  • Cathartidae gen. et sp. indet. Late Oligocene of Mongolia[27]
  • Brasilogyps Late Oligocene/Early Miocene of Brazil[27]
  • Hadrogyps ("American dwarf vulture") Middle Miocene of SW North America[27]
  • Cathartidae gen. et sp. indet. Late Miocene/Early Pliocene of Lee Creek Mine, USA[29]
  • Pliogyps ("Miocene vulture") Late Miocene – Late Pliocene of S North America[27]
  • Perugyps ("Peruvian vulture") Pisco Late Miocene/Early Pliocene of SC Peru[29]
  • Dryornis ("Argentinean vulture") Early – Late? Pliocene of Argentina; may belong to modern genus Vultur[27]
  • Cathartidae gen. et sp. indet. Middle Pliocene of Argentina[29]
  • Aizenogyps ("South American vulture") Late Pliocene of SE North America[27]
  • Breagyps ("long-legged vulture") Late Pleistocene of SW North America[27]
  • Geronogyps Late Pleistocene of Argentina and Peru[27]
  • Gymnogyps varonai Late Quaternary of Cuba[30]
  • Wingegyps Late Pleistocene of Brazil[31]
  • Pleistovultur Late Pleistocene/Early Holocene of Brazil[32]
  • Cathartidae gen. et sp. indet. Cuba[33]
  • Gymnogyps amplus Late Pleistocene – Holocene of W North America[34]

Description

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Naked head with long beak
The featherless head of the American black vulture, Coragyps atratus brasiliensis, reduces bacterial growth from eating carrion.

New World vultures are generally large, ranging in length from the lesser yellow-headed vulture at 56–61 centimeters (22–24 inches) up to the California and Andean condors, both of which can reach 120 centimeters (48 inches) in length and weigh 12 or more kilograms (26 or more pounds). Plumage is predominantly black or brown, and is sometimes marked with white. All species have featherless heads and necks.[35] In some, this skin is brightly colored, and in the king vulture it is developed into colorful wattles and outgrowths.

All New World vultures have long, broad wings and a stiff tail, suitable for soaring.[36] They are the best adapted to soaring of all land birds.[37] The feet are clawed but weak and not adapted to grasping.[38] The front toes are long with small webs at their bases.[39] No New World vulture possesses a syrinx,[40] the vocal organ of birds. Therefore, the voice is limited to infrequent grunts and hisses.[41]

The beak is slightly hooked and is relatively weak compared with those of other birds of prey.[38] This is because it is adapted to tear the weak flesh of partially rotted carrion, rather than fresh meat.[37] The nostrils are oval and set in a soft cere.[42] The nasal passage is perforate, not divided by a septum, so that when looking from the side, one can see through the beak.[43] The eyes are prominent, and, unlike those of eagles, hawks, and falcons, they are not shaded by a brow bone.[42] Members of Coragyps and Cathartes have a single incomplete row of eyelashes on the upper lid and two rows on the lower lid, while Gymnogyps, Vultur, and Sarcoramphus lack eyelashes altogether.[44]

New World vultures have the unusual habit of urohidrosis, or defecating on their legs to cool them evaporatively. As this behavior is also present in storks, it is one of the arguments for a close relationship between the two groups.[7]

Distribution and habitat

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Andean condor skeleton (Museum of Osteology)

New World vultures are restricted to the western hemisphere, ranging from southern Canada to South America.[45] Most species are mainly resident, but the turkey vulture breeds in Canada and the northern US and migrates south in the northern winter.[46] New World vultures inhabit a large variety of habitats and ecosystems, ranging from deserts to tropical rainforests and at heights of sea level to mountain ranges,[45] using their highly adapted sense of smell to locate carrion. These species of birds are also occasionally seen in human settlements, perhaps emerging to feed upon the food sources provided from roadkills.[citation needed]

Behavior and ecology

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Breeding

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New World vultures and condors do not build nests, but lay eggs on bare surfaces. On average one to three eggs are laid, depending on the species.[35] Chicks are naked on hatching and later grow down. Like most birds, the parents feed the young by regurgitation.[42] The young are altricial, fledging in 2 to 3 months.[41] California Condor chicks fledge anywhere from 5–6 months, while Andean condor chicks fledge anywhere from 6–10 months.[47][48]

Feeding

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King vulture and American black vultures feeding on a carcass

All living species of New World vultures and condors are scavengers. Their diet consists primarily of carrion, and they are commonly seen near carcasses. Other additions to the diet include fruit (especially rotten fruit) and garbage. The genus Cathartes locates carrion by detecting the scent of ethyl mercaptan, a gas produced by the bodies of decaying animals. The olfactory lobe of the brains in these species, which is responsible for processing smells, is particularly large compared to that of other animals.[49] Other species, such as the American black vulture and the king vulture, have weak senses of smell and find food only by sight, sometimes by following Cathartes vultures and other scavengers.[40]

Tolerance to bacterial toxins in decaying meat

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Vultures possess a very acidic digestive system, with their gut dominated by two species of anaerobic bacteria that help them withstand toxins present in decaying prey.[50] In a 2014 study of 50 (turkey and black) vultures, researchers analyzed the microbial community or microbiome of the facial skin and the large intestine.[51] The facial bacterial flora and the gut flora overlapped somewhat, but in general, the facial flora was much more diverse than the gut flora, which is in contrast to other vertebrates, where the gut flora is more diverse. Two anaerobic faecal bacteria groups that are pathogenic in other vertebrates stood out: Clostridia and Fusobacteriota (formerly Fusobacteria). They were especially common in the gut with Clostridia DNA sequence counts between 26% and 85% relative to total sequence counts, and Fusobacteriota between 0.2% and 54% in black vultures and 2% to 69% of all counts in turkey vultures. Unexpectedly, both groups of anaerobic bacteria were also found on the air-exposed facial skin samples, with Clostridia at 7%–40% and Fusobacteriota up to 23%. It is assumed that vultures acquire them when they insert their heads into the body cavities of rotten meat. The regularly ingested Clostridia and Fusobacteriota outcompete other bacterial groups in the gut and become predominant. Genes that encode tissue-degrading enzymes and toxins that are associated with Clostridium perfringens have been found in the vulture gut metagenome. This supports the hypothesis that vultures do benefit from the bacterial breakdown of carrion, while at the same time tolerating the bacterial toxins.[51]

Status and conservation

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The California condor is critically endangered. It formerly ranged from Baja California to British Columbia, but by 1937 was restricted to California.[52] In 1987, all surviving birds were removed from the wild into a captive breeding program to ensure the species' survival.[52] In 2005, there were 127 Californian condors in the wild. As of October 31, 2009 there were 180 birds in the wild.[53] The Andean condor is vulnerable.[23] The American black vulture, turkey vulture, lesser yellow-headed vulture, and greater yellow-headed vulture are listed as species of Least Concern by the IUCN Red List. The king vulture is also listed as Least Concern, although there is evidence of a decline in the population.[54]

In culture

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The American black vulture and the king vulture appear in a variety of Maya hieroglyphs in Mayan codices. The king vulture is commonly represented, with its glyph being easily distinguishable by the knob on the bird's beak and by the concentric circles that represent the bird's eyes.[55] It is sometimes portrayed as a god with a human body and a bird head.[55] According to Mayan mythology, this god often carried messages between humans and the other gods. It is also used to represent Cozcaquauhtli, the thirteenth day of the month in the Mayan calendar.[55] Meanwhile, the American black vulture is normally connected with death or shown as a bird of prey, and its glyph is often depicted attacking humans. This species lacks the religious connections that the king vulture has. While some of the glyphs clearly show the American black vulture's open nostril and hooked beak, some are assumed to be this species because they are vulture-like, painted black, and lack the king vulture's knob.[55]

See also

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Notes

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  1. ^ de Lafresnaye, Frédéric (1839). "Nouvelle classification des oiseaux de proie ou rapaces". Revue Zoologique (in French). 2: 193–196 [194].
  2. ^ Bock, Walter J. (1994). History and Nomenclature of Avian Family-Group Names. Bulletin of the American Museum of Natural History. Vol. 222. New York: American Museum of Natural History. pp. 112, 234. hdl:2246/830.
  3. ^ Myers (2008)
  4. ^ Brookes (2006)
  5. ^ Phillips (2000)
  6. ^ Johnson "et al." 2013
  7. ^ a b Sibley and Ahlquist (1991)
  8. ^ de Boer (1975)
  9. ^ Ligon (1967)
  10. ^ König (1982)
  11. ^ Griffiths (1994)
  12. ^ Fain & Houde (2004)
  13. ^ Avise (1994)
  14. ^ Brown (2009)
  15. ^ Cracraft et al. (2004)
  16. ^ Gibb et al. (2007)
  17. ^ Ericson et al. (2006)
  18. ^ a b c d Jarvis, E. D.; Mirarab, S.; Aberer, A. J.; Li, B.; Houde, P.; Li, C.; Ho, S. Y. W.; Faircloth, B. C.; Nabholz, B.; Howard, J. T.; Suh, A.; Weber, C. C.; Da Fonseca, R. R.; Li, J.; Zhang, F.; Li, H.; Zhou, L.; Narula, N.; Liu, L.; Ganapathy, G.; Boussau, B.; Bayzid, M. S.; Zavidovych, V.; Subramanian, S.; Gabaldon, T.; Capella-Gutierrez, S.; Huerta-Cepas, J.; Rekepalli, B.; Munch, K.; et al. (2014). "Whole-genome analyses resolve early branches in the tree of life of modern birds" (PDF). Science. 346 (6215): 1320–1331. Bibcode:2014Sci...346.1320J. doi:10.1126/science.1253451. hdl:10072/67425. PMC 4405904. PMID 25504713. Archived from the original (PDF) on 2015-02-24. Retrieved 2015-08-28.
  19. ^ a b Hackett et al. (2008)
  20. ^ Griffiths, C. S.; Barrowclough, G. F.; Groth, J. G.; Mertz, L. A. (2007-11-06). "Phylogeny, diversity, and classification of the Accipitridae based on DNA sequences of the RAG-1 exon". Journal of Avian Biology. 38 (5): 587–602. doi:10.1111/j.2007.0908-8857.03971.x.
  21. ^ IUCN (2019-12-03). "Cathartes burrovianus: BirdLife International: The IUCN Red List of Threatened Species 2021: e.T22697630A163511443". IUCN Red List of Threatened Species. 2019-12-03. doi:10.2305/iucn.uk.2021-3.rlts.t22697630a163511443.en.
  22. ^ IUCN (2020-08-28). "Gymnogyps californianus: BirdLife International: The IUCN Red List of Threatened Species 2020: e.T22697636A181151405". IUCN Red List of Threatened Species. 2020-08-28. doi:10.2305/iucn.uk.2020-3.rlts.t22697636a181151405.en.
  23. ^ a b BirdLife International (2020)
  24. ^ IUCN (2020-08-10). "Vultur gryphus: BirdLife International: The IUCN Red List of Threatened Species 2020: e.T22697641A181325230". IUCN Red List of Threatened Species. 2020-08-10. doi:10.2305/iucn.uk.2020-3.rlts.t22697641a181325230.en. S2CID 241274385.
  25. ^ IUCN (2016-10-01). "Sarcoramphus papa: BirdLife International: The IUCN Red List of Threatened Species 2016: e.T22697645A93627003". IUCN Red List of Threatened Species. 2016-10-01. doi:10.2305/iucn.uk.2016-3.rlts.t22697645a93627003.en.
  26. ^ Mayr (2006)
  27. ^ a b c d e f g h i j Emslie (1988)
  28. ^ Wetmore, A. (1927). "Fossil Birds from the Oligocene of Colorado" (PDF). Proceedings of the Colorado Museum of Natural History. 7 (2): 1–14.
  29. ^ a b c Stucchi (2005)
  30. ^ Suárez (2003)
  31. ^ Alvarenga (2004).
  32. ^ Alvarenga et al. (2008).
  33. ^ Suarez (2004)
  34. ^ Steverson, Valerie J.; Prothero, Donald R. (2010). "Evolutionary Patterns in Late Quaternary California Condors" (PDF). PalArch's Journal of Vertebrate Palaeontology.
  35. ^ a b Zim et al. (2001)
  36. ^ Reed (1914)
  37. ^ a b Ryser & Ryser (1985)
  38. ^ a b Krabbe (1990)
  39. ^ Feduccia (1999)
  40. ^ a b Kemp and Newton (2003)
  41. ^ a b Howell and Webb (1995)
  42. ^ a b c Terres (1991)
  43. ^ Allaby (1992)
  44. ^ Fisher (1942)
  45. ^ a b Harris (2009)
  46. ^ Farmer (2008)
  47. ^ "Andean Condor | The Peregrine Fund". peregrinefund.org. Retrieved 2020-09-29.
  48. ^ "Gymnogyps californianus". www.fs.fed.us. Retrieved 2020-09-29.
  49. ^ Snyder (2006)
  50. ^ Will Dunham (26 November 2014). "Gut check: how vultures dine on rotting flesh, and like it". Reuters. Thomson Reuters. Retrieved 27 November 2014.
  51. ^ a b Michael Roggenbuck; Ida Bærholm Schnell; Nikolaj Blom; et al. (25 November 2014). "The microbiome of New World vultures". Nature Communications. 5 (5498): 5498. Bibcode:2014NatCo...5.5498R. doi:10.1038/ncomms6498. PMID 25423494.
  52. ^ a b BirdLife International (2009a)
  53. ^ "San Diego Zoo's Animal Bytes: California Condor". The Zoological Society of San Diego's Center for Conservation and Research for Endangered Species. Retrieved 2009-12-29.
  54. ^ BirdLife International (2001)
  55. ^ a b c d Tozzer (1910)

References

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