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New World vulture

 
Animal Classification: New World vultures

(Cathartidae)

Class: Aves

Order: Ciconiiformes

Suborder: Cathartae

Family: Cathartidae

Thumbnail description
Medium large to very large, highly social birds that feed primarily on carrion with very few or no feathers on their heads; plumage color is generally black, gray, or brown with some portion of white in three of the largest species

Size
2.2–33.1 lb (1–15 kg)

Number of genera, species
5 genera; 7 species

Habitat
Forests, savannas, woodland, pastures, mountains, deserts, river ways, and seashores

Conservation status
Critically Endangered: 1 species; Near Threatened: 1 species

Distribution
Southern Canada to Tierra del Fuego

Evolution and systematics

The Cathartidae include seven species of vultures that range exclusively in North and South America. Also referred to as "Neotropical" or New World vultures, they were once thought to be closely related to Old World vultures, found in Europe, Asia, and Africa, but any similarities of the two groups can best be attributed to convergence or parallel evolution.

As recently as the last quarter of the twentieth century it was argued successfully on the basis of physiology, behavior, and genetics, that the cathartid vultures are not descended from the Accipitidiae as is accepted for the Old World vultures, but descended instead from a common ancestor with the Ciconidae, or storks. For instance, all of the cathartids and ciconids use "urohydrosis" or the method of cooling themselves by emitting liquid waste on the bare portion of their legs where densely packed blood vessels close to the skin are cooled by evaporation and, in turn, body core temperature is reduced. They also never rest on one foot as do birds of prey but instead lie down. While both New and Old World vultures express a distinct social hierarchy, that of Old World vultures is based more on the hunger level of a bird arriving at a carcass while the social status of New World vultures is based primarily on an individual's personal status that is determined by the individual's species, age, sex, and experience.

The evolutionary history of the Old and New World vultures is comparatively good. The earliest New World vulture was reported in England dating from late Paleocene deposits. Several "cathartid type" fossils are known from middle and late Eocene deposits in France and Germany but no remains after early Miocene have been located in the Old World. Fossil records show the New World vultures first appearing in America in the early Oligocene, flourishing along side Old World type vultures that became extinct toward the end of the Pleistocene, only 10,000–20,000 years ago. When the mass mammalian extinctions occurred, both Old and New World type vultures followed the fate of their prey.

The California condor appears to be the only large vulture to survive. Its Pleistocene range that spanned southern North America and included both coasts, was reduced to the west coast from British Columbia to northern Baja by modern times. Like the Andean condor today, it relied heavily on carrion found along the coast.

Physical characteristics

The family Cathartidae consists of five genera with seven species. The smallest species, by weight, is the 2.1 lb (0.94 kg) lesser yellow-headed vulture (Cathartes burrovianus). The other two species in the genus Cathartes, the turkey vulture (C. aura) weighing 3.3 lb (1.5 kg) and the greater yellow-headed vulture (C. melambrotus) at 2.6 lb (1.2 kg), are not much heavier yet all give the appearance of being much larger than their actual weight. The physical effect of their large flying surface area to weight ratio, called a "light wing loading," makes these three species comparatively more buoyant in air and able to take advantage of the slightest thermal to stay aloft close to the ground. The black vulture (Coragyps atratus) on the other hand has a relatively short wingspan, or flying surface, for its heavier weight of 4.4 lb (2.0 kg). These flight characteristics join with other behavioral and anatomical factors to help define niche separation in each species of this scavenger guild. The colorful king vulture (Sarcoramphus papa) is built more like a black vulture but larger, weighing about 7.5 lb (3.4 kg). The California condor (Gymnogyps californianus), like the smaller vultures, is sexually monomorphic in size and color, but is much heavier weighing 17–24 lb (7.7–10.9 kg) with a wingspan of 114 in (290 cm). The Andean condor (Vultur gryphus) is one of the largest flying birds in the world. It is sexually dimorphic in shape, color, and size. Females range in weight from 18 to 23 lb (8.3–10.5 kg), have dark gray skin on the head which has no caruncle, similar to the male. Their iris color in the adult is a deep red while that of the male is tan, plumages are the same. The larger male ranges from 24 to 33 lb (10.9–15 kg).

While the Cathartidae have only a rudimentary syrinx and cannot call or sing as other birds can they are able to communicate with a suprising array of grunts, growls, and hisses.

Distribution

The most widely distributed cathartid species is the turkey vulture, ranging from the Canadian border to the southern end of South America. There are also four subspecies of turkey vultures usually recognized based on slight differences in head color and distribution. The most migratory appears to be C. aura aura which has been extending its summer breeding range over the last few decades north though New England. It spends winters from the southern United States to northern South America competing with the more sedentary subspecies of that region. The movement patterns of the other cathartid species appear to be less latitudinally migratory but are more regional, associated with weather patterns, food supply, and breeding season.

Roosting areas are particularly important in influencing the distribution pattern in these vultures particularly for the rarer, larger condors where roosting conditions are more specific. Where hundreds of black and turkey vultures may roost in particular groves of trees or on the supports of man-made towers, it is more difficult for condors to find appropriate cliff roosts with the right climatic conditions. Dozens of condors may use a network of these traditional roosts as secure bases from which to forage in a particular area and they are as important as adequate nesting sites in delineating the distribution of these vultures.

Habitat

From the northern to southern portions of where cathartid vultures range they are found in every habitat where their carrion food supply can be effectively exploited, including but not limited to deserts, coastlines, water ways, open grasslands and savannas, forests, cities, and mountain and canyon regions.

Unlike Old World vultures that do not forage in closed forests, three cathartid species in the New World are well adapted to successfully exploit forest as well as open habitat. They accomplish this through a sense of smell that is so acute they can find even small bird, reptile, and mammal carcasses in a forest shortly after they begin to decompose. Both condor species use winds generated off mountain slopes and thermal activity to move distances of hundreds of kilometers at thousands of meters of altitude within a few days covering several habitat types.

Behavior

New World vultures are highly gregarious roosting nightly and foraging communally by day. Unless there is already wind, in which case they can and will fly before dawn, vultures and condors typically wait until later in the morning when rising thermals can assist soaring flight to where the most recent carcass is located. When the first rays of sun hit the roost or throughout the day when it peaks from behind a cloud, vultures will spread their wings and orient themselves at right angles to the sun. Seen also after bathing, this pose, called sunning, dries and straightens flight feathers and functions to assist with preening in reducing ectoparasites.

Like the accipitrid vultures in Africa, the Cathartidae partition the food resources of the available carrion in an area through timing, anatomical differences, and a relatively ordered hierarchy. Black vultures, with a heavier wing loading, are most efficient at foraging when flying at higher altitudes where they can best observe the behavior and activities of their own and other species. Lacking the ability to use olfaction, like the three smaller Carthartes species, they rely on vision to hunt. When flying conditions allow, a foraging flock of black vultures will disperse over miles, where they can effectively scan for resource opportunities to exploit over a large area. When activity of interest is noted by one or more individuals, the adjusted flying pattern appears to signal the attention of other flock mates. As more and more birds gather over a carcass, the inadvertent signal produced by the mass of large black birds at varying altitudes can persist over many days attracting the larger, less common species of scavengers such as king vultures and Andean condors. The same scenario is carried out on the west coast of North America involving turkey vultures, ravens, and California condors.

Where black vultures occur, their population levels can get quite large, even into the thousands sustained by some type of consistent and predictable food source like a city dump. So behaviorally astute are black vultures that they will adjust their flight distance to humans depending on the circumstances. The same marked bird feeding without fear within a few feet of people at the Panama City market place one day will flush away the next day when approached within 164 ft (50 m) in the countryside. When black vultures first arrive at a natural carcass, not part of the usual system, they have usually followed one of the Cathartes species. At low, equal numbers, black vultures generally cannot dominate turkey vultures, but as their group size increases, the Cathartes species often move to the side of the main feeding activity or leave altogether.

Feeding ecology and diet

King vultures, and especially condors, can go longer periods between meals than the smaller vultures as long as water is available. Reintroduced California condors re-trapped after 26 days of unsuccessful foraging showed no behavioral or clinical signs of stress. They generally fly higher, faster, and over larger areas, foraging away from roost site and nesting territories. When they arrive at a carcass, they dominate any of the smaller species. Observations of marked wild Andean condors indicate that birds with high status are less reluctant to approach the carcass. The smaller bills of the Cathartes and Coragyps vultures are not sufficient to tear through the hide of large carcasses and usually are confined to natural openings of the mouth, eyes, ears, and anus. Three to five of the most dominant individuals of the 100 member black vulture flock can defend the few holes in a fresh, large carcass. Sometimes hundreds of birds wait off to one side for the social dynamics to change. When condors or even king vultures arrive, the waiting, lower status vultures become alert and begin to crowd the carcass as the first condor approaches. With a bill every bit as powerful as the largest Old World vulture, one or more condors soon open several access points in the tough hide, making it difficult or impossible for a few dominant black vultures to successfully defend the carcass. With a breakdown in the hierarchy, a feeding frenzy ensues and even young, normally submissive birds can race into the confusion and successfully dash out with food.

Reproductive biology

The black vulture and the three Cathartes species all lay three eggs on a yearly basis. Although sexually mature by age two they may take several more years to acquire a mate and successfully breed. The king vulture and the two condors lay only one egg per season. Condors have a very slow reproductive rate and may take two or more years to produce one young. Parental dependence is months long in the larger species but short to non-existent in the genus Cathartes possibly due to their almost immediate success at finding food through olfaction. Monogamy is typical, with pair bonds that are life long, but shifts in mates may occur if the pair is unproductive over several years. Territory defense in condors is males against intruding males and females against females.

Conservation status

Where turkey vultures and black vultures are gradually expanding their range in some areas, the king vulture and condors have had significant declines. The California condor that narrowly escaped extinction at the end of the Pleistocene had been declining since the early 1800s when the growing human population reduced its coastal food supply and directly shot and poisoned the species. By the early 1980s, only 21 had survived and by 1987 all of the wild flock had to be brought into captivity to insure the species survival. Captive breeding programs at the Los Angeles Zoo and the San Diego Wild Animal Park and later, the World Center for Birds of Prey in Idaho have been highly successful in producing numbers of birds while preserving the remaining founding genetic lines. The reintroduction of the species into its former habitat began in 1992 and by the close of 2001 there were 183 birds with nearly 60 of those in the wild at three release sites; two in California and one in Arizona. Attempts to breed in the wild began in 2002. Andean condors have also been reintroduced into parts of their former range where they had been extirpated. North American Zoos, through their Species Survival Plan for Andean Condors, raised and released over 80 Andean condors in Venezuela and Colombia where they now breed in the wild.

Significance to humans

Condors were important in the mythology and featured in the rituals of the pre-Columbian cultures in the Andes. Their image is found incorporated in the textile designs, pottery, and carvings of these peoples. Even today the Andean condor appears on the coats of arms of Colombia, Bolivia, Ecuador, and Chile. Native North American groups greatly respected the California condor. It was buried with the dead, and its image was incorporated into their artistic motifs.

Many early human cultures associated vultures with death and these birds became important symbols in burial rituals. Today vultures are not respected as they were by earlier cultures, but they are tolerated for the valuable environmental service they provide. Vultures have suffered in recent years from human-generated pollution. Positioned as they are at the end of the food chain, vultures are likely to accumulate toxins and contaminants, which can kill them outright or damage their reproductive success. Other human-made hazards, such as power lines, can also pose a danger to flying birds.

Species accounts

Turkey vulture
Lesser yellow-headed vulture
Greater yellow-headed vulture
American black vulture
King vulture
California condor
Andean condor

Resources

Books:

Blake, E.R. Manual of Neotropical Birds. Vol. 1: Spheniscidae (Penguins) to Laridae (Gulls and Allies). Chicago & London: University of Chicago Press, 1977.

Snyder, N.F.R., and H.A. Snyder. Birds of Prey: Natural History and Conservation of North American Raptors. Stillwater, Minnesota: Voyageur Press, 1991.

Wallace, M.P., and W. Toone. "Captive Management for the Long Term Survival of the California Condor." In Wildlife 2001: Populations, edited by D. R. McCullough and R.H. Barrett. Elsevier Applied, 1992.

Wilbur, S.R., and J.A. Jackson, eds. Vulture Biology and Management. University of California Press, 1983.

Periodicals:

Audubon, J.J. "Account of the Habits of the Turkey Buzzard Vultur aura Particularly with the View of Exploding the Opinion Generally Entertained of Its Extraordinary Powers of Smelling." Edinb. New Phil. Journal 2(1826):172–184.

Bang, B.G. "The Nasal Organs of the Black and Turkey Vultures: A Comparative Study of the Cathartid Species Coragyps atratus atratus and Cathartes aura septentrionalis (With Notes on Cathartes aura falklandica, Pseudogyps bengalensis and Neophron percnopterus)." Journal of Morphology 115(1972): 153–184.

Bernal, L.G., D.C. Houston, and P. Cotton. "The Role of Greater Yellow-headed Vultures as Scavengers in Neotropical Forests." Ibis 136(1994).

Clinton-Eitniear, J. "King Vulture Research Report." Vulture News 6(1981):7–8.

Cox, C.R., V.I. Goldsmith, and H.R. Engelhardt. "Pair Formation in California Condors." Amer. Zool. 33(1993): 126–138.

Davis, D. "Morning and Evening Roosts of Turkey Vultures at Malheur Refuge, Oregon." Western Birds 10(1979): 125–130.

Gailey, J., and N. Bolwig. "Observations on the Behaviour of the Andean Condor Vultur gryphus." Condor 75(1973):60–68.

Graves, G.R. "Greater Yellow-headed Vulture (Cathartes melambrotus) Locates Food by Olfaction." Journal of Raptor Research 26(1992): 38–39.

Houston, D.C. "Competition for Food Between Neotropical Vultures in Forest." Ibis 130(1988): 402–417.

Kiff, L.F. "An Historical Perspective on the Condor." Outdoor California 44(1983): 5–6, 34–37.

Kiff, L.F. "To the Brink and Back: The Battle to Save the California Condor." Terra 28(1990): 6–18.

Meretsky, V., and N.F.R. Snyder. "Range Use and Movements of California Condors." Condor 94(1992): 313–335.

Pattee, O.H "The Role of Lead in Condor Mortality." Endangered Species Bulletin Vol. 12, No. 9 (1987): 6–7.

Rabenold, P.P. "Family Associations in Communally Roosting Black Vultures." Auk 103(1986): 32–41.

Snyder, N.F.R., R.R. Ramey, and F.C. Sibley. "Nest Site Biology of the California Condor. " Condor 88(1986): 228–241.

Snyder, N.F.R., and H.A. Snyder. "Biology and Conservation of the California Condor." Current Ornithology 6(1989): 175–267.

Stewart, P.A. "The Biology and Communal Behaviour of American Black Vultures." Vulture News 9/10(1983): 14–36.

Toone, W., and A.C. Risser. "Captive Management of the California Condor Gymnogyps californianus." International Zoology Yearbook 27(1988): 50–58.

[Article by: Michael Phillip Wallace, PhD]

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WordNet: New World vulture
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Note: click on a word meaning below to see its connections and related words.

The noun has one meaning:

Meaning #1: large birds of prey superficially similar to Old World vultures
  Synonym: cathartid


Wikipedia: New World vulture
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New World vultures
Large black bird with unfeathered head, standing looking to left
American Black Vulture
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Superclass: Tetrapoda
Class: Aves
Order: Accipitriformes
Family: Cathartidae
Lafresnaye, 1839
Genera

Coragyps
Cathartes
Gymnogyps
Vultur
Sarcoramphus

Approximate Cathartidae range map

Yellow - Summer-only range of Turkey Vulture 
Green - At least one species present year-round

The New World vulture family Cathartidae contains seven species in five genera, all but one of which are monotypic. It includes five vultures and two condors found in warm and temperate areas of the Americas.

New World vultures are not closely genetically related to the superficially similar family of Old World vultures; similarities between the two groups are due to convergent evolution. Just how closely related they are is a matter of debate (see Taxonomy and evolution). They were widespread in both the Old World and North America, during the Neogene.

Vultures are scavenging birds, feeding mostly from carcasses of dead animals. New World vultures have a good sense of smell, but Old World vultures find carcasses exclusively by sight. A particular characteristic of many vultures is a bald head, devoid of feathers.

Contents

Taxonomy and nomenclature

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

The New World vultures comprise seven species in five genera. The genera are Coragyps, Cathartes, Gymnogyps, Sarcoramphus, and Vultur. Of these, only Cathartes is not monotypic.[1] The family's scientific name, Cathartidae, comes from cathartes, Greek for "purifier".[2] Although New World vultures have many resemblances to Old World vultures they are not very closely related. Rather, they resemble Old World vultures because of convergent evolution.[3]

New World vultures were traditionally placed in a family of their own in the Falconiformes.[4] However, in the late 20th century some ornithologists argued that they are more closely related to storks on the basis of karyotype,[5] morphological,[6] and behavioral[7] data. Thus some authorities place them in the Ciconiiformes with the storks and herons; Sibley and Monroe (1990) even considered them a subfamily of the stork family. This has been criticized as an oversimplification,[8][9] and an early DNA sequence study[10] was based on erroneous data and subsequently retracted.[11][12][13] Consequently, there is a recent trend to raise the New World vultures to the rank of an independent order Cathartiformes not closely associated with either birds of prey or storks or herons.[14] In 2007 the American Ornithologists' Union's North American checklist moved Cathartidae back into the lead position in Falconiformes, but with an asterisk that indicates it is a taxon "that is probably misplaced in the current phylogenetic listing but for which data indicating proper placement are not yet available".[15] The AOU's draft South American checklist places the Cathartidae in their own order, Cathartiformes.[16] However, recent DNA study on the evolutionary relationships between bird groups also suggests that they are related to the other birds of prey and should be part of a new order Accipitriformes instead.[citation needed]

Extant species

Extinct species and fossils

A related extinct family were the Teratornithidae or Teratorns, essentially an exclusively (North) American counterpart to the New World vultures — the latter were, in prehistoric times, also present in Europe and possibly even evolved there. The Incredible Teratorn is sometimes called "Giant Condor" because it must have looked similar to the modern bird. They were, however, not very closely related but rather an example of parallel evolution, and the external similarity is less emphasized in recent times due to new information suggesting that the teratorns were more predatory than vultures.[19]

The fossil history of the Cathartidae is fairly extensive, but nonetheless confusing. Many taxa that may or may not have been New World vultures were considered to be early representatives of the family.[20] There is no unequivocal European record from the Neogene.

At any rate, 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 genera are:[citation needed]

  • Diatropornis Late Eocene/Early Oligocene -? Middle Oligocene of France[21]
  • Phasmagyps Early Oligocene of WC North America[21]
  • Cathartidae gen. et sp. indet. (Late Oligocene of Mongolia)[21]
  • Brasilogyps Late Oligocene - Early Miocene of Brazil[21]
  • Hadrogyps Middle Miocene of SW North America[21]
  • Cathartidae gen. et sp. indet. Late Miocene/Early Pliocene of Lee Creek Mine, USA[22]
  • Pliogyps Late Miocene - Late Pliocene of S North America[21]
  • Perugyps Pisco Late Miocene/Early Pliocene of SC Peru[22]
  • Dryornis Early - Late? Pliocene of Argentina; may belong to modern genus Vultur[21]
  • Cathartidae gen. et sp. indet. (Middle Pliocene of Argentina[22]
  • Aizenogyps Late Pliocene of SE North America[21]
  • Breagyps Late Pleistocene of SW North America[21]
  • Geronogyps Late Pleistocene of Argentina and Peru[21]
  • Wingegyps Late Pleistocene of Brazil [23]
  • Cathartidae gen. et sp. indet. (Cuba)[24]

Description

The featherless head of the American Black Vulture, Coragyps atratus brasiliensis, reduces bacterial growth from eating carrion.

These birds are generally large, ranging in length from the Lesser Yellow-headed Vulture at 56–61 centimeters (22–24 in) up to the California and Andean Condors, both of which can reach 120 centimeters (48 in) in length and weigh 12 or more kilograms (26 or more lb). Plumage is predominantly black or brown, and is sometimes marked with white. All species have featherless heads and necks.[25] In some, this skin is brightly colored, and in the King Vulture it is developed into colorful wattles and outgrowths.

All species have long, broad wings and a stiff tail, suitable for soaring.[26] They are the best adapted to soaring of all land birds.[27] The feet are clawed but weak and not adapted to grasping.[28] The front toes are long with small webs at their bases.[29] No New World vulture possesses a syrinx,[30] the vocal organ of birds, therefore the voice is limited to infrequent grunts and hisses.[31]

The beak is slightly hooked and is relatively weak when compared those of other birds of prey.[28] It is weak because it is adapted to tear the weak flesh of partially rotted carrion, rather than fresh meat.[27] The nostrils are oval and are set in a soft cere.[32] The nasal passage is not divided by a septum (they are "perforate"), so from the side one can see through the beak,[33] as in the Turkey Vulture. The eyes are prominent, and unlike those of eagles, hawks and falcons, they are not shaded by a bony brow bone.[32] 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.[34]

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.[4]

Distribution and habitat

New World vultures are restricted to the Western Hemisphere. They can be found from southern Canada to South America.[35] Most species are mainly resident, but the Turkey Vulture populations breeding in Canada and the northern US migrate south in the northern winter.[36] 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,[35] 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]

Behaviour

Feeding

American Black Vultures on a cow carcass

All living species of New World vultures and condors are scavengers. Though their diet is overwhelmingly composed of carrion, some species, such as the American Black Vulture, have been recorded as killing live prey. Other additions to the diet include fruit, eggs, and garbage. An unusual characteristic of the species in genus Cathartes is a highly developed sense of smell, which they use to find carrion. They locate carrion by detecting the scent of ethyl mercaptan, a gas produced by the beginnings of decay in dead 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.[37] 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.[30] The head and neck of New World Vultures are featherless as an adaptation for hygiene; this lack of feathers prevents bacteria from the carrion it eats from ruining its feathers and exposes the skin to the sterilizing effects of the sun.[38]

Breeding

New World vultures and condors do not build nests. Instead, they lay eggs on bare surfaces. One to three eggs are laid, depending on the species.[25] Chicks are naked at hatching and later grow down. The parents feed the young by regurgitation.[32] The young are altricial and fledge in 2 to 3 months.[31]

Status and conservation

The California Condor is critically endangered. It formerly ranged from Baja California to British Columbia, but by 1937 was restricted to California.[17] In 1987, all surviving birds were removed from the wild into a captive breeding program to ensure the species' survival.[17] In 2005, there were 127 Californian Condors in the wild. The Andean Condor is near threatened.[18] 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. This means that populations appear to remain stable, and they have not reached the threshold of inclusion as a threatened species, which requires a decline of more than 30 percent in ten years or three generations.[39]

In culture

The American Black Vulture and the King Vulture appear in a variety of Maya hieroglyphics in Mayan codices. The King Vulture is one of the most common species of birds represented in the Mayan codices.[40] Its glyph is easily distinguishable by the knob on the bird’s beak and by the concentric circles that represent the bird’s eyes.[40] It is sometimes portrayed as a god with a human body and a bird head.[40] 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.[40] In Mayan codices, 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 and painted black, but lack the King Vulture’s knob.[40]

Notes

  1. ^ Myers (2008)
  2. ^ Brookes (2006)
  3. ^ Phillips (2000)
  4. ^ a b Sibley and Ahlquist (1991)
  5. ^ de Boer (1975)
  6. ^ Ligon (1967)
  7. ^ König (1982)
  8. ^ Griffiths (1994)
  9. ^ Fain & Houde (2004)
  10. ^ Avise (1994)
  11. ^ Brown (2009)
  12. ^ Cracraft et al. (2004)
  13. ^ Gibb et al. (2007)
  14. ^ Ericson et al. (2006)
  15. ^ American Ornithologists' Union (2009)
  16. ^ Remsen et al. (2008)
  17. ^ a b c BirdLife International (2009a)
  18. ^ a b BirdLife International (2009)
  19. ^ Campbell & Tonni 1983
  20. ^ Mayr (2006)
  21. ^ a b c d e f g h i j Emslie (1988)
  22. ^ a b c Stucchi (2005)
  23. ^ Alvarenga (2004).
  24. ^ Suarez (2004)
  25. ^ a b Zim et al. (2001)
  26. ^ Reed (1914)
  27. ^ a b Ryser & Ryser (1985)
  28. ^ a b Krabbe (1990)
  29. ^ Feduccia (1999)
  30. ^ a b Kemp and Newton (2003)
  31. ^ a b Howell and Webb (1995)
  32. ^ a b c Terres (1991)
  33. ^ Allaby (1992)
  34. ^ Fisher (1942)
  35. ^ a b Harris (2009)
  36. ^ Farmer (2008)
  37. ^ Snyder (2006)
  38. ^ Stone (1992)
  39. ^ BirdLife International (2001)
  40. ^ a b c d e Tozzer (1910)

References

External links


 
 
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