Cuban macaw
The Cuban macaw or Cuban red macaw is an extinct species of macaw native to the main island of Cuba and the nearby Isla de la Juventud. It became extinct in the late 19th century. Its relationship with other macaws in its genus was long uncertain, but it was thought to have been closely related to the scarlet macaw, which has some similarities in appearance. It may also have been closely related, or identical, to the hypothetical Gosse's macaw. A 2018 DNA study found the Cuban macaw to be the sister species of two red and two green species of extant macaws.
At about long, the Cuban macaw was one of the smallest macaws. It had a red, orange, yellow, and white head, and a red, orange, green, brown, and blue body. Little is known of its behaviour, but it is reported to have nested in hollow trees, lived in pairs or families, and fed on seeds and fruits. The species' original distribution on Cuba is unknown, but it may have been restricted to the central and western parts of the island. It was mainly reported from the vast Zapata Swamp, where it inhabited open terrain with scattered trees.
The Cuban macaw was traded and hunted by Native Americans, and by Europeans after their arrival in the 15th century. Many individuals were brought to Europe as cagebirds, and 19 museum skins exist today. A few subfossil and archaeological remains have been found on Cuba. It had become rare by the mid-19th century due to pressure from hunting, trade, and habitat destruction. Hurricanes may also have contributed to its demise. The last reliable accounts of the species are from the 1850s on Cuba and 1864 on Isla de la Juventud, but it may have persisted until 1885.
Taxonomy
Early explorers of Cuba, such as Christopher Columbus and Diego Álvarez Chanca, mentioned macaws there in 15th- and 16th-century writings. Cuban macaws were described and illustrated in several early accounts about the island. In 1811, the German naturalist Johann Matthäus Bechstein scientifically named the species Psittacus tricolor.Bechstein's description was based on the bird's entry in the French naturalist François Le Vaillant's 1801 book Histoire Naturelle des Perroquets, wherein it was referred to as l'Ara Tricolor. Le Vaillant's account was itself partially based on the late 18th century work Planches Enuminées by the French naturalists Comte de Buffon and Edme-Louis Daubenton, as well as a specimen in Paris; as it is unknown which specimen this was, the species has no holotype. The French illustrator Jacques Barraband's original watercolour painting, which was the basis of the plate in Le Vaillant's book, differs from the final illustration in showing bright red lesser wing covert feathers, but the significance of this is unclear.
File:Ara tricolor Berlin 9.jpg|thumb|left|Study skin in Museum für Naturkunde, Berlin, one of 19 specimens in existence
Today, 19 skins of the Cuban macaw exist in 15 collections worldwide, but many are of unclear provenance. Several were provided by the Cuban naturalist Juan Gundlach, who collected some of the last individuals that regularly fed near the Zapata Swamp in 1849–50. Some of the preserved specimens are known to have lived in captivity in zoos or as cagebirds. The single specimen at World Museum, National Museums Liverpool died in Edward Smith-Stanley, 13th Earl of Derby's aviaries at Knowsley Hall in 1846. Several more skins are known to have existed, but have been lost. There are no records of its eggs.
Four subfossil specimens have been discovered: half a carpometacarpus from a possibly Pleistocene spring deposit in Ciego Montero, identified by extrapolating from the size of Cuban macaw skins and bones of extant macaws, a rostrum from a Quaternary cave deposit in Caimito, a worn skull from Sagua La Grande, which was deposited in a waterfilled sinkhole possibly during the Quaternary and associated with various extinct birds and ground sloths, and a fragmentary carpometacarpus from Upper Pleistocene layers of the El Abrón Cave in Pinar del Río, the first physical evidence from the western part of Cuba. In 2021, the first archaeological remains were reported, a tarsometatarsus and upper beak from two sites in Old Havana dated to the 17th and 18th centuries.
Related Caribbean macaws
As many as 13 now-extinct species of macaw have variously been suggested to have lived on the Caribbean islands, but many of these were based on old descriptions or drawings and only represent hypothetical species. Only three endemic Caribbean macaw species are known from physical remains: the Cuban macaw, the Saint Croix macaw, which is known only from subfossils, and the Lesser Antillean macaw, which is known from subfossils and reports. Macaws are known to have been transported between the Caribbean islands and from mainland South America to the Caribbean both in historic times by Europeans and natives, and in prehistoric times by Palaeoamericans. Historical records of macaws on these islands, therefore, may not have represented distinct, endemic species; it is also possible that they were escaped or feral foreign macaws that had been transported to the islands. All the endemic Caribbean macaws were likely driven to extinction by humans in historic and prehistoric times. The identity of these macaws is likely to be further resolved only through fossil finds and examination of contemporary reports and artwork.A macaw shot on Jamaica around 1765 was seen stuffed by a Dr. Robertson, but was since lost. He stated it was different from any macaw he had seen before, and sent a description of it to the British naturalist Philip Henry Gosse, who published his own description in 1847, calling it yellow-headed macaw. It was described as being similar to the Cuban macaw, mainly differing in having a yellow forehead. On this basis, the British zoologist Walter Rothschild named it Ara gossei in 1905, and it was subsequently referred to by common names such as Gosse's macaw, the Jamaican yellow-headed macaw, and the Jamaican red macaw. Some researchers believe the specimen described may have been a feral Cuban macaw. A stylised 1765 painting of a macaw by the British Lieutenant L. J. Robins, published in a volume called The Natural History of Jamaica, matches the Cuban macaw, and may show a specimen that had been imported there; however, it has also been claimed that the painting shows Gosse's macaw. In 2017, the Australian ornithologist Joseph M. Forshaw found the Cuban, the Lesser Antillean, and the supposed Gosse's macaw so similar that they could have been the same species transported between islands, or, with macaws that possibly occurred on other islands, been closely related and possibly derived from the scarlet macaw.
Rothschild's 1907 book Extinct Birds included a depiction of a specimen in the Liverpool Museum which was presented as a Cuban macaw. In a 1908 review of the book published in The Auk, the American ornithologist Charles Wallace Richmond claimed that the picture looked sufficiently dissimilar from known Cuban macaws that the specimen may actually be of one of the largely unknown species of macaw, such as a species from Haiti. This suggestion has not been accepted.
The name Ara tricolor haitius was coined for a supposed subspecies from Hispaniola by the German ornithologist Dieter Hoppe in 1983, but is now considered to have been based on erroneous records. In 1985, the American ornithologist David Wetherbee suggested that extant specimens had been collected from both Cuba and Hispaniola, and that the two populations represented distinct species, differing in details of their colouration. Whetherbee stated the name Ara tricolor instead applied to the supposed Hispaniolan species, as he believed Cuba had no bird collectors prior to 1822, and that the illustration and description published by Le Vaillant were based on a specimen collected during a 1798 expedition to Hispaniola. As the Cuban species was thereby in need of a new specific name, Wetherbee coined Ara cubensis for it. He also suggested that Gosse's macaw was based on a "tapiré"; a specimen whose colouration was altered through a Native American technique whereby developing feathers can be changed to red and yellow by painting them with body fluids of the dyeing dart frog. The idea that the name Ara tricolor applied to a Hispaniolan species had gained acceptance by 1989, but in 1995, the British ornithologist Michael Walters pointed out that birds had indeed been described from Cuba prior to 1822, that the supposed differences in colouration were of no importance, and that the basis of Wetherbee's argument was therefore invalid. There is no clear evidence for a species of macaw on Hispaniola, and later researchers concurred with Walters.
Evolution
Since detailed descriptions of extinct macaws exist only for the species on Cuba, it is impossible to determine their interrelationships. It has been suggested that the closest mainland relative of the Cuban macaw is the scarlet macaw, due to the similar distribution of red and blue in their plumage, and the presence of a white patch around the eyes, naked except for lines of small red feathers. Furthermore, the range of the scarlet macaw extends to the margins of the Caribbean Sea. The two also share a species of feather mite, which supports their relationship. The American ornithologist James Greenway suggested in 1967 that the scarlet macaw and the Cuban macaw formed a superspecies with the other extinct species thought to have inhabited Jamaica, Hispaniola and Guadeloupe.A 2018 DNA study by the Swedish biologist Ulf S. Johansson and colleagues analysed the mitochondrial genome of two Cuban macaw specimens in the Swedish Museum of Natural History. Though it was expected the Cuban species would form a clade with the likewise predominantly red scarlet macaw and the red-and-green macaw, they instead found it to be basal to those two large red macaws, as well as to the two large green macaws, the military macaw and the great green macaw. The cladogram below follows the 2018 study:
The Cuban macaw was smaller than the related extant species, and one of the smallest Ara species, which suggests smaller size may have been the ancestral state of the group, though it may also have become smaller after becoming established in the Antilles. Johansson and colleagues estimated that the Cuban macaw had diverged from its mainland relatives around 4 million years ago, during the early Pliocene. Since this is after the land bridge that is thought to have connected the Greater Antilles with South America ceased to exist, the ancestors of the Cuban macaw must have dispersed to the Antilles over open water. Therefore, the Cuban macaw was not a recent offshoot of the scarlet macaw, having a long independent history on Cuba. Johansson and colleagues therefore noted that though many of the extinct species of Caribbean macaws that had been described in the past are probably dubious, there would have been ample time for a radiation of macaws there, based on how long the Cuban species had been separated from the mainland species. A 2020 genetic study of the scarlet macaw by the American ecologist Kari L. Schmidt and colleagues resulted in a similar cladogram to that of Johansson and colleagues.