Lesser Antillean macaw

Lesser Antillean macaw
A sepia line drawing showing five birds sitting in a tree, a black bird in flight and a tortoise or turtle on the ground underneath.
Du Tertre's 1667 illustration of three Guadeloupe amazons (8) and one Lesser Antillean macaw (7)
Conservation status
Not recognized (IUCN 3.1)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Order:	Psittaciformes
Superfamily:	Psittacoidea
Family:	Psittacidae
Subfamily:	Arinae
Tribe:	Arini
Genus:	Ara
Species:	A. guadeloupensis
Binomial name
Ara guadeloupensis (Clark, 1905)
A map showing the location of Guadeloupe, an island region in the Caribbean Sea.
Location of Guadeloupe

The Lesser Antillean macaw or Guadeloupe macaw (Ara guadeloupensis) is a hypothetical extinct species of macaw that is thought to have been endemic to the Lesser Antillean island region of Guadeloupe. In spite of the absence of conserved specimens, many details about the Lesser Antillean macaw are known from several contemporary accounts, and the bird may be the subject of some illustrations. It is therefore one of the best-documented species of extinct Caribbean macaw. Based on these accounts, Austin Hobart Clark named the species in 1905. Due to the lack of credible physical remains, doubts have been raised about the existence of this species. A Pleistocene phalanx bone since unearthed from the island of Marie-Galante confirmed the existence of a similar-sized macaw inhabiting the island prior to the arrival of humans, and was correlated with the Lesser Antillean macaw in 2015.

According to contemporary descriptions, the body was red and the wings were red, blue and yellow. The tail feathers were between 15 and 20 inches (38 and 51 cm) long. Apart from the smaller size and the all-red colouration of the tail feathers, it resembled the scarlet macaw of tropical South America and may therefore have been a close relative of that species. The bird ate fruit – including those of the poisonous manchineel tree, was monogamous, nested in trees and laid two eggs twice a year. Early writers described it as being abundant in Guadeloupe, but that it was becoming rare by 1760, and only survived in uninhabited areas. Disease and hunting by humans are thought to have eradicated it shortly after.

The Lesser Antillean macaw is one of thirteen extinct macaw species that have been proposed to have lived in the Caribbean islands. Many of these species are now considered dubious because only three are known from physical remains, and there are no extant endemic macaws on the islands today. Macaws were frequently transported between the Caribbean islands and the South American mainland in both prehistoric and historic times, so it is impossible to know whether contemporaneous reports refer to imported or native species.

Taxonomy

File:Lesser Antillean Macaw.jpg

Possible depiction by François-Nicolas Martinet, 1765

The Lesser Antillean macaw is well-documented compared to most other extinct Caribbean macaws, since it was mentioned and described by several contemporaneous writers. Parrots thought to be the Lesser Antillean macaw were first mentioned by the Spanish historian Gonzalo Fernández de Oviedo y Valdés in 1553, referring to a 1496 account by Ferdinand Columbus, who mentioned parrots as big as chickens, which the Caribs called "Guacamayas", in Guadeloupe. In 1774, Comte de Buffon also stated that Christopher Columbus had found macaws in Guadeloupe. Jean-Baptiste Du Tertre gave the first detailed descriptions in 1654 and 1676, and illustrated the bird and other animals found in Guadeloupe, in the island group of the Lesser Antilles. Jean Baptiste Labat also described the bird in 1742.

Austin Hobart Clark gave the Lesser Antillean macaw its scientific name, Ara guadeloupensis, in 1905, based on the contemporaneous accounts, but he also cited a 1765 colour plate as possibly depicting this species. He wrote that it was different in several features from the superficially similar scarlet macaw (Ara macao), as well as the green-winged macaw (Ara chloropterus) and the Cuban macaw (Ara tricolor). In 1967, ornithologist James Greenway wrote that the macaws reported from Guadeloupe could have been imported to the island from elsewhere by the natives, but this is impossible to prove. Greenway also suggested that the scarlet macaw and the Cuban macaw formed a superspecies with the Lesser Antillean macaw and other hypothetical extinct species suggested for Jamaica and Hispaniola. According to the palaeontologist Julian Hume, its similarity to the scarlet macaw indicates that they are close relatives, and that the Guadeloupe species may have descended from the mainland macaw.

A small parrot ulna found on the Folle Anse archaeological site on Marie-Galante, an island in the Guadeloupe region, was assigned to the Lesser Antillean macaw by the ornithologists Matthew Williams and David Steadman in 2001. In 2008, ornithologists Storrs Olson and Edgar Maíz López cast doubt upon this identification, and proposed that the bone instead belonged to the extant imperial amazon (Amazona imperialis). The size and robustness of the bone was similar to ulnae of the imperial amazon, and though it was worn, the authors identified what appeared to be a notch, which is also present on ulnae of the genus Amazona, but not in the genus Ara. In the same paper, they also argued that another hypothetical extinct, sympatric parrot, the Guadeloupe amazon (Amazona violacea), was identical to the imperial amazon. Subfossil remains from the island of Montserrat have also been suggested to belong to the Lesser Antillean macaw. The species was recognised by Birdlife International and the IUCN Red List until 2013, but was not considered valid thereafter.

In 2015, a terminal phalanx bone or ungual (claw bone) attributable to the genus Ara from Marie-Galante was described by Monica Gala and Arnaud Lenoble. It was discovered in the Blanchard Cave during excavations in 2013-2014, in a fossil-bearing deposit dating to the late Pleistocene epoch. The deposit was radiocarbon dated to about 10,690 years ago; the earliest evidence of human settlement in the area has been dated to 5.300 years ago. This confirmed that the Guadeloupe region once had an endemic macaw, which was not brought there as a result of human colonisation. All other macaw bones from the Lesser Antillean islands are from archaeological contexts, and could therefore have been the remains of birds brought there by Amerindians. The size of the phalanx bone matched what was described for the Lesser Antillean macaw by contemporary writers, and the authors therefore correlated the two, while admitting this could only be tentative, as there were no other remains of the bird to compare with.

Extinct Caribbean relatives

Edwards' Dodo, a 1626 painting by Roelant Savery, possibly showing this macaw on the left, and a Martinique macaw on the right.

Macaws are known to have been transported between the Caribbean islands and from mainland South America both in historic times by Europeans and natives, and prehistoric times by Paleoamericans. Parrots were important in the culture of native Caribbeans, were traded between islands, and were among the gifts offered to Christopher Columbus when he reached the Bahamas in 1492. It is therefore difficult to determine whether the numerous historical records of macaws on these islands mention distinct, endemic species, since they could have been escaped individuals or feral populations of foreign macaws of known species that had been transported there. As many as thirteen extinct macaws have at times been suggested to have lived on the islands until recently. In addition to the bone assigned to the Lesser Antillean macaw, only two endemic Caribbean macaw species are known from physical remains; the Cuban macaw is known from nineteen museum skins and subfossils, and the Saint Croix macaw (Ara autochthones) is known from subfossils. No endemic Caribbean macaws remain today; they were likely driven to extinction by humans in historic and prehistoric times.

1626 painting by Savery, showing similar red macaws with all-red tails

Many hypothetical extinct macaws were based only on contemporaneous accounts, but these species are considered dubious today. Several of them were named in the early 20th century by Walter Rothschild, who had a tendency to name species based on little tangible evidence. The red-headed macaw (Ara erythrocephala) and the Jamaican red macaw (Ara gossei) were named for accounts of macaws on Jamaica, the Martinique macaw (Ara martinica) from Martinique island and the Dominican green-and-yellow macaw (Ara atwoodi) was supposedly from Dominica island. The violet macaw (Anodorhynchus purpurascens), which was named for accounts of blue parrots also supposedly from Guadeloupe, is now thought to have been incorrectly based on the Guadeloupe amazon.

Other species of macaw have also been mentioned, but many never received binomials, or are considered junior synonyms of other species. Williams and Steadman defended the validity of most named Caribbean macaw species, and wrote that each Greater and Lesser Antillean island probably had its own endemic species. Olson and Maíz doubted the validity of the hypothetical macaws, and that all Antillean islands once had endemic species, but wrote that the island of Hispaniola would be the most likely place for another macaw species to have existed because of the large land area, though no descriptions or remains of such are known. They wrote that such a species could have been driven to extinction before the arrival of Europeans. The prehistoric distribution of indigenous macaws in the Caribbean can only be determined through further palaeontological discoveries.

Description

The Lesser Antillean macaw was described as having similar colouration to the scarlet macaw, but with shorter tail feathers between 15 and 20 inches (38 and 51 cm) long. The tail feathers of the scarlet macaw are 2 feet (61 cm) long and have blue tips, and the outer feathers are almost entirely blue. In spite of the tail feathers being shorter, it is not certain whether the Lesser Antillean macaw was smaller than the scarlet macaw overall, as the relative proportions of body parts vary between macaw species. The tail feathers were longer than those of the Cuban macaw, which were 12 inches (30 cm) long. The morphology of the fossil phalanx bone from Marie-Galante was most similar to the second or third ungual of the scarlet macaw, though slightly smaller at 15.3 mm (0.60 in) compared to 15–17 mm (0.59–0.67 in).

File:Ara Rouge.jpg

Variety of the 1765 plate

Du Tertre described the Lesser Antillean macaw as follows in 1654:

The Macaw is the largest of all the parrot tribe; for although the parrots of Guadeloupe are larger than all other parrots, both of the islands and of the main land, the Macaws are a third larger than they ... The head, neck, underparts, and back are flame color. The wings are a mixture of yellow, azure, and scarlet. The tail is wholly red, and a foot and a half long.

Two scarlet macaws, note the longer, blue tail feathers

Labat described the Lesser Antilles macaw in much the same way in 1742, while adding several details:

The feathers of the head, neck, back and underparts are flame colour; the wings are of a mixture of blue, yellow and red; the tail, which is from fifteen to twenty inches in length is wholly red. The head and the beak are very large, and it walks gravely; it talks very well, if it is taught when young; its voice is strong and distinct; it is amiable and kind, and allows itself to be caressed ...

Both authors wrote that the macaws were the largest parrots of Guadeloupe, and stressed that the parrot species of each Caribbean island was distinct and could be distinguished from each other visually and vocally. According to Hume, this means that the birds described could not simply have been escaped South American macaws. Furthermore, the docile and amiable nature described by Du Tertre and Labat does not match the behaviour of South American macaws.

Apart from Du Tertre's crude 1667 drawing and Labat's 1722 derivative, a few contemporaneous paintings depict red macaws that may be the Lesser Antilles macaw. A colour plate from Buffon's 1765 book of natural history (no. 12, entitled L'Ara Rouge) shows a red macaw with entirely red tail feathers and more red on the tertial and scapular feathers of the wing than are present on the scarlet macaw. Copies of the plate differ in the nuances used, but are identical in pattern. The painting suggests that a specimen may have been present in Europe at the time. Linnaeus cited the plate in his 1766 description of the scarlet macaw, but his description does not match the bird shown. A 1626 painting by Roelant Savery, which also includes a dodo, shows a red macaw that agrees with the descriptions of the Lesser Antillean macaw. A second macaw in the painting has been identified as being the hypothetical extinct Martinique macaw, but though many parrots were imported into Europe at the time from all over the world, it is impossible to determine the accuracy of such paintings today.

Behaviour and ecology

File:Guadeloupe Psittaciformes.Labat.jpg

Labat's 1722 illustration of a Guadeloupe amazon, a Guadeloupe parakeet, and a Lesser Antillean macaw

Du Tertre gave a detailed account of the behaviour of the Lesser Antillean macaw in 1654:

This bird lives on berries, and on the fruit of certain trees, but principally on the apples of the manchioneel (!), which is a powerful and caustic poison to other animals. It is the prettiest sight in the world to see ten or a dozen Macaws in a green tree. Their voice is loud and piercing, and they always cry when flying. If one imitates their cry, they stop short. They have a grave and dignified demeanor, and so far from being alarmed by many shots fired under a tree where they are perched, they gaze at their companions who fall dead to the ground without being disturbed at all, so that one may fire five or six times into the same tree without their appearing to be frightened.

Manchineel fruit, which was part of the diet of this parrot

In a 1667 work, Du Tertre gave a similar account, and also said that the macaw only ate the poisonous manchineel fruits in times of necessity. He also described its reproductive behaviour:

The male and the female are inseparable companions and it is rare that one is seen singly. When they wish to breed (which they do once or twice a year) they make a hole with their beaks in the stump of a large tree, and construct a nest with feathers from their own bodies. They lay two eggs, the size of those of a partridge (Perdix cinerea). The others of the parrot kind make their nests in the same way, but lay green eggs ... The Macaws are much larger than the large parrots of Guadeloupe or Grenada, and live longer than a man; but they are almost all subject to a falling sickness.

The twice-yearly breeding may have actually been staggered breeding, which is practised by some tropical birds.

Clark suggested that the Lesser Antillean macaw also occurred on Dominica and Martinique, but there is no evidence for this. Instead, it probably existed on other islands close to Guadeloupe. The fossil phalanx bone from Marie-Galante was deposited in a time when the distance between that island and the rest of the Guadeloupe archipelago was very reduced compared to today, due to lower sea-levels. The areas were separated by three channels, the largest of which was 6 kilometres (3.7 miles) wide. This would not have been a hindrance to flying animals, and the macaws of the Guadeloupe Islands would probably have been a single population during the Pleistocene.

Extinction

File:Ara guadeloupensis.jpg

Variety of the 1765 plate

In 1534, Johann Huttich wrote that the forests of Guadeloupe were full of this macaw, which was apparently as abundant as grasshoppers, and the natives of the island cooked it together with human flesh and that of other birds. In 1654, Du Tertre stated that the flesh was tough to eat, and that some considered it unpalatable and even poisonous. He wrote that he and the other inhabitants of the island often consumed it, and that he experienced no ill effects from it. He also stated that the natives wore the feathers as head decoration and as moustaches through the septum of the nose. He described how it was hunted by the natives:

The natives make use of a stratagem to take them alive; they watch for a chance to find them on the ground, eating the fruit which has fallen from the trees, when they approach quietly under cover of the trees, then all at once run forward, clapping their hands and filling the air with cries capable not only of astounding the birds, but of terrifying the boldest. Then the poor birds, surprised and distracted, as if struck with thunderbolt, lose the use of their wings, and, making a virtue of necessity, throw themselves on their backs and assume defensive with the weapons nature has given them - their beaks and claws - with which they defend themselves so bravely that not one of the natives dares to put his hand on them. One of the natives bring a big stick which he lays across the belly of the bird, who seizes it with beak and claws; but while he is occupied in biting it, the native ties him so adroitly to the stick that he can do with him anything he wishes ...

Since Du Tertre wrote that the macaws were prone to sickness, an outbreak of a disease, along with hunting, may have contributed to its demise. In 1760, Mathurin Jacques Brisson quoted a letter by M. de la Borde, which stated that macaws had become very rare in the Antillean islands because they were hunted for food. By then they could by only be found in areas not frequented by man, and were probably extinct soon after. Parrots are often among the first species to be exterminated from a given locality, especially islands.