Rodrigues solitaire
Rodrigues Solitaire Temporal range: Recent
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The only known picture drawn by someone who saw the bird in life, François Leguat, 1708 | |
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Genus: | Pezophaps Strickland, 1848
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Species: | P. solitaria
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Pezophaps solitaria (Gmelin, 1789)
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The Rodrigues Solitaire (Pezophaps solitaria) is an extinct, flightless bird that was endemic to the Mascarene island of Rodrigues, east of Madagascar in the Indian Ocean. It was genetically related to pigeons and doves, and its closest relative was the likewise extinct Dodo of Mauritius, the two forming the Raphinae subfamily. The closest extant relative is the Nicobar Pigeon.
It was the size of a swan and showed pronounced sexual dimorphism. Males were much larger than females, measuring up to 90 in length and weighing 28 kg, whereas females measured 70 cm and weighed 17 kg. It was grey and brown, but the female was lighter than than males. Both sexes were highly territorial and possessed large bony knobs on their wings, used in combat.
The Rodrigues Solitaire was first mentioned in the 17th century, and was described in detail by François Leguat, leader of a group of French Huguenot refugees who were marooned on Rodrigues from 1691. It was hunted by humans and species introduced by them, and was extinct by the mid-1700s as a result. Apart from Leguat's account and drawing, as well as a few other contemporary descriptions, nothing was known about the bird until a few subfossil bones were found in caves in 1789. Thousands of bones were subsequently excavated. It is the only extinct bird with a star constellation named after it, Turdus Solitarius.
Taxonomy and evolution
François Leguat was the first to refer to the bird as the "Solitaire", but it has been suggested he borrowed the name from a tract mentioning the Réunion Solitaire.[2] The bird was first scientifically named as a species of Dodo, Didus solitarius, based on Leguat's description, by Johann Friedrich Gmelin in his edition of Systema Naturae.[3] The specific name is a reference to the bird's solitary habits.
Hugh Edwin Strickland suggested the common descent of the Solitaire and the Dodo in 1848 with Alexander Gordon Melville, after dissecting the only known Dodo specimen with soft tissue, and comparing them with the few Solitaire remains then available. Strickland recognized its generic distinction and named the new genus Pezophaps, meaning "pedestrian pigeon" in Greek.[4] The difference between the sexes of the bird were so large that Strickland thought they belonged to two species, naming the smallest Pezophaps minor.[5]
For a long time, the Dodo and the Rodrigues Solitaire, collectively termed "didines", were placed in a family of their own, the Raphidae. This was because their relationships to other groups of birds, such as rails, was unresolved. Later, more evidence led to Raphidae's demotion to a subfamily, within the Columbidae.[6]
Comparison of mitochondrial cytochrome b and 12S rRNA sequences isolated from a femur of a Rodrigues Solitaire and tarsal of a Dodo confirms their close relationship.[7] Genetic evidence has also been interpreted to show that the Nicobar Pigeon is their closest living relative, followed by the Crowned Pigeons of the genus Goura, and the superficially Dodo-like Tooth-billed Pigeon from Samoa.[8] The Mascarene Islands, a group of islands in the Indian Ocean east of Madagascar, which includes Mauritius and Réunion, are of volcanic origin and less than 10 million years old, so the ancestors of both birds likely remained capable of flight for some considerable time after the separation of their lineages.[9]
There exist a few other reports of "Solitaires" from the Mascarenes, as the term was also used for other species with "solitary" habits, such as the enigmatic Oiseau Bleu and the Réunion Solitaire. Some scientists believed that Réunion was the home of not only a white Dodo, but also a white Solitaire, both of which are now believed to be results of misinterpretations of old reports.[10] Walter Rothschild had a restoration made of the Réunion Solitaire which was basically a white Rodrigues Solitaire, based on this speculation.[11] An atypical 17th century description of a Dodo and bones found on Rodrigues, now known to have belonged to the Rodrigues Solitaire, led to the naming of a new species, Didus nazarenus.[12] Being based on Solitaire remains, it is now a junior synonym of this species.[13]
Description
One observer described the Rodrigues Solitaire as the size of a swan.[14] Sexual size dimorphism in this species is perhaps the greatest in any carinate bird.[15] Males were considerably larger than the females, measuring 90 cms in length (3 ft) and weighing up to 28 kg, whereas the females were 70 cms and weighed 17 kg.[16] The weight may have varied substantially due to fat-cycles, and may have been as low as 21 kgs in males and 13 in females.[15] The plumage was grey and brown, and the females were paler than males, having light elevations on the lower neck. There was a black band near the base of the beak.[14] The beak was slightly hooked, and its neck and legs were long.[11] The skull was 170 mms long, flattened at the top, with the fore and hind part elevated into two bony ridges of cancellous bone.[17]
Both sexes possessed a large callus-like knob of bone on their wrists at the base of the carpometacarpus, on its otherwise small wings.[3] It was largest in males, up to the size of a musket bullet, and very little developed in females. In life this knob would have been covered by keratinous integument and used as a weapon.[5] Similar features are also known from other pigeon species.[18]
Most of the skeletal features of the Solitaire that are unique among pigeons have been attributed to its flightlessness. The leg bones were broader and the digits were shorter than those of flighted pigeons, and the skull, trunk and pelvic limbs were peramorphic, meaning that they changed considerably with age. However, the pectoral region and the wings were paedomorphic, meaning they were underdeveloped and retained juvenile features.[15]
Contemporary descriptions
Apart from François Leguat's rather crude depiction, the life appearance of the Rodrigues Solitaire is only known from a handful of descriptions, as no soft tissue remains are known.[16] Clearly impressed by the bird, Leguat devoted three pages of his memoirs to the Solitaire.[9] He described the appearance of the bird as follows:
Of all the Birds in the Island the most remarkable is that which goes by the name of the solitary, because it is very seldom seen in company, tho’ there are abundance of them. The Feathers of the Males are of a brown grey Colour: the Feet and Beak are like a Turkey’s, but a little more crooked. They have scarce any Tail, but their Hind-part covered with Feathers is roundish, like the Crupper [rump] of a horse; they are taller than Turkeys. Their Neck is straight, and a little longer in proportion than a Turkey’s when it lifts up his Head. Its Eye is Black and lively, and its Head without Comb or Cop. They never fly, their Wings are too little to support the Weight of their Bodies; they serve only to beat themselves, and flutter when they call one another. They will whirl about for twenty or thirty times together on the same side, during the space of four or five minutes. The motion of their Wings makes then a noise very like that of a Rattle; and one may hear it two hundred Paces off. The Bone of their Wing grows greater towards the Extremity, and forms a little round Mass under the Feathers, as big as a Musket Ball. That and its Beak are the chief Defence of this Bird. ‘Tis very hard to catch it in the Woods, but easie in open Places, because we run faster than they, and sometimes we approach them without much Trouble. From March to September they are extremely fat, and taste admirably well, especially while they are young, some of the Males weigh forty-five Pounds.[11]
Several of Leguat's observations have been confirmed through study of subfossil Solitaire remains; apart from the wrist-knob described by him, there is also a rugose surface at the base of the beak indicating the position of the caruncular ridge, which Leguat described as a "widow's peak". The curved contour lines of the pelvis also support the roundness of its hind parts he compared to that of a horse.[3]
Leguat continued with an elaborate description of the female Solitaire, which also appears to be the sex depicted in his illustration of the bird:
The Females are wonderfully beautiful, some fair, some brown; I call them fair, because they are the colour of fair Hair. They have a sort of Peak, like a Widow’s upon their Breasts [Beaks], which is of a dun colour. No one Feather is straggling from the other all over their Bodies, they being very careful to adjust themselves, and make them all even with their Beaks. The Feathers on their Thighs are round like Shells at the end, and being there very thick, have an agreeable effect. They have two Risings on their Craws [crop] and the Feathers are whiter than the rest, which livelily represents the fine neck of a Beautiful Woman. They walk with so much Stateliness and good Grace, that one cannot help admiring them and loving them; by which means their fine Mein often saves their Lives.[11]
It has been proposed that Leguat's comparison between the crop of the female Solitaire and the "beautiful bosom of a woman" (changed to "fine neck" in some editions of his memoirs) was out of longing for female companionship.[18]
Another description of appearance and behaviour is found in an anonymous document rediscovered in 1874 called Relation de l'Ile Rodrigue, which has been attributed to a man named Julien Tafforet, who was marooned on Rodrigues in 1726:
The solitaire is a large bird, which weighs about forty or fifty pounds. They have a very big head, with a sort of frontlet, as if of black velvet. Their feathers are neither feathers nor fur; they are of light grey colour, with a little black on their backs. Strutting proudly about, either alone or in pairs, they preen their plumage or fur with their beak, and keep themselves very clean. They have their toes furnished with hard scales, and run with quickness, mostly among the rocks, where a man, however agile, can hardly catch them. They have a very short beak, of about an inch in length, which is sharp. They, nevertheless, do not attempt to hurt anyone, except when they find someone before them, and, when hardly pressed, try to bite him. They have a small stump of a wing, which has a sort of bullet at its extremity, and serves as a defence.[11]
Behaviour and ecology
Observations of the solitaire indicate that breeding pairs were highly territorial; presumably they settled disputes by striking each other with the wings. To aid this purpose, they used the knobs on their wrists. Fractures in their wing-bones also support that they were used in combat.[15] It has been suggested that the knobs was perhaps formed through continuous injuries, as they resemble diseased bone.[3] Several accounts also state that they defended themselves with a powerful bite. The species perhaps lived primarily in the island's woodlands, rather than on the shores.[14]
The most detailed account of their habits is that of Leguat. He described the mating and nesting as follows:
When these Birds build their Nests, they choose a clean Place, gather together some Palm-Leaves for that purpose, and heap them up a foot and a half high from the Ground, on which they sit. They never lay but one Egg, which is much bigger than that of a Goose. The Male and Female both cover it in their turns, and the young is not hatch’d till at seven Weeks’ end: All the while they are sitting upon it, or are bringing up their young one, which is not able to provide itself in several Months, they will not suffer any other Bird of their Species to come within two hundred Yards round of the Place; But what is very singular, is, the Males will never drive away the Females, only when he perceives one he makes a noise with his Wings to call to the Female, and she drives the unwelcome Stranger away, not leaving it till ‘tis without her Bounds. The Female do’s the same as to the Males, whom she leaves to the Male, and he drives them away. We have observ’d this several Times, and I affirm it to be true.
The Combats between them on this occasion last sometimes pretty long, because the Stranger only turns about, and do’s not fly directly from the Nest. However, the others do not forsake it till they have quite driven it out of their Limits. After these Birds have rais’d their young One, and left it to itself, they are always together, which the other Birds are not, and tho’ they happen to mingle with other Birds of the same Species, these two Companions never disunite. We have often remark’d, that some Days after the young leaves the Nest, a Company of thirty or forty brings another young one to it, and the now fledg’d Bird, with its Father and Mother joyning with the Band, march to some bye Place. We frequently follow’d them, and found that afterwards the old ones went each their way alone, or in Couples, and left the two young ones together, which we call’d a Marriage.[11]
Due to the clutch being described as consisting of a single egg, and the large size of the bird itself, it has been proposed that the Solitaire was K-selected, meaning that it produced a low number of altricial offspring, which required extensive parental care until they matured. The gathering of unrelated juveniles suggests that they formed crèches, which may have followed foraging adults as part of the learning process.[15]
Due to the size difference it has been suggested that the Solitaire was not monogamous as stated by Leguat, but that he, as a deeply religious man, attributed the trait to the bird for moral reasons.[5] It has instead been proposed that it was polygynous, and that the wing-rattling behaviour described for males suggests lek-mating, where males gather for competitive mating display.[15] However, it has been pointed out that size dimorphism does occur in some monogamous birds, and most other pigeons are monogamous as well.[16]
Tafforet's account of the behaviour confirms Leguat's description, adding that Solitaires would even attack humans approaching their chicks:
They do not fly at all, having no feathers to their wings, but they flap them, and make a great noise with their wings when angry, and the noise is something like thunder in the distance. They only ly, as I am led to suppose, but once in the year, and only one egg. Not that I have seen their eggs, for I have not been able to discover where they lay. But I have never seen but one little one alone with them, and, if any one tried to approach it, they would bite him very severely. These birds live on seeds and leaves of trees, which they pick up on the ground. They have a gizzard larger than the fist, and what is surprising is that there is found in it a stone of the size of a henn's egg, of oval shape, a little flattened, although this animal cannot swallow anything larger than a small cherry-stone. I have eaten them: they are tolerably well tasted.[11]
Pierre-André d'Héguerty, writing about his time at the island around 1735, described how the Solitaire behaved in captivity:
One finds there [in Rodrigues] birds of various species that are commonly taken by giving chance, and amongst others Solitaires which have almost no wing feathers. It is a bird bigger than a swan, with a melancholy appearance. Tamed [probably meaning 'captive'], one sees them always walking the same line till they run out of space, then retracing their steps without deviating. Cut open, one normally finds Bézpards which are valued and useful in medicine.[9]
Diet
Leguat stated the Rodrigues Solitaire fed on dates, whereas Tafforet mentioned seeds and leaves. Apart from those, no other accounts mention the diet.[14] It has been suggested it ate latan palm fruits, and competed with the now extinct Cylindraspis tortoises for these. It is not known how the young were fed, but related pigeons provide crop milk.[9]
Several of the contemporary accounts state that the Solitaire used gizzard stones. Dodos also did this, which may imply a similar diet.[9] Leguat described the stones in the following passage, while also mentioning that Solitaires refused to feed in captivity:
Tho’ these Birds will sometimes very familiarly come up near enough to one, when we do not run after them, yet they will never grow Tame. As soon as they are caught they shed Tears without Crying, and refuse all sustenance till they die.
We find in the Gizzards of both Male and Female, a brown Stone, of the bigness of a Hen’s Egg, ‘tis somewhat rough, flat on one side and round on the other, heavy and hard. We believe this Stone was there when they were hatched, for let them be never so young, you meet with it always. They never have but one of ‘em, and besides, the Passage from the Craw to the Gizard is so narrow, that a like Mass of half Bigness cou’d not pass. It serv’d to whet our Knives better than any other Stone Whatsoever.[11]
In 1877, three stones were found in a cavern on Rodrigues, each in close association with a Solitaire skeleton. These were inferred to be the gizzard stones mentioned by Leguat. One of the stones was examined and found to be dolerite, somewhat rough, hard and heavy (ca. 50 g.), but hardly flat on one side as described by Leguat. This could be due to it being associated with a young individual.[5] Though Leguat stated the bird hatched with the gizzard stone already inside, in reality, the adults most likely fed stones to the hatchlings.[9]
Relationship with humans
Sir Thomas Herbert was the first to mention that "Dodos" lived on Rodrigues in 1634, most likely referring to the Solitaire, and "Dodos" were again mentioned as being present there in 1700.[18]
The next account, and the first referring to the bird as the "Solitaire", was published in the memoirs of François Leguat in 1708, titled A New Voyage to the East Indies.[19] Leguat was the leader of a group of nine French Huguenot refugees who were the first to colonise the island, from 1691 to 1693, after having been abandoned there by their captain. He described the bird in some detail, including its solitary nesting behaviour. The Huguenots praised the birds for their flavour, especially the young ones, and used their gizzard stones as knife sharpeners.[20] Leguat's observations are considered some of the first cohesive accounts of animal behaviour in the wild. He later left for Mauritius, but too late to observe Dodos there.[9]
Many of the old accounts mention that Solitaires were hunted by man. Writing in 1735, Gennes described the capture and consumption of two specimens as follows:
Our men told of having seen goats and a large quantity of birds of different kinds: they brought, amongst others, two of which were bigger by a third than the largest turkey; they appeared, nevertheless to be still quite young, still having down on the neck and head; their wingtips were but sparsely feathered, without any proper tail. Three sailors told me of having seen two others, of the same species, as big as the biggest ostrich. The young ones that were brought had the head made more or less like the latter animal, but their feet were similar to those of turkeys, instead of that of the ostrich which is forked and cloven in the shape of a hind's foot. These two birds, when skinned, had an inch of fat on the body. One was made into a pie, which turned out to be so tough that it was uneatable.[9]
Unlike the Dodo, no Rodrigues Solitaires are known to have been sent to Europe alive. However, it has been claimed that Bertrand-François Mahé de La Bourdonnais sent a "Solitaire" to France from the nearby island of Réunion around 1740. Since the Réunion Solitaire is believed to have gone extinct by this date, the bird may actually have been a Rodrigues Solitaire.[9]
Extinction
When Joseph-François Charpentier de Cossigny attempted to get a specimen in 1755, none could be found:
For 18 months I have been trying without success to procure a solitaire from Rodrigues Island... I have promised all one could want, in spirits or piastres, to whoever brings me at least one alive. It is claimed that cats, which have gone wild on this little island, have destroyed this species of bird that only has stumps for wings, but I am strongly inclined to believe that these cats are the men of the post who have eaten all those they have found, as they are good to eat. At last, I have been given hope of obtaining one, which, so it is said, has been spotted.[9]
The Rodrigues Solitaire probably became extinct sometime between the 1730s and 1760s, the exact date is difficult to determine. Its disappearance can be correlated with the tortoise trade from 1730-1750, as the traders burnt off vegetation, hunted the Solitaires, and brought cats and pigs, which preyed on eggs and chicks.[9]
Alexandre Guy Pingré did not encounter any Solitaires when he visited Rodrigues to observe the 1761 transit of Venus, though he had been assured they still survived. His friend Pierre Charles Le Monnier named the star constellation Turdus Solitarius after the bird, to commemorate the journey. The Solitaire is therefore the only extinct bird with a constellation named after it, but celestial mapmakers did not know what it looked like, so star maps depict other birds.[18]
Some later scholars doubted Leguat's story, and that the Solitaire had ever existed.[18] In 1789, sub-fossil Solitaire bones encrusted in stalagmite were first discovered in a cave.[21] Fossils were also recovered in the 1860s, but more complete remains were found during the 1874 transit of Venus, since an observation station was located on Rodrigues.[5] Many of these excavations were requested by Edward and Alfred Newton, who proceeded to describe the osteology of the bird in detail. Thousands of bones were excavated, and mounted skeletons were composited from the remains of several specimens.[22] These finds confirmed Leguat's descriptions, but at this time, no living residents of Rodrigues remembered having seen living Solitaires.[9] In 1831, a man who had lived on Rodrigues for 40 years stated he had never seen birds large enough to be Solitaires. Rodrigues is only 104 square km large, so it is implausible that the bird would have survived for so long undetected.[14]
See also
References
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instead. - ^ Strickland, H.E.; Melville, A. G. (1848). The Dodo and Its Kindred; or the History, Affinities, and Osteology of the Dodo, Solitaire, and Other Extinct Birds of the Islands Mauritius, Rodriguez, and Bourbon. London: Reeve, Benham and Reeve.
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- ^ a b c d e f g h i j k l Cheke, A. S.; Hume, J. P. (2008). Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues. T. & A. D. Poyser. ISBN 978-0-7136-6544-4.
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instead. - ^ a b c d e Fuller, E. (2002). Dodo – From Extinction To Icon. London: HarperCollins.
- ^ Leguat, F. (1708): Voyages et Avantures de François Leguat & de ses Compagnons, en Deux Isles Desertes des Indes Orientales, etc. 2: 71. Jean Louis de Lorme, Amsterdam. [1]
- ^ Ellis, Richard (2004). No Turning Back: The Life and Death of Animal Species. New York: Harper Perennial. p. 161. ISBN 0-06-055804-0.
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instead. - ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1080/00222937808682294, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
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instead.