Hoopoe starling
| Hoopoe starling | |
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| Only known life drawing, showing the natural position of the crest, by Paul Jossigny, early 1770s | |
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| Genus: | Fregilupus Lesson, 1831
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| Species: | F. varius
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| Binomial name | |
| Fregilupus varius (Boddaert, 1783)
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| Location of Réunion (encircled) | |
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The hoopoe starling (Fregilupus varius), also known as the Réunion starling, Bourbon crested starling, or crested starling, is an extinct species of starling that lived on the Mascarene island of Réunion. Its closest relatives were the Rodrigues starling and the Mauritius starling from nearby islands, and the three appear to be of Southeast Asian origins. The bird was first discovered in the 17th century, and was long thought related to the hoopoe, from which its name is derived. Various affinities were proposed since, but it was confirmed to be a starling in a DNA study.
The hoopoe starling was 30 cm (12 in) in length, its plumage was mainly white and grey, with the back, wings and tail being a darker brown and grey. It had a light, mobile crest, which curled forwards. The bird is thought to have been sexually dimorphic, with males being larger and having a more curved beak. The juveniles were browner than adults. Little is known about the behaviour of the hoopoe starling. It was reported to have lived in large flocks, and inhabited humid areas and marshes. It was omnivorous, and fed on plant matter and insects. Its pelvis was robust, its feet and claws were large, and its jaws strong, which indicates it foraged near the ground.
The bird was hunted by settlers on Réunion, who also kept them as cagebirds. Today, 19 specimens exist in various museums around the world. The hoopoe starling was reported to be in decline by the early 19th century, and it was probably extinct before the 1860s. Various factors have been proposed, including competition and predation by introduced species, disease, deforestation, and persecution by humans, who hunted it for food and for allegedly being a crop pest.
Taxonomy
The first account thought to mention the hoopoe starling is a 1658 list of the birds of Madagascar, written by French governor Étienne de Flacourt. He mentioned a black and grey "tivouch", or hoopoe, but some later authors have doubted whether this referred to the hoopoe starling, or the Madagascan subspecies of the hoopoe (Upupa epops marginata), though that bird does not look much different from the Eurasian subspecies. The hoopoe starling was first mentioned from the island of Réunion (then called "Bourbon") by Père Vachet in 1669, but not in detail until the following account by Sieur Dubois in 1674:[2]
Hoopoes or 'Calandres', having a white tuft on the head, the rest of the plumage white and grey, the bill and the feet like a bird of prey; they are a little larger than the young pigeons. This is another good game [i.e., to eat] when it is fat.[3]
Early settlers on Réunion referred to the bird as "huppe", due to the similarity of its crest and curved bill with that of the hoopoe. Little was recorded about the hoopoe starling the next hundred years, but specimens began to be transported to Europe during the 18th century. The species was first scientifically described by French naturalist Philippe Guéneau de Montbeillard in the 1779 edition of Comte de Buffon's Histoire Naturelle, but it did not receive its scientific name until named by Dutch naturalist Pieter Boddaert in the 1783 edition of the book. Boddaert named it Upupa varia; the genus name is that of the hoopoe, and the specific name means variegated, in reference to the black and white colouration. Boddaert provided Linnean binomial names for plates in Buffon's works, so the accompanying 1770s plate of the hoopoe starling by François-Nicolas Martinet is considered the holotype, or type illustration. The plate itself could have been based on a specimen in the Muséum National d'Histoire Naturelle de Paris, but this is impossible to determine today. The Paris museum originally had five skins of the hoopoe starling, some which only arrived there in the 19th century. The possibly female specimen MNHN 2000-756, which is one of the most often illustrated skins, has an artificially trimmed crest, resulting in an unnaturally semi-circular shape, unlike its appearance in life; the type illustration has a similarly shaped crest.[2]
De Flacourt's "tivouch" led early writers to believe that variants of the bird were found on Madagascar and the Cape of Africa. They were thought to be hoopoes of the Upupa genus, which received names such as Upupa capensis and Upupa madagascariensis. In addition, various authors allied it with different bird groups, such as birds of paradise, bee-eaters, cowbirds, Icteridae, and choughs, which resulted in the bird being reassigned to other genera, thereby receiving new names such as Coracia cristata and Pastor upupa in the process. In 1831, René-Primevère Lesson placed the bird in its own monotypic genus, Fregilipus, a composite word of Upupa and Fregilus, the latter a now defunct genus name of the chough. Only in 1868 did Auguste Vinson establish that the bird was restricted to the island of Réunion, and proposed the new binomial Fregilupus borbonicus, in reference to the island.[2]
In 1857, Hermann Schlegel first proposed that the species belonged to the starling family (Sturnidae), and reclassified it as part of the Sturnus genus; S. capensis. The reclassification was followed by other writers, and in 1872, Carl Jakob Sundevall proposed the new genus name Lophopsarus, meaning "crested starling", but Fregilupus varius remains the valid binomial today, as it is oldest, and all other scientific names are synonyms.[2] In 1874, after detailed analysis of the only known skeleton, held at the Cambridge University Museum of Zoology, James Murie agreed that it was a starling.[4] In 1890, Richard Bowdler Sharpe stated the hoopoe starling was similar to the starling genus Basilornis, but did not note any similarities other than both having a crest.[5] In 1941, Malcolm R. Miller dissected a specimen preserved in spirits at the Cambridge Museum, and found its musculature similar to that of the common starling (Sturnus vulgaris), but noted that the tissue was very degraded, and that these similarities did not necessarily confirm a relationship with starlings.[6] In 1957, Andrew John Berger cast doubt on its affinity with starlings, due to subtle anatomical differences, after dissecting a spirit specimen at the American Museum of Natural History.[7] Some authors also proposed a relationship with vangas (Vangidae), but Hiroyuki Morioka rejected this in 1996, after a comparative study of skulls.[8]
In an 1875 article, the British ornithologist Alfred Newton attempted to identify a black and white bird mentioned in a 18th century manuscript, describing a marooned sailors stay on the Mascarene island of Rodrigues in 1726-1727, and hypothesised it was related to the hoopoe starling.[9] subfossil bones were later found on Rodrigues, which were correlated with the bird mentioned in the old manuscript. In 1879, the bones became the basis for a new species, Necropsar rodericanus, the Rodrigues starling, named by ornithologists Albert Günther and Edward Newton. They found the Rodrigues bird to be closely related to the hoopoe starling, and only kept it in a separate genus due to what they termed "present ornithological practice".[10] In 1967, American ornithologist James Greenway suggested that the Rodrigues starling belonged in the same genus as the hoopoe starling, due to their similarity.[11] Further subfossils found in 1974 confirmed that the Rodrigues bird was a distinct genus of starling.[12] The stouter bill is mainly what warrants generic separation from Fregilupus.[13] In 2014, British palaeontologist Julian P. Hume described a new extinct species, the Mauritius starling (Cryptopsar ischyrhynchus), based on subfossils from Mauritius. It was shown to be closer to the Rodrigues starling than to the hoopoe starling in skull, sternum and humerus features.[2]
Evolution
In 1943, Dean Amadon suggested that Sturnus-like species could have arrived in Africa, and given rise to the wattled starling (Creatophora cinerea) and the Mascarene species. According to Amadon, the Rodrigues and hoopoe starlings were related to Asiatic starlings such as some species of Sturnus, rather than the glossy starlings (Lamprotornis) of Africa and the Madagascan starling (Saroglossa aurata), based on their colouration.[14][15] A 2008 study which analysed the DNA of various starlings confirmed that the hoopoe starling belonged in a clade of Southeast Asian starlings, but as an isolated lineage with no close relatives. The following cladogram shows its position:[16]
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An earlier attempt by another team could not extract viable hoopoe starling DNA.[17] The authors of the successful study suggested that the ancestors of the hoopoe starlings reached Réunion from Southeast Asia by using island chains as "stepping stones" to cross the Indian Ocean, a scenario which has also been suggested for other Mascarene birds. The hoopoe starling lineage diverged from other starlings 4 million years ago, which is about two million years before Réunion emerged from the sea, so its initial evolution may have happened on now partially submerged landmasses.[16] Extant starlings such as the Bali myna (Leucopsar rothschildi) and the white-headed starling (Sturnia erythropygia), have similarities with the extinct Mascarene species in colouration and other features. As the Rodrigues and Mauritius starlings seem to be morphologically closer to each other than to the hoopoe starling, which instead appears closer to Southeast Asian starlings, there may have been two separate colonisations of starlings from Asia to the Mascarenes, with the hoopoe starling being the latest arrival. Apart from Madagascar, the Mascarenes were the only islands in the Southwest Indian Ocean with native starlings. This is probably due to the isolation, varied topography, and vegetation of these islands.[2]
Description
The hoopoe starling was 30 cm (12 in) in length. The culmen was 41 mm (1.6 in) long, the wing 147 mm (5.8 in), the tail 114 mm (4.5 in), and the tarsus was about 39 mm (1.5 in) long.[18] It was the largest of the three Mascarene starlings. A supposed adult male (NHMUK 1889.5.30.15) in the Paris museum has a light ash-grey head and back of the neck, lighter on the hind-neck, and the elongate crest is the same colour with white shafts. The back and tail are ash-brown, the wings darker with a greyish wash, and the uppertail covert feathers and rump with a rufous wash. The primary coverts are white with brown tips, though the bases are brown instead of the tips in some other specimens. The superciliary stripe, lore, and most of the underside of the specimen is white, with a pale rufous wash on the flanks and undertail coverts. The extent of the light rufous on the underside varies across specimens. The beak and legs are lemon-yellow with yellowish-brown claws. It has a bare, triangular area of skin around the eye, which may have been yellow in life. The iris of the species was described as bluish brown, but has been depicted as brown, yellow or orange.[2]
There has been historical confusion over which characteristics that differed between the sexes. Only three specimens were sexed, all males, with age and individual variation not being taken into account. The male was generally thought largest, with a longer and more curved beak. In 1911, Réunion resident Eugène Jacob de Cordemoy recalled his observations of the bird about 50 years before, and suggested that only the male had a white crest, but this is thought to be an error. A supposed female (MNHN 2000-756) in the Paris museum appears to have a smaller crest, with a smaller and less curved beak, and reduced primary coverts. A juvenile specimen has a smaller crest and primary coverts, with a brown wash instead of ash-grey on the crest, lore, and superciliary stripe, and light brown back instead of ash-brown. The juveniles of some southeast Asian starlings are also browner than the adults.[2]
Vinson, who had observed live hoopoe starlings when residing on Réunion, described the the crest as being composed of flexible, disunited, forwards curled barbs of various lengths, highest at the middle. The crest was mobile, and could be erected according to the mood of the bird. Vinson compared the crest to that of a cockatoo and the tail feathers of a bird of paradise. Most mounted specimens also have erect crests, which indicates this was the natural position. The only illustration of the hoopoe starling now thought to have been made from life is one by Paul Philippe Sauguin de Jossigny drawn in the early 1770s. Under the drawing, Jossigny noted to engravers that for accuracy, they should depict the crest as angled forwards from the head, not straight up. Hume believes that this was followed when the type illustration was drawn by Martinet, and that it was derivative of Jossigny's image, therefore not a life drawing. Murie had suggested that only the illustrations by Martinet and Jacques Barraband were drawn from life, but he was unaware of Jossigny's drawing. Jossigny also made the only known life drawing of the now extinct Newton's parakeet (Psittacula exsul), a specimen which had been sent to him on Mauritius, so this is perhaps also where he drew the hoopoe starling.[2]
Behaviour and ecology
Little is known about the behaviour of the hoopoe starling. In his 1807 account of the bird, François Levaillant included information received from a resident of Réunion, and stated it was abundant, lived in large flocks, and inhabited humid areas and marshes. In 1831, René Lesson claimed that its habits were similar to those of a crow, but without clarifying what was meant.[2] Vinson's 1877 account describes his experience with the bird more than fifty years earlier:
Now these daughters of the wood, when they were numerous, flew in flocks and went thus in the rain forests, while deviating little from one another, as good companions or as nymphs taking a bath: they lived on berries, seeds and insects, and the créoles, disgusted by the latter fact, held them for an impure game. Sometimes, coming from the woods to the littoral [coast], always flying and leaping from tree to tree, branch to branch, they often alighted in swarms on coffee trees in bloom, and there was in the past the testimony of an inhabitant of the Island of Bourbon, said the naturalist Levaillant, that they caused big damage in coffee trees by making the flowers fall prematurely. But it is not the white flowers of coffee that the hoopoes were searching for and thus behaving so, it was for the caterpillars and insects that devoured them; and in this they made an important service to the silviculture of the Island of Bourbon and the rich coffee plantations, with which this land was then covered, the golden age of the country![2]
Like most other starlings, the hoopoe starling was omnivorous, and fed on fruits, seeds, and insects. Its tongue was long, slender, sharp, and frayed, the bird may have been able to move it rapidly, which helped when feeding on fruit, nectar, pollen, and invertebrates. Its pelvic elements were robust and its feet and claws were large, which indicates it foraged near the ground. Its jaws were strong, and Morioka compared its skull to that of the hoopoe, so it may have foraged in a similar way, probing and opening holes in substrate by inserting the beak and gaping. De Montbeillard received information about the stomach contents of a dissected specimen, which consisted of seeds and the berries of "Pseudobuxus", which may refer to Eugenia buxifolia, a small tree or bush which has sweet, orange berries. He added that the bird weighed 4 oz (113 g), but that it was fatter around June and July. Several accounts suggest that the hoopoe starling migrated within Réunion, spending six months in the lowland, and six months in the mountains. Food may have been easier to obtain in the lowlands during austral winter, whereas the birds may have bred in the mountain forests during austral summer. The bird probably nested in tree cavities. Its song was described as a "bright and cheerful whistle", and as "clear notes", which indicates it was similar to that of other starlings.[2]
Many other endemic species of Réunion became extinct after the arrival of man and the resulting disruption of the island's ecosystem. The Mascarene parrot lived alongside other recently extinct birds such as the Réunion ibis, the Mascarene parrot, the Réunion parakeet, the Réunion swamphen, the Réunion owl, the Réunion night heron, and the Réunion pink pigeon. Extinct Réunion reptiles include the Réunion giant tortoise and an undescribed Leiolopisma skink. The small Mauritian flying fox and the snail Tropidophora carinata lived on Réunion and Mauritius but vanished from both islands.[19]
Relationship with humans
The hoopoe starling was described as being tame and easily hunted; in 1704, Feuilley explained how birds were caught, and that they were also predated by cats:
Hoopoes and merles [Hypsipetes borbonicus] are the same fatness as those in France, and are of a marvellous taste, which are fat at the same time as parrots, living on the same foods. In order to catch them, hunting was done with staffs or long thin poles from six to seven feet in length, though this hunt is infrequently seen. The marrons [escaped] cats destroy many. These birds allow themselves to be approached very closely, so the cats take them without leaving their places.[2]
The hoopoe starling was kept as a cagebird on both Réunion and Mauritius, and numerous specimens were procured during the early 19th century, though the bird was becoming scarcer. It is unknown whether any live specimens were ever transported away from the Mascarenes. Cordemoy recalled that captive birds could be fed a wide variety of food, such as bananas, potatoes, and chayote, and that wild birds would never enter inhabited areas. Many individuals survived on Mauritius after escaping there, and it was thought possible that a feral population would be established. The Mauritian population lasted less than a decade, and the last specimen on the island was taken in 1836, the last definite record of a live specimen anywhere. Specimens could still be collected on Réunion in the 1830s, possibly into the early 1840s.[2]
Today, there are 19 surviving hoopoe starling skins scattered across various museums around the world (including the single skeleton and two specimens preserved in spirit), two are in the Paris museum and four in Troyes. Thee additional skins, in Turin, Livorno, and Caen, were destroyed during World War 2, and four skins seem to have disappeared from Réunion and Mauritius, which now only have one each. Specimens were sent to Europe beginning in the second half of the 18th century, but most were collected during the first half of the 19th century. It is unclear exactly when each specimen was acquired, and specimens were frequently traded and moved between collections. It is also unclear which specimens that served as basis for the various descriptions and illustrations.[2] The only known subfossil hoopoe starling specimen is a femur, discovered in a grotto on Réunion in 1993.[20]
Extinction
Several causes for the decline and sudden disappearance of the hoopoe starling have been proposed, all relating to human activities on Réunion. An often suggested reason is that the introduction of the common myna (Acridotheres tristis) lead to competition between these two starling species. That bird had been introduced to Réunion in 1759 for the purpose of combating locusts, but later became a pest. Yet the hoopoe starling coexisted with the myna for nearly a hundred years, so they may not have competed nor lived in the same habitats. The black rat (Rattus rattus) arrived on Reunion in the 1670s, and the brown rat (Rattus norvegicus) in 1735; they multiplied rapidly and soon became a problem for agriculture as well as native species. Like the hoopoe starling, the rats inhabited tree cavities, and they would have preyed on the eggs, juveniles, as well as nesting birds. During the mid 19th century, the Réunion slit-eared skink (Gongylomorphus borbonicus) became extinct due to predation by the introduced wolf snake (Lycodon aulicum), which may have deprived the hoopoe starling of an important food source.[2] It is possible that the hoopoe starlings received avian diseases from introduced birds, a factor which is known to have caused decline and extinctions among endemic birds of Hawaii. The ecologist Anthony S. Cheke believed this to be the main reason for the hoopoe starling's extinction, as the species had survived for generations in spite of various other threats.[21]
From the 1830s and onwards, Réunion was heavily deforested to make way for plantations, and slaves were emancipated in 1848, which lead to further cultivation of areas that had remained pristine until then. The hoopoe starling was thereby pushed to the fringes of its former habitat. Hume believes that over-hunting was the final blow to the hoopoe starling; once the forests became more accessible, hunting by the rapidly growing human population may have driven the remaining birds to extinction. In 1821, a law ordering the extermination of birds that did damage to grain was implemented, and the hoopoe starling had a reputation of damaging crops. During the 1860s, various writers noted that the bird had almost disappeared, but it was probably already extinct by this time. In 1877, Vinson lamented that the last individuals could had been killed during recent forest fires. No attempts to preserve the species in captivity appear to have been made. The hoopoe starling survived longer than many other now extinct Mascarene species, and it was the last of the starlings of these islands to go extinct; the species on Rodrigues and Mauritius probably became extinct due to the arrival of rats. At least five species of Aplonis starlings have gone extinct in islands of the Pacific Ocean, and rats contributed to their demise as well. The hoopoe starling may have survived longer due to the topography of Réunion being more rugged, and because it spent much of the year in highland areas.[2]
References
- ^ Template:IUCN
- ^ a b c d e f g h i j k l m n o p q Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.11646/zootaxa.3849.1.1, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.11646/zootaxa.3849.1.1instead. - ^ Rothschild, W. (1907). Extinct Birds. London: Hutchinson & Co. pp. 3–4.
- ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1111/j.1096-3642.1874.tb02508.x, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1111/j.1096-3642.1874.tb02508.xinstead. - ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1038/040177a0, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1038/040177a0instead. - ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.2307/4078667, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.2307/4078667instead. - ^ Berger, A. J. (1957). "On the anatomy and relationships of Fregilupus varius, an extinct starling from the Mascarene Islands" (PDF). Bulletin of the American Museum of Natural History. 113: 225–272.
- ^ Morioka, H. (1996). "The skull of the Réunion starling Fregilupus varius and the Sickle-billed Vanga Falculea palliata, with notes on their relationships and feeding habits". In Yamagishi (ed.), S. (ed.). Social evolution of birds in Madagascar, with special respect to vangas. Osaka: Osaka City University. pp. 5–16. ISBN 9781408190494.
{{cite book}}:|editor1-last=has generic name (help) - ^ Newton, A. (1875). "Additional Evidence as to the Original Fauna of Rodriguez". Proceedings of the Zoological Society of London: 39–43.
- ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1098.2Frstl.1879.0043, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1098.2Frstl.1879.0043instead. - ^ Greenway, J. C. (1967). Extinct and Vanishing Birds of the World. New York: American Committee for International Wild Life Protection 13. pp. 129–132. ISBN 0-486-21869-4.
- ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1017/CBO9780511735769.004, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1017/CBO9780511735769.004instead. - ^ Hume, J. P.; Walters, M. (2012). Extinct Birds. London: A & C Black. pp. 271–273. ISBN 1-4081-5725-X.
- ^ Amadon, D. (1943). "Genera of starlings and their relationships" (PDF). American Museum Novitates. 1247: 1–16.
- ^ Amadon, D. (1956). "Remarks on the starlings, family Sturnidae" (PDF). American Museum Novitates. 1803: 1–41.
- ^ a b Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1111/j.1463-6409.2008.00339.x, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1111/j.1463-6409.2008.00339.xinstead. - ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1016/j.ympev.2008.01.020, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1016/j.ympev.2008.01.020instead. - ^ Fuller, E. (2001). Extinct Birds (revised ed.). New York: Comstock. pp. 364–365. ISBN 978-0-8014-3954-4.
{{cite book}}: Invalid|ref=harv(help) - ^ Cheke, A. S.; Hume, J. P. (2008). Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues. T. & A. D. Poyser. pp. 49–52. ISBN 978-0-7136-6544-4.
- ^ Mourer-Chauvire, C.; Bour, R.; Ribes, S.; Moutou, F. (1999). "Avian paleontology at the close of the 20th century: The avifauna of Réunion Island (Mascarene Islands) at the time of the arrival of the first Europeans". Smithsonian Contributions to Paleobiology. 89: 8–11.
- ^ Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1017/CBO9780511735769.003, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with
|doi=10.1017/CBO9780511735769.003instead.