Hoopoe starling

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Hoopoe starling
Specimen in
Musée Cantonal de Zoologie, Lausanne

Extinct (ca. 1850)  (IUCN 3.1)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Family: Sturnidae
Genus: Fregilupus
Lesson
, 1831
Species:
F. varius
Binomial name
Fregilupus varius
(Boddaert, 1783)
Location of Réunion (circled)
Synonyms
List
  • Upupa varia Boddaert, 1783
  • Upupa capensis Gmelin, 1788
  • Upupa madagascariensis Shaw, 1811
  • Coracia cristata Vieillot, 1817
  • Pastor upupa Wagler, 1827
  • Fregilupus capensis Lesson, 1831
  • Coracia tinouch Hartlaub, 1861
  • Fregilupus borbonicus Vinson, 1868
  • Fregilupus varia Gray, 1870
  • Sturnus capensis Schlegel, 1872
  • Lophopsarus varius Sundeval, 1872
  • Coracias tivouch Murie, 1874

The hoopoe starling (Fregilupus varius), also known as the Réunion starling or Bourbon crested starling, is a species of

extinct in the 1850s. Its closest relatives were the also-extinct Rodrigues starling and Mauritius starling from nearby islands, and the three apparently originated in south-east Asia. The bird was first mentioned during the 17th century and was long thought to be related to the hoopoe, from which its name is derived. Some affinities have been proposed, but it was confirmed as a starling in a DNA
study.

The hoopoe starling was 30 cm (12 in) in length. Its

omnivorous
, feeding on plant matter and insects. Its pelvis was robust, its feet and claws large, and its jaws strong, indicating that it foraged near the ground.

The birds were hunted by settlers on Réunion, who also kept them as pets. Nineteen specimens exist in museums around the world. The hoopoe starling was reported to be in decline by the early 19th century and was probably extinct before the 1860s. Several factors have been proposed, including competition and predation by introduced species, disease, deforestation, and persecution by humans, who hunted it for food and as an alleged crop pest.

Taxonomy

The first account thought to mention the hoopoe starling is a 1658 list of birds of

Mascarene island of Réunion (then called "Bourbon") by Père Vachet in 1669, and first described in detail by French traveller Sieur Dubois's 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", because of the similarity of its crest and curved bill with that of the hoopoe. Little was recorded about the hoopoe starling during the next 100 years, but specimens began to be brought to Europe during the 18th century. The species was first scientifically described by

scientific name from Dutch naturalist Pieter Boddaert for the book's 1783 edition. Boddaert named the bird Upupa varia; its genus name is that of the hoopoe, and its specific name means "variegated", describing its black-and-white colour.[2]

National Museum of Natural History, Paris
, with a similar crest

Boddaert provided

National Museum of Natural History in Paris, this is impossible to determine today; the Paris museum originally had five hoopoe starling skins, some of which only arrived during the 19th century. The possibly female specimen MNHN 2000-756, one of the most-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

monotypic genus, Fregilipus, a composite of Upupa and Fregilus, the latter a defunct genus name of the chough. French naturalist Auguste Vinson established in 1868 that the bird was restricted to the island of Réunion and proposed a new binomial, Fregilupus borbonicus, referring to the former name of the island.[4]

German ornithologist Hermann Schlegel first proposed in 1857 that the species belonged to the starling family, (Sturnidae), reclassifying it as part of the genus Sturnus, S. capensis. This reclassification was observed by other authors; Swedish zoologist Carl Jakob Sundevall proposed the new genus name Lophopsarus ("crested starling") in 1872, yet Fregilupus varius—the oldest name—remains the bird's binomial, and all other scientific names are synonyms.[2] In 1874, after a detailed analysis of the only known skeleton (held at the Cambridge University Museum of Zoology), British zoologist James Murie agreed that it was a starling.[5] English zoologist Richard Bowdler Sharpe said in 1890 that the hoopoe starling was similar to the starling genus Basilornis, but did not note any similarities other than their crests.[6]

lithographs of the only known skeleton and individual bones, by Philibert Charles Berjeau

In 1941, American ornithologist Malcolm R. Miller found the bird's musculature similar to that of the common starling (Sturnus vulgaris) after he dissected a specimen preserved in spirits at the Cambridge Museum, but noted that the tissue was very degraded and the similarity did not necessarily confirm a relationship with starlings.[7] In 1957, American ornithologist Andrew John Berger cast doubt on the bird's affinity with starlings because of subtle anatomical differences, after dissecting a spirit specimen at the American Museum of Natural History.[8] Some authors proposed a relationship with vangas (Vangidae), but Japanese ornithologist Hiroyuki Morioka rejected this in 1996, after a comparative study of skulls.[9]

In 1875, British ornithologist

humeral features.[15]

Evolution

In 1943, American ornithologist

glossy starlings (Lamprotornis) of Africa and the Madagascar starling (Saroglossa aurata), based on their colouration.[16][17] A 2008 study by Italian zoologist Dario Zuccon and colleagues analysing the DNA of a variety of starlings confirmed that the hoopoe starling belonged in a clade of Southeast Asian starlings as an isolated lineage, with no close relatives. The following cladogram shows its position:[18]

White bird with dark wing and tail feathers and a blue mask
The related Bali myna, which is similarly coloured and also has a crest

Sturnornus albofrontatus
(white-faced starling)

Leucopsar rothschildi
(Bali myna)

Fregilupus varius (hoopoe starling)

Sturnus sinensis
(white-shouldered starling)

Sturnus pagodarum
(brahminy starling)

Sturnus erythropygia
(white-headed starling)

Sturnus malabaricus
(chestnut-tailed starling)

An earlier attempt by another team could not extract viable hoopoe starling DNA.[19] Zuccon and colleagues suggested that ancestors of the hoopoe starling reached Réunion from Southeast Asia by using island chains as "stepping stones" across the Indian Ocean, a scenario also suggested for other Mascarene birds. Its lineage diverged from that of other starlings four million years ago (about two million years before Réunion emerged from the sea), so it may have first evolved on landmasses now partially submerged.[18]

Extant relations, such as the Bali myna (Leucopsar rothschildi) and the white-headed starling (Sturnia erythropygia), have similarities in colouration and other features with the extinct Mascarene species. According to Hume, since the Rodrigues and Mauritius starlings seem morphologically closer to each other than to the hoopoe starling—which appears closer to Southeast Asian starlings—there may have been two separate migrations of starlings from Asia to the Mascarenes, with the hoopoe starling the latest arrival. Except for Madagascar, the Mascarenes were the only islands in the southwestern Indian Ocean with native starlings, probably because of their isolation, varied topography, and vegetation.[15]

Description

Turnaround video of specimen RMNH 110.050, Naturalis Biodiversity Center, Leiden

The hoopoe starling was 30 cm (12 in) in length. The bird's

superciliary stripe, lore, and most of the specimen's underside is white, with a pale rufous wash on the flanks and undertail coverts. The extent of light rufous on the underside varies by specimen. The beak and legs are lemon-yellow, with yellow-brown claws. It has a bare, triangular area of skin around the eye, which may have been yellow in life. Though the species' iris was described as bluish-brown, it has been depicted as brown, yellow, or orange.[21]

There has been confusion about which characteristics were sexually dimorphic in the species. Only three specimens were sexed (all males), with age and individual variation not considered. The male is thought to have been largest with a longer, curvier beak. In 1911, Réunion resident Eugène Jacob de Cordemoy recalled his observations of the bird about 50 years before, suggesting that only males had a white crest, but this is thought to be incorrect. A presumed female (MNHN 2000-756) in the Paris museum appears to have a smaller crest, a smaller and less-curved beak, and smaller primary coverts. A juvenile specimen (MHNT O2650) has a smaller crest and primary coverts, with a brown wash instead of ash grey on the crest, lore, and superciliary stripe, and a light-brown (instead of ash-brown) back. The juveniles of some southeast Asian starlings are also browner than adults.[4]

life drawing
, showing the natural position of the crest, by Paul Jossigny, early 1770s

Vinson, who observed live hoopoe starlings when he lived on Réunion, described the crest as flexible, disunited and forward-curled barbs of various lengths, highest in the centre, and able to be erected at will. He compared the bird's crest to that of a

life drawing. Jossigny also made the only known life drawing of the now-extinct Newton's parakeet (Psittacula exsul) after a specimen sent to him from Rodrigues to Mauritius, so this is perhaps also where he drew the hoopoe starling. Murie suggested that only the illustrations by Martinet and Jacques Barraband were "original", since he was unaware of Jossigny's drawing, but noted a crudeness and stiffness in them which made neither appear lifelike.[4][5]

The hoopoe starling can be distinguished skeletally from other Mascarene starlings by its

mandibular fenestra (opening). The sternum (breast-bone) was short and wide, particularly at the hind end. The coracoid was relatively reduced in length, and its shaft was robust. The humerus (upper arm bone) was robust with a straight shaft, with the upper and lower ends flattened from front to back. The radius of the lower arm was robust. The pelvis was extremely robust. The femur (thigh bone) was robust, especially at the upper and lower ends, and the shaft was straight. The tibiotarsus (lower leg bone) was long and robust, with a broad and expanded shaft, especially near the lower end. The tarsometatarsus (ankle bone) was long and robust, with a relatively straight shaft.[2]

Behaviour and ecology

Painting of bright-eyed hoopoe starling on a tree branch
1860s illustration by Albert Roussin

Little is known about the behaviour of the hoopoe starling. According to François Levaillant's 1807 account of the bird (which included observations from a Réunion resident) it was abundant, with large flocks inhabiting humid areas and marshes. In 1831, Lesson, without explanation, described its habits as similar to those of a crow. Its song was described as a "bright and cheerful whistle" and "clear notes", indicating a similarity to the songs of other starlings.[4] Vinson's 1877 account relates his experiences with the bird more than 50 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![4]

Like most other starlings, the hoopoe starling was

omnivorous, feeding on fruits, seeds, and insects. Its tongue—long, slender, sharp, and frayed—may have been able to move rapidly, helpful when feeding on fruit, nectar, pollen, and invertebrates. Its pelvic elements were robust and its feet and claws large, indicating that it foraged near the ground. Its jaws were strong; Morioka compared its skull to that of the hoopoe, and it may have foraged in a similar way, probing and opening holes in substrate by inserting and opening its beak.[4]

De Montbeillard was informed of the stomach contents of a dissected specimen, consisting of seeds and the berries of "Pseudobuxus" (possibly

ultraviolet irradiation in this and other black-skinned birds.[22]

Many other endemic species on Réunion became extinct after the arrival of humans and the resulting disruption of the island's

Réunion parakeet, the Réunion swamphen, the Réunion scops 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 before vanishing from both islands.[23]

Relationship with humans

The hoopoe starling was described as tame and easily hunted. In 1704, French pilot engineer Jean Feuilley explained how the birds were caught by humans and 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.[4]

The hoopoe starling was kept as a pet on Réunion and Mauritius, and although the bird was becoming scarcer, some specimens were obtained during the early 19th century. It is unknown whether any live specimens were ever transported from the Mascarenes. Cordemoy recalled that captive birds could be fed a wide variety of food, such as bananas, potatoes, and chayote, and wild birds would never enter inhabited areas. Many individuals survived on Mauritius after escaping there, and it was thought that a feral population could be established. The Mauritian population lasted less than a decade; the final specimen on the island (the last definite record of a live specimen anywhere) was taken in 1836. Specimens could still be collected on Réunion during the 1830s and, possibly, the early 1840s.[4]

There are nineteen surviving hoopoe starling specimens in museums around the world (including one skeleton and two specimens preserved in spirit), with two in the Paris museum and four in Troyes. Additional skins in Turin, Livorno, and Caen were destroyed during World War II, and four skins have disappeared from Réunion and Mauritius (which now have one each). Specimens were sent to Europe beginning in the second half of the 18th century, with most collected during the first half of the 19th century. It is unclear when each specimen was acquired, and specimens were frequently moved between collections. It is also unclear which specimens were the basis for which descriptions and illustrations.[4] The only known subfossil hoopoe starling specimen is a femur, discovered in 1993 in a Réunion grotto.[24]

Extinction

Painting of tufted, brown-and-white bird with a curved beak on a branch
1807 illustration by Jacques Barraband

Several causes for the decline and sudden extinction of the hoopoe starling have been proposed, all connected to the activities of humans on Réunion, who it survived alongside for two centuries. An oft-repeated suggestion is that the introduction of the

wolf snake (Lycodon aulicum), which may have deprived the bird of a significant food source.[4] Hoopoe starlings may have contracted diseases from introduced birds, a factor known to have triggered declines and extinctions in endemic Hawaiian birds. According to British ecologist Anthony S. Cheke, this was the chief cause of the hoopoe starling's extinction; the species had survived for generations despite other threats.[25]

Beginning in the 1830s, Réunion was deforested for plantations. Former slaves joined white peasants in cultivating pristine areas after slavery was abolished in 1848, and the hoopoe starling was pushed to the edges of its former habitat. According to Hume, over-hunting was the final blow to the species; with forests more accessible, hunting by the rapidly growing human population may have driven the remaining birds to extinction. In 1821, a law mandating the extermination of grain-damaging birds was implemented, and the hoopoe starling had a reputation for 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 might have been killed by recent forest fires. No attempts to preserve the species in captivity seem to have been made.[4][26]

The hoopoe starling survived longer than many other extinct Mascarene species, and was the last of the Mascarene starling species to face extinction. The Rodrigues and Mauritius species probably disappeared with the arrival of rats; at least five species of

Pacific Islands, with rats contributing to their extinction. The hoopoe starling may have survived longer because of Réunion's rugged topography and highlands, where it spent much of the year.[15]

References

  1. . Retrieved 13 November 2021.
  2. ^ a b c d e Hume, J. P. (2014). pp. 8–14.
  3. Hutchinson & Co
    . pp. 3–4.
  4. ^ a b c d e f g h i j k l Hume, J. P. (2014). pp. 29–44.
  5. ^ from the original on 13 August 2018. Retrieved 13 January 2018.
  6. .
  7. .
  8. from the original on 26 July 2020. Retrieved 26 October 2021.
  9. .
  10. ^ Newton, A. (1875). "Additional evidence as to the original fauna of Rodriguez". Proceedings of the Zoological Society of London. 1875: 39–43. Archived from the original on 12 October 2018. Retrieved 13 January 2018.
  11. .
  12. .
  13. ISBN 978-0-511-73576-9.{{cite book}}: CS1 maint: location missing publisher (link
    )
  14. .
  15. ^ a b c Hume, J. P. (2014). pp. 55–58.
  16. ^ Amadon, D. (1943). "Genera of starlings and their relationships" (PDF). American Museum Novitates (1247): 1–16. Archived from the original (PDF) on 11 June 2007. Retrieved 26 March 2013.
  17. ^ Amadon, D. (1956). "Remarks on the starlings, family Sturnidae" (PDF). American Museum Novitates (1803): 1–41. Archived from the original (PDF) on 12 June 2007. Retrieved 26 September 2014.
  18. ^
    S2CID 56403448
    .
  19. .
  20. .
  21. ^ Hume, J. P. (2014). pp. 14–29.
  22. S2CID 218652238
    .
  23. .
  24. .
  25. .
  26. ^ Vinson, A. (1877). "Faune détruite. Les Aepiornidés et les Huppes de l'île Bourbon". Bulletin Hebdomadaire de l'Association Scientifique de France (in French). 20: 327–331.

Works cited

External links

Media related to Fregilupus varius at Wikimedia Commons