| Hoopoe starling | |
|---|---|
 | |
| Specimen inMusée Cantonal de Zoologie,Lausanne | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Aves |
| Order: | Passeriformes |
| Family: | Sturnidae |
| Genus: | †Fregilupus Lesson, 1831 |
| Species: | †F. varius |
| Binomial name | |
| †Fregilupus varius (Boddaert, 1783) | |
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| Location ofRéunion (circled) | |
| Synonyms | |
List
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Thehoopoe starling (Fregilupus varius), also known as theRéunion starling orBourbon crested starling, is a species ofstarling that lived on theMascarene island ofRéunion and becameextinct in the 1850s. Its closest relatives were the also-extinctRodrigues starling andMauritius 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 thehoopoe, from which its name is derived. Some affinities have been proposed, but it was confirmed as a starling in aDNA study.
The hoopoe starling was 30 cm (12 in) in length. Itsplumage was primarily white and grey, with its back, wings and tail a darker brown and grey. It had a light, mobilecrest, which curled forwards. The bird is thought to have beensexually dimorphic, with males larger and having more curved beaks. The juveniles were more brown than the adults. Little is known about hoopoe starling behaviour. Reportedly living in large flocks, it inhabited humid areas and marshes. The hoopoe starling wasomnivorous, 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 alsokept 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.
The first account thought to mention the hoopoe starling is a 1658 list of birds ofMadagascar written by French governorÉtienne de Flacourt. He mentioned a black-and-grey "tivouch" orhoopoe; later authors have wondered whether this referred to the hoopoe starling or the Madagascan subspecies of hoopoe (Upupa epops marginata), which resembles the Eurasian subspecies. The hoopoe starling was first noted on theMascarene island ofRéunion (then called "Bourbon") by Père Vachet in 1669, and first described in detail by French travellerSieur 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 byPhilippe Guéneau de Montbeillard in the 1779 edition ofComte de Buffon'sHistoire Naturelle, and received itsscientific name from Dutch naturalistPieter Boddaert for the book's 1783 edition. Boddaert named the birdUpupa varia; its genus name is that of the hoopoe, and its specific name means "variegated", describing its black-and-white colour.[2]
Boddaert providedLinneanbinomial names for plates in Buffon's works, so the accompanying 1770s plate of the hoopoe starling by French engraverFrançois-Nicolas Martinet is considered theholotype or type illustration. Though the plate may have been based on a specimen in theNational 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 theCape of Africa; they were thought to be hoopoes of the genusUpupa, which received names such asUpupa capensis andUpupa madagascariensis. Some authors also allied the bird with groups such asbirds-of-paradise,bee-eaters,cowbirds,Icteridae, andchoughs, resulting in its reassignment to other genera with new names, such asCoracia cristata andPastor upupa. In 1831, French naturalistRené-Primevère Lesson placed the bird in its ownmonotypic genus,Fregilipus, a composite ofUpupa andFregilus, the latter a defunct genus name of the chough. French naturalistAuguste 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 ornithologistHermann Schlegel first proposed in 1857 that the species belonged to thestarling family, (Sturnidae), reclassifying it as part of the genusSturnus,S. capensis. This reclassification was observed by other authors; Swedish zoologistCarl Jakob Sundevall proposed the new genus nameLophopsarus ("crested starling") in 1872, yetFregilupus 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 theCambridge University Museum of Zoology), British zoologist James Murie agreed that it was a starling.[5] English zoologistRichard Bowdler Sharpe said in 1890 that the hoopoe starling was similar to the starling genusBasilornis, but did not note any similarities other than their crests.[6]
In 1941, American ornithologist Malcolm R. Miller found the bird's musculature similar to that of thecommon 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 ornithologistAndrew John Berger cast doubt on the bird's affinity with starlings because of subtle anatomical differences, after dissecting a spirit specimen at theAmerican Museum of Natural History.[8] Some authors proposed a relationship withvangas (Vangidae), but Japanese ornithologist Hiroyuki Morioka rejected this in 1996, after a comparative study of skulls.[9]
In 1875, British ornithologistAlfred Newton attempted to identify a black-and-white bird mentioned in an 18th-century manuscript describing a marooned sailor's stay on the Mascarene island ofRodrigues in 1726–27, hypothesising that it was related to the hoopoe starling.[10]Subfossil bones later found on Rodrigues were correlated with the bird in the manuscript; in 1879, these bones became the basis for a new species,Necropsar rodericanus (theRodrigues starling), named by British ornithologistsAlbert Günther andEdward Newton. Günther and Newton found the Rodrigues bird closely related to the hoopoe starling, but kept it in a separate genus owing to "present ornithological practice".[11] American ornithologistJames Greenway suggested in 1967 that the Rodrigues starling belonged in the same genus as the hoopoe starling, owing to their close similarity.[12] Subfossils found in 1974 confirmed that the Rodrigues bird was a distinct genus of starling; primarily, its stouter bill warrantsgeneric separation fromFregilupus.[13][14] In 2014, British palaeontologistJulian P. Hume described a new extinct species, theMauritius starling (Cryptopsar ischyrhynchus), based on subfossils fromMauritius, which was closer to the Rodrigues starling than to the hoopoe starling in its skull,sternal, andhumeral features.[15]
In 1943, American ornithologistDean Amadon suggested thatSturnus-like species could have arrived in Africa and given rise to thewattled starling (Creatophora cinerea) and the Mascarene starlings. According to Amadon, the Rodrigues and hoopoe starlings were related to Asiatic starlings—such as someSturnus species—rather than to theglossy starlings (Lamprotornis) of Africa and theMadagascar 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 aclade of Southeast Asian starlings as an isolated lineage, with no close relatives. The followingcladogram shows its position:[18]
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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 theBali myna (Leucopsar rothschildi) and thewhite-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]
The hoopoe starling was 30 cm (12 in) in length. The bird'sculmen was 41 mm (1+5⁄8 in) long, its wing147 mm (5+13⁄16 in), its tail114 mm (4+1⁄2 in), and itstarsus about39 mm (1+9⁄16 in) long.[20] It was the largest of the three Mascarene starlings. A presumed 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), with a long crest the same colour with white shafts. Its back and tail are ash-brown, its wings darker with a greyish wash, and its uppertail covert feathers and rump have a rufous wash. Itsprimarycoverts are white with brown tips; the bases (instead of the tips) are brown in other specimens. Thesuperciliary 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 weresexually 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]
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 acockatoo and to the tail feathers of abird-of-paradise. Most mounted specimens have an erect crest, indicating its natural position. The only illustration of the hoopoe starling now thought to have been made from life was drawn by French artist Paul Philippe Sauguin de Jossigny during the early 1770s. Jossigny instructed engravers under the drawing that for accuracy, they should depict the crest angled forward from the head (not straight up). Hume believes that Martinet did this when he made the type illustration, and it was derivative of Jossigny's image rather than alife drawing. Jossigny also made the only known life drawing of the now-extinctNewton'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 andJacques 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 itscranium being rounded when seen from above, bulbous towards the back. Thefrontal bone was narrow, and theforamen magnum (the opening for thespinal cord) was large. Therostrum was long and narrow, with narrow, ovalnarial-openings (bony nostrils). The upper beak was narrow and strongly decurved, and the lower beak was narrow and sharply pointed. The mandible had a distinct retroarticular process (which connected with the skull), and there was a single largemandibular fenestra (opening). Thesternum (breast-bone) was short and wide, particularly at the hind end. Thecoracoid was relatively reduced in length, and its shaft was robust. Thehumerus (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. Thefemur (thigh bone) was robust, especially at the upper and lower ends, and the shaft was straight. Thetibiotarsus (lower leg bone) was long and robust, with a broad and expanded shaft, especially near the lower end. Thetarsometatarsus (ankle bone) was long and robust, with a relatively straight shaft.[2]
Little is known about the behaviour of the hoopoe starling. According toFrançois 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 acrow. 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 wasomnivorous, 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]
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De Montbeillard was informed of the stomach contents of a dissected specimen, consisting of seeds and the berries of "Pseudobuxus" (possiblyEugenia buxifolia, a bush with sweet, orange berries). He noted that the bird weighed 110 grams (4 oz), and was fatter around June and July. Several accounts suggest that the hoopoe starling migrated on Réunion, spending six months in the lowlands and six months in the mountains. Food may have been easier to obtain in the lowlands during winter, with the birds breeding in the mountain forests during summer. The hoopoe starling probably nested in tree cavities.[4] The Belgian biologist Michaël P. J. Nicolaï and colleagues pointed out in 2020 that the hoopoe starling had black skin combined with light plumage and lived in highirradiation zones, which may have evolved for protection againstultraviolet 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'secosystem. The Hoopoe starling lived with other now-extinct birds, such as theRéunion ibis, theMascarene parrot, theRéunion parakeet, theRéunion swamphen, theRéunion scops owl, theRéunion night heron, and theRéunion pink pigeon. Extinct Réunion reptiles include theRéunion giant tortoise and an undescribedLeiolopismaskink. Thesmall Mauritian flying fox and the snailTropidophora carinata lived on Réunion and Mauritius before vanishing from both islands.[23]
The hoopoe starling was described as tame and easily hunted. In 1704, French pilot engineerJean 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 waskept 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, andchayote, 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 inTroyes. Additional skins in Turin, Livorno, and Caen were destroyed duringWorld 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 Réuniongrotto.[24]
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 thecommon myna (Acridotheres tristis) led to competition between these two starling species. The myna was introduced to Réunion in 1759 to combat locusts, and became a pest itself, but the hoopoe starling coexisted with the myna for nearly 100 years and they may not have shared habitat. Theblack rat (Rattus rattus) arrived on Reunion in the 1670s, and thebrown rat (Rattus norvegicus) in 1735, multiplying rapidly and threatening agriculture and native species. Like the hoopoe starling, the rats inhabited tree cavities and would have preyed on eggs, juveniles, and nesting birds. During the mid-19th century theRéunion slit-eared skink (Gongylomorphus borbonicus) became extinct because of predation by the introducedwolf 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 ecologistAnthony 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 speciesin 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 ofAplonis starlings have disappeared from thePacific Islands, with rats contributing to their extinction. The hoopoe starling may have survived longer because of Réunion's ruggedtopography and highlands, where it spent much of the year.[15]
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