Red rail

Red rail
painting of red rail
Painting of a possibly stuffed specimen attributed to Jacob Hoefnagel, ca. 1610

Extinct  (around 1693)  (IUCN 3.1)[1]
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Gruiformes
Family: Rallidae
Genus: Aphanapteryx
Frauenfeld, 1868
A. bonasia
Binomial name
Aphanapteryx bonasia
( Selys, 1848)
Mauritius island location.svg
Location of Mauritius (in blue)

The red rail (Aphanapteryx bonasia) is an extinct species of flightless rail. It was endemic to the Mascarene island of Mauritius, east of Madagascar in the Indian Ocean. It had a close relative on Rodrigues island, the likewise extinct Rodrigues rail (Erythromachus leguati), with which it is sometimes considered congeneric. Its relationship with other rails is unclear. Rails often evolve flightlessness when adapting to isolated islands, free of mammalian predators. The red rail was a little larger than a chicken and had reddish, hairlike plumage, with dark legs and a long, curved beak. The wings were small, and its legs were slender for a bird of its size. It was similar to the Rodrigues rail, but was larger, and had proportionally shorter wings. It has been compared to a kiwi or a limpkin in appearance and behaviour.

It is believed to have fed on invertebrates, and snail shells have been found with damage matching an attack by its beak. Human hunters took advantage of an attraction red rails had to red objects by using coloured cloth to lure the birds so that they could be beaten with sticks. Until subfossil remains were discovered in the 1860s, scientists only knew the red rail from 17th century descriptions and illustrations. These were thought to represent several different species, which resulted in a large number of invalid junior synonyms. It has been suggested that all late 17th-century accounts of the dodo actually referred to the red rail, after the former had become extinct. The last mention of a red rail sighting is from 1693, and it is thought to have gone extinct around 1700, due to predation by humans and introduced species.


Photo of bones
First known remains, 1866

The red rail was long known only from a few contemporary descriptions referring to red "hens" and names otherwise used for grouse or partridges in Europe, as well as the sketches of the travellers Pieter van den Broecke and Sir Thomas Herbert from 1617 and 1634. These were thought to depict separate species of birds by some authors, but were regarded as one by the English naturalist Hugh Edwin Strickland in 1848.[2] The German ornithologist Hermann Schlegel thought Broecke's sketch depicted a smaller dodo species from Mauritius, and that the Herbert sketch showed a dodo from Rodrigues, and named them Didus broecki and Didus herberti in 1854.[3] The Flemish artist Jacob Hoefnagel's 1610 painting, the 1601 sketch from the Gelderland ship's journal, and the English traveller Peter Mundy's 1638 description and sketch later surfaced, but there was still uncertainty about the identity of the birds depicted.[4]

In the 1860s, subfossil foot bones and a lower jaw were found along with remains of other Mauritian animals in the Mare aux Songes swamp, and were identified as belonging to a rail by the French zoologist Alphonse Milne-Edwards in 1866.[5] He also determined they belonged to the birds in the 17th century descriptions and illustrations. In 1869, Milne-Edwards combined the genus name of Aphanapteryx imperialis, which had been coined the previous year by the Austrian naturalist Georg Ritter von Frauenfeld for the Hoefnagel painting, with the older specific name broecki.[6] Due to nomenclatural priority, the genus name was later combined with the oldest species name bonasia, which was coined by the Belgian scientist Edmond de Sélys Longchamps in 1848. Sélys Longchamps had originally named the genus Apterornis, wherein he also included the Réunion solitaire and the Réunion swamphen, but the name was preoccupied by Aptornis, a bird described by the English biologist Richard Owen in 1844.[7][8] Aphanapteryx means "invisible-wing", but the meaning of bonasia is unclear. Some early accounts refer to red rails by the vernacular names for the hazel grouse, Tetrastes bonasia, so the name evidently originates there. The name itself perhaps refers to bonasus, meaning "bull" in Latin, or bonum and assum, meaning "good roast". It has also been suggested to be a Latin form of the French word bonasse, meaning simple-minded or good-natured.[8]

More fossils were later found by Theodore Sauzier, who had been commissioned to explore the "historical souvenirs" of Mauritius in 1889.[9] Around the end of the 19th century, a complete specimen was found by the barber Louis Etienne Thirioux, who also found important dodo remains.[10]


Further remains described in 1893

Apart from being a close relative of the Rodrigues rail, the relationships of the red rail are uncertain. The two are commonly kept as separate genera, Aphanapteryx and Erythromachus, but have also been united as species of Aphanapteryx at times.[4] They were first generically synonymised by the British ornithologists Edward Newton and Albert Günther in 1879, due to skeletal similarities.[11] In 1892, the Scottish naturalist Henry Ogg Forbes described Hawkins's rail, an extinct species of rail from the Chatham Islands, as a new species of Aphanapteryx; A. hawkinsi. He found the Chatham Islands species more similar to the red rail than the latter was to the Rodrigues rail, and proposed that the Mascarene Islands had once been connected with the Chatham Islands, as part of a lost continent he called "Antipodea". Forbes moved the Chatham Islands bird to its own genus, Diaphorapteryx, in 1893, on the recommendation of Newton, but later reverted to his older name. The idea that the Chatham Islands bird was closely related to the red rail and the idea of a connection between the Mascarenes and the Chatham Islands were later criticised by the British palaeontologist Charles William Andrews due to no other species being shared between the islands, and the German ornithologist Hans F. Gadow explained the similarity between the two rails as parallel evolution.[12][13][14]

An illustration of a bird with a long neck, a long, sharp, red beak, red legs and feet, mid-grey to black feathers and a large, red, naked area around its eye
1907 restoration of the similar Rodrigues rail by Frederick William Frohawk, based on old accounts

In 1945, the French palaeontologist Jean Piveteau found skull features of the red and Rodrigues rail different enough for generic separation, and in 1977, the American ornithologist Storrs L. Olson stated that though the two species were similar and derived from the same stock, they had also diverged considerably, and should possibly be kept separate. Based on geographic location and the morphology of the nasal bones, Olson suggested that they were related to the genera Gallirallus, Dryolimnas, Atlantisia, and Rallus.[8] The American ornithologist Bradley C. Livezey was unable to determine the affinities of the red and Rodrigues rail in 1998, stating that some of the features uniting them and some other rails were associated with the loss of flight rather than common descent. He also suggested that the grouping of the red and Rodrigues rail into the same genus may have been influenced by their geographical distribution.[15] The French palaeontologist Cécile Mourer-Chauviré and colleagues also considered the two as belonging to separate genera in 1999.[16]

Rails have reached many oceanic archipelagos, which has frequently led to speciation and evolution of flightlessness. According to the British researchers Anthony S. Cheke and Julian P. Hume in 2008, the fact that the red rail lost much of its feather structure indicates it was isolated for a long time. These rails may be of Asian origin, like many other Mascarene birds.[4] In 2019, Hume supported the distinction of the two genera, and cited the relation between the extinct Mauritius owl (Otus sauzieri) and the Rodrigues owl (Otus murivorus) as another example of the diverging evolutionary paths on these islands. He stated that the relationships of the red and Rodrigues rails was more unclear than that of other extinct Mascarene rails, with many of their distinct features being related to flightlessness and modifications to their jaws due to their diet, suggesting long time isolation. The speed of which these features evolved may also have been affected by gene flow, resource availability, and climate events, and flightlessness can evolve rapidly in rails, as well as repeatedly within the same groups, as seen in for example Dryolimnas, so the distinctness of the red and Rodrigues rails may not have taken long to evolve (some other specialised rails evolved in less than 1–3 million years). Hume suggested that the two rails were probably related to Dryolimnas, but their considerably different morphology made it difficult to establish how. In general, rails are adept at colonising islands, and can become flightless within few generations in suitable environments, for example without predators, yet this also makes them vulnerable to human activities.[17]


drawing of red rail
1907 illustration by Frohawk

From the subfossil bones, illustrations and descriptions, it is known that the red rail was a flightless bird, somewhat larger than a chicken. Subfossil specimens range in size, which may indicate sexual dimorphism, as is common among rails.[8] It was about 35–40 cm (14–16 in) long, and the male may have weighed 1.3 kg (2.9 lb) and the female 1 kg (2.2 lb).[18] The pelvis was 60 mm (2.4 in) in length, the femur was 69–71 mm (2.7–2.8 in), the tibia was 98–115 mm (3.9–4.5 in), the tarsometatarsus was 79 mm (3.1 in), and the humerus was 60–66 mm (2.4–2.6 in).[9] Its plumage was reddish brown all over, and the feathers were fluffy and hairlike; the tail was not visible in the living bird and the short wings likewise also nearly disappeared in the plumage.[19] It had a long, slightly curved, brown bill, and some illustrations suggest it had a nape crest.[10] It perhaps resembled a lightly built kiwi, and it has also been likened to a limpkin, both in appearance and behaviour.[4][19]

The sternum and humerus were small, indicating that it had lost the power of flight. Its legs were long and slender for such a large bird, but the pelvis was compact and stout.[9] It differed from the Rodrigues rail, its closest relative, in having a proportionately shorter humerus, a narrower and longer skull, and having shorter and higher nostrils. They differed considerably in plumage, based on early descriptions.[8] The red rail was also larger, with somewhat smaller wings, but their leg proportions were similar.[11] The pelvis and sacrum was also similar.[9] The Dutch ornithologist Marc Herremans suggested in 1989 that the red and Rodrigues rails were neotenic, with juvenile features such as weak pectoral apparatuses and downy plumage.[20][18] Hume examined all available upper beaks in 2019, but found no differences in curvature.[17]

Contemporary descriptions

Mundy visited Mauritius in 1638 and described the red rail as follows:

A Mauritius henne, a Fowle as bigge as our English hennes, of a yellowish Wheaten Colour, of which we only got one. It hath a long, Crooked sharpe pointed bill. Feathered all over, butte on their wings they are soe Few and smalle that they cannot with them raise themselves From the ground. There is a pretty way of taking them with a red cap, but this of ours was taken with a stick. They bee very good Meat, and are also Cloven footed, soe that they can Neyther Fly nor Swymme.[21]

drawing of red rail
Drawing by Peter Mundy, 1638

The yellowish colouration mentioned by Mundy instead of the red of other accounts was used by the Japanese ornithologist Masauji Hachisuka in 1937 as an argument for this referring to a distinct species, Kuina mundyi, but it has also been suggested it was due to the observed bird being a juvenile.[8] Another English traveller, John Marshall, described the bird as follows in 1668:

Here are also great plenty of Dodos or red hens which are larger a little than our English henns, have long beakes and no, or very little Tayles. Their fethers are like down, and their wings so little that it is not able to support their bodies; but they have long leggs and will runn very fast, and that a man shall not catch them, they will turn so about in the trees. They are good meate when roasted, tasting something like a pig, and their skin like pig skin when roosted, being hard.[4]

Contemporary depictions

An oil painting depicting a red-feathered parrot with yellow wing tips; a large, ungainly, duck-like bird with grey, white and yellow feathers; a parrot with a black back, yellow breast and a yellow and black tail; and a brown-feathered bird with a long bill eating a frog
Edwards' Dodo, a 1626 painting by Roelant Savery, possibly showing a red rail (or a bittern) in the lower right

Much information about the bird's appearance comes from a painting attributed to Hoefnagel, based on a bird in the menagerie of Emperor Rudolph II around 1610.[22] It is the only coloured depiction of the species, showing the plumage as reddish brown, but it is unknown whether it was based on a stuffed or living specimen.[10] The bird had most likely been brought alive to Europe, as it is unlikely that taxidermists were on board the visiting ships, and spirits were not yet used to preserve biological specimens. Most tropical specimens were preserved as dried heads and feet. It had probably lived in the emperor's zoo for a while together with the other animals painted for the same series.[4] The painting was discovered in the emperor's collection and published in 1868 by Georg von Frauenfeld, along with a painting of a dodo from the same collection and artist.[6] This specimen is thought to have been the only red rail that ever reached Europe.[23]

The travel journal of the Dutch East India Company ship Gelderland (1601–1603), rediscovered in the 1860s, contains good sketches of several now-extinct Mauritian birds attributed to the artist Joris Laerle, including an unlabelled red rail. The bird appears to have been stunned or killed, and the sketch is the earliest record of the species, but was only rediscovered in the 1860s. The image was sketched with pencil and finished in ink, but details such as a deeper beak and the shoulder of the wing are only seen in the underlying sketch.[24] In addition, there are three rather crude black-and-white sketches, but differences in them were enough for some authors to suggest that each image depicted a distinct species, leading to the creation of several scientific names which are now synonymous with Aphanapteryx bonasia.[19]

Jacopo Bassano's 1570 painting Arca di Noè, perhaps showing a red rail in the lower right

There are also depictions of what appears to be a red rail in three of the Dutch artist Roelant Savery's paintings.[25] In his famous Edwards' Dodo painting from 1626, a rail-like bird is seen swallowing a frog behind the dodo, but this identification has been doubted, and it may instead show a bittern.[10] A bird resembling a red rail is also figured in the Italian artist Jacopo Bassano's painting Arca di Noè ("Noah's Ark") from c. 1570.[19] It has been pointed out that it is doubtful that a Mauritian bird could have reached Italy this early, but the attribution may be inaccurate, as Bassano had four artist sons who used the same name.[26] A similar bird is also seen in the Flemish artist Jan Brueghel the Elder's Noah's Ark painting.[4]

Behaviour and ecology

Sketch showing red rail, a dodo and a parrot
Labeled sketch from 1634 by Sir Thomas Herbert, showing a broad-billed parrot, a red rail, and a dodo

Contemporary accounts are repetitive and do not shed much light on the bird's life history. The shape of the beak indicates it could have captured reptiles and invertebrates. There were many endemic land snails on Mauritius, including the extinct Tropidophora carinata, and subfossil shells have been found with damage matching attacks from the beak of the red rail.[4][27] No contemporary accounts were known to mention the red rail's diet, until the 1660s report of Johannes Pretorius about his stay on Mauritius was published in 2015, where he mentioned that the bird "scratches in the earth with its sharp claws like a fowl to find food such as worms under the fallen leaves."[28]

A 1631 Dutch letter (long thought lost, but rediscovered in 2017) uses word-play to refer to the animals described, with red rails supposedly being an allegory for soldiers:[29]

The soldiers were very small and moved slowly, so that we could catch them easily with our hands. Their armor was their mouth, which was very sharp and pointed; they used it instead of a dagger, were very cowardly and nervous; they did not behave as soldiers at all, and walked in a disorderly manner, one here, the other there, and did not show any faithfulness towards one another.[29]

While it was swift and could escape when chased, it was easily lured by waving a red cloth, which they approached to attack; a similar behaviour was noted in its relative, the Rodrigues rail. The birds could then be picked up, and their cries when held would draw more individuals to the scene, as the birds, which had evolved in the absence of mammalian predators, were curious and not afraid of humans.[19]

1869 adaptation of the Hoefnagel painting

The English traveller Sir Thomas Herbert described its behaviour towards red cloth in 1634:

The hens in eating taste like parched pigs, if you see a flocke of twelve or twenties, shew them a red cloth, and with their utmost silly fury they will altogether flie upon it, and if you strike downe one, the rest are as good as caught, not budging an iot till they be all destroyed.[4]

Many other endemic species of Mauritius became extinct after the arrival of man heavily damaged the ecosystem, making it hard to reconstruct. Before humans arrived, Mauritius was entirely covered in forests, but very little remains today due to deforestation.[30] The surviving endemic fauna is still seriously threatened.[31] The red rail lived alongside other recently extinct Mauritian birds such as the dodo, the broad-billed parrot, the Mascarene grey parakeet, the Mauritius blue pigeon, the Mauritius owl, the Mascarene coot, the Mauritian shelduck, the Mauritian duck, and the Mauritius night heron. Extinct Mauritian reptiles include the saddle-backed Mauritius giant tortoise, the domed Mauritius giant tortoise, the Mauritian giant skink, and the Round Island burrowing boa. The small Mauritian flying fox and the snail Tropidophora carinata lived on Mauritius and Réunion, but became extinct in both islands. Some plants, such as Casearia tinifolia and the palm orchid, have also become extinct.[4]

Relationship with humans

drawing that includes a red rail
Pieter van den Broecke's 1617 drawing of a dodo, a one-horned sheep, and a red rail; after the dodo became extinct, visitors may have confused the red rail with it

Though Mauritius had previously been visited by Arab vessels in the Middle Ages and Portuguese ships between 1507 and 1513, they did not settle on the island.[32] The Dutch Empire acquired the island in 1598, renaming it after Maurice of Nassau, and it was used from then on for the provisioning of trade vessels of the Dutch East India Company.[33] To the sailors who visited Mauritius from 1598 and onwards, the fauna was mainly interesting from a culinary standpoint. The dodo was sometimes considered rather unpalatable, but the red rail was a popular gamebird for the Dutch and French settlers. The reports dwell upon the varying ease with which the bird could be caught according to the hunting method and the fact that when roasted it was considered similar to pork.[19]

Johann Christian Hoffmann, who was on Mauritius in the early 1670s, described a red rail hunt as follows:

... [there is also] a particular sort of bird known as toddaerschen which is the size of an ordinary hen. [To catch them] you take a small stick in the right hand and wrap the left hand in a red rag, showing this to the birds, which are generally in big flocks; these stupid animals precipitate themselves almost without hesitation on the rag. I cannot truly say whether it is through hate or love of this colour. Once they are close enough, you can hit them with the stick, and then have only to pick them up. Once you have taken one and are holding it in your hand, all the others come running up as it [sic] to its aid and can be offered the same fate.[4]

Hoffman's account refers to the red rail by the German version of the Dutch name originally applied to the dodo, "dod-aers", and John Marshall used "red hen" interchangeably with "dodo" in 1668. British ornithologist Alfred Newton (brother of Edward) suggested in 1868 that that the name of the dodo was transferred to the red rail after the former had gone extinct. Cheke suggested in 2008 that all post 1662 references to "dodos" therefore refer to the rail instead.[34][35] A 1681 account of a "dodo", previously thought to have been the last, mentioned that the meat was "hard", similar to the description of red hen meat.[4] Errol Fuller has also cast the 1662 "dodo" sighting in doubt, as the reaction to distress cries of the birds mentioned matches what was described for the red rail.[19] Milne-Edwards suggested that early travellers may have confused young dodos with red rails.[25]

drawing of red rail
1601 sketches of a hunted specimen by Joris Laerle

230 years before Charles Darwin's theory of evolution, the appearance of the red rail and the dodo led Mundy to speculate:

Of these 2 sorts off fowl afforementionede, For oughtt wee yett know, Not any to bee Found out of this Iland, which lyeth aboutt 100 leagues From St. Lawrence. A question may bee demaunded how they should bee here and Not elcewhere, beeing soe Farer From other land and can Neither fly or swymme; whither by Mixture off kindes producing straunge and Monstrous formes, or the Nature of the Climate, ayer and earth in alltring the First shapes in long tyme, or how.[19]


In addition to hunting pressure by humans, the fact that the red rail nested on the ground made it vulnerable to pigs and other introduced animals, which ate their eggs and young, probably contributing to its extinction, according to Cheke.[26] Hume pointed out that the red rail had coexisted with introduced rats since at least the 14th century, which did not appear to have affected them (as they seem to have been relatively common in the 1680s), and they were probably able to defend their nests (Dryolimnas rails have been observed killing rats, for example). They also seemed to have managed to survive alongside humans as well as introduced pigs and crab-eating macaques.[17]

Since the red rail was referred to by the names of the dodo in the late 1600s, it is uncertain which accounts that mention the species are the latest.[17] When the French traveller François Leguat, who had become familiar with the Rodrigues rail in the preceding years, arrived on Mauritius in 1693, he remarked that the red rail had already become rare. He was the last source to mention the bird, so it is assumed that it became extinct around 1700.[19][36] Feral cats, which are effective predators of ground-inhabiting birds, were established on Mauritius around the late 1680s (to control rats), and this has been cause for rapid disappearance of rails elsewhere, for example on Aldabra Atoll. Being inquisitive and fearless, Hume suggested the red rail would have been easy prey for cats, and was thereby driven to extinction.[28][17]

See also


  1. ^ BirdLife International (2012). "Aphanapteryx bonasia". IUCN Red List of Threatened Species. 2012: e.T22728884A39099824. doi:10.2305/IUCN.UK.2012-1.RLTS.T22728884A39099824.en.
  2. ^ 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. p. 21.
  3. ^ Schlegel, H. (1854), "Ook een Woordje over den Dodo (Didus ineptus) en zijne Verwanten", Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen (in Dutch), 2: 232–256
  4. ^ 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. London: T. & A. D. Poyser. pp. 25–38. ISBN 978-0-7136-6544-4.
  5. ^ Milne-Edwards, A. (1866). Recherches sur la faune ornithologique éteinte des iles Mascareignes et de Madagascar (in French). Paris: G. Masson. pp. 61–83.
  6. ^ a b Milne-Edwards, A. (1869). "Researches into the zoological affinities of the bird recently described by Herr von Frauenfeld under the name of Aphanapteryx imperialis". Ibis. 11 (3): 256–275. doi:10.1111/j.1474-919X.1869.tb06880.x.CS1 maint: ref=harv (link)
  7. ^ de Sélys Longchamps, E. (1848), "Résumé concernant les oiseaux brévipennes mentionnés dans l'ouvrage de M. Strickland sur le Dodo", Revue Zoologique Par la Société Cuvierenne (in French), 11: 292–295
  8. ^ a b c d e f Olson, S. (1977). "A synopsis on the fossil Rallidae". In Ripley, S. D. (ed.). Rails of the World – A Monograph of the Family Rallidae. Boston: Codline. pp. 357–363. ISBN 978-0-87474-804-8.
  9. ^ a b c d Newton, E.; Gadow, H. (1893). "IX. On additional bones of the Dodo and other extinct birds of Mauritius obtained by Mr. Theodore Sauzier". The Transactions of the Zoological Society of London. 13 (7): 281–302. doi:10.1111/j.1469-7998.1893.tb00001.x.CS1 maint: ref=harv (link)
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  12. ^ Forbes, H. O. (1893). "Mr. H. O. Forbes's Discoveries in the Chatham Islands". Nature. 48 (1230): 74–75. doi:10.1038/048074c0.
  13. ^ Forbes, H. O. (1893). "Mr. H. O. Forbes's Discoveries in the Chatham Islands". Nature. 48 (1232): 126. doi:10.1038/048126a0.
  14. ^ Andrews, C. W. (1896). "On the extinct birds of the Chatham Islands. Part I.: The osteology of Diaphorapteryx hawkinsi". Novitates Zoologicae. 73–84: 72.
  15. ^ Livezey, B. C. (1998). "A phylogenetic analysis of the Gruiformes (Aves) based on morphological characters, with an emphasis on the rails (Rallidae)". Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences. 353 (1378): 2077–2151. doi:10.1098/rstb.1998.0353. PMC 1692427.
  16. ^ 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: 23. hdl:10088/2005.
  17. ^ a b c d e Hume, J. P. (2019). "Systematics, morphology and ecology of rails (Aves: Rallidae) of the Mascarene Islands, with one new species". Zootaxa. 4626 (1): 9–37, 63–67. doi:10.11646/zootaxa.4626.1.1. PMID 31712544.
  18. ^ a b Livezey, B. C. (2003). "Evolution of flightlessness in rails (Gruiformes: Rallidae): phylogenetic, ecomorphological, and ontogenetic perspectives". Ornithological Monographs (53): iii–654. doi:10.2307/40168337. JSTOR 40168337.
  19. ^ a b c d e f g h i Fuller, E. (2001). Extinct Birds (revised ed.). New York: Comstock. pp. 96–97. ISBN 978-0-8014-3954-4.
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