View source for Red rail

{{Short description|Extinct species of flightless rail which was endemic to Mauritius}}
{{For|the place in England|Red Rail, Herefordshire}}
{{featured article}}
{{speciesbox
| name = Red rail
| status = EX
| status_system = IUCN3.1
| status_ref = <ref>{{cite journal | author = BirdLife International | author-link = BirdLife International | title = ''Aphanapteryx bonasia'' | journal = [[IUCN Red List of Threatened Species]] | volume = 2012 | page = e.T22728884A39099824 | year = 2012 | doi = 10.2305/IUCN.UK.2012-1.RLTS.T22728884A39099824.en | doi-access = free }}</ref>
| image = Red Rail.jpg
| image_alt = painting of red rail
| image_caption = Painting of a possibly stuffed specimen by [[Jacob Hoefnagel]], ca. 1610
| extinct = shortly after 1693
| genus = Aphanapteryx
|parent_authority=[[Georg von Frauenfeld|Frauenfeld]], 1868
| species = bonasia
| authority = ([[Edmond de Sélys Longchamps|de Sélys-Longchamps]], 1848)
|range_map= Mauritius island location.svg
|range_map_caption= Location of [[Mauritius]] (in blue)
| synonyms ={{collapsible list|bullets = true|title=<small>List</small>
|''Apterornis bonasia'' <small>[[Edmond de Sélys Longchamps|Selys]], 1848</small>
|''Didus broecki'' <small>[[Hermann Schlegel|Schlegel]], 1854</small>
|''Didus herbertii'' <small>Schlegel, 1854</small>
|''Aphanapteryx imperialis'' <small>[[Georg von Frauenfeld|Frauenfeld]], 1868</small>
|''Aphanapteryx imperatoris'' <small>Frauenfeld, 1868</small>
|''Aphanapteryx broeckei'' <small>([[Alphonse Milne-Edwards|Milne-Edwards]], 1868)</small>
|''Didus herberti'' <small>Milne-Edwards, 1868</small>
|''Didus broeckii'' <small>Milne-Edwards, 1868</small>
|''Aphanapteryx broeckii'' <small>Milne-Edwards, 1869</small>
|''Pezophaps broeckei'' <small>(Schlegel, 1873)</small>
|''Pezophaps herbertii'' <small>(Schlegel, 1873)</small>
|''Aphanapteryx broecki'' <small>Günther & Newton, 1879</small>
|''Aphanapteryx broekei'' <small>Newton & Gadow, 1893</small>
|''Kuina mundyi'' <small>[[Masauji Hachisuka|Hachisuka]], 1937</small>
|''Pezocrex herberti'' <small>(Hachisuka, 1953)</small>}}
}}

The '''red rail''' ('''''Aphanapteryx bonasia''''') is an [[extinct]] species of [[Rallidae|rail]] that 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 [[Conspecificity|congeneric]], but their relationship with other rails is unclear. Rails often evolve [[flightlessness]] when adapting to isolated islands, free of mammalian predators, and that was also the case for this species. 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 (bird)|kiwi]] or a [[limpkin]] in appearance and behaviour. 

This bird 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]].

==Taxonomy==
The red rail was first mentioned as "Indian river woodcocks" by the Dutch ships' pilot Heyndrick Dircksz Jolinck in 1598.<ref name="Hume2019"/> By the 19th century, the bird was 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 Dutch merchant [[Pieter van den Broecke]] and the English traveller [[Sir Thomas Herbert]] from 1646 and 1634. While they differed in some details, they were thought to depict a single species by the English naturalist [[Hugh Edwin Strickland]] in 1848.<ref name="Strickland">{{cite book
| last1 = Strickland
| first1 = H.E.
| author-link = Hugh Edwin Strickland
| last2 = Melville
| first2 = A. G.
| author2-link = Alexander Gordon Melville
| title = 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
| publisher = Reeve, Benham and Reeve
| location = London
| year = 1848
| page =  [https://archive.org/details/dodoitskindredor00stri/page/21 21]
| url = https://archive.org/details/dodoitskindredor00stri}}</ref><ref name="Hume2019"/>  The Belgian scientist [[Edmond de Sélys Longchamps]] coined the [[scientific name]] ''Apterornis bonasia'' based on the old accounts mentioned by Strickland. He also included two other Mascarene birds, at the time only known from contemporary accounts, in the genus ''Apterornis'': the [[Réunion ibis]] (now ''Threskiornis solitarius''); and the [[Réunion swamphen]] (now ''Porphyrio caerulescens''). He thought them related to the [[dodo]] and [[Rodrigues solitaire]], due to their shared rudimentary wings, tail, and the disposition of their digits.<ref name="Longchamps">{{cite journal |last1=de Sélys Longchamps |first1=E.|oclc=84482084 |title=Résumé concernant les oiseaux brévipennes mentionnés dans l'ouvrage de M. Strickland sur le Dodo |journal=Revue Zoologique |date=1848 |volume=1848 |pages=292–295 |url=https://www.biodiversitylibrary.org/item/19656#page/304/mode/1up|language=fr}}</ref><ref name="OlsonB"/><ref name="Hume2019"/> 

[[File:Aphanapteryx.jpg|thumb|left|upright|First known [[subfossils]] which were used to identify this bird as a [[rallidae|rail]] in 1868]] 
The name ''Apterornis'' had already been used for a different extinct bird genus from [[New Zealand]] (originally spelled ''[[Aptornis]]'', the adzebills) by the British biologist [[Richard Owen]] earlier in 1848. 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.<ref name="OlsonB"/> It is also possible that the name  alludes to bulls due the bird being said to have had a similar attraction to the waving of red cloth.<ref name="Hume2019"/>

The German ornithologist [[Hermann Schlegel]] thought van den 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.<ref>{{Citation  | last = Schlegel | first = H.  | title = Ook een Woordje over den Dodo (''Didus ineptus'') en zijne Verwanten  | journal = Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen  | volume = 2  | pages = 232–256  | language = nl  | year = 1854 | url =https://archive.org/stream/verslagenenmeded02koni#page/232/mode/2up }}</ref> In 1968, the Austrian naturalist [[Georg Ritter von Frauenfeld]] brought attention to paintings by the Flemish artist [[Jacob Hoefnagel]] depicting animals in the royal menagerie of [[Emperor Rudolph II]] in Prague, including a dodo and a bird he named ''Aphanapteryx imperialis''. ''Aphanapteryx'' means "invisible-wing", from Greek ''aphanēs'', unseen, and ''pteryx'', wing. He compared it with the birds earlier named form old accounts, and found its beak similar to that of a [[Kiwi (bird)|kiwi]] or [[ibis]].<ref name="Frauenfeld">{{cite journal |last1=von Frauenfeld |first1=G. R. |title=Auffindung einer bisher unbekannten abbildung des dronte und eines zweiten kurzflügeligen wahrscheinlich von den Maskarenen stammenden vogels |journal=Journal für Ornithologie |date=1868 |volume=1 |issue=92 |language=de |pages=138–140 |doi=10.1007/BF02261457 |bibcode= 1868JOrn...16..138V|s2cid=34186343 |url=https://www.biodiversitylibrary.org/page/33009718#page/150/mode/1up}}</ref><ref name="OlsonB"/><ref name="Hume2019"/>

In 1865, [[subfossil]] foot bones and a lower jaw were found along with remains of other Mauritian animals in the [[Mare aux Songes]] swamp, and were sent by the British ornithologists [[Edward Newton]] to the French zoologist [[Alphonse Milne-Edwards]], who identified them as belonging to a [[rallidae|rail]] in 1868. Milne-Edwards correlated the bones with the bird in the Hoefnagel painting and the old accounts, and combined the [[genus]] name ''Aphanapteryx'' with the older specific name ''broecki''.<ref>{{cite book|last1=Milne-Edwards|first1=A.|title=Recherches sur la faune ornithologique éteinte des iles Mascareignes et de Madagascar|date=1868|publisher=G. Masson|location=Paris|language = fr|pages=61–83|url=https://archive.org/stream/recherchessurlaf01miln#page/60/mode/2up}}</ref><ref name="Milne-Edwards"/><ref name="Hume2019"/>  Due to [[nomenclatural priority]], the genus name was later combined with the oldest species name ''bonasia''.<ref>{{Citation
 | last = de Sélys Longchamps
 | first = E.
 | title = Résumé concernant les oiseaux brévipennes mentionnés dans l'ouvrage de M. Strickland sur le Dodo
 | journal = Revue Zoologique par la Société Cuvierienne
 | volume = 11
 | pages = 292–295
 | language = fr
 | year = 1848 }}</ref><ref name="OlsonB"/> In the 1860s, the travel journal of the [[Dutch East India Company]] ship ''Gelderland'' (1601–1603) was rediscovered, which contains good sketches of several now-extinct Mauritian birds attributed to the Dutch artist Joris Laerle, including an unlabelled red rail.<ref name="Gelderland"/> 

More fossils were later found by the French naturalist Theodore Sauzier, who had been commissioned to explore the "historical souvenirs" of Mauritius in 1889, and these were described by Newton and the German ornithologist [[Hans F. Gadow]] in 1893.<ref name="Newton & Gadow">{{cite journal| doi = 10.1111/j.1469-7998.1893.tb00001.x| last1 = Newton | first1 = E.| author-link1 = Edward Newton| last2 = Gadow | first2 = H.| year = 1893| title = IX. On additional bones of the Dodo and other extinct birds of Mauritius obtained by Mr. Theodore Sauzier| journal = The Transactions of the Zoological Society of London| volume = 13| issue = 7| pages = 281–302| url = https://www.biodiversitylibrary.org/page/31083700#page/379/mode/1up}}</ref> In 1899, an almost complete specimen was found by the French barber Louis Etienne Thirioux, who also found important dodo remains, in a cave in the Vallée des Prêtres; this is the most completely known red rail specimen, and is catalogued as MI 923 in the [[Mauritius Institute]]. The second most complete individual (specimen CMNZ AV6284) also mainly consists of bones from the Thirioux collection. More material has since been found in various settings.<ref name="Hume2019"/><ref name="Provenance">{{cite journal |last1=Claessens |first1=L. P. A. M. |last2=Hume |first2=J. P. |title=Provenance and history of the Thirioux dodos |journal=Journal of Vertebrate Paleontology |date=2016 |volume=35 |issue=sup1 |pages=21–28 |doi=10.1080/02724634.2015.1111896|s2cid=87212166 }}</ref> The yellowish colouration mentioned by English traveller [[Peter Mundy]] in 1638 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 genus and species, ''Kuina mundyi'' (the generic name means "water-rail" in Japanese), but the American ornithologist [[Storrs L. Olson]] suggested in 1977 that it was possibly due to the observed bird being a juvenile red rail.<ref name="Hachisuka1937">{{cite journal |last1=Hachisuka |first1=M. |title=Untitled [Description of ''Kuinia mundyi'']|journal=Bulletin of the British Ornithologists' Club |date=1937 |volume=57 |pages=54–157 |url=https://www.biodiversitylibrary.org/item/125290#page/196/mode/1up}}</ref><ref name="OlsonB"/> 

===Evolution===
[[File:Aphanapteryx broeckei.jpg|thumb|Additional subfossils, 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.<ref name="Lost Land">{{cite book
| last1 = Cheke | first1 = A. S.
 | first2 = J. P. | last2 = Hume
|year=2008
|title=Lost Land of the Dodo: an Ecological History of Mauritius, Réunion & Rodrigues
| pages = 25–38
 | location = London
|publisher=T. & A. D. Poyser
|isbn=978-0-7136-6544-4}}</ref> They were first generically synonymised by Newton and [[Albert Günther]] in 1879, due to skeletal similarities.<ref name="Günther& Newton">{{cite journal| doi = 10.1098/rstl.1879.0043| last1 = Günther | first1 = A.| last2 = Newton | first2 = E.| year = 1879| title = The extinct birds of Rodriguez| journal = [[Philosophical Transactions of the Royal Society of London]]| volume = 168| pages = 423–437| url = https://archive.org/details/philtrans07832595/mode/2up?view=theater | bibcode = 1879RSPT..168..423G 
| doi-access = free}}</ref> In 1892, the Scottish naturalist [[Henry Ogg Forbes]] described [[Hawkins's rail]], an extinct species of rail from the [[Chatham Islands]] located east of New Zealand, 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 Gadow explained the similarity between the two rails as [[parallel evolution]].<ref>{{cite journal|last1=Forbes|first1=H. O.|title=Mr. H. O. Forbes's Discoveries in the Chatham Islands|journal=Nature|date=1893|volume=48|issue=1230|pages=74–75|doi=10.1038/048074c0|bibcode=1893Natur..48...74F|s2cid=3981755|url=https://zenodo.org/record/1429351}}</ref><ref>{{cite journal|last1=Forbes|first1=H. O.|title=Mr. H. O. Forbes's Discoveries in the Chatham Islands|journal=Nature|date=1893|volume=48|issue=1232|pages=126|doi=10.1038/048126a0|bibcode=1893Natur..48..126F|s2cid=46553514|url=https://zenodo.org/record/1429351|doi-access=free}}</ref><ref>{{cite journal  | last =  Andrews | first = C. W.  | title = On the extinct birds of the Chatham Islands. Part I.: The osteology of ''Diaphorapteryx hawkinsi'' | journal = Novitates Zoologicae  | volume = 73–84| url = https://archive.org/stream/novitateszoologi03lond#page/n91/mode/2up | pages = 72 | year =1896 }}</ref>

In 1945, the French palaeontologist [[Jean Piveteau]] found skull features of the red and Rodrigues rail different enough for generic separation, and in 1977, 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]]''.<ref name="OlsonB">{{cite book  | last = Olson  | first = S.  | chapter = A synopsis on the fossil Rallidae  | title = Rails of the World&nbsp;– A Monograph of the Family Rallidae  | editor = Ripley, S. D. | publisher = Codline  | location = Boston  | year = 1977  | pages = 357–363 | isbn = 978-0-87474-804-8}}</ref> 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.<ref name="Livezey1998">{{cite journal |last1=Livezey |first1=B. C. |title=A phylogenetic analysis of the Gruiformes (Aves) based on morphological characters, with an emphasis on the rails (Rallidae) |journal=Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences |date=1998 |volume=353 |issue=1378 |pages=2077–2151 |doi=10.1098/rstb.1998.0353|pmc=1692427 }}</ref> The French palaeontologist Cécile Mourer-Chauviré and colleagues also considered the two as belonging to separate genera in 1999.<ref name="Avifauna">{{Cite journal | last1 = Mourer-Chauvire | first1 = C. | last2 =  Bour | first2 = R. | last3 =  Ribes | first3 = S. | last4 =  Moutou | first4 = F. | title = 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 | journal = Smithsonian Contributions to Paleobiology | volume = 89 | pages = 23 | year = 1999 | hdl = 10088/2005 }}</ref>

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.<ref name="Lost Land"/> In 2019, Hume supported the distinction of the two genera, and cited the relation between the extinct [[Mauritius scops owl]] and the [[Rodrigues scops owl]] 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. He suggested their ancestors could have arrived on the Mascarenes during the middle [[Miocene]] at the earliest, but it may have happened more recently. 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.<ref name="Hume2019">{{cite journal |last1=Hume |first1=J. P. |title=Systematics, morphology and ecology of rails (Aves: Rallidae) of the Mascarene Islands, with one new species |journal=Zootaxa |date=2019 |volume=4626 |issue=1 |pages=9–37, 63–67 |doi=10.11646/zootaxa.4626.1.1|pmid=31712544 |s2cid=198258434 }}</ref>

==Description==
[[File:Extinctbirds1907 P29 Aphanapteryx bonasia0349AA.jpg|thumb|alt=drawing of red rail|1907 restoration 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.<ref name="OlsonB"/> It was about {{convert|35|-|40|cm|abbr=on}} long, and the male may have weighed {{convert|1.3|kg|abbr=on}} and the female {{convert|1|kg|abbr=on}}.<ref name="Livezey2003"/> Its plumage was reddish brown all over, and the [[feather]]s were fluffy and hairlike; the tail was not visible in the living bird and the short wings likewise also nearly disappeared in the [[plumage]]. It had a long, slightly curved, brown [[beak|bill]], and some illustrations suggest it had a [[nape]] crest.<ref name="Fuller Extinct"/> The bird perhaps resembled a lightly built kiwi, and it has also been likened to a [[limpkin]], both in appearance and behaviour.<ref name="Lost Land"/><ref name="Fuller Extinct">{{cite book
 | last = Fuller
 | first = E.
 | author-link = Errol Fuller
 | title = Extinct Birds
 | publisher = Comstock
 | edition = revised
 | location = New York
 | year = 2001
 | pages = 96–97
 | isbn = 978-0-8014-3954-4 }}</ref> 

The cranium of the red rail was the largest among Mascarene rails, and was compressed from top to bottom in side view. The [[premaxilla]] that comprised most of the upper bill was long (nearly 47% longer than the cranium) and narrow, and ended in a sharp point. The [[narial]] (nostril) openings were 50% of the [[Rostrum (anatomy)|rostrum]]'s length, and prominent, elongate [[foramina]] (openings) ran almost to the front edge of the narial opening. The mandibular rostrum of the lower jaw was long, with the length of the [[mandibular symphysis]] (where the halves of the mandible connect) being about 79% of the cranium's length. The mandible had large, deep set foramina, which ran almost up to a deep [[Sulcus (morphology)|sulcus]] (furrow). Hume examined all available upper beaks in 2019, and while he found no differences in curvature, he thought the differences in length was most likely due to sexual dimorphism.<ref name="Hume2019"/> 

The [[scapula]] (shoulder blade) was wide in side view, and the [[coracoid]] was comparatively short, with a wide shaft.<ref name="Hume2019"/> The [[sternum]] (breast bone) and [[humerus]] (upper arm bone) were small, indicating that it had lost the power of flight. The humerus was {{convert|60|-|66|mm|abbr=on}}, and its shaft was strongly curved from top to bottom. The [[ulna]] (lower arm bone) was short and strongly arched from top to bottom. Its legs were long and slender for such a large bird, but the [[pelvis]] was very wide, robust, and compact, and was {{convert|60|mm|abbr=on}} in length. The [[femur]] (thigh-bone) was very robust, {{convert|69|-|71|mm|abbr=on}} long, and the upper part of the shaft was strongly arched. The [[tibiotarsus]] (lower leg bone) was large and robust, especially the upper and lower ends, and was {{convert|98|-|115|mm|abbr=on}} long. The [[fibula]] was short and robust. The [[tarsometatarsus]] (ankle bone) was large and robust, and {{convert|79|mm|abbr=on}} long.<ref name="Newton & Gadow"/><ref name="Hume2019"/> The red rail differed from the Rodrigues rail 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.<ref name="OlsonB"/> The red rail was also larger, with somewhat smaller wings, but their leg proportions were similar.<ref name="Günther& Newton"/> The pelvis and [[sacrum]] was also similar.<ref name="Newton & Gadow"/> 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.<ref>{{cite conference |last1=Herremans |first1=M. |title=Trends in the evolution of insular land birds, exemplified by the Comoros, Seychelles and Mascarenes. |url=https://www.researchgate.net/publication/311907872 |conference=Proceedings International Symposium on Vertebrate Biogeography and Systematics in the Tropics | pages = 249–260 | location = Bonn}}</ref><ref name="Livezey2003">{{cite journal |last1=Livezey |first1=B. C. |title=Evolution of flightlessness in rails (Gruiformes: Rallidae): phylogenetic, ecomorphological, and ontogenetic perspectives |journal=Ornithological Monographs |date=2003 |issue=53 |pages=iii–654 |doi=10.2307/40168337 |jstor=40168337}}</ref> 

===Contemporary descriptions===
[[File:Gelderland1601-1603 Aphanapteryx bonasia.jpg|thumb|upright|alt=drawing of red rail|1601 sketches of a killed or stunned specimen, attributted to Joris Laerle]]
Mundy visited Mauritius in 1638 and described the red rail as follows:

{{Quotation|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.<ref>{{Cite journal | last1 = Sclater | first1 = W. l.| title = The "Mauritius Hen" of Peter Mundy | doi = 10.1111/j.1474-919X.1915.tb08192.x | journal = Ibis | volume = 57 | issue = 2 | pages = 316–319 | year = 1915| url = https://www.biodiversitylibrary.org/part/378278}}</ref>}}

Another English traveller, John Marshall, described the bird as follows in 1668:
{{Quotation|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.<ref name="Lost Land"/>}}

===Contemporary depictions===
[[File:Edwards' Dodo.jpg|thumb|left|''Edwards' [[Dodo]]'', a 1626 painting by [[Roelant Savery]], possibly showing a red rail (or a [[bittern]]) in the lower right|alt=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]]

The two most realistic contemporary depictions of red rails, the Hoefnagel painting from ca. 1610 and the sketches from the ''Gelderland'' ship's journal from 1601 attributed to Laerle, where brought to attention in the 19th century.<ref name="Hume2019"/> Much information about the bird's appearance comes from Hoefnagel's painting, based on a bird in the [[menagerie]] of [[Rudolf II, Holy Roman Emperor|Emperor Rudolph II]] around 1610.<ref name="Rothschild">{{Cite book
  | last = Rothschild
  | first = W.
  | author-link = Walter Rothschild, 2nd Baron Rothschild
  | title = Extinct Birds
  | publisher = Hutchinson & Co
  | year = 1907
  | location = London
  | page = 131
  | url = https://archive.org/stream/extinctbirdsatte00roth#page/130/mode/2up
}}</ref> It is the only unequivocal 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.<ref name="Extinct Birds">{{cite book
| last1 = Hume
| first1 = J. P.
| first2 = M.
| last2 = Walters
|year= 2012
|title= Extinct Birds
|publisher= A & C Black
|location= London
|pages= 108–109
|isbn=978-1-4081-5725-1}}</ref> 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.<ref name="Lost Land"/> 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.<ref name="Milne-Edwards">{{cite journal| doi = 10.1111/j.1474-919X.1869.tb06880.x| last = Milne-Edwards | first = A.| year = 1869| title = Researches into the zoological affinities of the bird recently described by Herr von Frauenfeld under the name of ''Aphanapteryx imperialis''| journal = Ibis| volume = 11| issue = 3| pages = 256–275| url = https://archive.org/stream/ibisns05brit#page/256/mode/2up}}</ref><ref>{{cite journal |last1=Milne-Edwards |first1=M. A. |title=XV.— Observations on the ancient fauna of the Mascarene Islands |journal=Annals and Magazine of Natural History |date=1869 |volume=4 |issue=20 |pages=129–132 |doi=10.1080/00222936908696012 |url=https://www.biodiversitylibrary.org/item/88433#page/145/mode/1up}}</ref> This specimen is thought to have been the only red rail that ever reached Europe.<ref>{{cite journal
|last1=Hume|first1=J. P.|last2=Prys-Jones|first2=R. P.
|year=2005
|title=New discoveries from old sources, with reference to the original bird and mammal fauna of the Mascarene Islands, Indian Ocean
|journal=[[Zoologische Mededelingen]]
|volume=79
|issue=3
|pages=85–95
|url=http://www.repository.naturalis.nl/document/42166
|format=PDF}}</ref>

[[File:The Animals Entering Noah's Ark 1570s Jacopo Bassano.jpg|thumb|[[Jacopo Bassano]]'s 1570 painting ''Arca di Noè'', perhaps showing a red rail (or a bittern) in the lower right]] 
The red rail depicted in the ''Gelderland'' journal appears to have been stunned or killed, and the sketch is the earliest record of the species. It is the only illustration of the species drawn on Mauritius, and according to Hume, the most accurate depiction. 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.<ref name="Gelderland">{{cite journal| doi = 10.3366/anh.2003.30.1.13| last = Hume | first = J. P.| year = 2003| title = The journal of the flagship ''Gelderland''&nbsp;– dodo and other birds on Mauritius 1601| journal = Archives of Natural History| volume = 30| issue = 1| pages = 13–27}}</ref><ref name="Hume2019"/> 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 synonyms.<ref name="Fuller Extinct"/> An illustration in van den Broecke's 1646 account (based on his stay on Mauritius in 1617) shows a red rail next to a dodo and a one-horned goat, but is not referenced in the text. An illustration in Herbert's 1634 account (based on his stay in 1629) shows a red rail between a broad-billed parrot and a dodo, and has been referred to as "extremely crude" by Hume. Mundy's 1638 illustration was published in 1919.<ref name="Hume2019"/>

As suggested by Greenway, there are also depictions of what appears to be a red rail in three of the Dutch artist [[Roelant Savery]]'s paintings.<ref name =Greenway>{{cite book
 | last = Greenway
 | first = J. C.
 | title = Extinct and Vanishing Birds of the World
 | publisher = American Committee for International Wild Life Protection 13
 | location = New York
 | year = 1967
 | pages = 117–119
 | isbn = 978-0-486-21869-4}}</ref> In his famous ''Edwards' Dodo'' painting from 1626, a rail-like bird is seen swallowing a frog behind the dodo, but Hume has doubted this identification and that of red rails in other Savery paintings, suggesting may instead show [[Eurasian bitterns]].<ref name="Extinct Birds"/><ref name="Hume2019"/> In 1977, the American ornithologist [[Sidney Dillon Ripley]] noted a bird resembling a red rail figured in the Italian artist [[Jacopo Bassano]]'s painting ''Arca di Noè'' ("[[Noah's Ark]]") from ca. 1570. Cheke 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.<ref name="Cheke87"/> A similar bird is also seen in the Flemish artist [[Jan Brueghel the Elder]]'s ''Noah's Ark'' painting.<ref name="Lost Land"/> Hume concluded that these paintings also show Eurasian bitterns rather than red rails.<ref name="Hume2019"/>

==Behaviour and ecology==
[[File:Aphanapteryx bonasiaIbis1869P007AA.jpg|thumb|upright|1869 adaptation of the Hoefnagel painting]] 

Contemporary accounts are repetitive and do not shed much light on the life history of the red rail. Based on fossil localities, the bird widely occurred on Mauritius, in montane, lowland, and coastal habitats.<ref name="Hume2019"/> The shape of the beak indicates it could have captured [[reptile]]s and [[invertebrate]]s, and the differences in bill length suggests the sexes foraged on items of different sizes. It may also have scavenged breeding colonies of birds and nesting-sites of [[tortoises]], as the Rodrigues rail did.<ref name="Lost Land"/><ref name="Hume2019"/> 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."<ref name="Pretorius">{{Cite journal | doi = 10.1080/08912963.2015.1036750| title = Captive birds on Dutch Mauritius: Bad-tempered parrots, warty pigeons and notes on other native animals| journal = Historical Biology| volume = 28| issue = 6| pages = 1| year = 2015| last1 = Hume | first1 = J. P. | last2 = Winters | first2 = R. | s2cid = 84473440}}</ref> 

Milne-Edwards suggested that since the tip of the red rail's bill was sharp and strong, it fed by crushing [[molluscs]] and other shells, like [[oystercatchers]] do. There were many endemic land-snails on Mauritius, including the large, extinct ''[[Tropidophora carinata]]'', and subfossil shells have been found with puncture holes on their lower surfaces, which suggest predation by birds, probably matching attacks from the beak of the red rail. The similarly sized [[weka]] of New Zealand punctures shells of land-snails to extract meat, but can also swallow ''[[Powelliphanta]]'' snails; Hume suggested the red rail was also able to swallow snails whole. Since Pretorius mentioned the red rail searched for worms in leaf-litter, Hume suggested this could refer to [[nemertean]] and [[planarian]] worms; Mauritius has endemic species of these groups which live in leaf-litter and rotten wood. He could also have referred to the now extinct worm-snake ''[[Madatyphlops cariei]]'', which was up to {{convert|200|mm|abbr=on}} long, and probably lived in leaf-litter like its relatives do.<ref name="Hume2019"/><ref name="Lost Land"/><ref>{{cite book
 | last1 = Griffiths
 | first1 = O. L.
 | first2 = V. F. B.
 | last2 = Florens
 | title = Non-Marine Molluscs of the Mascarene Islands
 | publisher = Bioculture Press
 | location = Mauritius
 | year = 2006
 | page = 185
 | isbn = 978-99949-22-05-5 }}</ref> 
[[File:Tropidophora carinata Reunion.jpg|thumb|left|Shells of the extinct land-snail ''[[Tropidophora carinata]]'' have been found with puncture holes possibly made by feeding red rails]] 
Hume noted that the front of the red rail's jaws were pitted with numerous foramina, running from the nasal aperture to almost the tip of the premaxilla. These were mostly oval, varying in depth and inclination, and became shallower hindward from the tip. Similar foramina can be seen in probing birds, such as kiwis, ibises, and [[sandpipers]]. While unrelated, these three bird groups share a foraging strategy; they probe for live food beneath substrate, and have elongated bills with clusters of [[mechanoreceptors]] concentrated at the tip. Their bill-tips allow them to detect buried prey by sensing cues from the substrate. The foramina on the bill of the red rail were comparable to those in other probing rails with long bills (such as the extinct [[snipe-rail]]), though not as concentrated on the tip, and the front end of the bill's curvature also began at the front of the nasal opening (as well as the same point in the mandible). The bill's tip was thereby both strong and very sensitive, and a useful tool for probing for invertebrates.<ref name="Hume2019"/> 

A 1631 letter probably by the Dutch lawyer Leonardus Wallesius (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:<ref name="LostManuscript">{{cite journal|last1=Winters|first1=R.|last2=Hume|first2=J. P.|last3=Leenstra|first3=M.|title=A famine in Surat in 1631 and Dodos on Mauritius: a long lost manuscript rediscovered|journal=Archives of Natural History|date=2017|volume=44|issue=1|pages=134–150|doi=10.3366/anh.2017.0422}}</ref><ref name="Hume2019"/> 
{{Quotation|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.<ref name="LostManuscript"/>}}
[[File:Lophopsittacus.mauritianus.jpg|thumb|alt=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]]
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 [[predator]]s, were curious and not afraid of humans.<ref name="Fuller Extinct"/> Herbert described its behaviour towards red cloth in 1634:
{{Quotation|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.<ref name="Lost Land"/>}}
Many other endemic species of Mauritius became extinct after the arrival of humans to the island 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]].<ref>{{cite journal| doi = 10.1017/S0030605300020457| last = Cheke | first = A. S.| year = 1987| title = The legacy of the dodo—conservation in Mauritius| journal = Oryx| volume = 21| issue = 1| pages = 29–36
| doi-access = free}}</ref> The surviving endemic [[fauna]] is still seriously threatened.<ref>{{cite journal| doi = 10.1017/S0030605300012643| last = Temple | first = S. A.| year = 1974| title = Wildlife in Mauritius today| journal = Oryx| volume = 12| issue = 5| pages = 584–590 | doi-access = free}}</ref> 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 scops 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.<ref name="Lost Land"/>

==Relationship with humans==
[[File:View of the Mauritius roadstead - engraving.jpg|thumb|left|alt=Men working in a wooded area on a 16th-century illustration|Depiction of Dutch activities on Mauritius in 1598, with various birds]]
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]].<ref name="Fuller Extinct"/> The last detailed account of the red rail was by the German pastor Johann Christian Hoffmann, on Mauritius in the early 1670s,<ref name="Hume2019"/> who described a hunt as follows:
{{Quotation|...&nbsp;[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.<ref name="Lost Land"/>}}

[[File:Aphanapteryx bonasia.JPG|thumb|alt=drawing that includes a red rail|[[Pieter van den Broecke]]'s 1617 drawing of a dodo, a one-horned goat, and a red rail; after the dodo became extinct, its name may have been transferred to the red rail]]
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.<ref name="Extinction date"/> Milne-Edwards suggested that early travellers may have confused young dodos with red rails.<ref name =Greenway/> The British ornithologist [[Alfred Newton]] (brother of Edward) suggested in 1868 that the name of the dodo was transferred to the red rail after the former had gone extinct.<ref name="NewtonA.">{{cite journal |last1=Newton |first1=A. |title=Recent ornithological publications |journal=Ibis |date=1868 |volume=4 |issue=2 |pages=479–482 |url=https://www.biodiversitylibrary.org/item/55161#page/507/mode/1up}}</ref> Cheke suggested in 2008 that all post 1662 references to "dodos" therefore refer to the rail instead.<ref name="Extinction date">{{cite journal| doi = 10.1111/j.1474-919X.2006.00478.x| last = Cheke | first = A. S.| year = 2006| title = Establishing extinction dates&nbsp;– the curious case of the Dodo ''Raphus cucullatus'' and the Red Hen ''Aphanapteryx bonasia''| journal = Ibis| volume = 148| pages = 155–158
}}</ref> 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.<ref name="Lost Land"/> The British writer [[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.<ref name="Fuller Extinct"/> 

In 2020, Cheke and the British researcher Jolyon C. Parish suggested that all mentions of dodos after the mid-17th century instead referred to red rails, and that the dodo had disappeared due to predation by [[feral pigs]] during a hiatus in settlement of Mauritius (1658–1664). The dodo's extinction therefore was not realised at the time, since new settlers had not seen real dodos, but as they expected to see flightless birds, they referred to the red rail by that name instead. Since red rails probably had larger clutches than dodos (as in other rails) and their eggs could be incubated faster, and their nests were perhaps concealed like those of the Rodrigues rail, they probably bred more efficiently, and were less vulnerable to pigs. They may also have foraged from the digging, scraping and rooting of the pigs, as does the weka.<ref name="Saga">{{cite journal |last1=Cheke |first1=A. S. |last2=Parish |first2=J. C. |title=The Dodo and the Red Hen, A Saga of Extinction, Misunderstanding, and Name Transfer: A Review |journal=Quaternary |date=2020 |volume=3 |issue=1 |pages=4 |doi=10.3390/quat3010004|doi-access=free |bibcode=2020Quat....3....4C }}</ref>

230 years before [[Charles Darwin]]'s [[theory of evolution]], the appearance of the red rail and the dodo led Mundy to speculate:
{{Quotation|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.<ref name="Fuller Extinct"/>}}

===Extinction===
[[File:Aphanapteryx bonasia by Mundy.jpg|thumb|alt=drawing of red rail|Drawing by [[Peter Mundy]], 1638]]
Many terrestrial rails are flightless, and island populations are particularly vulnerable to anthropogenic (human-caused) changes; as a result, rails have suffered more extinctions than any other family of birds. All six endemic species of Mascarene rails are extinct, all caused by human activities.<ref name="Hume2019"/> 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.<ref name="Cheke87">{{Cite book| last1 = Cheke | first1 = A. S. | editor1-last = Diamond| editor1-first = A. W.| doi = 10.1017/CBO9780511735769.003 | chapter = An ecological history of the Mascarene Islands, with particular reference to extinctions and introductions of land vertebrates | title = Studies of Mascarene Island Birds | url = https://archive.org/details/studiesmascarene00diam_318 | url-access = limited | pages = [https://archive.org/details/studiesmascarene00diam_318/page/n11 5]–89 | year = 1987 | isbn = 978-0-521-11331-1| location = Cambridge | publisher = Cambridge University Press }}</ref> 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]].<ref name="Hume2019"/>

Since the red rail was referred to by the names of the dodo in the late 1600s, it is uncertain which is the latest account of the latter.<ref name="Hume2019"/> 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 become rare. He was the last source to mention the bird, so it is assumed that it became extinct around 1700.<ref name="Fuller Extinct"/><ref name="Leguat">{{cite book
 | last = Leguat
 | first = F.
 | author-link = François Leguat
 | title = The voyage of François Leguat of Bresse, to Rodriguez, Mauritius, Java, and the Cape of Good Hope
 | publisher = Hakluyt Society
 | location = London
 | year = 1891
 | page = [https://archive.org/details/voyageoffranoi01legu/page/71 71]
 | url = https://archive.org/details/voyageoffranoi01legu }}</ref> [[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.<ref name="Pretorius"/><ref name="Hume2019"/>

==See also==
* [[Holocene extinction]]
* [[List of extinct birds]]

{{clear}}

==References==
{{Reflist|30em}}

==External links==
* {{Commons category-inline|Aphanapteryx}}
* {{Wikispecies-inline|Rallidae}}

{{Birds|state=collapsed}}
{{Gruiformes|R.|state=collapsed}}
{{Portal bar|Birds|Animals|Biology|Africa|Madagascar|Paleontology}}
{{Taxonbar|from=Q844868}}

[[Category:Birds described in 1848]]
[[Category:Bird extinctions since 1500]]
[[Category:Extinct animals of Mauritius]]
[[Category:Extinct birds of Indian Ocean islands]]
[[Category:Extinct flightless birds]]
[[Category:Rallidae]]
[[Category:Endemic birds of Mauritius|†]]