Rail (bird)

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Rails
Temporal range: Early Eocene-Recent [1]
Dusky moorhen442.jpg
Dusky moorhen, Gallinula tenebrosa
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Gruiformes
Family: Rallidae
Rafinesque, 1815
Genera

Some 40 living, and see below.

Rails (avian family Rallidae) are a large, cosmopolitan family of small- to medium-sized terrestrial and/or semi-amphibious birds. The family exhibits considerable diversity in its forms, and includes such ubiquitous species as the crakes, coots, and gallinule; other rail species are extremely rare or endangered. Many are associated with wetland habitats, some being semi-aquatic like waterfowl (such as the coot), but many more are wading birds or shorebirds. The ideal rail habitats are marsh areas, including rice paddies, and flooded fields or open forest. They are especially fond of dense vegetation for nesting. [2] The rail family is found in every terrestrial habitat with the exception of dry desert, polar or freezing regions, and alpine areas (above the snow line). Members of Rallidae occur on every continent except Antarctica. Numerous unique island species are known.

Contents

Name

"Rail" is the anglicized respelling of the French râle, from Old French rasle. It is named from its harsh cry, in Vulgar Latin *rascula, from Latin rādere ("to scrape"). [3]

Morphology

South Island takahe (Porphyrio hochstetteri) from behind, showing the short, soft, and fluffy remiges typical of flightless rails Takahe.jpg
South Island takahē (Porphyrio hochstetteri) from behind, showing the short, soft, and fluffy remiges typical of flightless rails

The rails are a family of small to medium-sized, ground-living birds. They vary in length from 12 to 63 cm (5 to 25 in) and in weight from 20 to 3,000 g (0.7 oz to 6 lb 10 oz). Some species have long necks and in many cases are laterally compressed. [4]

The bill is the most variable feature within the family. In some species, it is longer than the head (like the clapper rail of the Americas); in others, it may be short and wide (as in the coots), or massive (as in the purple gallinules). [5] A few coots and gallinules have a frontal shield, which is a fleshy, rearward extension of the upper bill. The most complex frontal shield is found in the horned coot. [6]

Rails exhibit very little sexual dimorphism in either plumage or size. Two exceptions are the watercock (Gallicrex cinerea) and the little crake (Zapornia parva). [7]

Flight and flightlessness

The wings of all rails are short and rounded. The flight of those Rallidae able to fly, while not powerful, can be sustained for long periods of time, and many species migrate annually. The weakness of their flight, however, means they are easily blown off course, thus making them common vagrants, a characteristic that has led them to colonize many isolated oceanic islands. Furthermore, these birds often prefer to run rather than fly, especially in dense habitat. Some are also flightless at some time during their moult periods. [8]

Flightlessness in rails is one of the best examples of parallel evolution in the animal kingdom. Of the roughly 150 historically known rail species, 31 extant or recently extinct species evolved flightlessness from volant (flying) ancestors. [9] This process created the endemic populations of flightless rails seen on Pacific islands today.

Many island rails are flightless because small island habitats without mammalian predators eliminate the need to fly or move long distances. Flight makes intense demands, with the keel and flight muscles taking up to 40% of a bird's weight. [9] Reducing the flight muscles, with a corresponding lowering of metabolic demands, reduces the flightless rail's energy expenditures. [10] For this reason, flightlessness makes it easier to survive and colonize an island where resources may be limited. [11] This also allows for the evolution of multiple sizes of flightless rails on the same island as the birds diversify to fill niches. [12]

In addition to energy conservation, certain morphological traits also affect rail evolution. Rails have relatively small flight muscles and wings to begin with. [13] In rails, the flight muscles make up only 12–17% of their overall body mass. [9] This, in combination with their terrestrial habits and behavioral flightlessness, is a significant contributor to the rail's remarkably fast loss of flight; [14] as few as 125,000 years were needed for the Laysan rail to lose the power of flight and evolve the reduced, stubby wings only useful to keep balance when running quickly. [15] Indeed, some argue that measuring the evolution of flightlessness in rails in generations rather than millennia might be possible. [11]

Another factor that contributes to the occurrence of the flightless state is a climate that does not necessitate seasonal long-distance migration; this is evidenced by the tendency to evolve flightlessness at a much greater occurrence in tropical islands than in temperate or polar islands. [16]

American coot (Fulica americana) skeleton on display at the Museum of Osteology. American Coot skeleton.jpg
American coot (Fulica americana) skeleton on display at the Museum of Osteology.

It is paradoxical, since rails appear loath to fly, that the evolution of flightless rails would necessitate high dispersal to isolated islands. [13] Nonetheless, three species of small-massed rails, Gallirallus philippensis , Porphyrio porphyrio , and Porzana tabuensis , exhibit a persistently high ability to disperse long distances among tropic Pacific islands, [13] though only the latter two gave rise to flightless endemic species throughout the Pacific Basin. [17] In examining the phylogeny of G. philippensis, although the species is clearly polyphyletic (it has more than one ancestral species), it is not the ancestor of most of its flightless descendants, revealing that the flightless condition evolved in rails before speciation was complete. [17]

A consequence of lowered energy expenditure in flightless island rails has also been associated with evolution of their "tolerance" and "approachability". [16] For example, the (non-Rallidae) Corsican blue tits exhibit lower aggression and reduced territorial defense behaviors than do their mainland European counterparts, [18] but this tolerance may be limited to close relatives. [19] The resulting kin-selecting altruistic phenomena reallocate resources to produce fewer young that are more competitive and would benefit the population as an entirety, rather than many young that would exhibit less fitness. [16] Unfortunately, with the human occupation of most islands in the past 5,000 to 35,000 years, selection has undoubtedly reversed the tolerance into a wariness of humans and predators, causing species unequipped for the change to become susceptible to extinction. [16]

Behaviour and ecology

In general, members of the Rallidae are omnivorous generalists. Many species eat invertebrates, as well as fruit or seedlings. A few species are primarily herbivorous. [2] The calls of Rallidae species vary and are often quite loud. Some are whistle-like or squeak-like, while others seem unbirdlike. [20] Loud calls are useful in dense vegetation, or at night where seeing another member of the species is difficult. Some calls are territorial. [5]

The most typical family members occupy dense vegetation in damp environments near lakes, swamps, or rivers. Reed beds are a particularly favoured habitat. Those that migrate do so at night.

Most nest in dense vegetation. In general, they are shy, secretive, and difficult to observe. Most species walk and run vigorously on strong legs, and have long toes that are well adapted to soft, uneven surfaces. They tend to have short, rounded wings, and although they are generally weak fliers, they are, nevertheless, capable of covering long distances. Island species often become flightless, and many of them are now extinct following the introduction of terrestrial predators such as cats, foxes, weasels, mongooses, rats, and pigs.

Many reedbed species are secretive (apart from loud calls), crepuscular, and have laterally flattened bodies. In the Old World, long-billed species tend to be called rails and short-billed species crakes. North American species are normally called rails irrespective of bill length. The smallest of these is Swinhoe's rail, at 13 cm (5.1 in) and 25 g. The larger species are also sometimes given other names. The black coots are more adapted to open water than their relatives, and some other large species are called gallinules and swamphens. The largest of this group is the takahē, at 65 cm (26 in) and 2.7 kg (6.0 lb).

The rails have suffered disproportionally from human changes to the environment, and an estimated [21] [22] [23] several hundred species of island rails have become extinct because of this. Several island species of rails remain endangered, and conservation organisations and governments continue to work to prevent their extinction.

Reproduction

The breeding behaviors of many Rallidae species are poorly understood or unknown. Most are thought to be monogamous, although polygyny and polyandry have been reported. [24] Most often, they lay five to 10 eggs. Clutches as small as one or as large as 15 eggs are known. [24] Egg clutches may not always hatch at the same time. Chicks become mobile after a few days. They often depend on their parents until fledging, which happens around 1 month old. [6]

Rallidae and humans

The Guam rail is an example of an island species that has been badly affected by introduced species. GuamRail02.jpg
The Guam rail is an example of an island species that has been badly affected by introduced species.

Some larger, more abundant rails are hunted and their eggs collected for food. [25] The Wake Island rail was hunted to extinction by the starving Japanese garrison after the island was cut off from supply during World War II. [26] At least two species, the common moorhen and the American purple gallinule, have been considered pests. [25]

Threats and conservation

Due to their tendencies towards flightlessness, many island species have been unable to cope with introduced species. The most dramatic human-caused extinctions occurred in the Pacific Ocean as people colonised the islands of Melanesia, Polynesia, and Micronesia, during which an estimated 750–1800 species of birds became extinct, half of which were rails. [27] Some species that came close to extinction, such as the Lord Howe woodhen, and the takahē, have made modest recoveries due to the efforts of conservation organisations. The Guam rail came perilously close to extinction when brown tree snakes were introduced to Guam, but some of the last remaining individuals were taken into captivity and are breeding well, though attempts at reintroduction have met with mixed results. [28] [29] [30]

Systematics and evolution

The family Rallidae was introduced (as Rallia) by the French polymath Constantine Samuel Rafinesque in 1815. [31] [32] The family has traditionally been grouped with two families of larger birds, the cranes and bustards, as well as several smaller families of usually "primitive" midsized amphibious birds, to make up the order Gruiformes. The alternative Sibley-Ahlquist taxonomy, which has been widely accepted in America, raises the family to ordinal level as the Ralliformes. Given uncertainty about gruiform monophyly, this may or may not be correct; it certainly seems more justified than most of the Sibley-Ahlquist proposals. However, such a group would probably also include the Heliornithidae (finfoots and sungrebes), an exclusively tropical group that is somewhat convergent with grebes, and usually united with the rails in the Ralli.

The cladogram below showing the phylogeny of the living and recently extinct Rallidae is based on a study by Juan Garcia-R and collaborators published in 2020. [7] The genera and number of species are taken from the list maintained by Frank Gill, Pamela Rasmussen and David Donsker on behalf of the International Ornithological Committee (IOC). [33] The names of the subfamilies and tribes are those proposed by Jeremy Kirchman and collaborators in 2021. [1]

Rallidae 
Rallinae
Pardirallini

Neocrex – crakes (2 species)

Pardirallus – rails (3 species)

Amaurolimnas – uniform crake

Aramides – rails (8 species)

Rallini

Rallus – rails (14 species)

Crecopsis – African crake

Rougetius – Rouget's rail

Dryolimnas – rails (2 species of which 1 is extinct)

Crex – corn crake

Lewinia – rails (4 species)

Aptenorallus – Calayan rail

Habroptila – invisible rail

Gallirallus – weka

Eulabeornis – chestnut rail

Hypotaenidia – rails and woodhen (12 species of which 4 are extinct)

Himantornithinae
Himantornithini

Himantornis – Nkulengu rail

Fulicini

Porzana – crakes and sora (3 species)

Tribonyx – nativehens (2 species)

Paragallinula – lesser moorhen

Gallinula – moorhens (7 species of which 2 are extinct)

Fulica – coots (11 species of which 1 is extinct)

Porphyrionini

Porphyrio – swamphens, gallinules and takahes (12 species of which 2 are extinct)

Laterallini

Micropygia – ocellated crake

Rufirallus – crakes (2 species)

Coturnicops – rails and a crake (3 species)

Laterallus – crakes and rails (13 species)

Zapornini

Zapornia – rails and crakes (15 species of which 5 are extinct)

Rallina – crakes (4 species)

Amaurornithini

Megacrex – New Guinea flightless rail

Poliolimnas – white-browed crake

Aenigmatolimnas – striped crake

Gallicrex – watercock

Amaurornis – waterhen and bush-hens (5 species)

Extant genera

The list maintained on behalf of the International Ornithological Committee (IOC) contains 152 species divided into 43 genera. [33] For more detail, see List of rail species.

Additionally, many prehistoric rails of extant genera are known only from fossil or subfossil remains, such as the Ibiza rail (Rallus eivissensis). These have not been listed here; see the genus accounts and the articles on fossil and Late Quaternary prehistoric birds for these species.

Pieter van den Broecke's 1617 drawing of the red rail of Mauritius, Aphanapteryx bonasia Aphanapteryx bonasia.JPG
Pieter van den Broecke's 1617 drawing of the red rail of Mauritius, Aphanapteryx bonasia

Recently extinct genera

1888 color lithograph of a rail Rail, from the Game Birds series (N40) for Allen & Ginter Cigarettes MET DP839177.jpg
1888 color lithograph of a rail

The undescribed Fernando de Noronha rail, genus and species undetermined, survived to historic times. The extinct genus Nesotrochis from the Greater Antilles was formerly considered to be a rail, but based on DNA evidence is now known to be an independent lineage of gruiform more closely related to Sarothruridae and adzebills.

Fossil record

Fossil species of long-extinct prehistoric rails are richly documented from the well-researched formations of Europe [34] and North America, as well from the less comprehensively studied strata elsewhere:

Doubtfully placed here

These taxa may or may not have been rails:

The presumed scolopacid wader Limosa gypsorum (Montmartre Late Eocene of France) is sometimes considered a rail and then placed in the genus Montirallus. [45]

See also

Related Research Articles

<span class="mw-page-title-main">Charadriiformes</span> Order of birds

Charadriiformes is a diverse order of small to medium-large birds. It includes about 390 species and has members in all parts of the world. Most charadriiform birds live near water and eat invertebrates or other small animals; however, some are pelagic (seabirds), others frequent deserts, and a few are found in dense forest. Members of this group can also collectively be referred to as shorebirds.

<span class="mw-page-title-main">Gruiformes</span> Order of birds

The Gruiformes are an order containing a considerable number of living and extinct bird families, with a widespread geographical diversity. Gruiform means "crane-like".

<span class="mw-page-title-main">Anatidae</span> Biological family of water birds

The Anatidae are the biological family of water birds that includes ducks, geese, and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica. These birds are adapted for swimming, floating on the water surface, and in some cases diving in at least shallow water. The family contains around 174 species in 43 genera.

<span class="mw-page-title-main">Inaccessible Island rail</span> Small flightless bird in the family Rallidae endemic to an island in the Tristan Archipelago

The Inaccessible Island rail is a small bird of the rail family, Rallidae. Endemic to Inaccessible Island in the Tristan Archipelago in the isolated south Atlantic, it is the smallest extant flightless bird in the world. The species was described by physician Percy Lowe in 1923 but had first come to the attention of scientists 50 years earlier. The Inaccessible Island rail's affinities and origin were a long-standing mystery; in 2018 its closest relative was identified as the South American dot-winged crake, and it was proposed that both species should be nested within the genus Laterallus.

<span class="mw-page-title-main">Moorhen</span> Genus of birds

Moorhens—sometimes called marsh hens—are medium-sized water birds that are members of the rail family (Rallidae). Most species are placed in the genus Gallinula, Latin for "little hen". They are close relatives of coots. They are often referred to as (black) gallinules. Recently, one of the species of Gallinula was found to have enough differences to form a new genus Paragallinula with the only species being the lesser moorhen.

<span class="mw-page-title-main">Lord Howe woodhen</span> Species of bird

The Lord Howe woodhen also known as the Lord Howe Island woodhen or Lord Howe (Island) rail, is a flightless bird of the rail family, (Rallidae). It is endemic to Lord Howe Island off the Australian coast. It is currently classified as endangered by the IUCN.

<span class="mw-page-title-main">Swamphen</span> Genus of birds

Porphyrio is the swamphen or swamp hen bird genus in the rail family. It includes some smaller species of gallinules which are sometimes separated as genus Porphyrula or united with the gallinules proper in Gallinula. The Porphyrio gallinules are distributed in the warmer regions of the world. The group probably originated in Africa in the Middle Miocene, before spreading across the world in waves from the Late Miocene to Pleistocene.

<i>Gallirallus</i> Genus of birds

Gallirallus is a genus of rails that live in the Australasian-Pacific region. The genus is characterised by an ability to colonise relatively small and isolated islands and thereafter to evolve flightless forms, many of which became extinct following Polynesian settlement.

<span class="mw-page-title-main">Tahiti rail</span> Extinct species of bird from Tahiti

The Tahiti rail, Tahitian red-billed rail, or Pacific red-billed rail is an extinct species of rail that lived on Tahiti. It was first recorded during James Cook's second voyage around the world (1772–1775), on which it was illustrated by Georg Forster and described by Johann Reinhold Forster. No specimens have been preserved. As well as the documentation by the Forsters, there have been claims that the bird also existed on the nearby island of Mehetia. The Tahiti rail appears to have been closely related to, and perhaps derived from, the buff-banded rail, and has also been historically confused with the Tongan subspecies of that bird.

<i>Rallus</i> Genus of birds

Rallus is a genus of wetland birds of the rail family. Sometimes, the genera Lewinia and Gallirallus are included in it. Six of the species are found in the Americas, and the three species found in Eurasia, Africa and Madagascar are very closely related to each other, suggesting they are descended from a single invasion of a New World ancestor.

<i>Porzana</i> Genus of birds

Porzana is a genus of birds in the crake and rail family, Rallidae. Its scientific name is derived from Venetian terms for small rails. The spotted crake is the type species.

<span class="mw-page-title-main">Pelagornithidae</span> Extinct family of seabirds

The Pelagornithidae, commonly called pelagornithids, pseudodontorns, bony-toothed birds, false-toothed birds or pseudotooth birds, are a prehistoric family of large seabirds. Their fossil remains have been found all over the world in rocks dating between the Early Paleocene and the Pliocene-Pleistocene boundary.

<span class="mw-page-title-main">Invisible rail</span> Large flightless bird endemic to Indonesia

The invisible rail, Wallace's rail, or drummer rail is a large flightless rail that is endemic to the island of Halmahera in Northern Maluku, Indonesia, where it inhabits impenetrable sago swamps adjacent to forests. Its plumage is predominantly dark slate-grey, and the bare skin around its eyes, the long, thick bill, and the legs are all bright red. Its call is a low drumming sound which is accompanied by wing-beating. The difficulty of seeing this shy bird in its dense habitat means that information on its behaviour is limited.

<i>Laterallus</i> Genus of birds

Laterallus is a genus of birds in the rail family Rallidae. These small, relatively short-billed terrestrial rails are found among dense vegetation near water in the Neotropics, although a single species, the black rail, also occurs in the United States.

<span class="mw-page-title-main">Dot-winged crake</span> Species of bird

The dot-winged crake is a Vulnerable species of bird in subfamily Rallinae of family Rallidae, the rails, gallinules, and coots. It is found in Argentina, Brazil, Chile, and Uruguay.

<i>Rallina</i> Genus of birds

Rallina is a genus of bird in the rail family, Rallidae. It contains four species found in forest and marshland in Asia and Australasia. They are 18–34 cm long and mainly chestnut or brown, often with black and white markings. They are four species that are now placed in the genus Rallicula that were previously included in the genus Rallina. In fact, some taxonomic authorities continue to place them there. A fifth species, the Great Nicobar crake was proposed but not accepted as a separate species.

The Mangaia crake is an extinct species of flightless bird in the rail family, Rallidae.

<span class="mw-page-title-main">Hova gallinule</span> Extinct species of bird

The Hova gallinule is an extinct bird in a monotypic genus in the rail family. It was a large gallinule that was endemic to Madagascar. About the size of the Tasmanian nativehen, it was originally placed in the same genus, Tribonyx. Hovacrex may also be synonymous with Gallinula. The lack of wing bones in the subfossil material found at Sirabé in central Madagascar makes it uncertain as to whether the species was flightless.

References

  1. 1 2 Kirchman, J.J.; Rotzel McInerney, N.; Giarla, T.C.; Olson, S.L.; Slikas, E.; Fleischer, R.C. (2021). "Phylogeny based on ultra-conserved elements clarifies the evolution of rails and allies (Ralloidea) and is the basis for a revised classification". Ornithology. 138 (4): ukab042. doi: 10.1093/ornithology/ukab042 .
  2. 1 2 Horsfall & Robinson (2003): pp. 206–207
  3. Reedman, Ray (2016). Lapwings, Loons and Lousy Jacks: The How and Why of Bird Names. Exeter, UK: Pelagic Publishing Ltd. ISBN   9781784270933 via Google Books.
  4. "Rail | bird". Encyclopedia Britannica. Archived from the original on 2021-06-10. Retrieved 2021-06-03.
  5. 1 2 Horsfall & Robinson (2003): p. 208
  6. 1 2 Horsfall & Robinson (2003): p. 210
  7. 1 2 Garcia-R, J.C.; Lemmon, E.M.; Lemmon, A.R.; French, N. (2020). "Phylogenomic reconstruction sheds light on new relationships and timescale of rails (Aves: Rallidae) evolution". Diversity. 12 (2): 70. doi: 10.3390/d12020070 .
  8. Horsfall & Robinson (2003): p. 209
  9. 1 2 3 Kirchman (2012)
  10. McNab & Ellis (2006)
  11. 1 2 McNab (1994)
  12. Trewick (1997)
  13. 1 2 3 McNab and Ellis 2006
  14. Livezey 2003
  15. Slikas et al. (2002)
  16. 1 2 3 4 McNab 2002
  17. 1 2 Kirchman 2012
  18. Perret and Blondel 1993
  19. Granjon and Cheylan 1989
  20. Horsfall & Robinson (2003): p. 207
  21. Duncan, Richard P.; Boyer, Alison G.; Blackburn, Tim M. (2013). "Magnitude and variation of prehistoric bird extinctions in the Pacific". PNAS. 110 (16): 6436–6441. Bibcode:2013PNAS..110.6436D. doi: 10.1073/pnas.1216511110 . PMC   3631643 . PMID   23530197.
  22. Platt, John R. (May 2018). "Memorializing the Wake Island Rail: An Extinction Caused by War". TheRevelator.org. Archived from the original on 2019-06-02. Retrieved 2018-07-29.
  23. Steadman, David W. (2006). Extinction and Biogeography of Tropical Pacific Birds. University of Chicago Press. p. 296. ISBN   9780226771427.
  24. 1 2 Horsfall & Robinson (2003): pp. 209–210
  25. 1 2 Horsfall & Robinson (2003): p. 211
  26. BLI (2007)
  27. Steadman (2006)
  28. "GUAM: BROWN TREE SNAKE RESPONSIBLE FOR EXTINCTION OF 5 SPECIES". AP Archive. February 1996. Archived from the original on 2018-07-29. Retrieved 2018-07-29.
  29. "Guam Rail Gallirallus owstoni". San Diego Zoo.org. Archived from the original on 2018-07-29. Retrieved 2018-07-29.
  30. Hurrell, Shaun. "Is this Guam bird coming back from extinction in the wild?". BirdLife International.org. Archived from the original on 2018-07-29. Retrieved 2018-07-29.
  31. Rafinesque, Constantine Samuel (1815). Analyse de la nature ou, Tableau de l'univers et des corps organisés (in French). Palermo: Self-published. p. 70. Archived from the original on 2018-08-19. Retrieved 2018-02-21.
  32. Bock, Walter J. (1994). History and Nomenclature of Avian Family-Group Names. Bulletin of the American Museum of Natural History. Vol. Number 222. New York: American Museum of Natural History. pp. 136, 252. hdl:2246/830.
  33. 1 2 Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (August 2022). "Flufftails, finfoots, rails, trumpeters, cranes, limpkin". IOC World Bird List Version 12.2. International Ornithologists' Union. Archived from the original on 14 March 2023. Retrieved 14 November 2022.
  34. Mlíkovský (2002)
  35. OLSON, STORRS L. (1977). "A SYNOPSIS OF THE FOSSIL RALLIDAE" (PDF). Smithsonian Libraries - Smithsonian Research Online. Archived (PDF) from the original on 2018-07-29. Retrieved 2018-07-29.
  36. A small species of rail: Hugueney et al. (2003)
  37. Dozens of mostly broken isolated skull and limb bones of a rail or crake the size of a slaty-breasted or small buff-banded rail: Worthy et al. (2007)
  38. Quadrate (MNZ S.40957) and 2 femora (MNZ S.42658, S.42785) of a rail or crake the size of a large buff-banded rail: Worthy et al. (2007)
  39. Several limb bones of a smallish rail: Gál et al. (1998–99)
  40. Partial hand of a common moorhen-sized rail: Ballmann (1969)
  41. Olson, Storrs L. (1974). "A new species of Nesotrochis from Hispaniola, with notes on other fossil rails from the West Indies (Aves: Rallidae)". Proceedings of the Biological Society of Washington. 87(38): 439–450. hdl : 10088/8374 .
  42. Worthy, T.H. (1997). "A mid-Pleistocene rail from New Zealand". Alcheringa: An Australasian Journal of Palaeontology. 21 (1): 71–78. doi:10.1080/03115519708619186.
  43. "Pleistorallus flemingi; holotype". Collections Online. Museum of New Zealand Te Papa Tongarewa. Archived from the original on 24 October 2012. Retrieved 18 July 2010.
  44. Specimen QM F40203. A left carpometacarpus piece of a bird about the size of Lewin's rail. Probably from a rail, but it is too damaged to determine its affiliations more precisely: Boles (2005)
  45. Olson (1985), Mlíkovský (2002)

Further reading