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Zootaxa 4626 (1): 001–107 ISSN 1175-5326 (print edition)
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Copyright © 2019 Magnolia Press
Monograph ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4626.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:6CC12BAF-968F-4BDE-9315-340AF12A76EC
ZOOTAXA
4626
Systematics, morphology and ecology of rails (Aves: Rallidae)

of the Mascarene Islands, with one new species
JULIAN PENDER HUME

Correspondence Address: Bird Group, Department of Life Sciences, Natural History Museum, Akeman St, Tring, Herts HP23 6AP.
E-mail: j.hume@nhm.ac.uk
Magnolia Press
Auckland, New Zealand
Accepted by T. Worthy: 30 May 2019; published: 3 Jul. 2019

JULIAN PENDER HUME
Systematics, morphology and ecology of rails (Aves: Rallidae) of the Mascarene Islands,
with one new species
(Zootaxa 4626)
107 pp.; 30 cm.
3 Jul. 2019
ISBN 978-1-77670-699-0 (paperback)
ISBN 978-1-77670-700-3 (Online edition)
FIRST PUBLISHED IN 2019 BY

Magnolia Press
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ISSN 1175-5326 (Print edition)

ISSN 1175-5334 (Online edition)
2 · Zootaxa 4626 (1) © 2019 Magnolia Press HUME

Table of Contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Class Aves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Order Gruiformes Bonaparte, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Family Rallidae Rafinesque, 1815 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Genus Aphanapteryx Frauenfeld, 1868a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
†Mauritian Red Rail (Poule Rouge) Aphanapteryx bonasia (Sélys-Longchamps, 1848) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Genus Erythromachus Milne-Edwards, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
†Rodrigues Rail or Leguat’s Rail Erythromachus leguati Milne-Edwards, 1874. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Genus Hypotaenidia Reichenbach, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Buff-banded Rail Hypotaenidia philippensis (Linnaeus, 1766) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Genus Dryolimnas Sharpe, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
White-throated Rail Dryolimnas cuvieri (Pucheran, 1845) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
†Cheke’s Wood Rail Dryolimnas chekei sp. nov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
†Réunion Wood Rail Dryolimnas augusti Mourer-Chauviré et al. 1999 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Genus Porphyrio Brisson, 1760 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
†Réunion Gallinule (Oiseaux Bleu) ‘Porphyrio caerulescens’ (Sélys-Longchamps, 1848) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
African Swamphen Porphyrio porphyrio madagascariensis (Latham, 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Allen’s Gallinule Porphyrio alleni Thomson, 1842 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Genus Fulica Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
†Mascarene Coot Fulica newtonii Milne-Edwards, 1867b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Genus Gallinula Brisson, 1760 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Madagascan Moorhen Gallinula chloropus pyrrhorrhoa A. Newton, 1861b . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Géant or Leguat’s Giant Leguatia gigantea Schlegel, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Order Phoenicopteriformes Fürbringer, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Family Phoenicopteridae Bonaparte, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Genus Phoenicopterus Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Greater Flamingo Phoenicopterus roseus Pallas, 1811 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Sexual dimorphism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Extinction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Abstract
Five species in five genera of extinct endemic rails have been described from the Mascarene Islands of Mauritius, Réunion
and Rodrigues: the Mauritian Red Rail or Poule Rouge Aphanapteryx bonasia; Mascarene Coot or Poule d’eau Fulica
newtonii; which occurred on Mauritius and Réunion; Réunion Wood Rail Dryolimnas augusti; Réunion Gallinule or
Oiseaux bleu ‘Porphyrio caerulescens’; and Rodrigues or Leguat’s Rail Erythromachus leguati. All are known from fossil
remains and/or from contemporary accounts and illustrations. A sixth species of rail Dryolimnas sp. nov. is described
herein from fossils from Mauritius, but was not unequivocally previously reported in the contemporary literature. This
paper provides an analysis of the Rallidae of the Mascarene Islands based on existing and newly discovered fossil remains,
and details historical reports and accounts. Comprehensive osteological descriptions and synonymies are also included.
Their ecology and extinction chronologies are interpreted from historical evidence. The relationships of Aphanapteryx
and Erythromachus are unresolved, having clearly been isolated for a considerable time; the middle Miocene is the
earliest their ancestors could have arrived on the Mascarenes, but this may have happened more recently. Mascarene
derivatives of Fulica, Porphyrio and Dryolimnas are of much more recent origin, and appear to have originated in
Africa or Madagascar. All terrestrial rails on Mauritius, Réunion and Rodrigues, were probable victims of cat predation
following their historic introduction to the islands, whereas over-hunting by humans was probably the primary cause of
extinction of ‘Porphyrio caerulescens’ on Réunion. The only extant rail on the Mascarenes today, the Madagascar race
of Eurasian Moorhen Gallinula chloropus pyrrhorrhoa, is a recent arrival, having colonised Mauritius and Réunion after
the extinction of Fulica newtonii.
Key words: Mascarene rails, Rallidae, Aphanapteryx, Erythromachus, Dryolimnas, Porphyrio, Gallinula, Leguatia,
extinction, affinities, ecology, sexual dimorphism
RAILS (AVES: RALLIDAE) OF THE MASCARENE ISLANDS Zootaxa 4626 (1) © 2019 Magnolia Press · 3
Introduction
Rails crakes, coots and gallinules (Rallidae) comprise a diverse range of birds that have colonised every continent
except Antarctica, and numerous islands around the world (Olson 1973; Ripley 1977; Taylor 1996; Taylor & van
Perlo 1998); they occupy a variety of habitats, with a preference for marshland and dense forests. Terrestrial species
are generally cryptic and difficult to observe, and many island species have become flightless (Olson 1977; Ripley
1977; Taylor & van Perlo 1998; Livezey 2003; Steadman 2006). Island populations in particular are extremely vul-
nerable to anthropogenic changes and, as a result, Rallidae have suffered more extinctions than any other bird family
(Ripley 1977; Taylor & van Perlo 1998; Steadman 2006; Hume 2017).
Situated in the southwestern Indian Ocean (Fig. 1), the Mascarene Islands of Mauritius (20.25° S, 57.5° E) (Fig.
2), Réunion (21° S, 55.5° E) (Fig. 3) and Rodrigues (19.75° S, 63.5° E) (Fig. 4) were once inhabited by a number
of rallid genera, the majority of which were flightless (Table 1). They were considered tame and easy to catch when
first discovered by mariners and settlers, but due primarily to the introduction of predatory mammals, especially
cats, and over-hunting all species disappeared rapidly (Cheke & Hume 2008; Hume & Walters 2012; Hume 2013,
2017). Two endemic Mascarene genera, Aphanapteryx and Erythromachus, whose relationships remain obscure,
once occurred on Mauritius and Rodrigues respectively. They were large flightless rails with long, decurved bills
and hair-like plumage. They survived alongside the introduced Black Rat Rattus rattus (Linnaeus, 1758) for centu-
ries (Hume 2013), an invasive, predatory species which has proved so lethal to many other island birds (Hume &
Walters 2012; Hume 2017), but both rails were probable victims of introduced cats, which arrived later (Cheke &
Hume 2008; Hume 2017). Aphanapteryx was last recorded in 1693, whereas Erythromachus disappeared between
1726 and 1761 (Cheke & Hume 2008; Hume & Walters 2012; Hume 2017).
FIGURE 1. The Mascarene Islands of Réunion, Mauritius and Rodrigues with the distribution of Rallidae in the southwestern
Indian Ocean used in the analysis.

FIGURE 2. Detail of Mauritius and original vegetation, showing the place names mentioned in the text (adapted from Cheke
& Hume 2008).
The Indian Ocean endemic genus Dryolimnas (White-throated Rail) occurs widely on islands throughout the
south-western Indian Ocean (Fig. 1). Nominate D. c. cuvieri (Pucheran, 1845) occurs on Madagascar and is vagrant
to Mayotte, Comoros (Safford & Hawkins 2013), while flightless derivatives occur on Aldabra, D. c. aldabranus
(Günther, 1879), and formerly on Assumption, D. c. abbotti (Ridgway, 1894), where became extinct between 1908
and 1937 (Hume 2017). The holotype of D. cuvieri, a female, was collected on Mauritius in 1809 (Schlegel &
Pollen 1868), but appears to have been a vagrant (Hartlaub 1877); molecular work is long overdue to resolve this
long-standing conundrum. A large, flightless Dryolimnas derivative, D. augusti Mourer-Chauviré, Bour, Ribes &
Moutou, 1999 occurred on Réunion until at least the 1670s (Mourer-Chauviré et al. 1999). In addition, a poorly
volant or flightless Dryolimnas also once inhabited Mauritius, and probably also disappeared at the end of the 17th
century; it is described herein.
The Mascarene Coot Fulica newtonii Milne-Edwards, 1867b occurred on Mauritius and Réunion, and appears
to have moved between the islands (Mourer-Chauviré et al. 1999; Cheke & Hume 2008). It was last recorded on
Réunion in 1671-72 and on Mauritius in 1693 (Cheke & Hume 2008; Hume 2017), and seemingly disappeared due
to cat predation, but over-hunting by humans on Réunion may have also been a factor. Much has been written (e.g.
Milne-Edwards 1874; Rothschild 1907a; Lambrecht 1933; Hachisuka 1953) about the enigmatic Réunion Gallinule
or poule bleu, ‘Porphyrio caerulescens’ (Sélys-Longchamps, 1848), a large blue, probable Porphyrio gallinule,
which occurred on the montane region of the Plaine des Cafres on Réunion (Fig. 3), and at altitude elsewhere on the
island (Feuilley 1705). It was reported independently by a number of competent observers; however, no physical
evidence has yet been found to confirm its relationships (Mourer-Chauviré et al. 1999). Its extinction was due to over-
hunting and perhaps cat predation, being last recorded in 1730 but possibly persisting until the 1760s (Grant 1801).
FIGURE 3. Detail of Réunion and original vegetation, showing the place names mentioned in the text (adapted from Cheke &
Hume 2008).

The only surviving rallid on the Mascarenes today is the Madagascan subspecies of Eurasian Moorhen Gal-
linula chloropus pyrrhorrhoa A. Newton, 1861a, which occurs on Mauritius and Réunion; it appears to be a recent
arrival, having probably colonised remaining wetlands made vacant by the extinction of the Mascarene Coot (Cheke
& Hume 2008; Hume 2013).
This paper synthesizes available sources of data on Mascarene Rallidae, including historical observations and
accounts, and renewed osteological analyses of museum specimens and palaeontological material. Comparisons
presented here, using morphometric data based on osteology, show that an endemic species of Dryolimnas occurred
on Mauritius, alongside the above-mentioned Mascarene Rallidae genera. As these rails are almost exclusively
known from palaeontological or historical records (Table 1), the texts of relevant historical accounts are included,
particularly those for which no previous English translations exist.
TABLE 1. Nomenclatural status of real and supposed endemic Mascarene rails.

Species Distribution Types of evidence
†Aphanapteryx bonasia Mauritius Fossil record, contemporary illustrations and accounts
†Erythromachus leguati Rodrigues Fossil record and contemporary accounts
†Dryolimnas chekei nov. sp. Mauritius Fossil record
†Dryolimnas augusti Réunion Fossil record and contemporary account
†’Porphyrio caerulescens’ Réunion Accounts only
†Fulica newtonii Mauritius and Réunion Fossil record and contemporary accounts
Didus herbertii Mauritius Based on misinterpretation of account. Synonymised under
Aphanapteryx bonasia
Didus broeckei Mauritius Based on misinterpretation of account. Synonymised under
Pezocrex herberti Mauritius Based on misinterpretation of account. Synonymised under
Kuina mundyi Mauritius Based on misinterpretation of account. Synonymised under
Leguatia gigantea Mauritius and Rodrigues Based on misinterpretation of account. Synonymised under
Phoenicopterus roseus
Materials and methods
Abbreviations: Institutions: AMNH, American Museum of Natural History, New York; BMB, Booth Museum of
Natural History, Brighton, England; CMNZ, Canterbury Museum, Christchurch, New Zealand; DRP, Dodo Re-
search Project; FLMR, Francois Leguat Museum, Rodrigues; LWM, Liverpool World Museum; NHMUK, Natural
History Museum, London and Tring, (formerly BMNH, British Museum (Natural History)); MNHN, Muséum
National d’Histoire Naturelle, Paris, with the prefix MAD for Madagascar, e.g. MNHN MAD7142; MHNR (pre-
fix MHN-RUN-CT) Muséum d’Histoire Naturelle, Réunion; MI, Mauritius Institute; OUM, Oxford University
Museum, Oxford, England; UMZC, University Museum of Zoology, Cambridge, England; USNM, United States
National Museum of Natural History, Washington, D.C.
Listing of skeletal material: ‘u/r’ unregistered material, u/s = unsexed, left (L) or right (R) prefixed by ‘p’ proxi-
mal, ‘s’ shaft or ‘d’ distal, juv. = juvenile. † indicates an extinct taxon.
Comparative material examined: See Appendix 1.
Measurements: All measurements were made using dial callipers and rounded to the nearest 0.1mm. TL =
total length: cranium, from naso-frontal hinge to margin of crista occipitalis in a median plane; rostrum, from
naso-frontal hinge to tip of rostrum maxillare in dorsal aspect; mandible, from processus retroarticularis to rostrum
mandibulae in dorsal aspect; coracoid, measured on the medial side; scapula, from tip of acromion to extremitas
caudalis in lateral plane; sternum, from anterior tip of spina externa to margo caudalis in a median plane; humerus,
radius and ulna, measured in dorsal aspect, carpometacarpus, measured in ventral aspect; pelvis, from anterior edge
of ala preacetabularis to posterior edge of ala postacetabularis ilii; femur, measured in cranial aspect; tibiotarsus,
measured in cranial aspect from articulating surfaces and also from crista cnemialis cranialis; tarsometatarsus,
measured in dorsal aspect; WNF = width of naso-frontal hinge; LN = length of narial opening, taken in lateral
aspect; RML = rostrum maxillare length, taken in a dorsoventral plane at the anterior point of the nasal opening
to tip; mandible, from posterior edge of rostrum mandibulae to tip; RMW = rostrum maxillare width, taken in a
lateromedial plane at the anterior point of the nasal opening; mandible, taken between posterior edge of rostrum
mandibulae; RMD = rostrum maxillare depth, taken in a dorsoventral plane at the anterior point of the nasal open-
ing; mandible, taken in dorsoventral plane at posterior edge of rostrum mandibulae; RMaL = rostrum mandibulae
length, taken in a dorsoventral plane; RMaW = rostrum mandibulae width, taken in a lateromedial plane; RMaD =
rostrum mandibulae depth, taken in a dorsoventral plane; LC = greatest length of carina sterni, from apex carinae to
margo caudalis of sternum; DK = greatest depth of keel, taken from facies muscularis sterni to apex carinae; LW =
least width; sternum, taken between 1st processus costales; pelvis, measured between ala preacetabularis ilii anterior
to sulcus antitrochantericus; PW = proximal width, taken in a lateromedial plane; mandible, taken from processus
mandibulae medialis to cotyla medialis; femur, from the caput femoralis through mid-depth point of the neck to
lateral side when viewed in cranial/caudal aspect; GW = greatest width, taken in a dorsoventral plane; cranium,
taken between processus postorbitales; sternum, taken between processus craniolaterales; pelvis, taken between ala
postacetabularis ilii; GD = greatest depth; cranium, from medial point of parietal to medial ventral surface; pelvis,
taken between ventral surface of processus terminalis ischii and posterior dorsal surface; PD = proximal depth,
taken in a dorsoventral plane; carpometacarpus, proximal end from processus extensorius to caudal margin of
trochlea carpalis; pelvis, taken between crista iliaca dorsalis and extremitas cranialis synsacri; SW = shaft width,
taken in lateromedial plane; SD = shaft depth, in dorsoventral plane; DW = distal width, in lateromedial plane;
coracoid, taken from angulus medialis to processus lateralis using the articular facet as the baseline; ulna, taken in
dorsal aspect; DD = distal depth, in dorsoventral plane.
FIGURE 4. Detail of Rodrigues, showing the place names mentioned in the text (adapted from Cheke & Hume 2008).

Anatomical nomenclature follows Baumel & Witmer (1993), with general abbreviations: m = musculus; n =
nervus; Elzanowski & Stidham (2010) for quadrate; and Mayr (2016) for hypotarsus. Nomenclature fol-
lows Gill et al. (2010) for New Zealand taxa, Dickinson & Remsen (2013), wherein taxon authors and dates can
be found for extant or historically extant taxa, and with updates from del Hoyo & Collar (2014) and Stervander et
al. (2019). The endemic rails of Madagascar, Madagascar Flufftail Sarothrura insularis (Sharpe, 1870); Slender-
billed Flufftail S. watersi Bartlett, 1880; Madagascar Rail Rallus madagascariensis Verreaux, 1933; Tsingy Wood-
rail Mentocrex beankaensis Goodman, Raherilalao & Block, 2011; Madagascar Wood-rail M. kioloides (Pucheran,
1845); Sakalava Rail Zapornia olivieri (G. Grandidier & Berlioz, 1929) and Western Baillon’s Crake Zapornia
pusilla intermedia (Hermann, 1804) are not part of the Mascarene Rallidae colonisation events and so are excluded
from this monograph.
Etymology: Etymologies of Mascarene rallids are given in their entirety, either from Olson (1977), Jobling
(1991, 2009) or from lexicographical sources for fossil species.
Systematics
Class Aves
Order Gruiformes Bonaparte, 1854
Family Rallidae Rafinesque, 1815
Genus Aphanapteryx Frauenfeld, 1868a
Aphanapteryx Frauenfeld, 1868a, p.6. Type (by monotypy): Aphanapteryx imperialis Frauenfeld = Apterornis bonasia Sélys-
Longchamps.
Etymology: From Greek aphançs = unseen, invisible-wing, privative prefix a, and pteryx = wing.
Revised Diagnosis: Aphanapteryx bonasia (Fig. 5) is distinguished from other Mascarene rallids by the follow-
ing characters:
Cranium (Fig. 6): largest cranium in Mascarene Rallidae; in lateral view, cranium dorso-ventrally compressed;
small fonticulus orbitocranialis; large foramen n. optici; processus postorbitalis short, bluntly triangular; deep fossa
temporalis; os lacrimale wide rostrocaudally and straight extending past os ectethmoidale posteriorly; no foramen
in os ectethmoidale; crista nuchalis transversa distinct; crista temporalis indistinct; in dorsal view, frontal narrow,
triangular and deeply excavated; in posterior view, crista temporalis extends almost to top of cranium; processus zy-
gomaticus distinct; condylus occipitalis large; foramen magnum large and an inverted heart-shape; in ventral view,
tuba audivita communis deep; fossa subcondylaris deeply excavated.
Rostrum (Fig. 6): os premaxillare long and narrow and terminating in a sharp point, and total length nearly 47%
longer than cranium length; in dorsal view, os nasale wide, covering narial openings; in lateral view, narial opening
reaching 50% the length of the rostrum; foramina neurovascularia, prominent, elongate, running almost to rostral
edge of narial opening; os nasale narrow; in ventral view, shallow, indistinct sulcus.
Mandible (Fig. 6): rostrum mandibulae long, length of symphysis approximately 79% of cranium length; in
dorsal view, processus mandibulae medialis distinct, triangular; processus retroarticularis long and directed laterad;
sulcus on crista intercotylaris deeply excavated; cotyla lateralis distinctly U-shaped; os dentale long, narrow and
terminating in a pointed tip; in lateral view, foramina neurovascularia, prominent, deeply set and running almost to
a deep sulcus; large fenestra rostralis mandibulae and very small fenestra caudalis mandibulae.
Quadrate: in lateral view, processus orbitalis long, deep and rounded; processus oticus short; processus man-
dibularis rounded, and strongly directed laterad; fovea quadratojugalaris oval or rounded and deeply excavated; in
medial view, condylus pterygoideus deeply incised; in medial aspect, a single foramen pneumaticum rostromediale
present and situated anterior to the medial ridge (see Elzanowski & Stidham 2010), which occurs in the same posi-
tion and with no variation other than size in all Rallidae examined in this paper.
Vertebrae: configuration of vertebrae of Thirioux Aphanapteryx associated skeleton (see details below under
Referred fossil material); atlas (1); axis (2); presynsacral (cervical) (3–15); presynsacral (cervicodorsal) (16); notar-
ium (presynsacral thoracic vertebrae 17–19); presynsacral (thoracic) (20), plus free caudal (5); pygostyle (1) = (20
+ 5 + 1); in Gallinula (20 + 7 + 1); in Fulica (22 + 5 + 1); in Dryolimnas (22 + 6 + 1) and Porphyrio (20 + 7 + 1).
Scapula (Fig. 7): in lateral view, wide; margo dorsalis distinct; facies articularis humeralis shallow; acromion
blunt, rounded and directed dorsad; tuberculum coracoideum indistinct.
Coracoid (Fig. 8): comparatively short; shaft wide; in dorsal aspect, cotyla scapularis shallow and indistinct;
large oval foramen n. supracoracoidei distal to processus procoracoideus; processus procoracoideus short and weak-
ly directed dorsomediad; impressio m. sternocoracoidei deeply excavated.
FIGURE 5. An almost complete, associated skeleton of Aphanapteryx bonasia (MI 923) collected by Etienne Thirioux in
September 1899 in the Vallée des Prêtres, Moka Range, mid-west Mauritius. Scale bar = 200 mm.
Sternum (Fig. 9): carina sterni extremely reduced more than in any other Mascarene rallid (See appendix 2,
Table 6); in dorsal view, processus craniolateralis wide, directed slightly cranially; tuberculum labri externi pro-
nounced; foramen pneumaticum deeply excavated; sulcus medianus sterni not clearly defined; in lateral view, apex
carinae directed slightly ventrad; in lateral view, carina sterni extremely reduced.

FIGURE 6. Comparison of crania of Mascarene rails, lateral view. A. Aphanapteryx bonasia MU923 u/s; B. Erythromachus
leguati NHMUK u/r, u/s. Scale bar = 10mm.
Humerus (Fig. 10): extremely reduced; shaft strongly curved dorsoventrally; in caudal aspect, tuberculum ven-
trale pronounced, incisura capitis deeply excavated; fossa pneumotricipitalis extremely shallow; tuberculum dorsale
reduced; sulcus humerotricipitalis broad and deep; in cranial view, sulcus transversus shallow; crista bicipitalis,
short, weakly square-shaped; crista deltopectoralis long, extending much further distad than crista bicipitalis; fossa
m. brachialis deeply excavated; indistinct processus supracondylaris dorsalis.
Radius (Fig. 11): sulcus tendinosa deep extending proximad approximately 25% of total length radius length;
facies articularis radiocarpalis deflected ventrally; in ventral aspect, depressio ligamentosa deeply excavated.
Ulna (Fig. 12): short and strongly arched dorsoventrally proximally; in caudal aspect, olecranon indistinct;
processus cotylaris dorsalis short; in ventral aspect, impressio brachialis deep.
Carpometacarpus (Fig. 13): in dorsal aspect, processus extensorius short and rounded; os metacarpale majus
robust; os metacarpale minus weakly arched creating narrow spatium intermetacarpale.
Phalanges digiti alulae I (Fig. 13): short, equal in length to os metacarpale alulare.
Phalanges digiti proximalis majoris (Fig. 13): short; pina cranialis indistinct.
Pelvis (Fig. 14): extremely wide and robust; in dorsal view, sulcus antitrochantericus extend laterally beyond
the ala preacetabularis; distinct, caudally-open canalis iliosynsacralis either side of the crista iliaca dorsalis; distinct
foramina intertransversariae; ala postacetabularis ilii wide on lateral edge covering ala ischii; in lateral view, fora-
men acetabuli larger in size than foramen ilioischiadicum; crista iliaca dorsalis arched; in ventral view, two pro-
nounced processus costales; ala ischii directed ventrad.
Femur (Fig. 15): extremely robust; in cranial view, shaft curved medially and directed caudally at the proximal
end; crista trochanteris strongly deflected mediad; linea intermuscularis indistinct; sulcus patellaris deeply exca-
vated; epicondylus lateralis distinct; condylus medialis extends the same distance distally as condylus lateralis; in
medial aspect, crista trochanteris extends caudad, offsetting centrally facies articularis acetabularis; shaft strongly
arched dorsally; in caudal aspect, impressiones obturatoriae distinct and directed cranially; linea intermuscularis
caudalis distinct, forming a ridge; fossa poplitea deeply excavated; tuberculum m. gastrocnemialis lateralis rounded
and separated from trochlea fibularis; condylus medialis smaller than condylus lateralis; trochlea fibularis deeply
excavated.
FIGURE 7. Comparison of scapulae (medio-ventral aspect, left side; except B which is lateral aspect, right side) of Mascarene
rails with Dryolimnas, A. UMZC 579 Aphanapteryx bonasia u/s, right proximal UMZC 579 u/s; B. NHMUK u/r Erythromachus
leguati u/s; C. NHMUK 1882.2.28.2 Dryolimnas cuvieri cuvieri u/s; D. NHMUK S/1989.38.1 D. c. aldabranus ♂; E. NHMUK
1910.4.8.1 D. c. abbotti u/s. Scale bar = 10 mm.
Tibiotarsus (Fig. 15, 16a): large and robust, especially at proximal and distal ends; in cranial view, sulcus ex-
tensorius deeply excavated, extending proximad; canalis extensorius oval-shaped; pons supratendineus wide and
slightly arched, and situated mediad; crista fibularis extends short; crista cnemialis lateralis long; in caudal aspect,
crista cnemialis cranialis weakly pronounced; shaft deep; condylus lateralis small, weakly rounded; depressio epi-
condylaris lateralis shallow.
Fibula: short and robust; in dorsal view, caput fibulae narrow and weakly grooved caudally; tuberculum m.
iliofibularis pronounced and excavated cranially.
Tarsometatarsus (Fig. 15, 16a, b): large and robust; in dorsal aspect, trochleae metatarsorum distinctly splayed,
especially trochlea. metatarsi II; metatarsi II directed slightly dorsad; retinaculum extensorium tarsometatarsi indis-
tinct; in plantar aspect, foramen vasculare distale deeply excavated; fossa metatarsi I extends proximad; hypotarsi
deflected laterad; on proximal end, a monocanaliculate hypotarsus (see Mayr 2016) with crista medialis flexoris
digitorum longus; sulcus for tendon of musculus flexor perforatus digiti II and sulcus for tendon of musculus flexor
hallucis longus present.
Remarks: Aphanapteryx was a large flightless rail, approximately 35–40 cm in total length, with extremely
robust legs and reduced wings and tail. The bill was long, decurved and deep proximally, the plumage was hair-like
and dull brownish-red in colouration, with the feathers of the nape elongate; Livezey (2003) estimated a body mass
of 1.3 kg in the male and around 1 kg for the female. Size variation between skeletal specimens (Fig. 16a) is almost
certainly due to marked sexual dimorphism, with the male larger than the female, a common phenomenon in Ral-
lidae (Olson 1977; Livezey 2003) (see Appendix 2). The bill was reported to be variable in curvature (e.g. Greenway
1958, 1967), but I have examined all known maxillae and found this not to be the case; however, Aphanapteryx
exhibited extreme variation in the total length of the rostrum (see Appendix 2, Table 2).

FIGURE 8. Comparison of coracoids (ventral aspect, right side) of Mascarene rails with Dryolimnas, A. FLMR 1081
Aphanapteryx bonasia u/s; B. NHMUK u/r Erythromachus leguati u/s; C. NHMUK 1882.2.28.2 Dryolimnas cuvieri cuvieri
u/s; D. NHMUK S/1989.38.1 D. c. aldabranus ♂; E. NHMUK 1910.4.8.1 D. c. abbotti u/s. Scale bar = 10 mm.
FIGURE 9. Comparison of sterna, lateral aspect (left), ventral aspect (right) of Mascarene rails, A. C. UMZC 303BB
Aphanapteryx bonasia u/s; B. D. NHMUK u/r Erythromachus leguati u/s. Scale bar = 10 mm.
†Mauritian Red Rail (Poule Rouge) Aphanapteryx bonasia (Sélys-Longchamps, 1848)
Indise riviers houdt snippen [Indian river woodcocks] Jolinck, [in 1598] (in Keuning 1947, p.62); Payandee 2002, p.39
Velt-hoenders Cornelisz, [in 1602] (in Strickland & Melville 1848, p.13)
Feldhüner Verhuffen, [in 1611] (in Strickland & Melville 1848 p.18); Barnwell 1948, p.20
[illustration] van den Broecke, [in 1617] 1646, p.102, pl.7
Hen Herbert, [in 1627] 1634, pp.212 & 214, 1638, pp.347 & 349, 1658, p.183 (Dutch edition), 1663, p.546 (French edition);
1665, p.405, 1677, p.385; Strickland & Melville 1848, p.21
Poule rouge [Red hen] Cauche, [in 1638] 1651, p.132; Oustalet, 1874a, p.12, 1874b, p.61, 1874c, p.12; Renshaw, 1936, p.60
Mauritius henne Mundy, [in 1638] (in Temple 1919 pp.352–353); Barnwell, 1948, p.29
Perdrix [=Feldhüner] Gooyer, in [1638–39] (in Bonaparte 1890, p.10)
Dodaersen Pretorius, [in 1666–69] (in Hume & Winters 2016, p.5)
Dodo or Red hen Marshall, [in 1668] (in Khan 1927, p.47–48)
Toddärsche Hoffman, [in 1674] 1680, p.52
Water-quail Leguat, [in 1693] 1708, p.71 [English edition]
Gelinotes Leguat, [in 1693] 1708, p.71 [French edition]
Apterornis bonasia Sélys-Longchamps, 1848, p.292 (based on the accounts of Cornelisz, Verhuffen, Herbert, Cauche and
Leguat); Bonaparte, 1854, p.3; Milne-Edwards, 1866–73 [1874], p.79
Didus herbertii Schlegel, 1854a, p.256; 1854b, p.346 (based on Herbert’s ‘Hen’); Milne-Edwards, 1868, p.344; Salvadori, 1893,
p.629
Didus broeckei Schlegel, 1854a, p.256 (based on van den Broecke’s drawing); Frauenfeld, 1868a, p.16, fig.26; Milne-Edwards,
1868, p.344
Didus brouckei Schlegel, 1854b, p.345 (lapsus)
Aphanapteryx imperialis Frauenfeld, 1868a, p.6, tab.2 (based on a vellum painting attributed to Jacob Hoefnagel c.1610 in the
library founded by Emperor Francis I of Austria (Anker 1974; Jackson 1999); Frauenfeld 1868b, p.140, 1869, p.761–764;
Milne-Edwards, 1868, p.344, 1869a, p.275
Didus herberti Frauenfeld, 1868a, p.16, fig.25; Milne-Edwards, 1868, p.344, 1869a, p.275; Rowley, 1877, p.130; Cowles, 1987,
p.96; Hume & Walters, 2012, p.374; Hume, 2017, p.420 (lapsus)
Aphanapteryx imperalis Frauenfeld, 1868b, p.140 (lapsus or emendation).
Aphanapteryx broeckei: Milne-Edwards, 1868, p.345; 1866–1873 [1874], p.81; Forbes, 1892a, p.252; E. Newton & Gadow,
1893, pl.X (in plate), 1895, p.235; Milne-Edwards, 1896a, p.76; Oustalet, 1897, p.95; Grandidier & Grandidier, 1905,
pp.325,375; Carié, 1916, p.107; D’Emmerez de Charmoy, 1928, p.108; Piveteau, 1945, p.7; Verin, 1983, p.27
Aphanapteryx broeckii: Milne-Edwards, 1869a, p.275, 1896a, p.81; Anon, 1874, p.455; Pelzeln, 1876, p.45; Rowley, 1877,
p.129; E. Newton, 1888, p.552; Slater, 1881, p.27; Salvadori, 1893, p.629; Sharpe, 1894, p.68; Emerez de Charmoy, 1903,
p.7; Pitot, 1914, p.88; Sclater, 1915, p.318; Lambrecht, 1933, p.468; Hachisuka, 1937, p.156; Vinson, 1949, p.74; Guérin,
1950, p.404 (lapsus)
Didus broeckii: Milne-Edwards, 1869a, p.275; Frauenfeld, 1869, p.762; Salvadori, 1893, p.629 (lapsus)
Aphanapterix imperialis Frauenfeld, 1869, p.762–763 (lapsus)
Aphanapteryx bonasia: Milne-Edwards, 1869–1871, pp.614–618, pl.109–110, figs.1–15; Noll, 1889, pp.124–126; E. Newton &
Gadow, 1895, pp.293–294, pl.35, figs.12–20; Rothschild, 1907a, p.131, pl.29; Sclater, 1915, pp.316–319, fig.1, 1924, p.102;
Killermann, 1915, pp.376–377, fig.14; Lambrecht, 1933, pp.468–469; Greenway, 1958 p.117, 1967 p.117; Brodkorb, 1967,
p.133; Luther, 1970, p.187, 1986, p.183, 1986 [2005], p.183; Olson, 1977, p.357; Fuller, 1987, p.87, 2001, p.145, 2002,
p.24; Cheke, 1987, p.36; Cowles, 1987, p.96; Balouet, 1990, p.45; Day, 1991, p.89; Staub, 1993, p.21; van Hoof, 1995,
p.51; Ziswiler, 1996, p.40; Taylor & van Perlo, 1998 p.59; Mourer-Chauviré et al., 1998 p.98; Mourer-Chauviré et al., 1999
p.23; Adams et al., 2003, p.344; Moree 1998, p.9, 2001, p.50, pl.64; Livezey, 1998, p.2083, 2003, p.1; Hume et al., 2004,
(10 June); Hume & Prŷs-Jones, 2005, p.86; Cheke, 2006 p.155, 2007, p.14, 2013a, p.2, 2013b, p.13, 2016, p.5; Cheke &
Hume, 2008 pp.25, 127, pl.4; Hume, 2009 p.229, 2013, p.218, 2014a, pp.36–37, 2017, p.126; Rijsdijk et al., 2009, p.17,
tbl.2; Tyrberg, 2009, p.71; Hume & Walters, 2012, p.108; Parish, 2013, p.99; Safford & Hawkins, 2013, pp.45,348,pl.48;
del Hoyo & Collar, 2014, pp.342,772; Hume et al. 2015, p.5; Hume & Winters, 2016, p.5; Claessens & Hume, 2016, p.23;
Winters et al., 2017, p.134
Pezophaps broeckei: Schlegel, 1873, p.4
Pezophaps herbertii: Schlegel, 1873, p.4 Salvadori, 1893, p.629, footnote (Salvadori incorrectly places P. herbertii in the
synonymy of Miserythrus [Erythromachus] leguati)
Pezophaps broeckii: Anon, 1874, p.455; Lambrecht, 1933, p.468
Aphanapteryx herberti Pelzeln, 1876, p.45
Aphanapteryx broecki: Günther & E. Newton, 1879, p.282; Forbes, 1893b, p.66; A. Newton, 1905, p.142; Rothschild, 1907b,
p.198; Renshaw, 1933, p.17; Staub, 1976, p.14 (lapsus)
Poulet Rouge Renshaw, 1933, p. 16, 1936, p.60
Kuina mundyi Hachisuka, 1937, p.156
Pezocrex herberti: Hachisuka, 1953, p.148
Kuinia mundyi Hume & Walters, 2012, p.373; Hume, 2017, p.418 (lapsus)
Holotype: None designated. Baron de Sélys-Longchamps (1848) based formal scientific names on three of the
birds mentioned by Strickland (in Strickland & Melville 1848), all known from historical accounts only (see Olson
1977). He described them as Apterornis solitarius, the dodo-like bird of Réunion; A. caerulescens, the above-

mentioned poule bleu ‘Porphyrio caerulescens’, also of Réunion; and A. bonasia, based on the accounts of the
velt-hoenders of Cornelisz; feldhüner of Verhuffen; hen of Herbert; poule rouge of Cauche and gelinotte of Leguat.
Olson (1977) noted that Apterornis solitarius was said by Milne-Edwards (1868, 1869a) to have been designated as
the type of Apterornis by Sélys-Longchamps (1848), but his and my examination of Sélys-Longchamps paper can
find no internal evidence of this. Regardless, Apterornis Sélys-Longchamps (19 November 1848) is preoccupied by
Apterornis Owen (13 April 1848), now a junior synonym of Aptornis G. A. Mantell (2 February 1848), an extinct
genus of unrelated birds (adzebills) of New Zealand (Gill et al. 2010).
FIGURE 10. Comparison of humeri (caudal aspect, left side) of Mascarene rails, A. UMZC 579 Aphanapteryx bonasia u/s; B.
NHMUK u/r Erythromachus leguati u/s. Scale bar = 10 mm.
FIGURE 11. Comparison of radii (ventral aspect, right side) of Mascarene rails with Dryolimnas, A. UMZC 579.1 Aphanapteryx
bonasia u/s; B. NHMUK u/r Erythromachus leguati u/s; C. NHMUK 1882.2.28.2 Dryolimnas cuvieri cuvieri u/s; D. NHMUK
S/1989.38.1 D. c. aldabranus ♂; E. NHMUK 1910.4.8.1 D. c. abbotti u/s. Scale bar = 10 mm.
Measurements: See Appendix 2.

Type locality: Mare aux Songes, Mauritius, Mascarenes.
Distribution: Mauritius, Mascarenes.
Etymology: Olson (1977: 357) provided a detailed description of the complicated etymology of Aphanapteryx,
and it is abridged above. The specific bonasia refers to the Eurasian Hazel Hen Tetrastes bonasia, as in early ac-
counts Aphanapteryx was referred to by vernacular names such as Gelinote or Gélinotte, which was elsewhere ap-
plied to the grouse. The derivation of bonasia itself is unclear, with the most frequent explanation being a derivative
from Greek bonasos or Latin bonasus (a bull), referring to the drumming of a male Ruffed Grouse Bonasa umbellus,
which sounds like the bellowing of a bull. Alternative explanations are: Latin bonum (good) and assum (a roast), in
reference to its culinary qualities; Greek bonasos, alluding to a mythical, timid bull; a reference to the tameness of
certain grouse, with Bonasa and bonasia being Latinised forms of French bonasse (simple-minded or good-natured).
Another possibility is that the etymology referring to a bull is alluding to the rails’ (and bulls’) similar attraction to
the waving of red cloth, as described for Aphanapteryx by a number of contemporary observers on Mauritius.
Referred fossil material: Unassociated subfossil material collected from the Mare aux Songes marsh, unless
otherwise stated. Many specimens are bulk numbered, so duplicate numbers exist for some of the skeletal elements:
Cranium, CMNZ AV6284 (Vallée des Prêtres); Rostrum, CMNZ AV6284 (Vallée des Prêtres); MNHN MAD u/r;
MNHN MAD6572; UMZC 303BB; UMZC 303BB: UMZC 303BB; Mandible, CMNZ AV6284; MNHN MAD
u/r; MNHN MAD u/r; UMZC 303BB; UMZC 303BB; UMZC 303BB; UMZC 303.B; Vertebrae CMNZ AV6284
(x25) (Vallée des Prêtres + Mare aux Songes); Scapula, UMZC579 (L); UMZC 579 (Lp); UMZC 579 (Rp); UMZC
579 (Rp); Coracoid, CMNZ AV6284 (R) (Vallée des Prêtres); MNHN MAD u/r (L); MNHN MAD u/r (R); MNHN
MAD u/r (L); MNHN MAD u/r (R); MNHN MAD u/r (R); MU 504(L); FLMR 1081 (R) (La Prairie); Sternum,
CMNZ AV6284 (Vallée des Prêtres); UMZC 303BB; Humerus, CMNZ AV6284 (R) (Vallée des Prêtres); CMNZ
AV6284 (L) (Vallée des Prêtres); MNHN MAD6579 (Rd); MNHN MAD6580 (R); MNHN MAD6578 (R); MNHN

MAD7212 (R); MNHN MAD7217 (L); MNHN MAD7210 (R); MNHN MAD7216 (Rd): MNHN MAD7216 (Rd);
MNHN MAD7218 (Lp); MNHN MAD7215 (L); MNHN MAD8856 (Ld); MNHN MAD7211 (L); UMZC 303BB
(L); UMZC 579 (Rd); UMZC 579 (L); UMZC 579 (Rp); UMZC 579 (L); UMZC 579 (Ld): UMZC 579 (Rd);
UMZC 579 (Ld); FLMR 609 (Rp) (S. Grand Baie); FLMR 610 (Ls) (S. Grand Baie); Radius, CMNZ AV6284 (R)
(Vallée des Prêtres); CMNZ AV6284 (L) (Vallée des Prêtres); UMZC 579.1 (L); Ulna, CMNZ AV6284 (R) (Vallée
des Prêtres); CMNZ AV6284 (Lp) (Vallée des Prêtres); FLMR 607 (Lp) (S. Grand Baie); MNHN MAD6967 (L);
MNHN MAD7190 (L); MNHN MAD7198 (R); UMZC 603 (R); Carpometacarpus, CMNZ AV6284 (Rp) (Vallée des
Prêtres); CMNZ AV6284 (L) (Vallée des Prêtres); Pelvis, CMNZ AV6284 (Vallée des Prêtres); MNHN MAD6841
(Vallée des Prêtres); UMZC 303BB; FLMR 608 (S. Grand Baie); Femur, CMNZ AV6284 (R) (Vallée des Prêtres);
CMNZ AV6284 (L) (Vallée des Prêtres); MNHN MAD6577 (R); MNHN MAD6576 (L); MNHN MAD7165 (R);
MNHN MAD7166 (Ld); MNHN MAD 7164 (Lp); MNHN MAD6818 (R) juv?; UMZC 303BB (R); UMZC 579
(L); UMZC 579 (Rd); UMZC 579 (Ls); FLMR 611 (Rp) (S. Grand Baie); FLMR 612 (Lp) (S. Grand Baie); Tibio-
tarsus, BMB 101750 (Ld); CMNZ AV6284 (R) (Vallée des Prêtres); CMNZ AV6284 (L) (Vallée des Prêtres); DRP
258.B1 (Rp); FLMR 613 (R) (S. Grand Baie); MNHN MAD6502 (L); MNHN MAD6561 (L); MNHN MAD6501
(R); MNHN MAD u/r (Rs); MNHN MAD u/r (Ld); NHMUK u/r (R); NHMUK u/r (L); NHMUK u/r (R); NHMUK
u/r (L); NHMUK u/r (L); NHMUK u/r (R); NHMUK u/r (R); NHMUK u/r (L); NHMUK u/r (Rp); NHMUK u/r
(R); NHMUK u/r (R); NHMUK u/r (R); NHMUK u/r (R); NHMUK u/r (R); NHMUK u/r (R); NHMUK u/r (L);
NHMUK u/r (Ld); NHMUK u/r (Ld); NHMUK u/r (Ld); NHMUK u/r (L); NHMUK u/r (Ld); NHMUK u/r (Ld);
NHMUK u/r (Ld); NHMUK u/r (Rs); NHMUK u/r (Rp); NHMUK u/r (Lp) (La Prairie); NHMUK u/r (Ld) (La
Prairie); NHMUK u/r (R); UMZC 579 (Rp); UMZC 579 (Lp); UMZC 579 (Rd); UMZC 579 (Rd); UMZC 579 (Rs);
UMZC 1003 (L); UMZC 1003 (L); UMZC 1003 (R); UMZC 1003 (R); UMZC 1003 (R); UMZC 1003 (R); UMZC
1003 (L); UMZC 1003 (R); UMZC 1003 (L); UMZC 1003 (R); UMZC 1003 (L); UMZC 303BB (Rd); Fibula,
CMNZ AV6284 (Rp) (Vallée des Prêtres); CMNZ AV6284 (Lp) (Vallée des Prêtres); Tarsometatarsus, CMNZ
AV6284 (R) (Vallée des Prêtres); CMNZ AV6284 (L) (Vallée des Prêtres); BMNH 101750 (Ld); DRP 237.B1 (R);
DRP 294.B2 (Rp); MNHN MAD6566 (R); MNHN MAD6565 (L); NHMUK u/r (L); NHMUK u/r (L); NHMUK
u/r (Lp); NHMUK u/r (R); NHMUK u/r (R); UMZC 579 (Lp); UMZC 579 (Rd); UMZC 579 (Ld); UMZC 303BB
(L); UMZC 303BB (L); UMZC 303BB (L); UMZC 303BB (R); UMZC 303BB (L); UMZC 303BB (L); UMZC
303BB (R); UMZC 303BB (L); UMZC 303BB (R); UMZC 303BB (R); UMZC 303BB (R); UMZC 303BB (R);
UMZC 303BB (R) (juv); Phalanges, CMNZ AV6284 (x6) (Vallée des Prêtres (4) + Mare aux Songes (2)); UMZC
580 (x50) (Vallée des Prêtres).
A unique, almost complete associated skeleton MI 923 (Fig. 5), collected by Etienne Thirioux in 1899 in the
Vallée des Prêtres, Moka Range, mid-west Mauritius, provides hitherto undescribed elements (see below) that are
not listed separately here. Thirioux was active from at least 1892–1908 and deposited some of his best material at
the Mauritius Institute, including the above-mentioned Aphanapteryx material, the most complete, associated dodo
known, and a unique associated Mauritian Giant Skink Leiolopisma mauritiana (Günther, 1877) skeleton (Claes-
sens & Hume 2016). Alfred Newton (1904: 626), and repeated later (A. Newton 1905: 142), provided details about
Thirioux’s discovery and provenance of Aphanapteryx:
…It was therefore with great interest that, just five years ago, October, 1899, I received a letter from M. E. Thirioux
informing me of his having found, in the preceding month of August, some remains of at least two dodos in a small,
partly collapsed cave, about 800 feet above the sea, and about two miles and a half from Port Louis. Encouraged
by this success M. Thirioux continued his operations, a matter of some difficulty, not to say danger, from time to
time, and was good enough to keep me acquainted with many of the results, sending me photographs of the bones
which he was fortunate in disinterring from the soil. They were not all dodos’ bones, but some belonged to other
extirpated forms of birds—as the brevipennate parrot (Lophopsittacus), the “Poule Rouge” (Aphanapteryx), and the
coot [Fulica newtonii]—and reptiles—as Didosaurus and one or more of the land tortoises—all of which are very
imperfectly known, while some of the small dodo bones are of great rarity, and at least one of them (the pygostyle)
had not been seen before.
A hand-written letter (UMZC Thirioux-Newton correspondence 1) from Thirioux to Newton dated 27 Septem-
ber 1899 provides further information: ‘I found these bones last month in mountain scree near Port Louis at a point
800 feet over the sea and 2 miles into the Vallée de Pailles open to the west (my translation)’ (See also Claessens &
Hume 2016). Another associated individual (CMNZ AV6284), the second most complete known, was exchanged in
1912 (along with dodo bones) with the Mauritius Institute for New Zealand moa material (P. Scofield pers. comm.
05.06.2018). All of the elements, other than the premaxilla, posterior portion of mandible and some vertebra, which
originate from the Mare aux Songes (MAS), are from the Thirioux collections. A partially associated individual
(FLMR 607-FLMR 613) (Fig. 17) was found, along with elements of giant tortoises Cylindraspis sp., by the author
and others at a depth of 500 mm in 1 m+ eroding sand dune deposits south of Grand Baie, in the north-west (Fig. 2).
An associated pelvis and femur (FLMR 614) was collected on the surface in a calcarenite overhang at Vieux Grand
Port, south-east Mauritius, whereas unassociated material (NHMUK u/r) was collected by the author in another
calcarenite overhang at La Prairie, south-west Mauritius, in >100 mm breccia at 200–300 mm depth, made up from
a combination of cave roof collapse with pockets of fine silt/coral sand. The age of the calcarenite is not known, but
may be similar in age to calcarenite deposits (Plaine Corail) on Rodrigues, which were inferred to have been depos-
ited in the Middle Pleistocene from several hundreds of thousands of years before present (Burney et al. 2015), or
up to one million years ago (Mylroie et al. 2016).
FIGURE 12. Comparison of ulnae (ventral aspect, left side) of Mascarene rails with Dryolimnas, A. MNHN MAD6967
Aphanapteryx bonasia u/s; B. NHMUK u/r Erythromachus leguati u/s; C. NHMUK 1882.2.28.2 Dryolimnas cuvieri cuvieri
u/s; D. NHMUK S/1989.38.1 D. c. aldabranus ♂; E. NHMUK 1910.4.8.1 D. c. abbotti u/s. Scale bar = 10 mm.
Diagnosis: As for genus.

Description and comparison: Aphanapteryx differs from other Mascarene rallid genera by the following char-
acters:
Cranium: in Erythromachus (Fig. 6), much smaller; in lateral view, processus postorbitalis shorter, more tri-
angular; shallower and less extensive fossae temporales; small foramen in os ectethmoidale; otherwise similar; in
Fulica, in lateral view, cranium less dorso-ventrally compressed; larger fonticuli orbitocraniales; but smaller than

foramen n. optici; processus postorbitalis short, triangular; fossae temporales wider, extending less posteriorly; os
lacrimale more deeper and straight extending passed os ectethmoidale; small foramen in os ectethmoidale; crista
nuchalis transversa distinct; crista temporalis indistinct; in dorsal view, frontal wider, square-shaped and much less
excavated (wider in Porphyrio); in posterior view, crista temporalis extends less dorsad (extremely reduced in Por-
phyrio and Gallinula); processus zygomaticus distinct; condylus occipitalis small and rounded; foramen magnum
large, but with a less inverted heart-shape; in ventral view, tuba auditiva communis less deep; fossa subcondylaris
much less deeply excavated; in Dryolimnas, overall similar, but smaller.
FIGURE 13. Comparison of carpometacarpi (ventral aspect, right side) of Mascarene rails, A. MI 925 Aphanapteryx bonasia
u/s with unique preservation of phalanx digiti alulae I and phalanx proximalis digitis majoris; B. NHMUK u/r Erythromachus
leguati u/s. Scale bar = 10 mm.
Rostrum: in Erythromachus, in dorsal view, os premaxillare narrower but longer (total length nearly 60% longer
than cranium length); os nasale narrower; in lateral view, overall less deep; narial opening extremely long, extending
for 66% the length of the rostrum; foramina neurovascularia less numerous, extending less to distal edge of narial
opening; os nasale narrower; in ventral view, deep, wide sulcus; in Fulica atra (F. newtonii unknown), (os premaxil-
lare equal in total length to cranium length), Gallinula chloropus (approximately 12% shorter than cranium length)
and Porphyrio porphyrio (around 14% longer than cranium length) rostrum shorter, wider and blunt; in lateral view,
narial opening short and oval; os nasale wide; in dorsal view, os nasale narrow exposing narial openings; in ventral
view, deeply excavated; in Dryolimnas, os premaxillare long and narrow, (approximately 14% longer than cranium
length), terminating in a sharp point; in dorsal view, os nasale narrow, not covering narial openings; in lateral view,
narial opening extremely long reaching 45% the length of the rostrum; foramina neurovascularia, prominent, run-
ning almost to distal edge of narial opening; os nasale extremely narrow; in ventral view, deep sulcus.
Mandible: in Erythromachus, in dorsal view, rostrum mandibulae long, but shorter (length symphysis 65% of
cranium length); processus mandibulae medialis less distinct; processus retroarticularis smaller; sulcus on crista
intercotylaris more deeply excavated; cotyla lateralis weakly U-shaped; os dentale long, narrow and terminating
in a pointed tip; foramina neurovascularia, prominent, deeply set and running almost to a deep sulcus; otherwise
similar; in Porphyrio porphyrio long, wide and blunt (length symphysis around 56% of cranium length); processus
retroarticularis reduced, in Gallinula chloropus (length symphysis 26% of cranium length) and Fulica atra (F. new-
tonii unknown) (length symphysis 26% of cranium length), rostrum mandibulae extremely short, wide and blunt;
processus retroarticularis reduced; in Dryolimnas, rostrum mandibulae comparatively short (length symphysis 34%
of cranium length).
FIGURE 14. Comparison of pelves (dorsal aspect) of Mascarene rails, A. UMZC 303BB Aphanapteryx bonasia u/s; B. UMZC
303c Erythromachus leguati u/s. Scale bar = 10 mm.
Quadrate: in Dryolimnas, Fulica, less robust; in Fulica, Gallinula, in lateral view, processus orbitalis shorter,
less deep, less rounded; processus oticus short; processus mandibularis less rounded, deeply indented and less
strongly directed laterad; in medial view, condylus pterygoideus less deeply incised; in Porphyrio, processus orbit-
alis long and more pointed.
Scapula: in Erythromachus, similar, but smaller; in Fulica newtonii, narrow; margo dorsalis more prominent;
facies articularis humeralis deeper, more pronounced; acromion more distinct, square-shaped and directed more

cranially; tuberculum coracoideum more distinct, more triangular; in Dryolimnas, margo dorsalis indistinct; facies
articularis humeralis more distinct, rounded; acromion triangular and directed dorsad; tuberculum coracoideum
indistinct; in Gallinula and Porphyrio, narrow, margo dorsalis indistinct; facies articularis humeralis shallow; acro-
mion square-shaped and strongly directed dorsad; tuberculum coracoideum more distinct, triangular.
Coracoid: in Erythromachus, much shorter; shaft comparatively wide; in dorsal aspect, cotyla scapularis deep-
er, oval-shaped and more distinct; large, but less oval foramen n. procoracoidei distal to processus procoracoideus;
impressio m. sternocoracoidei deeply excavated; in Fulica, Porphyrio and Gallinula, comparatively longer; more
gracile, shaft narrow; in dorsal aspect, cotyla scapularis deep, oval-shaped and distinct; small foramen n. procora-
coidei distal to processus procoracoideus (distinct projection of processus procoracoideus in Porphyrio); impressio
m. sternocoracoidei less deeply excavated (deeply excavated in Porphyrio); in Dryolimnas, similar to Fulica, but
much shorter; shaft narrower; in dorsal aspect, cotyla scapularis less shallow, oval-shaped and more distinct; small
foramen n. procoracoidei distal to processus procoracoideus; impressio m. sternocoracoidei deeply excavated.
FIGURE 15. Comparison of femora (caudal aspect, right side), tibiotarsi (cranial aspect, right side) and tarsometatarsi (left side,
dorsal aspect) of Mascarene rails. A., C., E. UMZC 303B Aphanapteryx bonasia u/s; B., D., F. UMZC 303c Erythromachus
leguati u/s. Scale bar = 10 mm.
Sternum: in Erythromachus, smaller, processus craniolateralis narrower, directed more laterad; in dorsal aspect;
tuberculum labri externi less pronounced; foramen pneumaticum less deeply excavated; in ventral view; spina ex-
terna variable, in NHMUK A1446 extremely reduced, or in NHMUK u/r, more distinct and square-shaped; in lateral
view, carina sterni extremely reduced; apex carinae directed more ventrally; otherwise similar; in Fulica, overall
much larger; in dorsal view; processus craniolateralis much narrower and triangular, directed more laterad; tuber-
culum labri externi larger, more pronounced; foramen pneumaticum more deeply excavated; sulcus medianus sterni
more clearly defined; in ventral view; spina externa more distinct and square-shaped with deep excavation at base;
in lateral view, carina sterni large with apex carinae directed strongly cranially; in Porphyrio, Gallinula and Dryo-
limnas; in dorsal view; strongly restricted caudad to processus costales, especially in Dryolimnas; processus cranio-
lateralis much narrower, directed more craniad; tuberculum labri externi extremely reduced; foramen pneumaticum
much more deeply excavated (open in Dryolimnas); sulcus medianus sterni more clearly defined; in ventral view;
spina externa forms a sharp point (less pointed, more rounded in Dryolimnas), with deep excavation at base (less in
Dryolimnas); in lateral view, carina sterni large with apex carinae directed strongly cranially.
Humerus: Erythromachus similar, but much smaller, shaft straighter, in caudal surface, fossae m. brachialis
less extensive, less deeply excavated; in cranial surface, fossa m. brachialis more deeply excavated; in Fulica, size
proximodistally longer, but proportionally much less robust, especially in shaft; caput humeri more pointed, more
bulbous; fossa pneumotricipitalis much deeper, extremely reduced in Aphanapteryx; margo caudalis more distinct;
crista deltopectoralis extends further distad; in Porphyrio, size smaller, in Gallinula, size smaller, proximal and
distal ends less robust; shaft straighter; fossa pneumotricipitalis much deeper; processus flexorius extends further
distad; in cranial aspect, epicondylus dorsalis more pronounced; fossae m. brachialis less deeply excavated; Dryo-
limnas similar but smaller; shaft more curved mediolaterally.
Radius: in Erythromachus, shorter, otherwise similar; in Fulica and Porphyrio, much longer proximodistally
and more craniocaudally thicker; in dorsal aspect, distinct ridge mediad to sulcus tendinosa, the latter of which is
more deeply excavated; in Gallinula and Dryolimnas, overall more gracile; shaft more curved mediolaterally; facies
articularis radiocarpalis deflected less ventrad.
FIGURE 16a. Sexual dimorphism in the skeleton of Aphanapteryx bonasia: tibiotarsi (UMZC 303B left side, cranial aspect);
tarsometatarsi (UMZC 303BB right side, dorsal aspect). Scale bar = 10 mm.
Ulna: smaller in Erythromachus, tuberculum ligamentosa collateralis ventralis more prominent; otherwise
similar; in Dryolimnas smaller, otherwise similar; in Porphyrio and Fulica, much larger, impressio brachialis more
deeply excavated; cotyla dorsalis shallower; olecranon more rounded; in Gallinula, overall more gracile; processus
cotylaris dorsalis more pronounced.
Carpometacarpus: Erythromachus and Dryolimnas similar, but smaller; in Fulica, Porphyrio and Gallinula,
longer and narrower; os metacarpale majus less robust; os metacarpale minus straighter, less arched; processus ex-
tensorius more pointed and directed more proximad.
Phalanges digiti alula I: in Dryolimnas, Fulica, Gallinula, Porphyrio long, twice length of os metacarpale
alulare.
Phalanges digiti proximalis majoris: in Dryolimnas c. cuvieri, Gallinula, Porphyrio pina cranialis distinct; in
Fulica, pina cranialis indistinct, but with distinct ridge situated ventrad.

Pelvis: in Erythromachus, in dorsal view, ala preacetabularis less expanded laterad; in lateral view, foramen
acetabuli equal or smaller in size than foramen ilioischiadicum; otherwise similar; in Fulica, Porphyrio and Gal-
linula, in dorsal view, sulcus antitrochantericus extend laterally equal to the ala preacetabularis giving a charac-
teristic narrow look to the pelvis; no canalis iliosynsacralis on either side of the crista iliaca dorsalis, the latter of
which terminates at proximal edge of the antitrochanter when held in lateral view (extends to distal edge of foramen
ilioischiadicum in Aphanapteryx and Erythromachus); reduced number of foramina intertransversariae; ala postac-
etabularis ilii short on lateral edge exposing ala ischii; in lateral view, foramen acetabuli smaller in size than foramen
ilioischiadicum; crista iliaca dorsalis less arched; foramen ilioischiadicum more deeply situated; in ventral view,
one distinct processus costalis, not two; ala postacetabularis ilii more deeply set; ala ischii directed more laterad; in
Dryolimnas, in dorsal view, canalis iliosynsacralis present; crista spinosa synsacri terminates at the proximal edge
of the antitrochanter; in lateral view, foramen acetabuli smaller in size than foramen ilioischiadicum; in ventral view,
one distinct processus costalis.
Femur: in Erythromachus, smaller, otherwise similar; in Fulica newtonii, extremely robust, largest of Mas-
carene endemic species; in cranial view, crista trochanteris larger and more strongly deflected mediad; facies articu-
laris acetabularis more pronounced, more rounded and directed more distad; sulcus patellaris extends less proximad;
epicondylus lateralis less distinct; in medial aspect, crista trochanteris more rounded; facies articularis acetabularis
and fovea ligamentosa capitis smaller; in caudal aspect, distal sulcus proximal to trochlea extends further proximad;
in Porphyrio, Gallinula and Dryolimnas, overall similar but much less robust, especially in shaft; in caudal aspect,
fossa poplitea shallower.
Tibiotarsus: in Erythromachus much smaller, sulcus extensorius less deeply excavated, extending less proxi-
mad; canalis extensorius more circular, less oval; otherwise similar; in Fulica; longer but less robust, especially at
proximal and distal ends; sulcus extensorius much more deeply excavated; pons supratendineus narrower and flat-
ter, less arched, and situated more mediad; crista fibularis extends further distad; crista cnemialis lateralis shorter; in
lateral view, crista cnemialis cranialis more pronounced; shaft less deep, especially at distal end; condylus lateralis
larger, more rounded; depressio epicondylaris lateralis more deeply excavated; in Gallinula, Porphyrio and Dryo-
limnas, smaller, otherwise similar to Fulica.
Fibula: in Erythromachus, similar, but smaller; in Porphyrio, Gallinula and Dryolimnas, smaller, less robust;
lacks caudal groove.
Tarsometatarsus: in Erythromachus, much smaller, but proportionally more robust; trochleae metatarsorum
more splayed, especially trochlea metatarsi II; retinaculi extensorii connected by an ossified bridge in some indi-
viduals (contra Mourer-Chauviré et al. 1999); otherwise similar; in Dryolimnas, smaller, trochleae less splayed;
retinaculi extensorii connected by an ossified bridge in nominate; on proximal end, open crista medialis flexoris
digitorum longus; sulcus for tendon of musculus flexor perforatus digiti II and sulcus for tendon of musculus flexor
hallucis longus shallower; in Fulica; greater in total length; in dorsal aspect; proximal and distal ends less robust;
trochleae metatarsorum less splayed, narrower; trochlea metatarsi II directed more plantad; shaft narrower; deeper
excavation around foramen vasculare distale; fossa metatarsi I extends much further proximad; in plantar aspect,
crista medialis hypotarsi centrad, not deflected laterad; on proximal end, canal for tendon of musculus flexor digi-
torum longus; sulci for tendon of musculus flexor perforatus digiti II and for tendon of musculus flexor halluces
longus shallower; Gallinula much smaller; on proximal end, canal for tendon of musculus flexor digitorum longus;
in Porphyrio, on proximal end, crista medialis flexoris digitorum longus present; sulci for tendon of musculus flexor
perforatus digiti II and for tendon of musculus flexor halluces longus more deeply incised (Fig. 16b).
Remarks. A chronological historical account: Much ink has been spilt over contemporary illustrations and
accounts of Aphanapteryx. There are a few early 17th century accounts from Mauritius that reference the rail at a
time when they were known as ‘Indise riviers houdt snippen’ (Indian river woodcocks) and ‘Velt-hoender or Feld-
hüner’ (Field-hens) (Strickland & Melville 1848), but no other information was provided. It was during the voyage
of Vice-Admiral Wybrandt Warwijck, who was equipped by the Oude Oost-Indische Compagnie (Old East India
Company), that Aphanapteryx was first mentioned. After the Dutch claimed Mauritius for the Netherlands in Sep-
tember, 1598, ships’ pilot, Heyndrick Dircksz Jolinck led two expeditions, one on 23–27 September, and the second
28 September–1 October, to explore the island to the north-east of Warwijck Harbour (now Vieux Grand Port) (Fig.
2) (Barnwell 1948; Moree 1998). There he encountered the first native fauna and remarked (Keuning 1947: 62—my
translation):
FIGURE 16b. Comparison of tarsometatarsi (proximal end, right side), of Mascarene rails. A. UMZC 303BB Aphanapteryx
bonasia u/s; B. UMZC 303C Erythromachus leguati u/s; C. NHMUK 1851.8.25.40 Porphyrio p. porphyrio u/s; D. UMZC
300AA Dryolimnas chekei sp. nov. u/s; E. NHMUK 1882.2.28.2 Dryolimnas cuvieri cuvieri u/s; F. NHMUK S/1989.38.1 D. c.
aldabranus ♂; G. NHMUK S/1977.23.1 Gallinula c. chloropus u/s; H. NHMUK 1930.3.24.63 Fulica atra u/s. In Aphanapteryx,
Erythromachus, Porphyrio and Dryolimnas, presence of (i) sulcus for tendon of musculus flexor perforatus digiti II; (ii) sulcus
for tendon of musculus flexor hallucis longus; in Gallinula and Fulica, (i) canal for tendon of musculus flexor perforatus digiti
II; (ii) sulcus for tendon of musculus flexor hallucis longus. Scale bar = 10 mm.
…Also we found here many kinds of birds such as pigeons, herons, water-snipes [waders], grey parrots [Psittacula
bensoni (Holyoak, 1973)], Indian river woodcocks [Aphanapteryx] as large as a hen; raboforcadoes [Frigatebirds
Fregata sp.] were many, but not very good to eat and its wings were so long that I could not measure them. We also
found handsome geese, ducks, cranes [flamingos] and other birds so tame that we caught almost all by hitting with
a stick.
Aphanapteryx was mentioned again during van der Hagen’s voyage in 1602 (Cornelisz in Strickland & Mel-
ville 1848). Cornelisz’s large and small field-hens may refer to more than one species, possibly Aphanapteryx and
Dryolimnas (Cheke 2013a; see below), sexual dimorphism, or adult/juvenile Aphanapteryx (translation from Cheke
2013a: 1):
[here is something] about the animals found there in that land, such as tortoises, dodos, flamingos, geese, ducks,
[birds] like velt-hoender [field-hens] large and small, Indian ravens, also doves, there are also red-tailed doves, there
[=through (eating) which] many men have been ill, here are also many grey parrots, finally also green [ones] with
long tails, some of which were captured.
During the voyage of Admiral Peter Wilhelm Verhuffen in 1611 (Verken 1613), the rails were also mentioned.
The name ‘Rebhüner [= partridge]’ was probably referable to introduced Perdix (translation from Strickland &
Melville 1848: 125):
There are also many Birds, as Turtle-doves, grey Parrots, Rabos forcados [Frigatebirds Fregata sp.], Feldhüner
[Field-hens], Rebhüner [Partridges], and other birds in size like Swans, with large heads [Dodo Raphus cucullatus
(Linnaeus, 1758)].

The English traveller and historian, Sir Thomas Herbert visited Mauritius in 1629 and described and depicted
in his travelogue a bird he simply called a ‘hen’. In some of Herbert’s six editions of ‘Some yeares travels’, which
comprise four in English, one in French and one in Dutch, the text differs slightly. His first edition (Herbert 1634:
214) account reads:
The Hens in eating taste like parched [roast] pigs, if you see a flocke of twelve or twentie, 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 [iota] till they be all destroyed.
Whereas in his account in Herbert (1638: 349), which is virtually identical to that in the expanded English and
French editions (Herbert 1663: 546, 1665: 405, 1677: 385), he remarks:
The birds, had here at no charge and little labour, are Hens, Bats, Herns [Herons], &c. The Hens flock together
twenty or forty in a company; if you catch one, take all: the surest way is to shew them a red cloth; that colour exas-
perates their spleen [vents their anger], they shew it by flying out-ragiously at it; and when wee struck downe one,
the residue budged not one iot till all were vanquished: they eat like parcht pigs if you roast them.
A rather enigmatic account was probably penned by the lawyer, Leonardus Wallesius, who was travelling with
two homeward bound Dutch ships ‘sGravenhage and Ter Veere (Winters et al. 2017); the ships arrived at Vieux
Grand Port, Mauritius on 29 April 1632 and stayed until May 19. Wallesius described the animals he encountered in
a ‘klucht’, a popular 17th century Dutch short comedy parody, in which Aphanapteryx was an allegory for a soldier
(ibid):
The soldiers were very small and moved slowly, so that we could catch them easily with our hands. Their ar-
mour 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 (translation from Winters et al. 2017: 149).
FIGURE 17. An associated, partial skeleton of Aphanapteryx bonasia found in an eroding sand dune south of Grand Baie,
north-east Mauritius. A femur can be seen in-situ to the left of the scale bar. Skeleton comprises right and left distal humerus;
left ulna shaft; proximal pelvis; complete right & left proximal femur; complete right tibiotarsus. Scale bar = 10 mm.
The English traveller Peter Mundy (in Temple 1919) visited Mauritius in 1634 on his way to Surat, India, and
saw a single ‘Mauritius henne’, but refers to its colouration as ‘yellowish wheaten’ rather than reddish-brown. His
description of other birds encountered during his voyages are accurate, so Mundy may have been referring to a juve-
nile Aphanapteryx (Olson 1977). If so, his unique description of colour and size (hen-sized) suggests that juveniles
were yellowish in colouration until they were at least as large as an adult; juvenile rallids reach adulthood within
one year (Ripley 1977). Mundy’s account reads (Temple 1919: 352–353):
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.
Hachisuka (1937), who was prone to overzealous mis-interpretation when describing Mascarene avifauna,
seized on the opportunity formally to describe a new taxon, erecting a new genus and species, Kuina mundyi, based
on Mundy’s ‘yellowish wheaten’ bird. The basis for this is completely unfounded.
Although his words cannot always be trusted (Cheke & Hume 2008), the mariner François Cauche (1651) in
1638 described in more detail than Peter Mundy the novel way of catching red rails with a red cloth, which suggests
that his observation was genuine. However, Cauche himself was unlikely to have disembarked on Mauritius and
probably obtained the information from other mariners (Cheke & Hume 2008).
Red hens, with a beak of a woodcock [Scolopax rusticola]; to take them one must only present them with a
piece of red cloth, [which] they follow, and let themselves be taken by hand: they are the size of our chickens, [and]
excellent to eat (my translation).
The Dutch settler, Johannes Pretorius, while on Mauritius in 1666–69 (Hume & Winters 2016: 5), provides the
only information about this rails’ feeding habits:
The dodaers [Aphanapteryx bonasia] is a red bird, as big as a fowl, has short wings and cannot fly. It scratches
in the earth with its sharp claws like a fowl to find food such as worms under the fallen leaves. This bird is unbe-
lievably stupid. When one waves a stocking cap and makes a sharp sound with the mouth, it immediately heads
towards that person, and if one carries a stick, all of them can be killed with it without any escaping. They are fatty
and greasy to eat. They have a long, sharp beak which is slightly curved at the end.
The English explorer John Marshall, who was on Mauritius in 1668, also called the red rail a ‘dodo’, which has
led to considerable debate about the date of extinction of the true Dodo (Cheke 2006; Cheke & Hume 2008) (see
below). Marshall’s account (Khan 1927: 47–48) reads:
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 downe, and their wings so little that it is not able to sup-
port their bodies; but they have long leggs and will runn very fast, that a man shall not take them, they will turne
so about the trees. They are good meate when roasted, tasting something like pig, and their skin like pig skin when
roosted, being hard.
The last detailed account of live birds was penned by the pastor, Johann Christian Hoffman (1680), who was on
Mauritius from 1673–75 (translation from Cheke & Hume 2008: 127):
[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 can-
not 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 to its aid and can be offered the same fate.
The red rail was illustrated on a number of occasions, with the two most realistic images only coming to light
in the 19th century. Joris Joostenszoon Laerle, who was on the flagship Gelderland when it visited Mauritius in
1601 (Milne-Edwards 1875a; Hume 2003), sketched a recently killed red rail (Fig. 18), providing by the far the
most accurate rendition and the only one drawn on Mauritius (Hume 2003). Frauenfeld (1868a) discovered hitherto
unknown paintings of Mauritian species, including Aphanapteryx, which were executed c.1610, and attributed them
to the Dutch artist Georg Hoefnagel. However, it was Jacob Hoefnagel (and not his father Georg or Joris Hoefnagel)
who, after becoming court painter in 1602, illustrated Aphanapteryx and a dodo (Anker 1974; Jackson 1999; Hume
2006); from 1602 until at least 1610, Emperor Rudolph II of Austria employed Jacob Hoefnagel to illustrate animals
housed in his zoological menageries. Rudolf II maintained the menageries established by Emperor Maximilian II at
Ebersdorf, Naugebau near Vienna, and at Prague (Anker 1974). The rail, probably the only specimen to reach Eu-

rope, arrived at the menagerie along with a dodo and the first Mauritian Fruit Bat Pteropus niger Kerr, 1792 (Cheke
2007; Cheke & Hume 2008). It is not known if any of these individuals arrived alive, but they were all certainly
illustrated as stuffed individuals; the red rail painting remains the only known colour depiction (Fig. 19). Frauenfeld
(1868a) gave the following diagnosis (translation in Rothschild 1907a: 131):
Of the size of a fowl, of a uniform brown red all over. Bill and legs dark, iris yellowish. Feathers decomposed,
as in the Apteryx, somewhat lengthened on the nape.
Frauenfeld’s mention of the yellow-coloured iris based on the painting suggests that the artist may have seen
the bird alive however. The red rail was illustrated, albeit crudely, in the account of Pieter van den Broecke (1646),
a VOC (Verenigde Oost-Indische Compagnie (Dutch East India Company)) employee who visited Mauritius 19
April–23 May 1617. The rail was figured beside a dodo and a single-horned goat (Fig. 20), but there is no reference
to it in the text.
FIGURE 18. A freshly killed specimen of Aphanapteryx bonasia drawn in 1601 by Joris Joostenszoon Laerle during the voyage
of the flagship Gelderland (From Hume 2003).
Another attributed illustration is in a c.1625 dodo painting by Roelandt Savery (Hachisuka 1953), but a detailed
examination of the original by the author disputes this assignation; the bird almost certainly depicts an Eurasian
Bittern Botaurus stellaris (Linnaeus, 1758) (Hume 2017). According to Greenway (1958, 1967), who provides no
reference to the paintings to which he was referring, two other Savery paintings show Aphanapteryx. One must be
that discussed and illustrated in Renshaw (1936: 60). However, this image is clearly not Aphanapteryx, as it is de-
picted the same size as a Peacock Pavo cristatus Linnaeus, 1758, at 100–115 cm in total length, excluding the tail
train, and with long wings and tail. My examination of Savery’s other faunal paintings has probably identified the
second red rail painting mention by Greenway, i.e. Savery’s ‘Dodo in a landscape with animals’. There is a large,
reddish-brown bird behind a pelican (centre left), which is clearly too long-winged to be a flightless rail, and again
looks more like a bittern.
Other examples of supposed Aphanapteryx illustrations exist. Ripley (1977) considered that a bird in the Ital-
ian artist Jacopo Bassano’s painting Arca di No (Noah’s Ark), executed c.1570, represents Aphanapteryx, but this is
much too early for a red rail to have arrived in Europe, and the bird itself looks more like another poorly reproduced
Eurasian bittern (see Cheke 1987; Cheke & Hume 2008). A 1613 painting by Jan Brueghel the Elder entitled ‘The
entry of the animals into Noah’s Ark’ also depicts a pair of rail-like birds, but one of the birds is in the exact same
pose as that of Bassano, and again both clearly represent Eurasian bitterns. I conclude that the supposed red rail
depictions of Bassano, Brueghel the Elder and all of those by Savery are not related to Aphanapteryx: the c.1610
painting by Jacob Hoefnagel is the only unequivocal coloured rendition.
FIGURE 19. A stuffed specimen of Aphanapteryx bonasia painted by Jacob Hoefnagel c.1610. This is the only unequivocal
colour depiction (From Frauenfeld 1868).

Accompanying the text of Thomas Herbert’s visit in 1629 (Herbert 1634), there is an extremely crude illustra-
tion of a red rail standing between a Mauritian Broad-billed Parrot Lophopsittacus mauritianus (Owen, 1866) and
a dodo (Fig. 20). The artist’s obvious inadequacies did not prevent Schlegel (1854a, b) and Hachisuka (1953: 138)
from placing scientific credence on it and applied the names Didus herbertii and Pezocrex herberti respectively;
Hachisuka further suggested that the straight bill and wattles were diagnostic characters. The absurdity of this as-
signation is highlighted by Olson (1977: 360), who pointed out that Hachisuka further failed to mention that the rail
has only two anterior toes, a much more remarkable and diagnostic character!
Peter Mundy (in Temple 1919) in 1634 illustrated his ‘Mauritius henne’ (Fig. 20), which evidently shows
Aphanapteryx, but was described with yellowish rather than reddish plumage, and was probably describing a juve-
nile (Olson 1977).
FIGURE 20. Three contemporary images of Aphanapteryx bonasia: (above) from van den Broecke (1646) in 1617, Aphana-
pteryx (right) with dodo and goat; (below left) from Herbert (1634) in 1629, Aphanapteryx (centre) with Broad-billed Parrot
Lophopsittacus mauritianus and dodo; (below right) from Mundy (in Temple 1919) in 1634.
It was the geologist and natural historian Hugh Edwin Strickland (in Strickland & Melville 1848: 21), when
compiling the literature about Aphanapteryx, who first concluded that all of the relevant contemporary accounts
and illustrations were referring to the same bird. However, its relationships were still unclear. It was not until sub-
fossil remains were discovered in the Mare aux Songes marsh in 1865 that the rallid affinities of Aphanapteryx
were finally resolved (Milne-Edwards 1868, 1869a). In 1889, Mauritian Governor John Pope Hennessy appointed
Théodore Sauzier to the position of president of the paleontological committee ‘Comité des Souvenirs Historiques’,
and instructed him to continue exploring the Mare aux Songes (Vinson 1968). Under Sauzier’s guidance, the marsh
was excavated from 1889 to 1891, and the discovery of further subfossil rail material permitted proof that Aphan-
apteryx was flightless (E. Newton & Gadow 1893). An associated individual was subsequently discovered by the
amateur natural historian Etienne Thirioux around the turn of the 20th century, probably at the Vallée des Prêtres,
Moka Range, central-west Mauritius (details taken from labels of other Thirioux material) (Fig. 2), and remains the
best-preserved and most complete specimen yet discovered (Claessens & Hume 2016) (Fig. 5). Further, similarly
preserved material has been found by the author in the Moka Range, under Snail Rock, all in cavities under large
boulders where the sediment is shallow, dry and without penetrating plant roots (Hume 2005). A partially articulated
specimen was found at Vieux Grand Port (south-east), with further individual elements collected by the author and
others in cave sites at La Prairie (south-west) and at coastal deposits south of Grand Baie (north-west) (Fig. 2, 17).
Ecology: The fossil deposits demonstrate that Aphanapteryx occurred widely on Mauritius in montane, lowland
and coastal habitats. Otherwise, virtually nothing was recorded about the ecology of Aphanapteryx, but a recently
discovered document written by Johannes Pretorius (see above) provides some insight (Hume & Winters 2016).
Pretorius’s account confirms that the rails were feeding on invertebrates amongst leaf litter by removing detritus
with their feet and probing under leaves with their bills. Furthermore, the maxillary and mandibular premaxilla of
Aphanapteryx are pitted with numerous foramina running from the nasal aperture to almost the tip of the premaxilla
(Fig. 6). They are mostly oval in shape and vary in depth and inclination, becoming shallower posteriorly (sub-
narial foramina). Similar foramina occur in other probing birds, especially kiwis (Apterygidae), ibises (Threskior-
nithidae) and sandpipers (Scolopacidae). These three unrelated bird families all share a common foraging strategy,
probing for live food buried beneath substrate, and have evolved elongated bills with clusters of mechanoreceptors
concentrated at the bill tip (Nebel & Thompson 2005; Cunningham et al. 2007, 2010, 2013); their specialized bill
tips allow detection of buried prey by picking up vibrotactile and/or pressure cues in the substrate. Although not as
concentrated at the tip, foramina occur on the elongate bill of Aphanapteryx, as they do in some other long-billed,
probing rails, e.g. New Zealand Snipe-rail Capellirallus karamu Falla, 1954, with the main point of bill curvature
beginning at the anterior end of the nasal opening, and mirrored by the mandible. This would not only have made the
tip strong, but also extremely sensitive, and a most useful tool when probing for invertebrates, as did Capellirallus
(Worthy & Holdaway 2002; Tennyson & Martinson 2006).
As Mauritius once had an abundant terrestrial gastropod fauna (Griffiths & Florens 2006), Aphanapteryx may
have also been an active snail predator, and perhaps scavenged seasonal sea bird breeding colonies and tortoise nest-
ing sites (as did Erythromachus, see below). Milne-Edwards (1868, 1869a, 1869b), followed by Hachisuka (1953),
who clearly plagiarised the original source, surmised that as the tip of the bill was sharp and strong, the rail probably
fed by crushing molluscs and other shells, rather like an oystercatcher Haematopus sp. Milne-Edward’s suggestion
may well have some credence, as many shells of some of the largest but now-extinct Mauritius land snails, e.g.
Tropidophora carinata (Born, 1780), have a puncture hole on the ventral surface, suggesting bird predation (Owen
Griffiths in Hume 2017) (Fig. 21). Similar-sized New Zealand Weka Gallirallus australis (Sparrman, 1786) will
puncture the shell of terrestrial gastropods to extract the meat (Meads et al. 1984), but they will also swallow Powel-
liphanta snails up to 20 mm in diameter (Climo 1975). No doubt Aphanapteryx also swallowed smaller snails whole.
Pretorius describes Aphanapteryx actively searching for worms in leaf-litter (Hume & Winters 2016: 5), which may
be in reference to predation of terrestrial nemertean and planarian worms, as a number of endemic species exist on
Mauritius and occur in leaf-litter as well as in rotting wood (Gulliver 1879; Kaburaki 1920). Pretorius may also have
been referring to a now-extinct endemic worm-snake, Typhlops cariei Hoffstetter, 1946, which reached lengths of
200mm and almost certainly occurred in leaf-litter as do other species (McDiarmid et al. 1999).
Livezey (2003) provided a principal component analysis (PCA) of detailed matrices of rallid skulls, which
showed a great diversity of size and bill and a probable reflection of foraging mode, with the most extreme bill shape
exhibited, amongst others, in Aphanapteryx and Erythromachus. The extreme variation in bill length of Aphan-
apteryx was almost certainly due to sexual dimorphism (see Livezey 2003), a characteristic seen in some other
rallid taxa, e.g. Cabalus modestus (Trewick 1997; Livezey 2003), which strongly suggests that the sexes foraged on
different size classes of food.

FIGURE 21. Possible Aphanapteryx predation puncture marks on the shells of the large, extinct terrestrial snail, Tropidophora
carinata. Snail specimens curtesy of Owen Griffiths. Scale bar = 10 mm.
Extinction: The accidental introduction of the black rat from at least the 14th century (Hume 2013) did not
appear to affect Aphanapteryx numbers, as the birds were still relatively common until the early 1680s (Hume et
al. 2004). Elsewhere, large rails can survive alongside rats or even habitually kill them, as observed in Dryolimnas
on Aldabra Atoll (Wanless 2003), and this species also aggressively defends its eggs and chicks against these ver-
min (Frith 1977). Presumably due to having a similar nature, Aphanapteryx survived alongside rats for centuries
(Hume 2013, 2014a). Furthermore, and despite being a ground nester, it also coped with the onslaughts of hunting
by humans and the probable predation of eggs and chicks by introduced pigs and the Crab-eating Macaque Macaca
fascicularis Raffles, 1821.
A much-maligned account was written by Benjamin Harry in 1681, chief mate of the ship Berkeley Castle, and
the first person to measure the inclination of the Earth’s magnetic field in the Southern Hemisphere (Jackson 2014).
While visiting Mauritius during his voyage from England to India, Harry wrote (from Strickland & Melville 1848:
26):
Now having a little respitt I will make a little description of the Island, first of its Producks then of its parts: ffirst
of winged and feathered fowle the less passant, are Dodos whose flesh is very hard, a small sort of Gees, reasonably
good Teele, Curleves, Pasca fflemingos, Turtle Doves, large Batts, many small Birdes which are good [to eat]
It is generally considered to be the last account mentioning dodos, but Cheke (2006) argued that it was referable
to Aphanapteryx, as the latter had inherited the dodo’s name after it became extinct (A. Newton 1868: 480; Cheke
1987, 2006). However, Dutch commander Hubert Hugo noted the capture of a ‘dodaersen’ on 16 August 1673,
whereas Opperhoofd (Chief) Isaac Joan Lamotius noted on a number of occasions the capturing of ‘dodaersen’, the
last as late as October 1688 (Moree 1998; Hume et al. 2004). Hugo knew the difference between a dodo and a rail,
having specifically questioned an escaped slave about it (Cheke 2006), and Lamotius was an educated man with
a great interest in natural history, so their observations are likely referring to the true dodo (see Hume et al. 2004;
Hume 2006; Hume & Winters 2016; Hume 2017), rather than Aphanapteryx (Cheke 2006).
If Harry’s report is referable to Aphanapteryx, the situation appears to have changed drastically over the next
decade, as in 1693, the Huguenot François Leguat (1708: 71 English edition) mentioned it for the last time, when
he stated:
This island [Mauritius] formerly abounded with wild geese and ducks, moor-hens [Fulica newtonii], water-
quails [Aphanapteryx], sea and land tortoises, but now all these are become scarce.
This decline coincides with the establishment of feral cats during the late 1680s (Cheke & Hume 2008), which
are formidable predators of flightless rails and other birds (Medina et al. 2011; Hume 2017). The rapid disappear-
ance of rails once cats became established has occurred elsewhere, most notably on Aldabra Atoll where D. c. al-
dabranus only survives on cat-free islands (Wanless 2003: Safford & Hawkins 2013). Being inquisitive and fearless,
both adult and young red rails must have fallen easy prey to feral cats, and as a result probably became extinct by
the end of the 17th century at the latest.
Genus Erythromachus Milne-Edwards, 1874
Erythromachus Milne-Edwards 1874, p.6, pls.11–12. Type by monotypy: E. leguati Milne-Edwards
Etymology: From Greek eruthros = red, and makhç = battle, combat, in reference to the rail’s aggression shown
towards red cloth.
Revised Diagnosis: Erythromachus differs from other Mascarene genera by the following characters:
Cranium (Fig. 6, 22): cranium medium-sized; in lateral view, dorso-ventrally compressed; small fonticuli orbi-
tocraniales; large foramen n. optici; processus postorbitalis short and triangular; fossae temporales shallow.
Rostrum (Fig. 6, 22): os premaxillare long, os nasale narrow, and total length nearly 60% longer than cranium
length; in lateral view, shallow; narial opening extremely long, extending for 66% the length of the rostrum; foram-
ina neurovascularia present, not extending to rostral edge of narial opening; os nasale narrow; in ventral view, deep,
wide sulcus.
Mandible (Fig. 6): rostrum mandibulae long and narrow terminating in a sharp point, length symphysis approxi-
mately 65% of cranium length; in dorsal view, processus mandibulae medialis indistinct; processus retroarticularis
small; sulcus on crista intercotylaris deeply excavated; cotyla lateralis weakly U-shaped; os dentale long, narrow
and terminating in a pointed tip; large foramina neurovascularia, deeply set and running almost to a deep sulcus
situated in centre of mandible.
Scapula (Fig. 7): small and narrow; in lateral view, margo dorsalis distinct; facies articularis humeralis shallow;
acromion blunt, rounded and directed dorsad; tuberculum coracoideum indistinct.
Coracoid (Fig. 8): short but shaft wide; in dorsal aspect, cotyla scapularis deep, oval-shaped; large foramen n.
supracoracoidei sternal to processus procoracoideus; impressio m. sternocoracoidei deeply excavated.
Sternum (Fig. 9): small; processus craniolateralis narrow, directed laterad; in dorsal aspect; tuberculum labri
externi indistinct; foramen pneumaticum on midline shallow; in ventral view; spina externa variable, reduced or
more distinct and square-shaped; in lateral view, carina sterni extremely reduced; apex carinae directed ventrad.
Humerus (Fig. 10): extremely reduced, shaft curved dorsoventrally; in caudal aspect, incisura capitis deeply
excavated; fossa pneumotricipitalis shallow; in cranial view, crista bicipitalis, short, square-shaped; fossa m. bra-
chialis shallowly excavated.
Radius (Fig. 11): short; sulcus tendinosa deep extending proximad approximately ¼ of total length radius length;
Ulna (Fig. 12, 22): short and strongly arched dorso-ventrally proximally; tuberculum ligamentosa collateralis
prominent; in caudal aspect, olecranon indistinct; processus cotylaris dorsalis short.
Carpometacarpus (Fig. 22): in dorsal aspect, processus extensorius short and rounded; os metacarpale majus
robust; os metacarpale minus weakly arched creating narrow spatium intermetacarpale; proximal synostosis long.
Pelvis (Fig. 14, 22): in dorsal view, ala preacetabularis weakly expanded laterally; in lateral view, foramen
ilioischiadicum is equal or slightly larger than foramen acetabuli; canalis iliosynsacralis extend to distal edge of
foramen ilioischiadicum.
Femur (Fig. 15, 22): extremely robust; in cranial view, shaft curved medially and directed caudally at the proxi-
mal end; crista trochanteris strongly deflected mediad; in medial aspect, shaft strongly arched dorsally; in caudal
aspect, condylus medialis larger than condylus lateralis.

Tibiotarsus (Fig. 15, 22): short, but robust; sulcus extensorius shallow and restricted proximally; canalis exten-
sorius circular, less oval; pons supratendineus arched.
Fibula: short and robust; in dorsal view, caput fibulae narrow and weakly grooved caudally; tuberculum m.
iliofibularis pronounced and excavated cranially.
Tarsometatarsus (Fig. 15): short, but extremely robust; in dorsal aspect, trochleae metatarsorum distinctly
splayed, especially trochlea metatarsi II; metatarsi II directed slightly dorsad; presence of an ossified bridge con-
necting retinaculi extensorii individually variable; in plantar aspect, crista medialis hypotarsi central, not deflected
laterally; in plantar aspect, surface area of foramen vasculare distale shallowly excavated; fossa metatarsi I extends
proximad; hypotarsi deflected laterad; on proximal end, crista medialis flexoris digitorum longus present; sulci for
tendon of musculus flexor perforatus digiti II and musculus flexor hallucis longus present.
Remarks: Milne-Edwards (1874), when describing Erythromachus, possessed both the original accounts and
the first subfossil remains then recently discovered in caves on the Plaine Corail, Rodrigues (Fig. 4). These, along
with elements of Rodrigues Night Heron Nycticorax megacephalus (Milne-Edwards, 1874), Rodrigues Turtle Dove
Nesoenas rodericanus (Milne-Edwards, 1874) and Rodrigues Parrot Necropsittacus rodricanus (Milne-Edwards,
1867a), were sent to Milne-Edwards in Paris by Edward Newton, brother of Alfred Newton, professor at the UMZC
(Milne-Edwards 1869b; A. Newton 1875b; Hume 2007, 2011a, 2014a, b) (data taken from labels). From these he
determined that a flightless rail, related to Aphanapteryx on Mauritius, had once occurred on Rodrigues, and pro-
posed a new genus, Erythromachus. As better material became available it was later referred to Aphanapteryx by
Günther & Newton (1879); however, Piveteau (1945) noted that distinct differences in the cranium justified the
generic separation of Erythromachus, which was an assignation followed by Olson (1977), Mourer-Chauviré et
al. (1999) and others, and applied here. As recently shown in the endemic scops owls of Mauritius and Rodrigues
(Louchart et al. 2018), there is precedent for differing evolutionary pathways for the two endemic genera of rails.
FIGURE 22. Associated partial skeleton of NHMUK u/r Erythromachus leguati collected in Caverne Poule Rouge. Clockwise
from top; cranium and rostrum; pelvis, right and left femur; right and left tibiotarsus, right carpometacarpus, left ulna. Scale bar
= 10 mm.
Erythromachus was smaller than Aphanapteryx, being approximately 35 cm in length, but had proportionately
longer wings (Appendix 2, Table 3, Fig. 23a); rare very young juvenile material, with bones incompletely ossified,
are also known (Fig. 23b). The plumage was hair-like and mottled grey and white, and the bill, bare orbital skin,
legs and feet bright red. The bill was similarly long and decurved, but the narial openings were much longer than
in Aphanapteryx. Erythromachus presumably exhibited marked size sexual dimorphism (Fig. 24), with the males
being presumably bigger than females, as in other Rallidae (Ripley 1977; Livezey 2003), and may have had a body
mass exceeding 500 g (Livezey 2003). The bill differed considerably in length, with one specimen being around a
third shorter (Günther & Newton 1879: pl. 43), and it was also reported to be variable in curvature (Günther & New-
ton 1879: 432), which the authors ascribe to probable sexual dimorphism. Livezey (2003: 190) also considered that
bill size variation was due probably to sexual dimorphism, but the small sample size was not adequate for a robust
statistical analysis. The idea of variable bill curvature also arose in part from the accounts of Leguat (1708), who
described the bill as being ‘straight’, whereas Tafforet (1725–26) compared it with the curved bill of an Eurasian
Curlew Numenius arquata (Linnaeus, 1758). However, I can find no basis in fact regarding this comment, as I have
examined all available premaxillae and found no differences in bill curvature.
FIGURE 23. FIGURE 23a (left). Post-cranial skeleton of Erythromachus leguati showing the proportions of the limbs; from
left to right, femur; tibiotarsus; fibula; tarsometatarsus; coracoid; humerus (right and left); ulna; radius (right and left); car-
pometacarpus; FIGURE 23b (right). Juvenile ulna (top right) and distal tarsometatarsus (bottom left) compared with an adult.

†Rodrigues Rail or Leguat’s Rail Erythromachus leguati Milne-Edwards, 1874
Gelinotes, Gelinottes or Gellinotes Leguat, 1708, p.71; Pingré, 1763 (ms 1804); Strickland & Melville, 1848, p.55: Studer, 1886,
p.30
Oiseaux Tafforet, 1725–26
Erythromachus leguati: Milne-Edwards, 1874, p.6, pls.11–12, 1866–73 [1874], p.122, 1875a, p.14, 1875b, p.1214;A. Newton,
1875b, p.41; Behm, 1880, p.289; Oliver, 1891, p.335; Sharpe, 1894, pp.69,333; Oustalet, 1897, p.96; Rothschild, 1907a,
p.135, pl.30, 1907b, p.198; Atkinson, 1922, p.52; Lambrecht, 1933, p.470; Piveteau, 1945, p.8; Staub, 1973a, p.24; North-
Coombes, 1971, p.70; Mourer-Chauviré et al., 1999, p.23; Worthy & Holdaway, 2002, p. 387; Livezey, 2003, p.1; Cheke &
Hume, 2008, p.127, tb.3.1, pl.14; Hume & Walters, 2012, p.110; Cheke, 2013b, p.13; Hume, 2013, p.218; 2014a, pp.36–37,
2017, p.128; Safford & Hawkins, 2013, p.45, pl.48; del Hoyo & Collar, 2014, pp.342,772; Burney et al., 2015, p.48
Miserythrus leguati: Milne-Edwards in Haast, 1874 (July), p.217; Anon, 1874, p.455; [A. Newton], 1875a, p.733 (incorrect
author attribution); Richmond, 1909, p.622
Erythromachus leguata Fürbringer, 1888, p.20 (lapsus)
Myserythrus leguati: Hachisuka, 1953, p.149 (lapsus)
Aphanapteryx leguati: Günther & Newton, 1879, p.431, pl.x; Slater, 1881, p.27; E. Newton, 1888, p.552; Milne-Edwards,
1896a, p.77; Atkinson, 1922, p.52; Greenway, 1958, p.117, 1967, p.117; Luther, 1970, p.187, 1986, p.183, 1986 [2005],
p.183; North-Coombes, 1971, p.70; Olson, 1977, p.361; Cheke, 1987, p.37; Cowles, 1987, p.96; Fuller, 1987, p.89, 2001,
p.149; Day, 1991, p.90; Livezey, 1998, p.2083; Taylor & van Perlo, 1998, p.60; Adams et al., 2003, p.344; Tyrberg, 2009,
p.71.
Aphanopteryx leguati: Rothschild, 1907a, p.135 (lapsus)
Erthyromachus leguati: Brodkorb, 1967, p.133 (lapsus)
Holotype: None designated.

Syntypes: As Milne-Edwards (1874) did not designate a holotype before returning the loan material to the
UMZC, presumably it was Alfred Newton who bulk-listed the following specimens as syntypes. None has precise
locality data other than collected in the caves of Rodrigues (Plaine Corail): Cranium: UMZC 303c; UMZC 303c;
Rostrum: UMZC 303c; Sternum: UMZC 303c; UMZC 303c: UMZC 303c; Humerus: UMZC 303c (L); UMZC 303c
(R); Ulna: UMZC 303c (L); UMZC 303c (L); UMZC 303c (R); UMZC 303c (R); Pelvis: UMZC 303c; UMZC
303c; Femur: UMZC 303c (R); UMZC 303c (R); Tibiotarsus: UMZC 303c (R); UMZC 303c (L); Tarsometatarsus:
UMZC 303c (L); UMZC 303c (R).
Type locality: Rodrigues, Mascarene Islands.
Distribution: Rodrigues, Mascarene Islands.
Etymology: After François Leguat (1637–1735), a French Huguenot and naturalist, who was marooned on
Rodrigues from 16 May 1691 to 21 May 1693, and who wrote so comprehensively about the original fauna of the
island.
Referred fossil material: Unassociated material, collected from specified or unspecified caves. Many speci-
mens are bulk numbered, so duplicate numbers exist for the same or different skeletal elements: Cranium: NHMUK
S/1978.14.46; Rostrum: NHMUK u/r (Caverne Bambara; Mandible: FLMR 10 (Caverne Poule Rouge); NHMUK u/
r (Caverne Bambara); Sternum: FLMR 12; NHMUK A1446; NHMUK A1446; NHMUK u/r; Scapula: NHMUK u/r
(L); Coracoid: NHMUK u/r (R); NHMUK u/r (Caverne Gastonia); Humerus: FLMR 10 (R) (Caverne Poule Rouge);
FLMR 12 (L) (Caverne Poule Rouge); Radius: FLMR 561 (R) (Caverne l’Affouche); NHMUK u/r (L) (Caverne
l’Affouche); NHMUK A1445 (L); NHMUK A1445 (R); NHMUK u/r (Ld) (Caverne l’Affouche); NHMUK u/r (R)
(Caverne Tortue); NHMUK u/r (L) (Caverne l’Affouche); Ulna: FLMR 10 (Lp) (Caverne Poule Rouge); Pelvis:
NHMUK A1447; NHMUK A1447; NHMUK A1447; NHMUK (Caverne Poule Rouge); Femur: FLMR 10 (R) (Cav-
erne Poule Rouge); FLMR 145 (Rp) (Caverne Tortue); FLMR 561 (R) (Caverne l’Affouche); MHNH MAD6477
(R); NHMUK 1448 (L); NHMUK 1448 (R); NHMUK 1448 (R); NHMUK 1448 (L); NHMUK 1448 (L); NHMUK
1448 (L); NHMUK 1448 (R); NHMUK 1448 (Ld); NHMUK 1448 (Ld); NHMUK 1448 (Ls); NHMUK u/c (Rd);
NHMUK u/c (L); NHMUK u/c (R); Tibiotarsus: FLMR 10 (Rd) (Caverne Poule Rouge); NHMUK S/1978.14.45
(Lp); NHMUK u/r (L); NHMUK u/r (R) (Caverne l’Affouche); NHMUK u/r (Rd); NHMUK u/r (R) (Caverne Poule
Rouge, associated with following); NHMUK u/r (Lp) (Caverne Poule Rouge, associated with previous); NHMUK
u/r (L) (Caverne Bambara); NHMUK A1449 (L); NHMUK A1449 (R); NHMUK A1449 (R); NHMUK A1449
(R); NHMUK A1449 (R); NHMUK A1449 (L); NHMUK A1449 (L); NHMUK A1449 (R); NHMUK A1449 (L);
NHMUK A1449 (R); NHMUK A1449 (Rd); NHMUK A1449 (Rd); NHMUK A1449 (Ld); NHMUK u/r (Lp);
Fibula: FLMR 607 (Caverne Poule Rouge) (R); FLMR 572 (R); NHMUK u/r (L); Tarsometatarsus: FLMR 10 (L)
(Caverne Poule Rouge); FLMR 10 (Lp) (Caverne Poule Rouge); FLMR 287 (Rp); NHMUK 1450 (L); NHMUK
1450 (R); NHMUK 1450 (R); NHMUK 1450 (L); NHMUK 1450 (R); NHMUK 1450 (L); NHMUK 1450 (L);
NHMUK 1450 (R); NHMUK 1450 (L); NHMUK 1450 (L); NHMUK u/r (Rp) (Caverne l’Affouche).
Associated individual: cranium; pelvis; tibiotarsus (R); tarsometatarsus (L), FLMR 609, collected by author
from Caverne Bambara at 300 mm depth in fining upwards calcarenite sand to silt, and found with a cranium of
Rodrigues Bulbul Hypsipetes cowlesi Hume, 2015, and associated dentaries of Phelsuma edwardnewtoni Vinson &
Vinson, 1969.
Second associated individual (Fig. 22): cranium; rostrum; ulna (L), carpometacarpus (R); pelvis; femur (R),
(L); tibiotarsus (R), (L), FLMR 608, collected by the author from Caverne Poule Rouge at ~100 mm depth in
fining upwards calcarenite sand to silt. On the surface directly above was a complete, associated skeleton of the
extinct Rodrigues Domed Tortoise Cylindraspis peltastes Dumeril & Bibron, 1835, which included preservation
of the cartilaginous scutes (see Hume 2014b). Erythromachus was found together with an associated skeleton of
Rodrigues Night Heron Nycticorax megacephala (Milne-Edwards, 1874); and individual elements of Rodrigues
Starling Necropsar rodericanus Günther & Newton, 1879; Rodrigues Blue Pigeon Alectroenas payandeei Hume,
2011a; Rodrigues Turtle-dove Nesoenas rodericanus (Milne-Edwards, 1874); Rodrigues Night Gecko Phelsuma
gigas (Liénard, 1842); Rodrigues Day Gecko P. edwardnewtoni; and White-tailed Tropicbird Phaethon lepturus
Daudin, 1802.
The femur and radius FLMR 561 were found in calcarenite silt at a depth of ~200 mm under a large boulder in
the main chamber of Caverne l’Affouche, along with individual elements of Phaethon lepturus; Solitaire Pezophaps
solitarius (Gmelin, 1789); Rodrigues Parakeet Psittacula exsul (A. Newton, 1872); Rodrigues Owl Otus murivorus
(Milne-Edwards, 1874); Rodrigues Warbler Acrocephalus rodericanus A. Newton, 1865; Phelsuma edwardnewtoni
and Cylindraspis peltastes, including scutes (see Hume 2014c). FLMR 145 was collected on the surface of Cav-
erne Tortue, along with individual elements of an undescribed petrel Pterodroma sp. (see Hume 2017); Phaethon
lepturus; Pezophaps solitarius; Rodrigues Parrot Necropsittacus rodricanus (Milne-Edwards, 1867a); Nesoenas
rodericanus; Acrocephalus rodericanus; Phelsuma edwardnewtoni and Cylindraspis sp..
Revised diagnosis: As for genus.
Description and comparison: See under Aphanapteryx.
Remarks: The Rodrigues Rail was first reported by Francois Leguat (1708: 71) in 1691–93, who noted the
rail’s similar and equally fatal attraction to red cloth as that observed in Aphanapteryx:
Our Wood-hens are fat all the year round, and of a most delicate taste. Their Colour is always of a bright Gray,
and there’s very little difference in the plumage between the two sexes. They hide their Nests so well, that we cou’d
not find ‘em out, and consequently did not taste their Eggs. They have a Red List about their Eyes, their Beaks are
straight and pointed, near two Inches long, and red also. They cannot fly, their fat makes ‘em too heavy for it. If you
offer them anything that’s red, they will fly at you to catch it out of your Hand, and in the heat of the Combat, we
had an opportunity to take them with ease.
They were further described in a manuscript by the Réunionnais mariner Julien Tafforet (1625–26), who stated
(my translation from the original document):
There is another kind of bird, the size of a young hen, which has the beak and legs red. Its beak is roughly like
that of a curlew, except that it is a little deeper and not quite so long. Its plumage is mottled with white and grey.
They generally feed on the eggs of the land tortoises, which they take from the ground, and makes them so fat that
they often have difficulty running. They are very good to eat, and their fat is of a yellowish-red, which is excellent
for pains. They have small pinions [wings], without feathers, so they cannot fly; but on the contrary, they run very
well. Their cry is a continual whistling. When they see somebody pursuing them they produce another sort of noise
from their bodies, like that of a person who has hiccups and with the stomach tensed.
Ecology: According to Tafforet’s (1725–26) account Erythromachus was an opportunist predator, as it clearly
took advantage of the breeding season of terrestrial tortoises to feed on the eggs, and perhaps hatchling young. Two
species of tortoise, a saddleback Cylindraspis vosmaeri (Suckow, 1798), and a smaller domed species, C. peltastes,
once occurred on Rodrigues. A recently discovered former tortoise breeding area shows that, in the Plaine Corail at
least, tortoises breeding sites were concentrated into small areas, and each site contained the remains of vast num-
bers of eggs (Hume et al. in prep); a phalanx of Erythromachus was also discovered amongst the egg remains. The
rails grew extremely fat during this seasonal glut of food, but at other times of the year probably fed on terrestrial

snails and other invertebrates, as well as scavenging seabird breeding colonies. Like Aphanapteryx, Erythromachus
probably fed on the endemic terrestrial nemertean worm Geonemertes rodericana Gulliver, 1879 and the endemic
planarian worm Kontikia whartoni (Gulliver, 1879), by probing for them in leaf-litter or in rotting wood; G. roderi-
cana has since become extinct (Moore et al. 1995). As Leguat failed to discover the nest and eggs of Erythromachus
during his two-year stay on Rodrigues (Leguat 1708), the birds may have nested away from the easily accessible,
open forest that was typical of coastal and lowland areas (Pingré 1763; Strahm 1989), and more likely nesting deep
in the forested valleys or in the mountainous hills of the interior, constructing nests well concealed in ground vegeta-
tion, as do many other flightless rails (e.g. Ripley 1977).
Extinction: Erythromachus may have lived alongside black rats for at least a century, but unlike on Mauritius
the exact time of their arrival is unknown. It is unlikely that they became established by Arab traders, as they did on
Mauritius (Hume 2013), or by the Portuguese who first encountered the island in 1507 (ibid). Visitors to Rodrigues
were extremely infrequent in the 17th century, and fewer still made a landfall (see Herbert 1663; Studer 1886;
North-Coombes 1971; Moree 2001; Hume 2003). It is more plausible that rats escaped during the poorly-docu-
mented marooning of Gerrit Andriesz, the helmsman of the ship Berckhout, who was abandoned on Rodrigues in
February 1644 for three months with 12 men (Heeringa 1895; North-Coombes 1971). Unlike Leguat and Tafforet,
however, Andriesz never wrote about his adventure, which is a great pity as some species of bird vulnerable to rat
predation, e.g. Alectroenas payandeei and Hypsipetes cowlesi, were never mentioned in the literature and may have
still been extant at this time.
Rats were certainly well established in 1691 by the time François Leguat and his companions had settled (Le-
guat 1708), and were also numerous during Tafforet’s visit in 1725–26 (Tafforet 1725–26); yet the rails remained
common, which was probably due to their aggressive nature. However, once the French settled in 1736 (Cheke
2010), and began a tortoise run to supply Mauritius, Réunion and passing ships with fresh tortoise meat, the rails ap-
pear to have declined rapidly (Cheke & Hume 2008). North-Coombes (1994) estimated that over 200,000 tortoises
were killed or exported alive from Rodrigues between 1736 and 1768, with around 10,000 taken annually for the
first two decades or so. It was only due to the collapse of the giant tortoise populations in the 1770s that the slaughter
ended, as it was then no longer viable to collect them (North-Coombes 1971; Cheke & Hume 2008). Consumption
by resident tortoise hunters no doubt took a heavy toll of rails, but it was the introduction of cats around c.1750 to
control rats that hastened their end (Cheke & Hume 2008; Hume 2017). Feral cats may have exterminated the birds
within a decade or so after their introduction, as Alexandre Guy Pingré, who arrived on May 28, 1761, to observe
the Transit of Venus (Pingré 1763; Nagapen 2004; Wulf 2012), noted that tortoises were still abundant, but the rails
had gone (my translation):
I heard said of neither gélinottes [Erythromachus leguati], nor butors [Rodrigues Night Heron], nor alouettes
[small waders], nor bécassines [snipe*]; there may have been some at the time of François Leguat, but they have
either retreated from their homes or, more likely, the races no longer survive, since the island has been populated
with cats.
*Leguat (1708; 160) named juvenile, nocturnal nesting shearwaters Puffinus sp. or petrels Pterodroma sp.
‘snipe’, but not in reference to true snipe Gallinago and Scolopax, which do not occur on the Mascarenes. Pingré
(1763) was presumably repeating Leguat’s terminology.
Genus Hypotaenidia Reichenbach, 1853
Buff-banded Rail Hypotaenidia philippensis (Linnaeus, 1766)
Rallus philippensis: Linnaeus, 1766, p.263

Hypotaenidia pectoralis (Lesson): E. Newton, 1888, p.552
Gallirallus philippensis: Olson, 1973, p.397
Hypotaenidia philippensis: Dickinson & Remsen, 2013, p. 155; del Hoyo & Collar, 2014, p.344
Remarks: There is a single record of Buff-banded Rail Hypotaenidia philippensis (Linnaeus, 1766), a male killed
at Champ des Mars, near Port Louis, in 1863 and held at UMZC, but the month is not known (Rountree et al. 1952:
180; Benson 1970: 171). E. Newton (1888: 552) listed it as accidental, and the genus has not been recorded again
in the Mascarenes, or anywhere else in the Indian Ocean (Safford & Hawkins 2013). Benson considered it to be
the race australis (now synonymised under H. p. mellori (Taylor & van Perlo 1998)), but noted that the individual
was heavily worn, with most of the feathers of the crown and rump missing and the wings incomplete. If a genuine
vagrant, it is a unique and extremely rare one, as the nearest source populations of H. philippensis are the Moluccas
and eastern Australia (Taylor & van Perlo 1998).
FIGURE 24. Selected post-cranial bones of Erythromachus leguati exhibiting extreme sexual size dimorphism in this species.
Clockwise from top: humerus, tarsometatarsus, tibiotarsus, and femur.
Genus Dryolimnas Sharpe, 1893
Dryolimnas Sharpe, 1893, p.26 = Rallus cuvieri Pucheran, 1845, p.279. Type by original designation
Etymology: From Greek drus, tree, and Modern Latin, limnas rail (> Gr. limnas of the marsh > limnç marsh)
Diagnosis of relevant skeletal anatomy: Rostrum (Fig. 25), short and straight; in dorsal view, os nasale nar-
rower than os premaxillare; in lateral view, narial opening long and approximately 45% of bill length; foramina
neurovascularia small and few in number; in ventral view, os premaxillare deeply excavated.
Humerus (Fig. 26): in caudal view, shaft dorsoventrally curved; caput humeri bulbous; tuberculum ventrale
prominent and directed caudad; incisura capitis deeply excavated; tuberculum dorsale prominent and directed lat-
erad; crista deltopectoralis short and weakly triangular; fossa pneumotricipitalis deeply excavated; crista bicipitalis
terminates in sharp angle to shaft; margo caudalis raised and extends distally to mid-shaft; processus flexorius

extends further distad than condylus dorsalis; indistinct processus supracondylaris dorsalis; fossa olecrani deeply
excavated; in cranial view, sulcus ligamentosa transversus shallow; sulcus n. coracobrachialis indistinct; fossa m.
brachialis deeply excavated; epicondylus ventralis prominent.
FIGURE 25. Comparison of crania and rostra of Dryolimnas cuvieri with D. chekei nov.sp., lateral view (left), dorsal view
(right). A., B. NHMUK 1882.2.28.2 Dryolimnas c. cuvieri u/s; C., D. UMZC 593.B u/s, I. UMZC 593.B u/s; †D. chekei u/s; E.,
F. NHMUK S/1989.38.1 D. c. aldabranus ♂; NHMUK 1910.4.8.1 G., H. †D. c. abbotti u/s. Scale bar = 10 mm.
Pelvis (Fig. 27): in dorsal aspect, ala preacetabularis ilii expanded proximally, lateral edges angular; crista
spinosa synsacri forms a single distinct ridge; proximal end of extremitas cranialis synsacri extends further craniad
than crista iliaca dorsalis; foramina intertransversariae large and oval-shaped; in lateral view, ala preacetabularis
deep with strongly arched crista iliaca dorsalis.
Femur (Fig. 28): proximal and distal ends incurved both posteriorly and internally, exhibiting two curvatures
in two different planes; in cranial aspect, crista trochanteris forms distinct, dorsally facing ridge; in caudal aspect,
deeply excavated medial to facies articularis acetabularis; condylus medialis and condylus lateralis connected with
distinct ridge.
Tibiotarsus (Fig. 29): robust, with wide and deep proximal and distal ends; in cranial aspect, crista cnemialis
distinct, directed cranio-dorsad; crista fibularis long and triangular distally; pons supratendineus wide; canalis ex-
tensorius deflected mediad; condylus lateralis larger than condylus medialis; sulcus m. fibularis deeply excavated;
in caudal aspect, trochlea cartilaginis tibialis deeply excavated.
Tarsometatarsus (Fig. 30): cotyla lateralis lower than cotyla medialis; retinaculum extensorium ossified; troch-
lea metatarsi II shorter than other trochlea and directed strongly plantad; in dorsal aspect, trochlea metatarsi III larg-
est with a deep sulcus extending onto facies dorsalis; foramen vasculare distale large and oval-shaped; on proximal
end, crista medialis flexoris digitorum longus present; sulci for tendon of musculus flexor perforatus digiti II and
musculus flexor hallucis longus present.
White-throated Rail Dryolimnas cuvieri (Pucheran, 1845)
Rallus cuvieri: Pucheran, 1845, p.279 (Mauritius)

Rallus gularis: Hartlaub, 1877, p.337; Milne-Edwards & Grandidier, 1879, p.569, pls.230b, 232; Guérin, 1942, 2(6) (auctorum
nec Horsfield, 1822, p.196)
Dryolimnas cuvieri: Sharpe, 1894, p.70
Dryolimnas cuvieri cuvieri: Sclater, 1924, p.101; Rountree et al. 1952; Dickinson & Remsen, 2013, p.155; del Hoyo & Collar,
2014, pp.344
Holotype: MNHN-ZO-2011-535 ♀ (Fig. 31). The holotype was described from a live specimen captured on Mauri-
tius in 1809 by Colonel M. Mathieu (Schlegel & Pollen 1868), who was attached to the naval artillery and supplied
an important collection of animals to the MNHN (Cheke & Hume 2008; Hume 2011a).
FIGURE 26. Comparison of humeri of Dryolimnas cuvieri and Gallinula chloropus with D. chekei nov. sp., caudal aspect
(above), cranial aspect (below). A. NHMUK S/1989.38.1 Dryolimnas c. aldabranus ♂; B. NHMUK 1910.4.8.1 †D. c. abbotti
u/s; C. UMZC 300AA4 †D. chekei u/s; D. NHMUK 1882.2.28.2 D. c. cuvieri u/s; E. NHMUK S/1977.23.1 Gallinula chloropus
u/s. Scale bar = 10 mm.

FIGURE 27. Comparison of pelves of Dryolimnas cuvieri and Gallinula chloropus with D. chekei nov. sp., dorsal view. A.
NHMUK S/1977.23.1 Gallinula chloropus u/s. B. NHMUK S/1989.38.1 Dryolimnas c. aldabranus ♂; C. UMZC 300AA5 †D.
chekei u/s; D. NHMUK 1882.2.28.2 D. c. cuvieri u/s; E. NHMUK 1910.4.8.1 †D. c. abbotti u/s. Scale bar = 10 mm.
FIGURE 28. Comparison of femora of Dryolimnas cuvieri with D. chekei nov. sp., caudal aspect. A. UMZC 300AA3 †Dry-
olimnas chekei u/s; B. NHMUK 1910.4.8.1 †D. c. abbotti u/s; C. NHMUK S/1989.38.1 D. c. aldabranus ♂; D. NHMUK
1882.2.28.2 D. c. cuvieri u/s. Scale bar = 10 mm.

Type locality: Mauritius, Mascarene Islands (vagrant).
Distribution: Madagascar, Mayotte, (Comoros), Mauritius, Aldabra Atoll (ssp aldabranus) and Assumption
Island (ssp abbotti).
Etymology: After Léopold Chrêtien Frédéric Dagobert Baron Cuvier (1769–1832), French anatomist, and bet-
ter known by his assumed name, Georges Cuvier (Jobling 2009).
Diagnosis: In skeleton, as for genus.
Description of skin (Fig. 31): In nominate, head, neck and breast vinous-chestnut; chin and throat white;
mantle and scapulars to upper-tail coverts greenish-olive, with blackish-brown streaks on lower mantle, scapulars
and back; tail olive-brown; flanks, belly and thighs greenish olive-brown; hind flanks and belly buffy-white barred
black; median undertail-coverts dark brown barred white and lateral undertail-coverts white; wings olive-brown
tinged greenish, with darker central streak on tertials; rostrum blackish, mandible red with blackish tip; iris reddish-
orange to reddish-brown; legs and feet greenish-grey or brownish-grey. Sexes alike. In D. c. aldabranus, smaller,

with much reduced wing than nominate (Table 2); top and sides of head more olive-brown; upperparts slightly paler,
streaking reduced or absent; wings dull grey-brown; lower flanks and belly paler with less distinct white barring;
bill blackish-brown, with pink on lower half of rostrum and basal third of mandible; iris orange-brown; legs and feet
dark brown or black. Sexes alike. In D. c. abbotti, smaller than nominate, with wing intermediate in length between
nominate and D. c. aldabranus (Table 3); upperparts paler, more greenish-olive; dark streaks variable, narrower,
fainter or absent; white on throat more extensive, reaching upper breast; broader white barring on underside; bill
black, basal third of mandible bright red; legs and feet dark brown.
Remarks: The White-throated Rail is divided into three subspecies, nominate and volant D. c. cuvieri, which is
found widely on Madagascar and as a vagrant on both Mayotte, Comoros, and Mauritius (Safford & Hawkins 2013),
while flightless and poorly volant forms, D. c. aldabranus and D. c. abbotti are endemic to Aldabra Atoll and As-
sumption Island respectively (Fig. 1); D. c. abbotti is now extinct (Safford & Hawkins 2013; Hume 2017). The ho-
lotype of the nominate collected on Mauritius is undifferentiated in colouration and wing and tarsus measurements
from those on Madagascar, which is the normal distribution of this species (Ripley 1977; Taylor & van Perlo 1998).
Dryolimnas cuvieri was considered to be either a former resident on Mauritius, or a vagrant, despite the holotype
having enlarged ovaries (Rountree et al. 1952; Cheke 1987; Cowles 1987), but no specimens have been recorded on
Mauritius since the type was collected. The discovery of subfossil remains have confirmed that an endemic, poorly
volant or flightless species once occurred on the island (see below).
TABLE 2. Wing chord and tarsus measurements (mm) of Dryolimnas cuvieri. For each entry, and where available, the
first is mean, second in parentheses range, and third standard deviation.
D. c. cuvieri Wing chord (mm) Tarsus
♂ (n = 9) 150.8 (141.0–163.0) SD =7.06 45.0 (39.1–49.3) SD=3.54
♀ (n = 16) 144.1 (139.0–153.0) SD=4.50 43.8 (41–47.3) SD=1.88
u/s (n = 8) 147.6 (142.0–158.0) SD=6.63 43.7 (40–47.5) SD=2.69
Holotype ♀ 140.00 48.3
D. c. aldabranus
♂ (n = 3) 119.0 (117.0–122.0) SD=2.64 39.5 (37.8–41.9) SD=2.10
♀ (n = 3) 116.0 (116.0–120.0) SD=4.00 40.1 (39.1–41.4) SD=1.16
u/s (n = 4) 114.7 (107.0–118.0) SD=5.18 40.7 (39.0–43.1) SD=1.71
D. c. abbotti
♂ (n = 3) 135.0 (133.0–137.0) SD=2.82 39.2 (39.2–40.0) SD=0.46
♂ Imm. (n = 1) 129.0 39.9
♀ (n = 1) 131.0 38.6
TABLE 3. Invasive probable rail predators and their dates of introduction

Species Mauritius Réunion Rodrigues Impact factor on Rallidae
Black Rat Rattus rattus 1369–1413 c.1672 c.1640 Minor
Cat Felis sylvestris c.1680 c.1700 c.1750 Major
Pig Sus scrofa 1610 c.1630 c.1695 Minor
Crab-eating Macaque Macaca fascicularis c.1607 - - Minor
†Cheke’s Wood Rail Dryolimnas chekei sp. nov
Dryolimnas cuvieri: Cowles, 1987, p.96; Cheke, 1987, p.37

Sauzier’s Wood Rail Cheke & Hume, 2008, p.38; Cheke, 2013a, p.2, 2013b, p.6
Dryolimnas sp.: Hume & Prŷs-Jones, 2005, p.89; Cheke & Hume, 2008, p.128; Hume & Walters, 2012, p.218; Cheke, 2013b,
p.6; Hume, 2013, p.218, 2014a, p.37, 2017, p.115; Safford & Hawkins, 2013, pp.45,348,358
Holotype: Tarsometatarsus: UMZC 300AA5 (R) (Fig. 30B) collected at the Mare aux Songes, south-east Mau-
ritius, by Théodore Sauzier in 1889–91. A. Newton referred it to Gallinula [chloropus] pyrrhorrhoa in error (see
Cowles 1987).
Type locality: Mare aux Songes, Mauritius, Mascarene Islands.
Distribution: Mauritius, Mascarene Islands.
Etymology: Named in honour of my friend and colleague, Anthony Cheke, in recognition of his unparalleled
lifelong contribution to Mascarene ecological history.
Paratypes: Collected at the Mare aux Songes by Théodore Sauzier in 1889–91. Mis-identified as Gallinula
[chloropus] pyrrhorrhoa by A. Newton (see Cowles 1987). Pelvis: UMZC 300AA6; tibiotarsus: UMZC 300AA3
(R); UMZC 300AA4 (L).
Referred fossil material: All collected at the Vallée des Prtres, Moka Range, central-west Mauritius. Some of
the following specimens are bulk-registered, so a few numbers are duplicated. Rostrum: UMZC 593.B; UMZC 593.
B; humerus: UMZC 300AA1 (L); UMZC 300AA2 (R) (these may be associated); MNHN Mad7136 (R); MNHN
MAD7136 (Rd); femur: MNHN MAD8867 (Lp); MNHN MAD7159 (Rp); tibiotarsus: MNHN MAD8867 (Lp);
MNHN MAD7159 (Rp); tarsometatarsus: UMZC 593b (R).
Diagnosis: Rostrum (Fig. 25): short and slightly decurved; in dorsal view, os nasale narrower than os premaxil-
lare; in lateral view, narial opening long and oval; foramina neurovascularia small and few in number; in ventral
view, os premaxillare deeply excavated.
Humerus (Fig. 26): proximal and distal ends reduced; in caudal view, shaft dorsoventrally curved; caput humeri
bulbous; tuberculum ventrale prominent and directed caudad; incisura capitis deeply excavated; tuberculum dorsale
prominent and directed laterad; crista deltopectoralis short and directed sharply cranially; fossa pneumotricipitalis
deeply excavated; crista bicipitalis terminates in sharp angle to shaft; margo caudalis raised and extends distally to
mid-shaft; processus flexorius extends further distad than condylus dorsalis; indistinct processus supracondylaris
dorsalis; fossa olecrani deeply excavated; in cranial view, sulcus ligamentosa transversus shallow; sulcus n. coraco-
brachialis indistinct; fossa m. brachialis deeply excavated; epicondylus ventralis prominent.
Pelvis (Fig. 27): in dorsal aspect, ala preacetabularis ilii expanded proximally, lateral edges triangular; crista
spinosa synsacri forms a single distinct ridge; proximal end of extremitas cranialis synsacri extends further craniad
than crista iliaca dorsalis; foramina intertransversariae large and oval-shaped; in lateral view, ala preacetabularis
deep with strongly arched crista iliaca dorsalis.
Femur (Fig. 28): proximal and distal ends incurved both posteriorly and internally, exhibiting two curvatures in
two different planes; in caudal aspect, deeply excavated medial to facies articularis acetabularis; condylus medialis
and condylus lateralis connected with distinct ridge.
Tibiotarsus (Fig. 25): robust, with wide and deep proximal and distal ends; in cranial aspect, pons supratendin-
eus wide; canalis extensorius deflected mediad; condylus lateralis larger than condylus medialis; sulcus m. fibularis
deeply excavated.
Tarsometatarsus (Fig. 29): extremely robust with widely spaced trochlea; in dorsal aspect, trochlea metatarsi III
largest with a deep sulcus extending on to facies dorsalis; foramen vasculare distale large and oval-shaped; in plan-
tar aspect, on proximal end, crista medialis flexoris digitorum longus present; sulci for tendon of musculus flexor
perforatus digiti II and musculus flexor hallucis longus present.
Size and colouration: The two flightless/poorly volant subspecies D. c. aldabranus and D. c. abbotti were
smaller in the skin than nominate, with a notably shorter humerus and tarsometatarsus (Figs. 26, 29), which is in
direct contrast to many flightless rails. This can probably be explained by the extremely reduced land surface area
of Aldabra and Assumption with corresponding restricted food resources (Hume & Heinen in prep). Volant D. c. cu-
vieri measures 30–33 cm in total length and has a body mass of up 276 g in the male, 223 g in female (Ripley 1977;
Taylor & van Perlo 1998), whereas D. c. aldabranus is smaller and has a body mass of up to 218 g in male, 223 g in
female; D. c. abbotti was intermediate in total length between nominate and Aldabra subspecies, but body mass is
unknown. The endemic, flightless Dryolimnas of Mauritius and Réunion were larger and heavier than recent Dryo-
limnas, possibly reaching 35+ cm in length and weighing up to 300 g, or even heavier. They also differ markedly in
proportions and characters of the skeleton, so likely have been isolated for a long period of time; hence the specific
rather than subspecific status applied here. Furthermore, and based on the total length of the tibiotarsus, D. chekei
exhibited the greatest size variation in all Rallidae used in this study (n = 2, TL 78.1–90.0, SD 8.41) (Appendix 2,
Table 13), which was almost certainly due to extreme sexual dimorphism (see Livezey 2003).
Although speculative, it is likely that Cheke’s Wood Rail and Réunion Wood rail D. augusti (see below) were
much duller in colouration than Dryolimnas cuvieri. This would be especially prevalent in worn individuals when

the contour feathers and remiges, due to flightlessness and movement through dense vegetation, would render the
plumage much abraded and more uniform in colour. This concurs with worn specimens of D. c. aldabranus (Penny
& Diamond 1971), and is also seen in the recently extinct, flightless Wake Island Rail Hypotaenidia wakensis
Rothschild, 1903 (see frontispiece in Olson & Rauzen 2011). No unequivocal account was made of D. chekei on
Mauritius, but a worn individual of D. chekei would be much harder to differentiate from Aphanapteryx in plumage
pattern, but as D. chekei was also much smaller, it may also have been considered a juvenile and thus overlooked
(see also Cheke 2013a). It is also interesting to note that Dryolimnas c. aldabranus aggressively responds to the
waving of a piece of cloth (Gillham 2000), as did Aphanapteryx and Erythromachus; if D. chekei also had this be-
haviour trait, it would further confuse its identification to an untrained eye.
Description and comparison: Dryolimnas chekei differs from other species of Dryolimnas by the following
characters:
Rostrum, os premaxillare, equal in size with D. c. cuvieri, slightly larger than D. c. abbotti and D. c. aldabranus;
otherwise similar.
FIGURE 29. Comparison of tibiotarsi of Dryolimnas cuvieri and Gallinula chloropus with D. chekei nov. sp., cranial aspect.
A. C. NHMUK S/1977.23.1 Gallinula chloropus u/s. B. UMZC 300AA4 u/s, D. UMZC 300AA3 †Dryolimnas chekei u/s; E.
NHMUK 1882.2.28.2 D. c. cuvieri u/s; F. NHMUK S/1989.38.1 D. c. aldabranus ♂; G. NHMUK 1910.4.8.1 †D. c. abbotti
Humerus, differs from D. c. cuvieri in being smaller and less robust; in caudal view, tuberculum ventrale com-
paratively larger; crista bicipitalis and crista deltopectoralis extend less distad and not at acute right angle with shaft;
incisura capitis wider and more deeply excavated; caput humeri smaller, less inflated; crus dorsale fossae raised;
fossa pneumotricipitalis shallower; in cranial aspect, impressio coracobrachialis less deeply excavated; fossa m.
brachialis comparatively larger and deeper set, and extends further proximad; from D. c. abbotti, in being equal in
total length or slightly larger, but comparatively straighter and more robust in shaft; distal end more robust; in caudal
view, tuberculum ventrale larger; crista deltopectoralis and crista bicipitalis more reduced, extending less distad;
incisura capitis comparatively narrower; tuberculum dorsale smaller; processus supracondylaris dorsalis indistinct;
in cranial view, fossa m. brachialis comparatively larger and deeper set, and extends further proximad; from D. c.
aldabranus in being overall much larger and more robust; in caudal view, crista deltopectoralis more angular, less
rounded; crus dorsale fossae extremely reduced; in cranial aspect, fossa m. brachialis comparatively larger and
deeper set; equal in total length with D. augusti, but medial fossa pneumotricipitalis deeper.
Pelvis, D. chekei larger than all Dryolimnas (D. augusti not available for comparison), being particularly robust
at proximal end; in lateral aspect, ala preacetabularis ilii deeper and more arched than other Dryolimnas; foramen
intervertebrale extend further ventrad; foramina intertransversariae smaller and distinctly open (similar in D. c. ab-
botti).
Femur, overall smaller in total length, less robust and straighter than D. c. cuvieri and D. augusti; larger than D.
c. aldabranus and D. C. abbotti, otherwise similar;
Tibiotarsus, larger and more robust than D. c. aldabranus and D. c. abbotti; but less so than D. augusti; in
cranial aspect, crista fibularis extends further distad than in D. c. cuvieri, D. c. abbotti and D. c. aldabranus; sulcus
extensorius deeper and extends more proximad; in proximal articular surface, crista cnemialis cranialis larger and
extends further caudad than D. augusti; otherwise morphologically similar.
Tarsometatarsus, more similar to D. c. abbotti and D. c. aldabranus than to D. c. cuvieri, but much more robust;
in dorsal aspect, retinaculi extensorii not connected by an ossified bridge (present in D. c. cuvieri and D. c. abbotti);
trochleae more splayed (similar to D. augusti); foramen vasculare distale larger and deeper set; in plantar aspect,
fossa metatarsi I oval-shaped, deeper, longer and closer proximad to trochlea metatarsi II; crista medialis hypotarsi
central, not deflected laterad; on proximal end, sulci for tendons of musculus flexor perforatus digiti II and musculus
flexor hallucis longus more deeply excavated.
Differs from Aphanapteryx and Fulica newtonii by much smaller size (the latter on proximal end has a canal
for tendon of musculus flexor digitorum longus; and from similar-sized Gallinula chloropus and Porphyrio alleni
by the following characters:
Humerus, in Gallinula, overall much more robust, shaft straighter; in caudal view, incisura capitis connected by
small ridge to tuberculum ventrale and caput humeri; crista deltopectoralis less angular; tuberculum ventrale more
square-shaped, less pointed; crus dorsale fossae less prominent; processus supracondylaris dorsalis smaller; epicon-
dylus ventralis directed less medially; in Porphyrio, proximal and distal ends more robust; in caudal view, crista
bicipitalis extends less distad; incisura capitis larger and deeper; tuberculum ventrale larger; processus flexorius
directed less medially, with no distinct sulcus on internal edge; in dorsal view, crista bicipitalis distinctly square-
shaped, extends more ventrally; in cranial aspect, fossa m. brachialis shallower, extending less proximad; condylus
ventralis and condylus dorsalis larger.
Pelvis, in Gallinula, ala preacetabularis ilii more expanded distally and directed more cranially; foramina inter-
transversariae small and indistinct (large and oval shaped in Dryolimnas); ala postacetabularis ilii less expanded; in
lateral aspect, crista iliaca dorsalis more arched and deeper; foramen ilioischiadicum much greater in size than fora-
men acetabuli; in dorsal view, ala postacetabularis extend less caudally and indented; foramina intertransversariae
small (large and distinctly open cranially in Dryolimnas); foramen extend much further cranially either side of crista
iliaca dorsalis; in Porphyrio, in dorsal view, ala preacetabularis ilii more expanded distally and directed more crani-
ally; foramina extremely reduced either side of crista iliaca dorsalis.
Femur, longer, but shaft straighter in Gallinula and Porphyrio and comparatively less robust, particularly in
mid-shaft width and depth; in cranial aspect, sulcus patellaris more deeply excavated; otherwise similar.
Tibiotarsus, much longer and more robust in Porphyrio and Gallinula; in cranial aspect, epicondylus medialis
more pronounced; pons supratendineus situated further proximad and sulcus extensorius narrower, more deeply
excavated, and situated more medially; in caudal aspect, crista fibularis extends less distad.
Tarsometatarsus, in Gallinula, longer with proximal and distal ends more robust, trochleae less splayed; reti-
naculi extensorii tarsometatarsi more ossified in some individuals; in dorsal aspect, fossa infracotylaris dorsalis less
deeply excavated; foramen vasculare distale smaller, situated less distad; in plantar aspect, crista medialis hypotarsi
situated centrad, not deflected laterad, extending further distad; in medial view, trochlea metatarsi II more rounded;
on proximal end, canal for tendon of musculus flexor digitorum longus; sulcus for tendon of musculus flexor perfo-
ratus digiti II and sulcus for tendon of musculus flexor hallucis longus more deeply excavated; in Porphyrio, longer
and gracile; sulcus extensorius deeply excavated extending over 50% along shaft; trochlea metatarsi widely splayed;

trochlea metatarsi II extends further distad; impressiones retinaculi extensorius not ossified; in plantar aspect, crista
medialis hypotarsi extends less distad; sulcus flexorius deeply excavated; in medial view, trochlea metatarsi II more
rounded; on proximal end, deeply incised crista medialis flexoris digitorum longus; sulci for tendons of musculus
flexor perforatus digiti II and musculus flexor hallucis longus more deeply excavated.
Remarks: Subfossil remains of D. chekei were originally referred to nominate Dryolimnas cuvieri by Cowles
(1987). A more detailed analysis of this material, including specimens not examined by Cowles (Hume 2013, 2017;
this paper), has shown that they represent an undescribed, poorly volant or totally flightless Dryolimnas collected
in 1889–91 by Théodor Sauzier at the Mare aux Songes in the south-east, and by Etienne Thirioux sometime before
April 30, 1908, in the Vallée des Prêtres, Moka Range, in the central-west (details from registration) (Fig. 2). D.
chekei is morphometrically most similar to the poorly volant D. augusti, and also to the flightless D. c. abbotti, but
noticeably much larger in pelvis, tibiotarsus and tarsometatarsus than the latter (Figs. 27, 29, 30; Appendix 2), sug-
gesting a greater degree of flightlessness.
FIGURE 30. Comparison of tarsometatarsi of Dryolimnas cuvieri and Gallinula chloropus with D. chekei nov. sp., (above) dor-
sal aspect, (below) plantar aspect. A. UMZC 593B u/s, B. UMZC 300AA5 †Dryolimnas chekei u/s; C. NHMUK 1897.5.10.47 D.
c. cuvieri u/s; D. NHMUK 1910.4.8.1 †D. c. abbotti u/s; E. NHMUK S/1989.38.1 D. c. aldabranus ♂; F. NHMUK S/1977.23.1
Gallinula chloropus u/s. Scale bar = 10 mm.
Ecology: On Aldabra, Dryolimnas feeds on a wide range of animal matter, including insects and other inverte-
brates in litter, marine and terrestrial molluscs, catching small, or scavenging large, land crabs, and consumes flies,
maggots and eggs associated with giant tortoise carcasses (Penny & Diamond 1971; Safford & Hawkins 2013).
Mauritius once harboured high densities of giant tortoises and land crabs, and a diverse terrestrial gastropod fauna
(Hume 2005; 2014c; Griffiths & Florens 2006), so Cheke’s Wood Rail almost certainly had a similar feeding niche,
and may have also taken advantage of seasonal breeding sites of giant tortoises and invertebrates associated with
seabird colonies. D. c. aldabranus also actively digs up and devours hatchling turtles and their eggs (Frith 1977),
so D. chekei may have also preyed on the progeny of terrestrial and marine tortoises. The extreme presumed sexual
dimorphism exhibited in D. chekei strongly suggests intersexual niche partitioning, and that the sexes foraged on
different-sized food items. This may have also reduced competition with Aphanapteryx. A mutualism reputedly ex-
ists between D. c. aldabranus and Aldabra Giant Tortoise Aldabrachelys gigantea (Schweigger, 1812), whereby the
tortoises respond to the rails pecking their soft underparts by raising the body (standing posture), allowing the birds
access to ectoparasites and other invertebrates on the skin (Huxley 1979; Safford & Hawkins 2013). Two sympatric
giant tortoise species, Cylindraspis triserrata Günther, 1873, and C. inepta Günther, 1873, once occurred on Mau-
ritius, so it is possible that D. chekei had a similar tortoise/rail mutualistic relationship.
Extinction: It is likely that D. chekei survived alongside rats on Mauritius, as does D. c. aldabranus on Aldabra,
but disappeared rapidly after the introduction of cats. If this was the case, and like all of the other endemic Mauritian
rails, Cheke’s Wood Rail probably became extinct due to feral cat predation by the end of the 17th century.
†Réunion Wood Rail Dryolimnas augusti Mourer-Chauviré et al. 1999
Ralles de bois Dubois 1674. p.172; Berlioz, 1946, p.5; Oliver, 1898, p.78
Dryolimnas cuvieri Probst, 1997, p.143
Dryolimnas augusti: Mourer-Chauviré et al. 1999, p.21 (Réunion); Mourer-Chauviré et al. 2006, p.43; Hume et al. 2006, p.16;
Cheke & Hume, 2008, p.128; Hume & Walters, 2012, p.99; Cheke, 2013b, p.13; Hume, 2013, p.218, 2014a, p.37, 2017,
p.115; Safford & Hawkins, 2013, pp.45,348,358; del Hoyo & Collar, 2014, pp.342,772
Holotype: Tarsometatarsi MHNR 13 (R), MHNR 14 (L), associated.

Type locality: Caverne de la Tortue, Saint-Paul, Réunion, Mascarene Islands.
Distribution: Réunion, Mascarene Islands.
Etymology: The scientific name commemorates French poet Auguste de Villèle (1858–1943), whose interest
in the history of Réunion and hospitality there made it possible for numerous naturalists to discover and explore the
caves of Réunion.
Paratypes: Fragment of left mandible, MHNR CT 1; sacrum, MHNR CT2; coracoid, MHNR CT3 (Ls); hu-
merus, MHNR CT4 (Rp); MHNR CT5 (Ld); ulna, MHNR CT6 (R); femur, MHNR CT7 (Lp); MHNR CT8 (Ld);
MHNR CT9 (Rd), MHNR CT10 (Rd); tibiotarsus, MHNR CT11 (Rp); MHNR CT12 (Ld); pedal phalanx 1 of digit
II, MHNR CT15; pedal phalanx 2 of digit II, MHNR CT16; MHNR CT17; pedal phalanx 1 of digit III, MHNR
CT18; pedal phalanx 2 of digit III, MHNR CT19; MHNR CT20; pedal phalanx 3 of digit III, MHNR CT21; pedal
phalanx 1 of digit IV, MHNR CT22; vertebrae, MHNR CT 23; MHNR CT 24; MHNR CT 25; MHNR CT 26;
MHNR CT 27.
Diagnosis: See Mourer-Chauviré et al. (1999: 22). Much larger and with stouter tarsometatarsus than recent
Dryolimnas species, and generally slightly longer and more robust in all elements than D. chekei (see Appendix
2).
Description and comparison: See Mourer-Chauviré et al. (1999: 21–24).
Remarks: The Réunion Wood Rail was a large bird, much larger than D. cuvieri, with particularly robust legs.
Wing elements are lacking, but the stout limbs suggest that it was flightless. Being slightly larger than D. chekei, it
may have reached 35–38 cm in length, with a body mass of up to 300–350 g.
Dubois (1674) mentions seeing a ‘Râle des Bois’ (wood rail) in 1671–72, which almost certainly refers to this
species (Mourer-Chauviré et al. 1999), and suggests that it was an inhabitant of forests. It is also likely that it was re-
stricted to lowland forests, because Dubois specifically mentioned the oiseaux bleu (see below) as the only montane
rallid. D. augusti probably had the same habits as D. chekei on Mauritius, nesting in dense vegetation deep in for-

ests, and feeding on insects, small crabs terrestrial molluscs, and perhaps tortoise eggs and hatchlings. It may have
also had a mutualistic relationship with the extinct Réunion Giant Tortoise Cylindraspis indica Schneider, 1783.
Dubois’s account confirms the Réunion Wood Rail’s survival at least until the early 1670s, but it was not men-
tioned again. It probably became extinct by the end of the 17th century, which seems extraordinary considering the
complex habitats on an island the size of Réunion. However, cats were introduced in the 1680s and had gone feral
by 1703 (Cheke & Hume 2008), so if the Réunion Wood Rail was confined to the lowlands, which appears to be the
case, it would have been rapidly exterminated by these voracious predators.
Genus Porphyrio Brisson, 1760
†Réunion Gallinule (Oiseaux Bleu) ‘Porphyrio caerulescens’ (Sélys-Longchamps, 1848)
Oyseaux bleus Dubois, 1674, p.170; Oustalet, 1897, p.99; Oliver, 1898, p.77; Cheke & Hume, 2008, p.43
Oiseaux bleu La Roque, [1701–10] (in Lougnon, 1992, p.209); Le Gentil, 1727, p.103
Oiseaux bleuff Feuilley, [1705] (in Lougnon 2005, p.128)
Gros oiseau Brown, 1773, p.332
Apterornis caerulescens Sélys-Longchamps, 1848, p.294;
Cyanornis erythrorhynchus Bonaparte, 1854, p.3; Hachisuka, 1953, p.155; Hume, 2013, p.219, 2014a, p.37
Porphyrio (Notornis?) coerulescens Schlegel, 1857, p.142, 1858, p.379; 1866a, p.167; Pitot, 1905, p.372 (orthographic error)
Porphyrio caerulescens Sharpe, 1894, p.193; Olson, 1977, p.365; Hume et al. 2006, p.15; Hume & Walters, 2012, p.113; Cheke,
2013b, p.13; Hume, 2013, p.219, 2017, p.131; Safford & Hawkins, 2013, p.45; del Hoyo & Collar, 2014, pp.352,773
Notornis sp. Oustalet, 1897, p.99
Porphyrio Notornis Pitot, 1914, p.89
Porphyrio madagascariensis Berlioz, 1946, p.8; Barré & Barau, 1982, p.34
Porphyrio porphyrio Barré et al. 1996, p.36
Cyanornis coerulescens Hachisuka, 1953, p.155; Hume, 2013, p.219; 2014a, p.37 (orthographic error)
Porphyrio sp. Strickland (in Strickland & Melville, 1848, p.59); Cheke & Hume, 2008, p.128, pl.8; Safford & Hawkins, 2013,
p.348
Apterornis coerulescens Rothschild, 1907a, p.145, pl.32, 1907b, p.198; Fuller, 1987, p.241; 2001, p.386 (orthographic error)
Porphyrio coerulescens Cheke, 1987, p.37; Adams et al. 2003, p.344 (orthographic error)
Cyanornis (?=Porphyrio) caerulescens Mourer-Chauviré et al. 2006, p.43

Measurements: None available.
Type locality: Plaine des Cafres, Réunion, Mascarene Islands.
Distribution: Réunion, Mascarene Islands.
Etymology: Inferred L. caerulescens, bluish, in reference to the all blue plumage colouration.
Remarks: Perhaps the most enigmatic of all rails, the Réunion Gallinule or Oiseaux bleu once occurred on
Réunion Island, but as yet not a shred of physical evidence of any kind has been found to resolve its taxonomy. There
can be no doubt that it was a large, terrestrial Porphyrio derivative (Olson 1977; Mourer-Chauviré et al. 1999); the
all blue colouration is only found in Porphyrio and not Fulica or Gallinula, and possessed characters—larger size
and more robust legs—usually associated with reduced powers of flight. There has been some disagreement regard-
ing the size of the Réunion Gallinule (see Cheke & Hume 2008), as Dubois (1674) stated that they were ‘the size of
solitaires’, and Feuilley (1705) that that they were ‘as large as a capon’. However, the Réunion Solitaire Threskiornis
solitarius (Sélys-Longchamps, 1848) would have been no more than 68 cm in total (the size of the African Sacred
Ibis Threskiornis aethiopicus (Latham, 1790) including tail), and capons (or domestic chickens) were approximately
the size of the ancestral Red Jungle Fowl Gallus gallus (Linnaeus, 1758), up to 70 cm in total length including the
long tail, so there is no real size discrepancy between these two contemporary accounts (Hume 2017).
The Réunion Gallinule was mentioned on a number of occasions by trustworthy observers, being considered
good game. Unlike Purple Gallinule, it could easily be caught and killed with sticks, despite being a fast runner,
and it was also reluctant to fly (Cheke & Hume 2008). The Réunion Gallinule appears to have been restricted to the
mountains, at least latterly, particularly on a montane plateau called the Plaine des Cafres at around 1,600–1,800 m
in altitude in south-central Réunion (Feuilley 1705; Cheke & Hume 2008) (Fig. 3). The area comprises open wood-
land in subalpine forest steppe with marshy pools (Cheke & Hume 2008). Very little was recorded about its ecology.
Dubois (1674: 170), while staying on Réunion in 1671–72, gave the first description (my translation):
Oyseaux bleus: As big as the solitaires [Threskiornis solitarius]; their plumage is entirely blue, the beak and the
feet red and made like those of hens; they do not fly, but run extremely quickly, so that a dog has difficulty catching
them in a chase; they are very good [to eat].
Feuilley (1705) in 1704 described them further (translation from Cheke & Hume 2008: 128):
The Oiseaux bleuff live in the plaines on top of the mountains, and especially on the Plaine des Cafres. They
are the size of a large capon, blue in colour. Those that are old are worth nothing to eat because they are so tough,
but when they are young they are excellent. Hunting them is not difficult because one kills them with sticks or with
stones.
FIGURE 31. Comparison of the skins, dorsal view above, ventral view below, of Dryolimnas cuvieri. From left to right: D. c.
cuvieri MNHN-Zo-2011-535 ♀ holotype; D. c. cuvieri NHMUK 1931.8.18.1000 ♂; D. c. cuvieri NHMUK 1931.8.18.999 ♀; D.
c. aldabranus NHMUK 1968.43.102 ♂; D. c. aldabranus NHMUK 1906.12.21.143 ♀; †D. c. abbotti NHMUK 1906.12.21.140
♂; †D. c. abbotti NHMUK 1906.12.21.141 ♀.

La Roque (in Lougnon 1992: 209) in 1708, while on the Plaine des Cafres, gave the only details about the nest-
ing behaviour (my translation):
One sees there [the Plaine des Cafres] a great numbers of oiseaux bleus which nest amongst grasses and aquatic
ferns.
Hébert, in 1708 (1940: 50–51), wrote a report for the East India Company and stated (my translation):
There are also oiseaux bleus, because their plumage is of a dark blue. They do not have another name. They are
as large as hens and are good to eat.
Le Gentil (1727) in 1717 and ‘Père Brown’ (1773) around 1730, gave the last unequivocal reports of the oiseaux
bleu (Cheke & Hume 2008). They referred to it as resembling a wood pigeon, which may have been in comparison
with the extinct, but undescribed Réunion Blue Pigeon Alectroenas sp., which was also almost all blue (Hume et al.
2006; Hume 2011a, 2017) (translation from Cheke & Hume 2008: 128):
Towards the east of the island there is a little plateau up a high mountain called the Plaine des Cafres where
one finds a large blue bird whose colour is very striking. It resembles a wood-pigeon [in colouration]. It flies but
rarely and always barely above the ground, but it walks with surprising speed. The inhabitants have never called it
anything other than oiseau bleu; its flesh is quite good and keeps well.
An anonymous account from 1763 (Anon 1763), but possibly written by Brigadier-General Richard Smith (Hill
1916), may have made the last mention of the Réunion Gallinule, but the writer provides no description of the bird.
A number of Mascarene birds were considered tame and without fear of humans; therefore the description cannot be
considered unequivocal. The account was further elaborated by Grant (1801), where it appears under ‘Observations
on the Isle of Bourbon, in 1763, by an officer in the British Navy’, and includes an insightful contemporary descrip-
tion of the Plaine des Cafres, the prime habitat of the gallinule (from Grant 1801: 167):
The plain des Caffres, is formed by the summits of mountains at a very considerable elevation above the sea: it
is said to be twenty miles in extent, and is very flat, and without stones. The access to it is very difficult in certain
places, though it may be ascended on horseback. The air is very pure, but as cold as winter’s day in England. When
the clouds pass over the surface of the plain, they have all the effect of a gentle rain. A brook runs through the middle
of it, which is broad but shallow, has a sandy bottom, and freezes in the winter….On this elevated plain there are
small trees, with a broom, furze, a kind of wild oat, and fern, which grows to the height of a shrub. There are also
some curious birds, which never descend to the sea-side, and who are so little accustomed to, or alarmed at, the
sight of man, that they suffer themselves to be killed by the stroke of a walking stick. It often rains on the sides of
the mountains, while this predominating plain is enlivened with the finest weather.
The Réunion Gallinule questionably survived until at least 1763 (Anon 1763; Grant 1801), long after many
of the other Réunion birds had become extinct. This was almost certainly due to the remoteness of its habitat (see
Feuilley 1705). Its extinction was due primarily to over-hunting, but cats, introduced at the end of the 17th century
(Cheke & Hume 2008), would have quickly eliminated the birds once they had turned feral and reached the remote
montane areas. Feral cats occur all over Réunion today, including the remotest, highest peaks, and are still serious
predators of native montane birds, especially breeding Barau’s Petrel Pterodroma baraui Jouanin, 1963 (Faulquier
et al. 2009).
African Swamphen Porphyrio porphyrio madagascariensis (Latham, 1801)
Gallinula porphyrio Latham, 1801, p.lxviii.

Porphyrio smaragdonotus Oustalet, 1897, p.98
Porphyrio porphyrio Carié, 1916, p.247
Porphyrio madagascariensis, Vinson, 1868, p.628; E. Newton, 1888, p.552; Peters, 1934, p.207; Rountree et al. 1952
Porphyrio porphyrio madagascariensis Safford & Hawkins, 2013, p.369; Dickinson & Remsen, 2013, p.160; del Hoyo & Col-
lar, 2014, pp.352
Remarks: Porphyrio swamphens are wide-ranging birds that have reached isolated islands and island archipelagos
on a number of occasions. Two species have been recorded in the Mascarenes as rare or accidental vagrants, but
neither species has ever naturally become established. The African Swamphen Porphyrio p. madagascariensis has
a wide distribution, being found in Egypt, Africa south of the Sahara and Madagascar (Safford & Hawkins 2013).
A specimen held at UMZC was collected on Mauritius and sent by Edward Newton to his brother, Alfred, at Cam-
bridge in 1861, and considered identical to the Madagascar species (A. Newton 1861a). E. Newton considered the
species as possibly resident, but later thought it was possibly introduced (E. Newton 1888); However, most authors
considered it exotic (see Rountree et al. 1952), as Milbert (1812) noted that some had been introduced from Mada-
gascar, and shipments of birds were introduced in the 1860s, after which the species became well established (A.
Newton 1861b; Meinertzhagen 1912; Carié 1916). It declined during the 20th century due to the active drainage
of wetlands on Mauritius to combat malaria (Cheke & Hume 2008), and seemingly disappeared completely by the
1950s (Safford & Basque 2007; Safford & Hawkins 2013). A single subsequent record in 1976 indicates that it is
an extremely rare vagrant, although Safford & Hawkins (2013) suggest that a tiny residue of the introduced birds
may have survived.
The African Swamphen was also occasionally reported on Réunion, with a flock of captive birds breeding on a
lake in the Jardin du Roi in Saint-Denis (Sganzin 1840; Cheke & Hume 2008), and it also occurred on the Étang de
St-Paul (Coquerel 1864); these latter-named birds may have been introduced birds or vagrants (Safford & Hawkins
2013). The population has since died out (Cheke & Hume 2008).
Allen’s Gallinule Porphyrio alleni Thomson, 1842
Porphyrio alleni Thomson, 1842, p.204; E. Newton, 1888, p.552
Remarks: Allen’s Gallinule Porphyrio alleni occurs in Africa, south of Sahara, Madagascar, and the Comoros, and
has been accidental to Europe and Atlantic islands (Safford & Hawkins 2013). It is an extremely rare visitor to the
Mascarenes, with only one record of four birds (one specimen collected and held at UMZC) in December 1873 at
Oyster Bay, Rodrigues (Hartlaub 1877; Bourne 1968; Safford & Hawkins 2013).
Genus Fulica Linnaeus, 1758
†Mascarene Coot Fulica newtonii Milne-Edwards, 1867b
Poules d’eau (waterhen) Pretorius, [1666] (in Hume & Winters, 2016, p.5) (Mauritius); Granaet, [1666] (in Barnwell, 1948,
p.40); François Martin in [1665 and 1667] (in Lougnon 1992, pp. 40, 43) (Réunion); Carpeau du Saussay, [1666] (in
Lougnon, 1992, p.51) (Réunion); Le Breton, [1671] (in Lougnon, 1992, p.135) (Réunion); Dubois, 1674, p.169 (Réunion);
Leguat, 1708 (French edition), p.71 (Mauritius); Oliver, 1898, p.77
Moor-hen Leguat, [in 1693] 1708 (English edition), p.71
Fulica newtonii: Milne-Edwards, 1867b, pp.203, 204(table), 211(table), 215, 216, 217, pl.10, 1866–1873 [1874], p.55; Oliver,
1891, p.369; Pitot, 1914, p.89; Olson, 1977, p.363; Cowles, 1987, p.96, 1994, p.92; Probst, 1997, p.48; Taylor & van Perlo,
1998, p.61; Probst & Brial, 2002, p.53; Mourer-Chauviré et al. 2006, p.43; Garcia et al. 2014, p.105; del Hoyo & Collar,
2014, pp.342,773; Hume, 2017, p.140
Fulica newtoni: Milne-Edwards, 1869–1871, p.139, fig.1–2, 1896a, p.76, 1896b, p.118; E. Newton, 1888, p.552; E. Newton &
Gadow, 1893, pp.282, 292–293, figs,1–11, 1895, p.233; Forbes, 1893a, p.544; Sharpe, 1894, p.209; Emerez de Charmoy,
1903, p.7; A. Newton, 1905, p.142; Carié, 1916, p.109; Sclater, 1924, p.109; Lambrecht, 1933, pp.470, 481; Guérin, 1942,
2(12), 1950, p.405; Berlioz, 1946, p.7; Greenway, 1958, p.119, 1967, p.119; Luther, 1970, p.187, 1986, p.183, 1986 [2005],
p.183; Barré & Barau, 1982, p.32; Cheke, 1987, p.35, 2013b, p.13; Balouet, 1990, p.45; Livezey, 2003, p.2; Adams et al.
2003, p.344; Hume et al. 2006, p.17; Cheke & Hume, 2008, p.135, tbls.3.1,4.1, pls.3,6; Hume & Walters 2012, p.120;
Hume, 2013, p.219, 2014a, p.38; Safford & Hawkins, 2013, pp.45,348 (lapsus)
Palaeolimnas newtoni: Forbes, 1893a, p.544; Rothschild, 1907a, p.150, 1907b, p.198; Brodkorb, 1967, p.130
Fulica (Palaeolimnas?) newtoni: Oustalet, 1897, p.99
Paludiphilus newtoni: Hachisuka, 1953, p.154

Syntypes: Milne-Edwards did not specify holotype material when describing Fulica newtonii, but three pre-
sumably unassociated elements, pelvis, tibiotarsus, tarsometatarsus, were figured (Milne-Edwards 1867b: pl.10).
He returned these specimens to UMZC, and by using details from the labels and by matching them to the illustra-
tions, I have been able to definitely assign the following as Milne-Edwards’ syntypes: Pelvis, UMZC 573 (pl.10,
1–4); Tibiotarsus, UMZC 573 (L) (pl.10, 11–15). There are two other tibiotarsi, right and left, with the same cata-

logue number (UMZC 573), but these are the largest known (see Appendix 2, Table 15) and may be associated.
They are too large to belong to the illustrated syntype series. Furthermore, the tarsometatarsus (pl.10, 5–10) in
Milne-Edward’s plate is from the left side, whereas a tarsometatarsus (UMZC 573) listed in the UMZC catalogue
as the possible holotype is from the right side. Another possibility is that the tarsometatarsus is figured in reverse, in
which the original plate is reproduced in a mirror image, i.e. the bone from the right side will appear as a left. This
was a standard procedure in lithography (Weaver 1964), but unfortunately there are no diagnostic erosional features
on the UMZC 573 bone to link it with the illustration. There are also four other left tarsometatarsi in the UMZC
collections all catalogued UMZC 298.aa, so the syntype tarsometatarsus cannot be confidently identified and is not
considered further.
Type locality: Mauritius, Mascarene Islands.
Distribution: Mauritius and Réunion, Mascarene Islands.
Etymology: After Sir Edward Newton (1832–1897), British assistant colonial secretary on Mauritius, and a
keen naturalist who organised the collection of many subfossil specimens on Mauritius and Rodrigues.
Referred fossil material: Rostrum, UMZC 298.aa; Vertebrae (cervical), NHMUK u/r; NHMUK u/r; NHMUK
u/r; Scapula, MNHN MAD8856 (Lp); MNHN MAD8856 (L); Sternum, UMZC 298.aa; Humerus, MNHN
MAD6571 (R); MNHN MAD6570 (R); NHMUK u/r (Lp); NMHUK u/r (Lp); NHMUK u/r (Lp); NHMUK u/r
(Rd); UMZC 298 (L); UMZC 298 (R); UMZC 298 (L); Radius, DRP 237.B4 (R); MNHN MAD8856 (R); Ulna,
UMZC 298 (R); UMZC 298 (L) juv.; Carpometacarpus, DRP 20.137 (R); Pelvis, MNHN MAD6505; NHMUK
u/r; NHMUK u/r; UMZC 298.aa; UMZC 298.aa; DRP 324.B1; Femur, UMZC 298.aa (L); LWM u/r (R); LWM u/r
(R); LWM u/r (R); LWM u/r (L); LWM u/r (Ld); Tibiotarsus, BMB 10179 (L); BMB 101748 (L); DRP294.B1 (Rd);
MNHN MAD6504 (R); MNHN MAD6503 (L); MNHN MAD6562 (L); MNHN MAD6823 (Rd); MNHN MAD
u/r (L); MNHN MAD u/r (L); NHMUK u/r (L); NHMUK u/r (Rd); NHMUK u/r (L) juv.; NHMUK u/r (L) juv.;
NHMUK u/r (L); UMZC 1004 (R); UMZC 1004 (R); UMZC 1004 (L); UMZC 1004 (R); UMZC 1004 (R); UMZC
1004 (L); UMZC 1004 (L); UMZC 1004 (R); UMZC 1004 (R); UMZC 1004 (L); UMZC 573 (R); UMZC 573 (L)
(largest known, possibly associated specimens); Tarsometatarsus, MNHN MAD6501 (R); MNHN MAD6508 (L);
NHMUK A1353 (R); NHMUK A1353 (L); NHMUK u/r (L); NHMUK u/r (R); NHMUK u/r (L); NHMUK u/r
(Ld); NHMUK u/r (Lp); NHMUK u/r (R); NHMUK u/r (R); NHMUK u/r (L); NHMUK u/r (R); NHMUK u/r (R);
NHMUK u/r (R); NHMUK u/r (Lp); NHMUK u/r (R); NHMUK u/r (Rd); NHMUK u/r (R) juv.; UMZC 573 (R);
Phalanx, MI u/r (Flacq). Additional Réunion material not examined by me (in Cowles 1994): Premaxillae LAC
1993.44; Pelvis LAC 1993.38; Sternum LAC 1993.39; Tibiotarsus LAC 1993.40 (L); LAC 1993.41 (R); Fibula
LAC 1993.43; Phalanges X5 LAC 1993.45; Vertebrae X2 LAC 1993.46. A recent discovery of a phalanx at a marsh
site in the Flacq area, north-east Mauritius (Fig. 2) in 2018 confirms its presence in the northern wetlands, but F.
newtonii probably occurred in all suitable wetland habitat on Mauritius.
Revised diagnosis: Rostrum (Fig. 32): os premaxillare short and weakly pointed; in dorsal view, os nasale nar-
row, not covering narial openings; in lateral view, foramina neurovascularia, present, small and circular with limited
extent; os nasale narrow; in ventral view, deep, distinct sulcus.
Scapula (Fig. 33): in lateral view, narrow; margo dorsalis distinct; facies articularis humeralis shallow; acro-
mion blunt, rounded and directed dorsad; tuberculum coracoideum distinct.
Sternum (Fig. 34): long and extremely narrow caudally; in dorsal view, processus craniolateralis broad and
project laterally; spina interna short, square-shaped, and slightly bifurcated; in lateral view, apex carinae extends
cranially less than rostrum sterni.
Humerus (Fig.35E): shaft strongly curved dorsoventrally; in caudal aspect, tuberculum ventrale pronounced,
incisura capitis shallow; fossa pneumotricipitalis variable, either with or without pneumaticum; tuberculum dorsale
oval shaped and reduced in size; sulcus humerotricipitalis shallow; in cranial view, sulcus ligamentosa transversus
shallow; crista bicipitalis, short, weakly square-shaped; fossa m. brachialis shallow; distinct processus supracondy-
laris dorsalis.
Radius (Fig. 35A): sulcus tendinosa deep extending proximad approximately ¼ of total length radius length;
Ulna (Fig. 35C): short and strongly arched dorso-ventrally proximally; in caudal aspect, olecranon distinct;
processus cotylaris dorsalis hooked with deep impressio scapulotricipitis; sulcus intercondylaris deep.
Pelvis (Fig. 35G): extremely long and narrow; in dorsal view, sulcus antitrochantericus extend laterally beyond
the ala preacetabularis; canalis iliosynsacralis opens cranially; few indistinct foramina intertransversariae; in lateral
view, foramen acetabuli smaller in size than foramen ilioischiadicum; crista iliaca dorsalis arched; in ventral view,
one pronounced processus costalis; ala ischii directed caudad.
Femur (Fig. 35I): extremely robust; in cranial view, shaft curved medially and directed caudally at the proximal
end; crista trochanteris strongly deflected mediad; linea intermuscularis indistinct; sulcus patellaris deeply exca-
vated; epicondylus lateralis distinct; condylus medialis extends the same distance distally as condylus lateralis; in
medial aspect, fovea ligamentosa capitis deep and circular; crista trochanteris extends caudad, offsetting centrally
facies articularis acetabularis; shaft strongly arched dorsally; in caudal aspect, impressiones obturatoriae indistinct;
linea intermuscularis caudalis distinct, forming a ridge; fossa poplitea shallow; condylus medialis smaller than con-
dylus lateralis; trochlea fibularis deeply excavated.
Tibiotarsus (Fig. 35K): long and robust; in cranial view, crista cnemialis forms a distinct proximal pointing
ridge; sulcus extensorius deeply excavated, extending proximad; canalis extensorius oval-shaped; pons supratendin-
eus wide and slightly arched, and situated mediad; crista cnemialis lateralis extends distad; in caudal aspect, crista
cnemialis cranialis strongly hooked; shaft shallow; condylus lateralis small, weakly rounded; depressio epicondy-
laris lateralis deep.
FIGURE 32. Comparison of rostra of Fulica newtonii with rostrum on cranium of F. atra, (above) dorsal view, (below) lateral
view. A. B. UMZC 298a Fulica atra u/s; C. D. UMZC 298.aa F. newtonii u/s. Scale bar = 10 mm.

FIGURE 33. Comparison of scapulae (ventrolateral aspect, left side) of Fulica newtonii with F. atra and Fulica cristata, A.
NHMUK S/1988.60.1 Fulica atra ♂; B. NHMUK 1897.5.10.25 F. cristata u/s; C. UMZC 8856 F. newtonii u/s; D. UMZC 8856
F. newtonii u/s. Scale bar = 10 mm.
Tarsometatarsus (Fig. 35M): long and robust; in dorsal aspect, trochleae metatarsorum weakly splayed; troch-
lea metatarsi II directed slightly plantad; impressiones retinaculi extensorii indistinct; two or more foramina vas-
cularia proximalia; in plantar aspect, surface area of foramen vasculare distale deeply excavated forming a large
circle; fossa metatarsi I deep and extends proximad; on proximal end, canal for tendon of musculus flexor digitorum
longus; deeply incised sulcus for tendon of musculus flexor perforatus digiti II and shallow sulcus for tendon of
musculus flexor hallucis longus present.
Description and comparison: For comparison with other Mascarene rallid genera, see under Aphanapteryx.
Fulica newtonii differs from F. atra by the following characters:
Rostrum: Larger, otherwise similar.
Scapula: Proportionately shorter; proximal end more robust; tuberculum coracoideum more pronounced; margo
dorsalis less defined; extremitas caudalis blunter.
Sternum: in dorsal view, processus craniolateralis more prominent, extending more laterad; rostrum sterni more
square-shaped, not bifurcated; in lateral view, apex carinae extends less craniad; otherwise similar.
Humerus: Comparatively short; incisura capitis shallower; in cranial view, fossa m. brachialis extends further
proximad; otherwise similar.
Radius: proportionately more robust, especially at proximal and distal ends; in caudal aspect, tuberculum apo-
neurosis more pronounced; in cranial aspect, caput radii more deeply incised; sulcus tendinous more deeply exca-
vated.
Ulna: comparatively short in total length; processus cotylaris dorsalis shorter, otherwise similar.
Pelvis: much larger, otherwise similar.
Femur: much larger, approximately 19% difference in total length and more robust, especially on proximal and
distal ends (see Appendix 2, Table 12); in cranial view, crista trochanteris larger and directed more distad; in caudal
aspect, sulcus more deeply excavated; otherwise similar.
Tibiotarsus: proportions differ greatly from F. atra, being closer to Porphyrio, i.e. long-legged; much larger,
approximately 32% difference in total length and more robust, especially on proximal and distal ends (see Appendix
2, Table 13); in cranial aspect, crista fibularis comparatively short; sulcus extensorius more deeply excavated.
Tarsometatarsus: proportions differ greatly from F. atra, being closer to Porphyrio, i.e. long-legged; much
larger, approximately 24% difference in total length, and more robust, especially on distal end.
Remarks: Milne-Edwards (1867b) described F. newtonii from a series of bones found at the Mare aux Songes
marsh, Mauritius. From my comparison with other similar-sized species, it measured around 35 cm in total length
and may have had a body mass of up to 1.2 kg. There is also a nearly complete growth series for juvenile tibiotarsi
(Fig. 36). Despite having a reduced sternum (see Appendix 2), Newton and Gadow (1893) noted that there was only
slight reduction in the carina sterni (Fig. 34), so it was clearly able to fly. F. newtonii has also been identified from
subfossil material discovered on Réunion and, because the two island populations are undifferentiated, Mourer-
Chauviré et al. (1999) suggested that it still had the capability to fly between Mauritius and Réunion, possibly until
comparatively recent times.
The extremely large pelvic and leg elements of F. newtonii, compared with the reduced pectoral and wing ele-
ments, are similar proportionately to the world’s largest coots, Chatham Island Coot F. chathamensis Forbes, 1892b
and New Zealand Coot F. prisca Hamilton, 1893. These coots had long necks, disproportionately long legs and
toes, and an upright stance (Worthy & Holdaway 2002; Tennyson & Martinson 2006), very similar to Porphyrio
gallinules. This is in complete contrast to all other extant Fulica, which are swimming birds with shorter legs, and
is most likely a result of the New Zealand birds being more terrestrial (Worthy & Holloway 2002). The Mascarene
Coot had a similar body shape with long neck and legs, which suggests that it was also probably more terrestrial.
The relationships of F. newtonii are unclear, but it is thought to be a derivative of the Eurasian Coot Fulica atra
Linnaeus, 1758 (Olson 1977) or Red-knobbed Coot F. cristata Gmelin, 1789, of Madagascar (Mourer-Chauviré
et al. 1999). However, due to the sister relationships of some other Madagascar avian taxa, e.g. Madagascar Teal
Anas bernieri (Hartlaub 1860) and Grey Teal Anas gracilis Buller, 1869 of Australasia (Mitchell et al. 2014a), and
Elephant birds Aepyornis sp. and New Zealand kiwi Apteryx sp. (Mitchell et al. 2014b), an African or Malagasy
origin may not be the case.
Ecology: Virtually nothing was recorded about the ecology of F. newtonii, other than it was universally called
Poule d’eau (waterhen) and considered variably good to eat. François Martin (in Lougnon 1992: 40) made the first
mention in 1665 at Saint-Gilles on Réunion, recording the abundance of coots and other birds, and the ease at which
they could be caught (my translation):
FIGURE 34. Comparison of sterna of Fulica newtonii with F. atra, (left) lateral view, (right) ventral view. A. C. UMZC 298a
Fulica atra u/s; B. D. UMZC 298.aa F. newtonii u/s. Scale bar = 10 mm.

FIGURE 35. Comparison of wing, pelvic and leg elements of Fulica newtonii with F. atra. Radii, left side, ventral aspect, A.
MNHN MAD8856 F. newtonii u/s; B. NHMUK S/1988.60.1 ♂; ulnae, right side, ventral aspect, C. UMZC 298.aa F. newtonii
u/s; D. UMZC 298a Fulica atra u/s; humeri, right side, caudal aspect, E. UMZC 298 F. newtonii u/s; F. UMZC 298a Fulica
atra u/s; pelves, dorsal view, G. UMZC 298.aa F. newtonii u/s; H. UMZC 298a Fulica atra u/s; left femora, cranial aspect, I.
UMZC 298.aa F. newtonii u/s; J. UMZC 298a Fulica atra u/s; left tibiotarsi, cranial aspect, K. UMZC 1004 F. newtonii u/s; L.
UMZC 298a Fulica atra u/s; left tarsometatarsi, dorsal aspect, M. UMZC 298.aa F. newtonii u/s; N. UMZC 298a Fulica atra
FIGURE 36. Juvenile Fulica newtonii NHMUK u/r showing ontogenetic changes. Note the elements are nearly as large as an
adult (far right), but the proximal ends are not ossified indicating subadult birds. Scale bar = 10 mm.

Various kinds of birds approached us, but so familiar that some even came to rest on our shoulders. The river
basin was covered in geese and waterhens, and the depths full of fish. We had plenty of everything and of our choice,
since a small baton was enough to kill parrots and other birds; that one could catch the pigs and goats by running
[after them], and the geese and the waterhens allowed one to approach [so closely] as to almost take them by hand.
We sent them all on board [ship].
But when Martin returned from Fort-Dauphin, Madagascar, two years later in 1667, he noted the destructive
carnage caused by the inhabitants (in Lougnon 1992: 43; my translation):
When some of the vessels of the fleet [of marquis de Mondevergue] were forced to sail from the Mascarene is-
land by the hurricane that surprised them there in February [1667], there remained many people left onshore, whom
were to be picked up afterwards. During the time that these people stayed on the island, they created unbelievable
commotion among the flocks, game and gardens. We saw neither geese nor waterhens on the Étang de St. Paul
which was formerly covered in them, and one was obliged to go to three or four miles from the dwelling to find
there goats and pigs.
Carpeau du Saussay in 1666 (in Lougnon 2002: 51) further commented on the abundance of birds, and stated
(my translation):
The birds are abundant and very familiar. Above all, we saw an infinity of turtle doves, pigeons, parrots, water-
hens, geese and ducks; only sticks and stones were needed to kill them.
The account from the log of Le Breton in 1671 (in Lougnon 1992: 135) noted that the Mascarene coot was still
abundant (my translation):
There are plenty of fish, especially mullet and eels, as well as all kinds of river birds. This is admirable to
see, also teals [Anas theodori Newton & Gadow, 1893] and wild geese [Réunion Sheldgoose Alopochen kervazoi
(Cowles, 1994)] which are smaller than those of Europe. There are also good numbers of waterhens, but they are
not good to eat.
Sieur Dubois (1674: 169), who was on Réunion from March 1671–September 4, 1672, not only gave the only
colour description, but also the last account of the coot on Réunion (my translation):
Waterhens, which are as large as fowls. They are always black, and have a large white crest on the head.
The ‘large white crest’ refers to the bare frontal shield that occurs in coot and gallinule genera, including Fulica
(Ripley 1977; Taylor & van Perlo 1998).
The coot was first mentioned on Mauritius by a VOC (Vereenigde Oost-Indische Compagnie [Dutch East India
Company]) book-keeper, Jacob Granaet, who arrived there on July 30, 1666 (Barnwell 1948; Hume & Winters
2016), and noted (from Barnwell 1948: 42):
Within the forests dwell parrots, turtle and other wild doves, mischievous and unusually large ravens [Lophop-
sittacus mauritianus], falcons [Mauritius Kestrel Falco punctatus Temminck, 1821], bats and other birds whose
names I do not know, never having seen such before….Water-fowls such as geese [Mauritius Sheldgoose Alopochen
mauritiana Newton & Gadow, 1893], teals, waterhens and flamingos are found among the marshes, very numerous,
especially the teals which are so tame they can be killed with sticks; all are fat and pleasant to eat.
They were briefly mentioned again in 1666 by Pretorius, who had stayed on Mauritius for three years (in Hume
& Winters 2016: 5), and wrote:
…the teals [Anas theodori] and waterhens are always in the water, and all live together in peace and quiet.
In reference to Pretorius’s remark about the Mascarene coots’ peaceful existence, even though flight must have
been impaired to some degree, one of a pair of associated humeri is pathological and exhibits a serious, but healed
fracture on the distal end (Fig. 37). I have also seen similar healed wing injuries in subfossil remains of Echo Para-
keet Psittacula echo (Newton & Newton, 1876) and Pink Pigeon Nesoenas mayeri (Prévost & Knip, 1843). This
suggests that adult birds could recover from injuries that would likely have otherwise ended in premature death if
predatory terrestrial mammals were present.
The Mascarene Coot probably had similar habits to F. atra (see Taylor & van Perlo 1998), nesting in aquatic
vegetation alongside water, and feeding on vegetable matter, aquatic seeds and invertebrates.
Extinction: Despite their sometimes inadequate culinary qualities, both the Mauritius and, especially Réunion
populations of Mascarene Coot were heavily hunted. Martin, in 1665, recorded an abundance of coots on the Étang
St. Paul on the northwest coast, the largest lake and one of the most densely human-populated areas on Réunion dur-
ing the 17th century (Lougnon 1992; Blanchard 2000), but when he returned two years later they had been hunted
out. They were still recorded as numerous elsewhere until the early 1670s, but not mentioned again thereafter. Black
rats were accidentally introduced after 1672, and reached plague proportions by 1676 (Cheke & Hume 2008; Hume
2013), so it is not known what effect, if any, they had on these birds, but the Mascarene Coot survived alongside
rats on Mauritius for centuries (Hume 2013). As cats were introduced in the 1680s (Cheke & Hume 2008), and that
F. newtonii was confined to lowland wetlands, it is likely that over-hunting and later cat predation exterminated the
species on Réunion at some point after 1672, as Feuilley (1705), who gave a detailed account about the avifauna,
including the native waterbirds, failed to mention the coot during his visit in 1704. In 1710 and just 40 years after
the first settlement, Antoine Boucher, the French East India Company’s store keeper and treasurer, condemned the
company’s management of Réunion and the wanton slaughter of the fauna (Cheke & Hume 2008; Hume 2014b).
He considered the settlers’ insatiable greed solely accountable for the faunal disappearance, and noted at the time of
writing that almost all of the larger land birds, including F. newtonii were either extinct or nearly so.
The Mascarene Coot was last mentioned in 1693 on Mauritius when it was considered rare (Leguat 1708), so pre-
sumably it succumbed to feral cat predation shortly thereafter, as did the other Mauritian endemic rails.
FIGURE 37. Associated pair of humeri of Fulica newtonii (UMZC 298 u/s) showing pathological specimen with a healed
fracture A, caudal aspect, C, cranial aspect, compared with normal specimen, B, D. Scale bar = 10 mm.
Genus Gallinula Brisson, 1760
Madagascan Moorhen Gallinula chloropus pyrrhorrhoa A. Newton, 1861b
Haretac Flacourt, 1661, p.164

Fulica chloropus: Linnaeus, 1758, p. 152, 1766, p. 258; Desjardins, 1831, p.45; Hartlaub, 1848, p. 392, 1860, p.173
Gallinula chloropus: Sganzin, 1840 [in 1831–32], p.45; Hartlaub, 1861, p.81; Pollen, 1863, p.325; Coquerel, 1864, p.23; Schle-
gel, 1865, p.47, 1866b, p.425; Schlegel & Pollen, 1868, p.136; Vinson, 1868, p.628; Gray, 1871, p.66; Sharpe, 1894, p.169;

Meinertzhagen, 1912, p.99
Gallinule or Moor-hen: Clark, 1859, p.lxxxv
Gallinula A. Newton, 1861a, p.116
Gallinula pyrrhorrhoa: A. Newton, 1861b, p.19 (Mauritius); E. Newton, 1861, p.275, 1863, p.458; Sclater, 1863, p.165; Roch
& Newton, 1863, p.174; Verreaux, 1865, p.4; Bartlett, 1875, p.68; Hartlaub, 1877, p.348; E. Newton, 1888, p.552; Guérin,
1950, p.2(1)
Gallinula chloropus, var. pyrrhorrhoa Grandidier, 1868, p.5; Finsch & Hartlaub, 1870, p.787; Milne-Edwards & Grandidier,
1882, p.594, pls.240, 241,241a; Oustalet, 1897, p.96; Carié, 1916, p.47
Gallinula chloropus pyrrhorrhoa Oustalet, 1897, p.96; Sclater, 1924, p.108; Rand, 1936, p.361; Guérin, 1939, p.187; Berlioz,
1946, p.37; Staub, 1973b, p.31; Rountree et al. 1952, p.181; Safford & Hawkins, 2013, 372; Dickinson & Remsen, 2013,
p.161; del Hoyo & Collar, 2014, pp.352
Remarks: The Madagascan race of the Eurasian Moorhen Gallinula chloropus (Linnaeus, 1758) is found through-
out Madagascar and on the Comoro Islands, and is also a resident breeder on Mauritius and Réunion (A. Newton
1861a; Cheke & Hume 2008; Safford & Hawkins 2013). The holotype of G. c. pyrrhorrhoa was collected in 1861
on Mauritius, when Alfred Newton received the skin from his brother, Edward (A. Newton 1861a, b). Another
specimen had been previously been collected in 1831 (Desjardins 1831), at the same time that the eggs were prob-
ably recorded (Cari 1904). Cossigny (Charpentier de Cossigny 1799) first observed them on Mauritius during the
1790s, and it appears that they were by then well established (Cheke & Hume 2008). Michel (1992) thought that it
had been introduced by the French, but provides no evidence to support this claim and it appears unlikely. It was
also first observed on Réunion in 1831 and was also considered common there at that time (Sganzin 1840). The lack
of records from the 17th century, plus its absence from the fossil-rich Mare aux Songes deposits, suggests that G.
c. pyrrhorrhoa is a recent colonist that may well have populated the niche of Fulica newtonii after it had become
extinct (Cheke & Hume 2008; Hume 2013).
The Madagascan Moorhen occurs in most remaining wetland areas of Mauritius today, especially in the north
where it is abundant (pers. obs) (Fig. 38), and is well established on the Étang St. Paul and other wetland areas up
to an altitude of 1,000 m on Réunion (Milon 1951; Safford & Hawkins 2013).
FIGURE 38. The last surviving Mascarene rail, Madagascan Moorhen Gallinula chloropus pyrrhorrhoa (centre on rock), at
the coastal Anse la Raie wetlands on the northern tip of Mauritius. This was once the habitat of Fulica newtonii. Photograph by
the author.
Géant or Leguat’s Giant Leguatia gigantea Schlegel, 1857
I include this fanciful, supposed Mascarene rail solely for completeness. It was the Mauritian historian Albert Pitot
(1914: 91), when discussing Leguatia, who best highlighted the absurd erroneous literature that resurrected it in the
first place:
This may serve as an example of how insufficient, superficial, and faulty information on a subject is liable to
give rise to most persistent misrepresentations. Such a fallacy might certainly be excused in old Leguat, who did
not boast of being either an expert zoologist or a clever artist; but it is surprising that such modern scholars such as
Dr. Schlegel and Mr. Oustalet, of the Paris Museum, should have deliberately shared in his error, without any other
reason than a badly-drawn sketch.
The origin of the Géant or Leguat’s Giant as it became known, stems from the account of François Leguat while
on Mauritius in 1693 (Leguat 1708: 209–210), who stated:
There is no whale fishery, but the great flights of Bitterns [Mauritian Night Heron Nycticorax mauritianus (E.
Newton & Gadow, 1893)], and many of those birds call’d Giants, because they are six foot high. They are extremely
high mounted, and have very long necks; Their Bodies are not bigger than that of a Goose. They are all white, ac-
cept a little place under their Wings, which is reddish. They have a Goose’s Bill, but a little sharper; their claws are
very long and divided. They feed in Marshy Places, and the Dogs frequently surprise them, because they require
a considerable time to get upon the Wing; We saw one one day at Rodrigo [Rodrigues], and we took him with our
Hands, he was so fat. That was the only one we observ’d there, which made me inclinable to believe he had been
carry’d thither by some Wind he could not resist. This Game is good enough [to eat].
Leguat illustrated the bird in his book (Leguat 1708: pl. X), but not based on what he saw, as he directly copied
an illustration of a Purple Gallinule Porphyrio porphyrio (Linnaeus, 1758) from that of Adriaen Collaert (c.1610),
which resulted in considerable confusion (A. Newton 1873; Stresemann 1923a, b). Based on Leguat’s testimony,
and ignoring the comments of Buffon (1781: 306) and Strickland (in Strickland & Melville 1848: 60) that Leguat
was describing a flamingo, Schlegel (1857) erected Gallinula (Leguatia) gigantea for the Géant, and placed it
amongst waterhens. The Géant was given further scientific credence by Oustalet (1874c), and when Rothschild
(1907a) and Hachisuka (1953) both reproduced elaborate illustrations depicting a six foot tall, giant water-rail, the
gallinule affinities of Leguatia became ingrained in the literature.
No giant rail bones have ever been found on Mauritius, but flamingo bones have been collected from the Mare
aux Songes marsh in 1865 and 1889 (E. Newton & Gadow 1893; A. Newton 1907), and also more recently in 2006
(Rijsdijk et al. 2009). On Réunion, a number of flamingo bones were found at Marais de l’Ermitage (Mourer-Chau-
viré et al. 1999). Furthermore, a partial tarsometatarsus held in the OUM, and collected from the Mare aux Songes
in 1865, is referred to Leguatia by Schlegel (details from label), but ironically belongs to Greater Flamingo Phoe-
nicopterus roseus Pallas, 1811 (pers. id). This specimen also has medullary bone deposits, which confirms that P.
roseus was a resident breeding species on Mauritius, as it was on Réunion, as some of the subfossil tarsometatarsi
found there also have medullary bone (Mourer-Chauviré et al. 1999). Irrespective of the subfossil remains, the most
damning evidence against the rallid affinities of Leguatia is the diagnostic red underwings described by Leguat,
which are characteristic of flamingos (Buffon 1781), but never occur in Rallidae (see Ripley 1977; Taylor & van
Perlo 1998).
Two species of flamingo occur in the Southwestern Indian Ocean, the Greater Flamingo and Lesser Flamingo
Phoeniconaias minor Saint-Hilaire, 1798, with the former breeding on Madagascar and Aldabra, and is a rare vagrant
to Seychelles, Comoros and some other smaller islands, whereas the latter occurs as a non-breeder on Madagascar
and is a rare vagrant to Comoros (Safford & Hawkins 2013). Both species have been recorded on the Mascarenes,
the Greater Flamingo, from which the Géant myth arose, was a breeding resident, while the Lesser Flamingo has
been observed only infrequently (Cheke & Hume 2008; Safford & Hawkins 2013). A single individual, from around
15 flamingos seen in 1870 on Mauritius, was later identified as Lesser Flamingo by Milne-Edwards (1874), two
were observed in 1923 at Baie aux Huitres, Rodrigues (Showler & Cheke 2002), and a much more recent sighting
of Lesser Flamingo was made in January 2006 at Grande Anse Beach on Réunion (Safford & Hawkins 2013). There
may have been other occurrences, but there was much confusion over field identification between the two species
(Oustalet 1897; Safford & Hawkins 2013). Rountree et al (1952) suggest that a resident, breeding population of
Lesser Flamingo may have once occurred, but provides no references to support this claim. All subfossil flamingo
material found at the Mare aux Songes is referable to the Greater Flamingo (pers. obs), as is subfossil material found
at the Marais de l’Ermitage, Réunion (Mourer-Chauviré et al. 1999).

Resident breeding populations of Greater Flamingos were extremely numerous on Mauritius and Réunion dur-
ing the mid-17th century, numbering around 3,000–4,000 in 1704 on the Étang du Gol, Réunion alone (Feuilley
1705; Barré & Barau 1982, Cheke & Hume 2008) (Fig. 3). However, they were systematically over-hunted for food,
and either died out or dispersed elsewhere between 1710 and 1730 on Réunion (Cheke 1987; Mourer-Chauviré et al.
2006), and around 1770 on Mauritius, having been reduced to just three individuals in 1768 (Bernardin 1773: 124).
Vagrants continued to sporadically arrive for the next century (Oustalet 1897), with a number of birds observed (and
immediately killed) in 1873 at Black River (Pitot 1914), which may have been part of the above-mentioned flock
of 15 observed in 1870. This appears to be the last record on Mauritius. The evidence against the validity of the
Géant is overwhelming, and in my opinion it must be permanently subsumed into the synonymy of Phoenicopterus
roseus. A much more interesting note based on Leguat’s observation is that it constitutes the first record of a Greater
Flamingo on Rodrigues. Here I include a revised systematics of Leguatia gigantea and include all of its permuta-
tions that I have managed to trace:
Order Phoenicopteriformes Fürbringer, 1888
Family Phoenicopteridae Bonaparte, 1831
Genus Phoenicopterus Linnaeus, 1758
Phoenicopterus Linnaeus, 1758, p.139. Type by monotypy P. ruber Linnaeus, 1758
Greater Flamingo Phoenicopterus roseus Pallas, 1811
Le Géant/Giant Leguat, 1708, ii, p.73 (French ed.), p.171 (English ed.); Oustalet, 1874c, pp.113–116
Peacock Leguat, 1708, p.41 (English ed.); Strickland (in Strickland & Melville, 1848, p.60); Oliver, 1891, p.44
Flamingo Buffon, 1781, p.306; Strickland (in Strickland & Melville, 1848, p.60); Clark, 1866, p.144; A. Newton, 1907, p.71
Phoenicopterus roseus: Pallas, 1811, p.207; Serverzow, 1875, p.184; Salvadori, 1895, p.12; Oustalet, 1897, p.101; Blandford,
1898, p.408; Guérin, 1942, p.1(54); Knox et al. 2002, p.708; Dickinson & Remsen, 2013, p.49; del Hoyo & Collar, 2014,
pp.152
Phoenicopterus antiquorum Temminck, 1820, p.587; Peters, 1931, p.141; Mayr & Cottrell, 1979, p.270 Nomen nudum
Geant (Strausartiger vogel) Hamel, 1848, pp. 65–96
Gallinula (Leguatia) gigantea Schlegel, 1857, p.142, 1858, p.378, 1866a, p.167; Rowley, 1877, p.131; Anon, 1892, p.365; A.
Newton, 1873, p.195, 1907, p.71
Phoenicopterus sp. Milne-Edwards, 1874, p.27; E. Newton & Gadow, 1893, p.282; Lambrecht, 1933, p.345
Leguatia gigantea Shufeldt, 1886, p.361; Oustalet, 1897, p.100; Rothschild, 1907a, p.151, pl.31, 1907b, p.198; Oudemans,
1917, pp.37–55; Stresemann, 1923a, p.451, 1923b, p.511; Carié, 1916, p.107, 1930, p.204; Rountree et al. 1952, p.157;
Hachisuka, 1953, p.161; North-Coombes, 1983, p.2; Holthuis et al. 1971, pp.7–8; Olson, 1977, p.373; Fuller, 1987, p.240,
2001, p.386, 2002, p.63; Balouet, 1990, p.47; Cheke & Hume, 2008, p.27; Hume & Walters, 2012, p.373; Hume, 2013,
p.211, 2014a, p.32, 2017, p.418
Leguatia Fürbringer, 1888, p.xliii
Gallinula (Leguatiana) gigantea Pitot, 1905, p.372 (Lapsus)
Giant Pitot, 1914, p.91
Gallinula gigantea Killermann, 1917, p.377; Lambrecht, 1933, p.479
Phoenicopterus ruber antiquorum Sclater, 1924, p.39; Rand, 1936, p.342; Roberts, 1940, p.33, pl.iv; Berlioz, 1946, p.6: Roun-
tree et al. 1952, p.171
Phoenicopterus ruber roseus: (Pallas, 1811) Mayr & Cottrell, 1979, p.270
Phoenicopterus ruber: Mourer-Chauviré et al. 1999, p.11; Hume et al. 2006, p.9 (Réunion)
Discussion
Due to their isolation, the Mascarene Islands were the last major island group to be discovered and settled by hu-
mans, so unparalleled documentation was made about the environment at a time when the islands were compara-
tively pristine (Cheke & Hume 2008). The three Mascarene Islands differ significantly from each other in age, geol-
ogy and ecology (Cheke & Hume 2008; Hume 2011b), so a rich diversity of niches was available for any colonising
species able to cross the vast distances of water in order to reach them. Réunion, the youngest island of the group,
has been dated at ~2 MY (McDougall 1971; Duncan 2009), Mauritius at 6.8 to 8 MY (Saddul 2002; Duncan 2009),
and the oldest rocks on Rodrigues are dated at 7.5 to 11 MY (Perroud 1982; Giorgi & Borchiellini 1998; Duncan
2009). An often cited potassium-argon dating gives 1.5 MY for Rodrigue’s oldest rocks (McDougall et al. 1965), but
this was based on material collected from the erosional remnant of a volcano that ceased activity 1.5 MYA (Duncan
2009); thus the Mascarene Islands have provided above-water land area since at least the mid-Miocene (but see
below). Amongst vertebrates, no terrestrial mammals or amphibians were able to establish, whereas high endemic-
ity occurs in Pteropus fruit bats (Pteropodidae), geckos (Gekkonidae), skinks (Scincidae), snakes (Bolyeriidae) and
giant tortoises (Testudonidae) (Cheke & Hume 2008). Birds are adept at populating isolated islands, and four bird
groups in particular have established multiple times on the Mascarenes: Rallidae (this paper, 7 species) (Fig. 39),
Columbiformes (10 species) (Hume 2011a), Psittaciformes (9 species) (Hume 2007) and Passeriformes (20+ spe-
cies) (Hume 2013, 2014a, 2015). The Mascarene endemic Rallidae Dryolimnas augusti and D. chekei, are clearly
part of modern Dryolimnas, and their origins lie in the Malagasy region (Cheke & Hume 2008). García et al. (2014)
have shown that Dryolimnas is basal in a clade of rails including Crex, Lewinia and Slaty-breasted Rail Lewinia
(Hypotaenidia) striatus (Linnaeus, 1766), which diverged in the Miocene around 21 MYA. Less clear are the ori-
gins of Fulica newtonii and ‘Porphyrio caerulescens’. These taxa are seemingly African or Malagasy derivatives,
but as molecular studies have shown a number of unexpected long-distance dispersal relationships, e.g. Laterallus
(Stervander et al. 2019), the origins of F. newtonii and ‘P. caerulescens’ are open to alternative interpretation. The
relationships of Aphanapteryx and Erythromachus are much more difficult to determine. Based on the widely-
spaced trochleae and narrow os nasale, Olson (1977) suggested that they had affinities with Dryolimnas, Rallus,
Gallirallus [Hypotaenidia] and Laterallus, but only the first two of these genera are indigenous to the Indian Ocean
region today, whereas the Hypotaenidia philippensis individual collected on Mauritius remains unique, and Lateral-
lus is restricted to the South Atlantic (Ripley 1977; Taylor & van Perlo 1998). The phylogenetic history of the two
Mascarene endemic genera appear to be clouded by significant apomorphies derived from morphologically changes
associated with flightlessness, and jaw modifications due to feeding niche, which suggests long isolation. However,
other factors such as gene flow, resource availability and climatic events may have also played a significant role in
affecting the intensity of directional selection and therefore the speed at which flightless and feeding related apo-
morphies evolved (see McNab 1994; Livezey 2003). Flightlessness can evolve rapidly in Rallidae as other studies
have shown, e.g. generations (Olson 1973); 125,000 years (Slikas et al. 2002); 400,000 years (Kirchman 2012).
Furthermore, it can happen on repeated occasions. A recent study has demonstrated that a Dryolimnas rail colonised
Aldabra Atoll and became flightless in the Middle Pleistocene, subsequently disappearing due to a high sea stand
inundation event, and then recolonised the atoll in the Late Pleistocene; thus eventually becoming flightless for a
second time (Hume & Martill 2019); this change towards flightlessness occurred in a period of less than ~20,000
years. Therefore, the derived condition of the Mascarene endemic rails may not have required long periods of time
to evolve and they likely colonised the Mascarenes more recently. Similarly, Dieffenbach’s Rail Hypotaenidia
dieffenbachii G.R. Gray, 1843, Chatham Rail Cabalus modestus Hutton, 1872 and Hawkins’ Rail Diaphorapteryx
hawkinsi (Forbes, 1892b) of the Chatham Islands are as equally specialised as the Mascarene Rallidae, yet evolved
their distinct apomorphies in less than 1–3 Ma (Trewick 1997a, b; Trewick et al. 2007).
The affinities of Aphanapteryx and Erythromachus are probably close to a Dryolimnas-type rail, but they now
differ considerably in osteology and thus their relationships remain unresolved.
Sexual dimorphism
Most flightless rails are characterised by an increase in body size (Livezey 2003), with the greatest increase exhib-
ited amongst a few genera of large, flightless rails including Aphanapteryx and Erythromachus (Appendix 2). As all
Rallidae referred to in this study show a degree of size variation, which is attributed to sexual dimorphism (Livezey
2003), who also demonstrated that in the skeleton the cranium and tibiotarsus are the best elements to illustrate
this fact. Interestingly, Aphanapteryx has the greatest size variation in the cranium in all of the examined Rallidae
(cranium n=2, TL 38.0–48.7 mm, SD 7.56), being much less in Erythromachus (cranium n=5, TL 37.3–40.4 mm,
SD 1.26) (Appendix 2, Table 1). Furthermore, nominate Dryolimnas cuvieri has the greatest variation in the rostrum

(n=2, TL 39.7–45.9,SD 4.38), followed closely by Aphanapteryx (n=6, TL 74.8–86.0 mm, SD 4.0) (Appendix 2,
Table 2). The paucity of cranial material for the other Mascarene rails compromises statistical interpretive conclu-
sions, but the tibiotarsus is reasonably well represented and so is used here.
Based on the total lengths of tibiotarsus, including the generally limited availability of comparative modern
specimens (Appendix 2, Table 13), and using the more robust data set of Livezey 2003), the highest variability in
tibiotarsus length in Mascarene rails is shown in Dryolimnas chekei (n = 2, TL mean = 84.0, 78.1–90.0 mm, SD
8.41), followed by Aphanapteryx (n = 31, TL mean = 111.5 mm, 100.2–121.3 mm, SD 6.08) and Erythromachus
(n=17, TL mean = 88.4 mm, 82.0–99.8 mm, SD 4.85); it is also high in Dryolimnas c. cuvieri (n=2, TL mean =
78.5, 75.3–81.8 mm, SD 4.59). What is interesting to note is that flightless D. c. aldabranus is less variable (n=15,
mean = 67.1 mm, 62.7–71.6 mm, SD 3.31) in both the skeleton (Appendix 2; Livezey 2003: Tbl. 33) and the skin
(Table 3) than the volant nominate, which is in direct opposition to the usual trends in insular flightless rails (see
Livezey 2003). Aquatic rails show greater size variation than terrestrial species (Appendix 2, table 13), with Fulica
cristata (n=7, TL mean = 105.3 mm, 94.3–115.0 mm, SD 8.56) exhibiting the greatest, whereas F. atra (n=9, TL
mean = 103.4 mm, 90.9–108.4 mm, SD 5.95) (n = 14, mean = 102.8 mm male; n = 21, mean = 100.3 mm female
(Livezey 2003) is much less dimorphic. The more terrestrial Fulica, F. chathamensis (n=17, TL mean = 152.8 mm,
142.6–166.5 mm, SD 7.67) and F. newtonii (n=26, TL mean = 127.4 mm, 119.5–150.0 mm, SD 6.95), also exhibit
great size variation. Livezey (2003), using the corpus length of 15 tibiotarsi of F. newtonii, concluded that the male
(n = 6, mean = 141.0 mm) was much larger in total length than the female (n = 9, mean = 128.8 mm); hence, this
species was extremely sexually dimorphic. The world’s largest gallinule, Porphyrio hochstetteri is surprisingly less
variable (n = 4, TL mean = 148.5 mm, 144.5–151.2 mm, SD 2.98); however, this taxon, according to the Pleistocene
fossil record, occurred widely in a range of coastal, lowland and highland habitats, and was not confined to the al-
pine zone as it is today (Trewick & Worthy 2001), so can be considered a terrestrial species.
Extreme sexual dimorphism can be explained in the aquatic rails, Porphyrio, Gallinula and Fulica, by their
habit of nesting and feeding in open, lacustrine environments where nest sites and food availability are at a premi-
um, and thus is a result of male-driven territoriality (Livezey 2003). It is difficult to determine what ecological and
behavioural factors were involved in the evolution of size sexual selection in the extinct terrestrial Rallidae of the
Mascarenes, but in other groups of birds, high levels of social polygamy, differences in parental care or intersexual
competition appear to be the root causes (Owens & Hartley 1998). As rails are mostly monogamous (Taylor & van
Perlo 1998), whereas intersexual competition is predominantly a result of food availability (Selander 1972; Shine
1989), it seems likely that the latter was the main evolutionary driving force in island terrestrial Rallidae, with each
sex exploiting a differing range of food items on islands with limited resources.
FIGURE 39. Author’s scaled illustration showing the morphologies of the endemic Mascarene Rallidae, from left to right:
Aphanapteryx bonasia (juvenile); Aphanapteryx bonasia (adult); Erythromachus leguati; Dryolimnas augusti; Dryolimnas
chekei sp. nov.; ‘Porphyrio caerulescens’; Fulica newtonii.
Ecology
Our knowledge of Mascarene rail ecology is based entirely on the inadequate documentation of visiting mariners
to the islands. But put into context, we have to be grateful for the nuances of detail left by a few learned travellers,
whose main concern after an onerous long sea voyage was not the luxury of observing bird behaviour, but the fun-
damental necessity of obtaining the next meal. Only one settler, Johannes Pretorius, when describing Aphanapteryx,
provided any detail of an extinct rails’ ecology on Mauritius (Hume & Winters 2016), whereas François Leguat and
Julien Tafforet gave precious few details about Erythromachus on Rodrigues (Leguat 1708; Tafforet 1725–26). The
rails of Réunion were best attested by Dubois (1674) and Feuilley (1705), but even their accounts leave much to be
desired.
From the available evidence it can be stated with some certainty that Aphanapteryx was an opportunist preda-
tor and molluscivore, scratching the ground with its strong feet when foraging, and probing with its long, sensitive
bill in leaf litter and soft substrates for invertebrates (Hume & Winters 2016). It actively hunted some of the largest
endemic, terrestrial snails, puncturing the ventral surface of large specimens to obtain the soft parts (Fig.21), and
perhaps swallowing smaller specimens whole. Aphanapteryx may have fed on tortoise eggs during their breeding
season, and possibly scavenged seabird colonies on the coasts; endemic terrestrial nemertean and planarian worms
and the now-extinct endemic worm-snake Typhlops cariei may have also been part of the diet. Sexual dimorphism
was seemingly extreme, with the male proposed to be larger than the female (Livezey 2003), so the sexes probably
had different feeding niches. Aphanapteryx presumably had a variety of loud calls, which if impersonated by hu-
mans and combined with the waving of red cloth, would attract the rails to within striking distance (Hume & Winters
2016). Nest and eggs were never recorded, but it is likely the species made a well-concealed nest in dense vegetation
in forests, as do many other flightless rails (Ripley 1977).
Erythromachus was also an opportunist predator, feeding on tortoise eggs when in season (Tafforet 1725–26),
and probably on invertebrates during other times of the year, including terrestrial molluscs and nemertean and
planarian worms. It may have also scavenged sea bird colonies. Sexual dimorphism was also probably extreme
(Livezey 2003), which suggests that the sexes had different feeding niches. Calls included a continued loud whis-
tling, and when pressed, a sound like someone with hiccups (Tafforet 1725–26). It was an aggressive species and
reacted in the same way as Aphanapteryx to the waving of a red cloth (Leguat 1708). The breeding biology of Eryth-
romachus was not recorded, but it no doubt chose forested areas of the island to breed, with the nest well concealed
in dense vegetation.
On Mauritius, Dryolimnas chekei was sympatric with Aphanapteryx, occurring in the same habitat but, being
smaller with a shorter, straight bill, presumably occupied a different niche. If the diet of D. chekei was similar to that
of other Dryolimnas, which feed on crabs, small marine and terrestrial snails, insects and other invertebrates (Saf-
ford & Hawkins 2013), and even eggs and hatchling testudines (Frith 1977), feeding competition between D. chekei
and Aphanapteryx was probably limited. Like other flightless rails, D. chekei probably nested in dense vegetation
deep in forests. On Réunion, D. augusti was the only lowland, terrestrial rail. Like other Dryolimnas, it presumably
nested deep in forests, concealing its nest in dense vegetation, and fed on various invertebrates; it may also have
seasonally taken tortoise eggs and hatchlings.
The montane regions of Réunion were inhabited by the poorly volant, but fast running Réunion Gallinule or
oiseaux bleu ‘Porphyrio caerulescens’, where it nested amongst reeds and ferns (La Roque in Lougnon 1992: 209).
It likely fed on vegetable matter and invertebrates, as do other Porphyrio (Ripley 1977).
The wetlands on Mauritius and Réunion were occupied by Fulica newtonii, a large, but volant coot that may
have moved between the islands (Mourer-Chauviré et al. 1999). The breeding biology of F. newtonii was not re-
corded, but it probably nested in aquatic vegetation alongside water, and like F. atra (Ripley 1977), fed on vegetable
matter, aquatic seeds and invertebrates.
Extinction
Rallidae are adept island colonisers, and where suitable environmental conditions occur, such as an absence of ter-
restrial mammals, can become flightless within a few generations (Olson 1973; McNab 1994; Kirchman 2012; Ster-
vander et al. 2019). However, this adaptability is not without consequence, as island rails are particularly vulnerable

once humans and their commensals become established; thus they have suffered extremely high extinction rates
(Olson 1977; Ripley 1977; Steadman 1995, 2006; Taylor & van Perlo 1998; Gill et al. 2010; Worthy et al. 2017;
Hume 2017). Steadman (1995, 2006) retrodicted that hundreds of rail populations must have disappeared during the
expansion of Polynesians across the Pacific Ocean; 21 of these have now been named (see Hume 2017), but for the
others, no physical or documented record exists. The isolation of the Mascarene Islands ensured their comparatively
pristine state until settlement by humans during the early 17th century, which resulted in an unprecedented chrono-
logical documentation of human-induced changes to the islands (Cheke & Hume 2008).
During the first century of human habitation, deforestation was negligible, and despite the presence of intro-
duced invasive species, especially black rats, rail populations did not appear to be overly affected. For most of the
Mascarene Rallidae it was the human-facilitated introduction of cats, which had been imported to control rats but
turned feral, that resulted in their rapid extinction (Cheke & Hume 2008; Hume 2017); cats arrived on Mauritius
and Réunion in the late 1680s, and c.1750 on Rodrigues (Table 3). However, over-hunting was particularly severe
on Réunion during the second half of the 17th century, and this is a reflection of a rapidly increasing human popula-
tion. In 1665, the European inhabitants numbered 12, then 734 by 1704, but rose rapidly to 61,000 by 1789 (Milon
1951; Hume 2014b), so it is no surprise that Réunion lost nearly all of its native, non-passerine land birds during this
period. Only two Mascarene rails unequivocally survived to the 18th century. The Réunion Gallinule survived until
at least 1730, or possibly the 1760s, primarily due to the remoteness of its montane habitat, whereas Erythromachus,
which occurred on the isolated island of Rodrigues, uninhabited until a tortoise hunting station was set up in the
1730s, fell victim to cats within a decade or so of their introduction around 1750.
The Mascarene Rallidae exemplify the competence of these birds to cross large distances of open-water to reach
isolated archipelagos and then evolve high endemicity. The varied topography and diverse vegetation zones of the
islands provided enough niches for sympatric populations of rails to co-exist, with Mauritius and Réunion each hav-
ing three taxa, whereas Rodrigues, the smallest and most remote of the group, harboured one highly size-variable,
sexually dimorphic species. Furthermore, the comparatively recent establishment on Mauritius and Réunion by Gal-
linula chloropus demonstrates the continuing ability of rails to colonise isolated island archipelagos.
Acknowledgements
I thank Robert Prŷs-Jones, Jacob Blokland, Marco Pavio and Trevor Worthy for their valued comments, and the
long discussions with Storrs Olson, which helped improve the paper; I thank Joel Cracraft and Paul Sweet (AMNH);
John Cooper, Lee Ismail and Beth Burr (BMB); Sandra Chapman, Mark Adams, Hein van Grouw, Judith White
and Joanne Cooper (NHMUK); Mike Brooke and Mathew Long (UMZC); Storrs Olson, Christopher Milensky and
Helen James (USNM); for access to specimens held in their care, especially Ronan Alain and Clare Sagne (MNHN)
who facilitated the long-term loan of the Thirioux fossil material; Paul Scofield (CMNZ) for details of speci-
mens held in his care and photography; Ben Warren (MNHN) for measurements; Harry Taylor (NHMUK),Justin
Jansen (RMNH Rijksmuseum van Natuurlijke Historie—Naturalis) and Mike Brooke (UMZC) for photography;
Alison Harding, Effie Warr and Angela Thresher (NHMUK) for help with reference material. I further thank Owen
Griffiths (MBC Mauritius Bioculture Ltd/FLMR); Greg Middleton, Steve Bourne, Liz Reed (AU Univeristy of
Adelaide/SAM South Australia Museum), Vincent Florens (University of Mauritius), Claudia Baider (National Her-
barium, Mauritius) and Julia Heinen (ZMUC Natural History Museum of Denmark) for field assistance and support;
and the Frank Chapman Award (AMNH); Dodo Research Foundation (DRF); DIF and Special Fund (NHMUK);
Synthesys (MNHN) and Percy Sladen Memorial Fund for financial assistance, which made the research for this
paper possible.
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APPENDIX 1.
The following skeletal specimens were used, all held in the AMNH, MNHN, NHMUK, UMZC and USNM bird skeleton col-
lections, with registration number and sex provided for each. Nomenclature follows Dickinson & Remsen (2013) and Gill et al.
(2010) for New Zealand taxa, wherein taxon authors and dates can be found for extant or historically extant taxa
Mentocrex kioloides NHMUK 1897.5.10.28 u/s; NHMUK 1929.8.4.10 u/s; NHMUK 1929.8.4.11 ♂;
Rallicula rubra AMNH 27133 ♂;
Gallirallus australis NHMUK 1863.7.30.10 u/s; NHMUK 1896.2.16.40 u/s; NHMUK 1955.5.16 u/s; NHMUK S/1952.2.363
u/s;
Hypotaenidia sylvestris NHMUK1939.12.9.3705 u/s; NHMUK1924.1.2.2 u/s;
Hypotaenidia owstoni USNM 501069 ♂; USNM 50168 ♂; USNM 501065 ♀; USNM 501067 ♀;
Hypotaenidia rovianae AMNH 30316 ♂; AMNH 28014 ♂; AMNH 30329 ♀; AMNH 30330 ♂;
Hypotaenidia insignis AMNH 27136 ♀;
Hypotaenidia woodfordi AMNH 27134 ♂;
Hypotaenidia wakensis USNM 289255 ♂; USNM 289256 ♀; AMNH 10803 ♂;
Rallus madagascariensis NHMUK 1897.5.10.26 ♀;
Dryolimnas c. cuvieri MNHN-ZO 2011-535 ♀; NHMUK 1897.5.10.47 u/s; NHMUK S/1882.2.28.2 u/s;
Dryolimnas c. abbotti NHMUK 1910.4.8.1 u/s;
Dryolimnas c. aldabranus NHMUK S/1989.38.7 ♂; NHMUK S/1989.38.1 ♂; NHMUK S/1989.38.6 ♂; NHMUK S/1989.38.3
♀; NHMUK S/1989.38.2 ♀; NHMUK S/1989.38.5 ♀; NHMUK S/1989.38.8 u/s; NHMUK S/1989.38.12 u/s; NHMUK
S/1993.6.1 u/s; NHMUK S/1993.6.3 u/s; NHMUK S/1993.6.4 u/s; NHMUK S/1993.6.2 u/s; NHMUK S/1989.38.10 u/s;
NHMUK S/1989.38.11 u/s; NHMUK S/1989.38.9 u/s; NHMUK S/1993.6.4 u/s; NHMUK S/1973.66.72 u/s; NHMUK
S/1986.27.1 u/s; NHMUK S/1979.5.25 u/s; NHMUK S/1981.13.35 u/s; NHMUK 1897.5.10.26 u/s; NHMUK 1868.9.12.19
u/s; NHMUK S/1961.11.1 u/s; NHMUK 1867.7.8.7 u/s; NHMUK S/1988.60.1 u/s; NHMUK S/1999.43.116 u/s; NHMUK
1897.5.10.25 u/s; USNM 615094 ♂;
Rougetius rougetii AMNH 27132 ♂;
Laterallus rogersi NHMUK 1924.1.2.2 ♂; USNM 319151 ♀; USNM 343633 u/s; AMNH 3931 ♀;
Amaurolimnas concolor concolor NHMUK 1847.6.16.86 u/s;
Amaurornis isabellina AMNH 27142 ♀;
Zapornia palmeri USNM 289242 u/s; USNM 289243 u/s; NHMUK S/1972.1.63 u/s; NHMUK 1927.2.3.3 u/s;
Eulabeornis castaneoventris AMNH 271138 #f;
Megacrex inepta AMNH 27137 ♀;
Porphyrio p. porphyrio NHMUK 1851.8.25.40 u/s;
Porphyrio p. melanotus NHMUK S/2002.46.3 ♂; NHMUK S/2002.46.4 ♀;
Porphyrio alleni NHMUK 1868.9.12.19 u/s; NHMUK S/1961.11.1 u/s; NHMUK 1867.7.8.7 u/s;
Porphyrio hochstetteri AMNH 26211 u/s; USNM 612797 u/s; NHMUK S/1961.17.1 u/s; NHMUK S/1974.9.3 u/s;
Gallinula c. chloropus NHMUK S/1977.23.1 u/s; NHMUK S/1973.66.72 ♂; NHMUK S/1986.27.1 ♂; NHMUK S/1979.5.25
♀; NHMUK S/1981.13.35 ♀; NHMUK S/1979.5.24 ♂; NHMUK S/1985.95.3 ♂; NHMUK S/1968.1.1 ♀; NHMUK
1894.9.14.1 ♀;
Gallinula c. pyrrhorrhoa NHMUK S/2001.50.357 u/s;
Gallinula c. galeata NHMUK 1904.8.5.33 u/s; NHMUK 1847.6.16.76 u/s;
Gallinula (Tribonyx) mortierii NHMUK S/1970.4.4 u/s; NHMUK S/1970.4.2 u/s; NHMUK S/1970.4.3 u/s;
Gallinula (Tribonyx) ventralis NHMUK S/1966.50.6 ♂; NHMUK S/1966.50.7 ♀; NHMUK 1870.6.22.5 u/s;
Gallinula nesiotis NHMUK S/1952.3.8 u/s;
Gallinula comeri NHMUK S/1958.15.1 ♂; NHMUK S/1974.19.3 u/s; NHMUK S/1974.19.2 ♂; NHMUK S/1974.19.1 ♀;
Pareudiastes pacificus AMNH 9700 u/s;
Fulica atra NHMUK S/1988.60.1 ♂; NHMUK S/1999.43.116 ♀; NHMUK 1859.9.6.420 u/s; NHMUK 1865.5.10.3 u/s;
NHMUK S/1966.52.44 u/s; NHMUK S/1979.12.2 u/s; NHMUK S/1987.3.1 u/s; NHMUK 1891.7.20.127 u/s; NHMUK
1930.3.24.63 u/s;
Fulica cristata AMNH 23329 ♀; NHMUK 1897.5.10.25 u/s; USNM 558475 ♂; USNM 558476 ♂; USNM 558477 ♂; USNM
321848 ♀; USNM 431788 ♀; USNM 430843 ♂;
Fulica americana NHMUK S/1984.77.16 ♂; NHMUK S/1977.81.3 ♀;
Fulica cornuta AMNH 10207 u/s;
Fulica gigantea AMNH 4542 u/s.
Subfossil material was referred to for Diaphorapteryx hawkinsi; Hypotaenidia dieffenbachii; Cabalus modestus; Laterallus
podarces (Wetmore, 1963), Hovacrex roberti (Andrews, 1897) and Fulica chathamensis Forbes, 1892a:
Diaphorapteryx hawkinsi AMNH 4667 u/s;
Hypotaenidia dieffenbachii USNM 559711 u/s;
Cabalus modestus NHMUK A966 u/s;
Laterallus podarces NHMUK S/1963.1.1 u/s; NHMUK S/1963.1.1 u/s;
Hovacrex roberti NHMUK A635 u/s;
Fulica chathamensis (all u/s), Cranium, NHMUK A1515; NHMUK A1515; NHMUK u/r; Rostrum, NHMUK A730; Mandible,
NHMUK A730; Scapula: NHMUK A730 (R); NHMUK A730 (L); Coracoid, NHMUK R3067 (R); NHMUK A968 (L);
NHMUK A730 (R); NHMUK A730 (L); Humerus: NHMUK A1602 (R); NHMUK A1602 (R); NHMUK A1602 (R);
NHMUK R3882 (L); NHMUK R3883 (L); NHMUK R3884 (L); Radius, NHMUK A730 (R); NHMUK A730 (L); Ulna:
NHMUK A730 (R); NHMUK A730 (L); Carpometacarpus, NHMUK A730 (L); NHMUK A730 (R); Sternum, NHMUK
R3073; NHMUK R3074; NHMUK R3075; NHMUK u/r; NHMUK u/r; NHMUK A961; Pelvis; NHMUK R3073;
NHMUK R3074; NHMUK R3076; NHMUK R1951; NHMUK R1952; NHMUK R1847; NHMUK R1848; NHMUK
R1849; NHMUK A1601; Femur: NHMUK u/r (L); NHMUK u/r L; NHMUK u/r (R); NHMUK u/r (L); NHMUK u/r (R;
NHMUK u/r (L); NHMUK R1953 (L); NHMUK R1954 (L); NHMUK R1955 (L); NHMUK R1956 (R); NHMUK R1957
(R); NHMUK R1958 (R); Tibiotarsus: NHMUK A296 (L); NHMUK A296 (R); NHMUK A1604 (R); NHMUK A1604
(R); NHMUK A1604 (L); NHMUK A1604 (R); NHMUK A1604 (L); NHMUK A1604 (R); NHMUK A1604 (L); NHMUK
A1604 (R); NHMUK A1604 (R; NHMUK R2069 (R); NHMUK R2070 (L); NHMUK R2071 (L); NHMUK R2072 (R);
NHMUK R2073 (L); NHMUK R2074 (L); NHMUK R2075 (L); Fibula, NHMUK A730 (L); NHMUK A730 (R); Tarso-
metatarsus: NHMUK R3025 (R); NHMUK R3026 (R); NHMUK R3027 (L); NHMUK R3028 (R); NHMUK R3029 (R);
NHMUK R3030 (L); NHMUK R3031 (R); NHMUK R3032 (R); NHMUK R3033 (R); NHMUK R3034 (R); NHMUK
R3035 (R); NHMUK R3036 (R); NHMUK R3037 (R); NHMUK R3038 (L); NHMUK R3039 (R); NHMUK A1605 (L);
NHMUK A1605 (R); NHMUK A1605 (R).
APPENDIX 2. Key to Abbreviations: TL = total length; WNF = width of naso-frontal hinge; LN = length of narial
opening; RML = rostrum maxillare length; RMW = rostrum maxillare width; RMD = rostrum maxillare depth;
RMaL = rostrum mandibulae length; RMaW = rostrum mandibulae width; RMaD = rostrum mandibulae depth;
LC = greatest length of carina sterni; DK = greatest depth of keel; LW = least width; PW = proximal width; GW =
greatest width; GD = greatest depth; PD = proximal depth; SW = shaft width; SD = shaft depth; DW = distal width;
DD = distal depth; (n) = number of specimens; (m) = mean; SD = standard deviation.

TABLE 1. Measurements (mm) of the cranium of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD GW (n) range GW (m) SD GD(n) range GD (m) SD WNF (n) range WNF (m) SD
†Aphanapteryx bonasia (2) 38.0–48.7 (43.3) 7.56 (2) 30.0–34.8 (32.4) 3.39 (2) 25.1–26.1 (25.6) 0.07 (2) 14.0–14.8 (14.4) 0.56
†Aphanapteryx bonasia Associated indi- (1) 48.7 (48.7) (1) 34.8 (34.8) (1) 26.1 (26.1) (1) 14.8 (14.8)
vidual MU 923
†Erythromachus leguati (5) 37.3–40.4 (39.4) 1.26 (6) 21.1–22.0 (21.4) 0.33 (6) 19.0–21.9 (20.6) 1.07 (5) 9.1–9.8 (9.4) 0.25
Mentocrex kioloides (1) 32.3 (32.3) (1) 19.7 (19.7) (1) 19.0 (19.0) (1) 9.3 (9.3)
Rallicula rubra (1) 24.7 (24.7) (1) 17.4 (17.4) - - (1) 7.1 (7.1)
Gallirallus australis (4) 39.3–44.9 (41.5) 2.36 (4) 24.7–27.9 (25.8) 1.42 (4) 22.5–24.9 (23.5) 1.10 (4) 9.2–10.9 (9.7) 0.79
Hypotaenidia sylvestris (2) 36.9–38.6 (37.7) 1.20 (2) 23.2–23.3 (23.2) 0.07 (2) 20.5–21.4 (20.9) 0.63 (2) 10.4–10.8 (10.6) 0.28
Hypotaenidia owstoni (4) 32.4–34.8 (33.7) 1.14 (4) 18.9–20.3 (19.6) 0.62 (4) 17.8–18.2 (17.9) 0.19 (4) 6.6–7.2 (6.8) 0.26
Hypotaenidia rovianae (4) 37.8–39.4 (38.6) 0.67 (4) 23.2–24.7 (24.0) 0.70 (4) 20.9–21.8 (21.2) 0.38 (2) 8.2–9.1 (8.6) 0.49
†Hypotaenidia wakensis (2) 25.4–26.6 (26.0) 0.84 (2) 17.0–17.8 (17.4) 0.56 (2) 15.4–16.4 (15.9) 0.70 (2) 6.5–6.8 (6.6) 0.21
†Hypotaenidia dieffenbachii (1) 37.9 (37.9) (1) 23.4 (23.4) (1) 20.4 (20.4) (1) 9.0 (9.0)
†Diaphorapteryx hawkinsi (1) 48.3 (48.3) (1) 44.1 (44.1) (1) 30.3 (30.3) (1) 17.6 (17.6)
Rallus madagascariensis (1) 27.4 (27.4) (1) 15.2 (15.2) (1) 15.9 (15.9) (1) 5.7 (5.7)
RAILS (AVES: RALLIDAE) OF THE MASCARENE ISLANDS

Dryolimnas cuvieri cuvieri (2) 32.6–33.2 (32.9) 0.42 (2) 19.4–20.3 (19.8) 0.63 (2) 19.7–19.9 (19.8) 0.14 (2) 7.4–8.2 (7.8) 0.56
Dryolimnas cuvieri aldabranus (13) 29.4–32.1 (30.7) 0.89 (12) 16.3–19.1 (17.7) 0.81 (13) 16.5–18.8 (17.8) 0.68 (11) 5.7–8.1 (7.0) 0.65
†Dryolimnas cuvieri abbotti (1) 31.6 (31.6) (1) 19.0 (19.0) (1) 18.6 (18.6) (1) 6.6 (6.6)
Rougetius rougetii (1) 32.6 (32.6) (1) 18.3 (18.3) (1) 20.0 (20.0) (1) 6.3 (6.3)
Laterallus rogersi (2) 20.8–22.0 (20.8) 0.84 (2) 12.9–13.4 (13.1) 0.35 (2) 11.7–12.4 (12.0) 0.49 (2) 3.7–3.9 (3.8) 0.14
Amaurornis isabellina - - (1) 21.0 (21.0) - - (1) 8.0 (8.0)
†Zapornia palmeri (4) 19.9–21.8 (20.8) 0.77 (4) 11.2–13.4 (12.7) 1.02 (4) 12.0–13.4 (12.4) 0.47 (4) 3.7–4.7 (4.2) 0.42
Eulabeornis castaneoventris - - (1) 22.0 (22.0) - - (1) 9.0 (9.0)
Porphyrio porphyrio melanotus (2) 39.5–39.6 (39.5) 0.07 (2) 26.8–27.1 (26.9) 0.21 (2) 25.2–25.9 (25.5) 0.49 (2) 17.6–18.5 (18.0) 0.63
Porphyrio alleni (3) 26.4–30.7 (28.1) 2.26 (3) 17.3–20.3 (18.4) 1.60 (3) 17.1–18.4 (17.7) 0.65 (3) 8.3–9.4 (8.7) 0.60
Porphyrio hochstetteri (4) 49.1–51.3 (49.9) 0.95 (4) 35.5–37.5 (36.6) 0.85 (4) 29.4–30.9 (30.5) 0.75 (4) 24.2–26.4 (25.1) 0.91
Gallinula chloropus (8) 29.3–33.7 (31.4) 1.41 (8) 17.4–19.4 (18.3) 0.56 (8) 16.8–20.3 (18.3) 1.12 (8) 7.1–8.8 (8.2) 0.49
Gallinula mortierii (3) 41.0–42.0 (41.6) 0.52 (3) 24.0–26.0 (25.0) 1.00 (3) 23.7–25.3 (24.3) 0.83 (3) 11.5–12.1 (11.7) 0.30
Gallinula ventralis (3) 32.5–34.9 (34.0) 1.33 (3) 20.4–21.1 (20.7) 0.36 (3) 18.5–21.3 (19.9) 1.40 (3) 8.2–8.3 (8.2) 0.05
Gallinula comeri (1) 34.3 (34.3) (1) 22.7 (22.7) (1) 21.1 (21.1) (1) 8.8 (8.8)
Fulica atra (9) 25.2–39.2 (36.0) 4.18 (9) 22.5–24.7 (23.5) 0.74 (9) 18.5–21.6 (20.5) 0.93 (9) 8.9–12.0 (10.8) 1.01
Fulica cristata (7) 33.7–38.6 (36.1) 1.79 (7) 22.3–24.4 (23.3) 0.85 (7) 20.5–23.6 (22.2) 1.20 (7) 9.7–11.9 (10.7) 0.76
Fulica americana (2) 32.9–34.4 (33.6) 1.06 (2) 20.7–22.6 (21.6) 1.34 (2) 18.9–20.3 (19.6) 0.98 (2) 9.3–10.8 (10.0) 1.06
Fulica cornuta (1) 42.1 42.1 (1) 29.8 29.8 (1) 24.0 24.0 (1) 14.6 14.6
Zootaxa 4626 (1) © 2019 Magnolia Press ·

†Fulica chathamensis (3) 40.1–42.4 (41.5) 1.22 (3) 27.0–27.7 (27.3) 0.35 (3) 21.5–22.6 (22.0) 0.55 (3) 12.0–13.1 (12.5) 0.55
83
TABLE 2. Measurements (mm) of the rostrum of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD PW (n) range PW (m) SD PD(n) range PD (m) SD RML (n) range
†Aphanapteryx bonasia (6) 74.8–86.0 (81.4) 4.0 (4) 9.0–14.3 (10.6) 2.48 (2) 8.5–12.0 (10.2) 1.92 (7) 35.6–40.2
†Aphanapteryx bonasia Associated (1) 86.0 (86.0) (1) 14.3 (14.3) (1) 12.0 (12.0) (1) 40.0
individual MU 923
†Erythromachus leguati (3) 76.6–80.8 (78.1) 2.29 (3) 7.9–8.5 (8.1) 0.30 (2) 7.7–8.1 (7.9) 0.28 (3) 28.1–29.0
Mentocrex kioloides (1) 27.4 (27.4) (1) 8.8 (8.8) (1) 6.8 (6.8) (1) 9.4
Rallicula rubra (1) 24.7 (24.7) (1) 7.4 (7.4) (1) 5.6 (5.6) (1) 8.2
Gallirallus australis (4) 44.7–49.2 (46.1) 2.10 (4) 8.9–10.0 (9.3) 0.47 (4) 10.4–10.9 (10.6) 0.23 (4) 14.8–16.3
Hypotaenidia sylvestris (2) 49.2–50.1 (49.6) 0.63 (2) 8.5–9.0 (8.7) 0.35 (2) 9.1–9.7 (9.4) 0.42 (2) 18.4–18.5
Hypotaenidia owstoni (4) 33.9–37.4 (35.7) 1.75 (4) 6.9–7.7 (7.3) 0.33 (4) 6.2–7.1 (6.6) 0.46 (4) 11.1–13.6
Hypotaenidia rovianae (4) 38.1–44.0 (41.6) 2.69 (4) 7.7–8.6 (8.1) 0.46 (4) 8.0–9.3 (8.8) 0.57 (4) 12.6–14.3
†Hypotaenidia wakensis (2) 23.6–28.7 (26.1) 3.60 (2) 5.3–6.1 (5.7) 0.56 (2) 5.–5.6 (5.3) 0.42 (2) 6.4–9.3
†Hypotaenidia dieffenbachii (1) 40.5 (40.5) (1) 7.4 (7.4) (1) 8.1 (8.1) (1) 12.6
Diaphorapteryx hawkinsi (1) 86.8 (86.8) (1) 15.8 (15.8) (1) 14.4 (14.4) (1) 47.1
Rallus madagascariensis (1) 38.4 (38.4) (1) 5.1 (5.1) (1) 5.0 (5.0) (1) 11.5
84 · Zootaxa 4626 (1) © 2019 Magnolia Press

Dryolimnas cuvieri cuvieri (2) 39.7–45.9 (42.8) 4.38 (2) 7.8–8.2 (8.0) 0.28 (2) 6.2–6.7 (6.4) 0.35 (2) 12.2–12.4
Dryolimnas cuvieri aldabranus (13) 36.8–47.1 (41.3) 3.69 (13) 6.1–8.0 (6.8) 0.54 (5) 5.3–7.0 (6.1) 0.81 (13) 11.4–17.5
†Dryolimnas cuvieri abbotti (1) 41.1 (41.7) (1) 7.1 (7.1) (1) 6.2 (6.2) (1) 12.4
†Dryolimnas chekei - - - - - - (1) 14.7
Rougetius rougetii (1) 35.2 (35.2) (1) 7.5 (7.5) (1) 6.1 (6.1) (1) 9.2
Laterallus rogersi (2) 20.6–22.5 (21.5) 1.34 (2) 3.5–3.7 (3.6) 0.14 (2) 3.6–4.1 (3.8) 0.35 (2) 6.0–7.0
Amaurornis isabellina (1) 33.5 (33.5) (1) 8.8 (8.8) (1) 9.1 (9.1) (1) 12.3
†Zapornia palmeri (4) 17.7–21.1 (19.7) 1.50 (4) 3.9–4.6 (4.3) 0.29 (4) 3.7–4.5 (3.9) 0.37 (4) 6.6–7.7
Eulabeornis castaneoventris (1) 49.2 (49.2) (1) 9.6 (9.6) (1) 8.8 (8.8) (1) 16.1
Porphyrio porphyrio melanotus (2) 44.0–47.4 (45.7) 2.40 (2) 13.1–14.4 (13.7) 0.91 (2) 16.5–17.7 (17.1) 0.84 (2) 22.6–23.5
Porphyrio alleni (3) 21.9–26.2 (23.8) 2.18 (3) 7.1–8.1 (7.6) 0.51 (3) 7.1–8.2 (7.7) 0.55 (3) 10.8–11.3
Porphyrio hochstetteri (4) 61.5–64.0 (62.8) 1.08 (4) 16.0–25.2 (20.7) 4.57 (4) 27.4–28.1 (27.9) 0.33 (4) 21.8–34.6
Gallinula chloropus (9) 24.8–31.7 (27.8) 1.96 (9) 7.6–9.0 (8.1) 0.57 (9) 7.0–8.7 (9.6) 1.24 (9) 8.5–11.9
Gallinula mortierii (3) 32.5–35.4 (34.3) 1.59 (3) 11.7–13.0 (12.4) 0.68 (3) 10.9–11.9 (11.3) 0.51 (3) 8.3–9.8
Gallinula ventralis (3) 26.4–29.4 (28.0) 1.52 (3) 8.6–9.6 (9.0) 0.51 (3) 8.6–8.8 (8.7) 0.11 (3) 7.3–8.5
Gallinula comeri (1) 31.4 (31.4) (1) 8.5 (8.5) (1) 9.3 (9.3) (1) 11.1
Fulica atra (9) 32.3–39.8 (36.7) 2.37 (9) 8.0–11.5 (10.2) 1.09 (9) 9.5–12.4 (10.5) 1.02 (9) 10.9–14.2
Fulica cristata (7) 29.7–38.9 (35.1) 3.62 (7) 8.6–11.4 (9.9) 1.01 (7) 9.3–10.5 (9.8) 0.44 (6) 10.9–14.1
Fulica americana (2) 34.1–35.6 (34.8) 1.06 (2) 8.6–9.2 (8.9) 0.42 (2) 10.0–10.7 (10.3) 0.49 (2) 10.0–10.9
Fulica cornuta (1) 54.0 (54.0) (1) 14.9 (14.9) (1) 12.2 (12.2) (1) 18.0
†Fulica newtonii - - - - - - (1) 14.9
†Fulica chathamensis (1) 49.0 (49.0) (1) 12.7 (12.7) (1) 12.1 (12.1) (1) 13.9
HUME
TABLE 2. (Continued)
Species RML (m) SD RMW (n) range RMW (m) SD RMD (n) range RMD (m) LNO (n) range LNO (m) SD
†Aphanapteryx bonasia (38.2) 1.92 (6) 4.7–9.5 (5.8) 1.81 (5) 5.1–6.5 (5.7) 0.54 (4) 33.0–41.0 (37.1) 3.65
†Aphanapteryx bonasia Associ- (40.0) (1) 5.9 (5.9) (1) 6.5 (6.5) (1) 35.3 (35.3)
ated individual MU 923
†Erythromachus leguati (28.5) 0.45 (4) 3.3–3.7 (3.5) 0.18 (4) 3.4–3.8 (3.6) 0.17 (3) 43.0–47.1 (44.8) 2.09
Mentocrex kioloides (9.4) (1) 4.3 (4.3) (1) 3.9 (3.9) (1) 15.5 (15.5)
Rallicula rubra (8.2) (1) 3.1 (3.1) (1) 3.0 (3.0) (1) 15.7 (15.7)
Gallirallus australis (15.5) 0.25 (4) 4.8–5.4 (5.1) 0.25 (4) 4.8–5.4 (5.1) 0.26 (4) 24.4–26.7 (25.7) 1.14
Hypotaenidia sylvestris (18.4) 0.07 (2) 3.8–3.9 (3.8) 0.07 (2) 4.1–4.2 (4.1) 0.07 (2) 25.6–26.9 (26.2) 0.91
Hypotaenidia owstoni (12.3) 1.05 (4) 4.0–4.4 (4.2) 0.17 (4) 3.2–3.5 (3.3) 0.17 (4) 19.1–20.9 (20.0) 0.93
Hypotaenidia rovianae (13.1) 0.78 (4) 3.8–4.5 (4.1) 0.35 (4) 3.7–4.3 (4.0) 0.25 (4) 21.2–25.9 (24.5) 2.22
†Hypotaenidia wakensis (7.8) 2.05 - - - - (2) 11.9–15.4 (13.6) 2.47
†Hypotaenidia dieffenbachii (12.6) (1) 4.2 (4.2) (1) 4.6 (4.6) (1) 24.9 (24.9)
Diaphorapteryx hawkinsi (47.1) (1) 8.8 (8.8) (1) 7.1 (7.1) (1) 29.9 (29.9)
Rallus madagascariensis (11.5) (1) 2.1 (2.1) (1) 1.8 (1.8) (1) 27.0 (27.0)
Dryolimnas cuvieri cuvieri (12.3) 0.14 (2) 3.0–3.2 (3.1) 0.14 (2) 3.0–3.1 (3.0) 0.07 (2) 23.9–29.5 (26.7) 3.95

Dryolimnas cuvieri aldabranus (13.8) 1.85 (13) 2.8–3.9 (3.4) 0.36 (13) 2.4–4.0 (3.3) 0.51 (13) 20.7–29.2 (25.2) 2.56
†Dryolimnas cuvieri abbotti (12.4) (1) 3.2 (3.2) (1) 3.1 (3.1) (1) 26.3 (26.3)
†Dryolimnas chekei (14.7) (1) 3.4 (3.4) - - (1) 28.0 (28.0)
Rougetius rougetii (9.2) (1) 3.1 (3.1) (1) 2.4 (2.4) (1) 22.3 (22.3)
Laterallus rogersi (6.5) 0.70 (1) 1.9 (1.9) (1) 2.0 (2.0) (2) 12.1–12.2 (12.1) 0.07
Amaurornis isabellina (12.3) (1) 5.4 (5.4) (1) 4.3 (4.3) (1) 16.8 (16.8)
†Zapornia palmeri (7.0) 0.45 (2) 2.0 (2.0) (2) 2.0–2.3 (2.1) 0.21 (4) 8.8–10.1 (9.3) 0.57
Eulabeornis castaneoventris (16.1) (1) 4.4 (4.4) (1) 4.5 (4.5) (1) 29.1 (29.1)
Porphyrio porphyrio melanotus (23.0) 0.63 (2) 7.4–8.8 (8.1) 0.98 (2) 11.0–11.4 (11.2) 0.28 (2) 12.6–12.9 (12.7) 0.21
Porphyrio alleni (11.1) 0.28 (3) 4.1–5.2 (4.5) 0.60 (3) 3.8–4.7 (4.3) 0.49 (3) 8.8–10.0 (9.3) 0.61
Porphyrio hochstetteri (29.7) 5.5 (2) 12.2–13.5 (12.8) 0.91 (2) 17.6–19.2 (18.4) 1.13 (4) 14.5–15.5 (14.8) 0.46
Gallinula chloropus (9.6) 1.24 (9) 3.4–5.1 (4.0) 0.58 (9) 3.4–4.7 (4.1) 0.49 (9) 13.4–16.9 (15.0) 1.03
Gallinula mortierii (9.1) 0.76 (3) 5.7–6.1 (5.9) 0.20 (3) 5.0–5.6 (5.3) 0.30 (3) 17.4–19.2 (18.4) 0.94
Gallinula ventralis (8.1) 0.69 (3) 5.0–5.1 (5.0) 0.05 (3) 4.0–4.8 (4.3) 0.40 (3) 13.8–16.2 (15.2) 1.24
Gallinula comeri (11.1) (1) 5.3 (5.3) (1) 5.3 (5.3) (1) 14.5 (14.5)
Fulica atra (12.8) 1.01 (9) 4.7–6.6 (5.7) 0.65 (9) 4.6–5.8 (5.2) 0.40 (9) 15.8–19.9 (17.6) 1.39
Fulica cristata (12.6) 1.04 (7) 5.0–6.1 (5.5) 0.42 (7) 4.2–5.4 (4.7) 0.42 (7) 15.3–18.8 (17.0) 1.19
Fulica americana (10.4) 0.63 (2) 4.9–5.9 (5.4) 0.70 (2) 4.8–5.0 (4.9) 0.14 (2) 17.9–18.2 (18.0) 0.21
Fulica cornuta (18.0) (1) 8.4 (8.4) (1) 6.8 (6.8) (1) 25.8 (25.8)
†Fulica newtonii (14.9) (1) 7.0 (7.0) (1) 5.5 (5.5) - -

†Fulica chathamensis (13.9) (1) 8.0 (8.0) (1) 6.5 (6.5) (1) 23.2 (23.2)
85
TABLE 3. Measurements (mm) of the mandible of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD PW (n) range PW (m) SD PD(n) range PD (m) SD
†Aphanapteryx bonasia (3) 105.7–122.0 (114.9) 8.36 (3) 11.0–12.5 (11.5) 0.81 (3) 9.5–11.9 (10.6) 1.20
†Aphanapteryx bonasia Associated individual MU 923 (1) 122.0 (122.0) (1) 11.0 (11.0) (1) 11.9 (11.9)
†Erythromachus leguati (1) 99.9 (99.9) (2) 7.4–9.2 (8.3) (1.27) (2) 5.7–7.8 (6.7) (1.48)
Mentocrex kioloides (1) 45.6 (45.6) (1) 6.1 (6.1) (1) 4.7 (4.7)
Rallicula rubra (1) 39.2 (39.2) (1) 3.4 (3.4) (1) 3.1 (3.1)
Gallirallus australis (3) 69.9–75.1 (72.1) 2.67 (3) 8.0–9.1 (8.5) 0.55 (3) 6.5–8.8 (8.0) 1.30
Hypotaenidia sylvestris (2) 71.0–71.3 (71.1) 0.21 (2) 8.3–8.7 (8.5) 0.28 (2) 6.1–6.4 (6.2) 0.21
Hypotaenidia owstoni (4) 52.8–56.7 (54.7) 2.11 (4) 6.1–6.9 (6.4) 0.33 (4) 4.5–5.3 (4.9) 0.36
Hypotaenidia rovianae (4) 61.8–68.2 (66.1) 2.90 (4) 7.6–7.7 (7.6) 0.05 (4) 5.8–6.4 (6.0) 0.25
†Hypotaenidia wakensis (2) 37.6–44.3 (40.9) 4.73 (2) 4.7–5.5 (5.1) 0.56 (2) 3.1–3.3 (3.2) 0.14
†Diaphorapteryx hawkinsi (1) 110.3 (110.3) (1) 14.5 (14.5) (1) 11.7 (11.7)

Rallus madagascariensis (1) 53.8 (53.8) (1) 5.4 (5.4) (1) 4.2 (4.2)
Dryolimnas cuvieri cuvieri (2) 59.8–65.8 (62.8) 4.24 (2) 6.4–7.0 (6.7) 0.84 (2) 4.5–5.7 (5.1) 0.14
Dryolimnas cuvieri aldabranus (13) 54.1–69.3 (61.5) 5.73 (13) 5.2–6.5 (5.9) 0.45 (13) 4.0–5.4 (4.6) 0.39
†Dryolimnas cuvieri abbotti (1) 60.9 (60.9) (1) 5.5 (5.5) (1) 4.9 (4.9)
Laterallus rogersi (2) 32.0–32.4 (32.2) 0.28 (2) 3.4–4.0 (3.7) 0.42 (2) 2.9 (2.9)
Amaurornis isabellina (1) 52.3 (52.3) (1) 6.9 (6.9) (1) 5.2 (5.2)
†Zapornia palmeri (4) 28.3–31.1 (29.7) 1.22 (4) 3.6–4.3 (3.9) 0.31 (4) 2.2–2.7 (2.4) 0.20
Eulabeornis castaneoventris (1) 72.0 (72.0) (1) 8.2 (8.2) (1) 6.6 (6.6)
Porphyrio porphyrio melanotus (2) 61.3–63.4 (62.3) 1.48 (2) 10.1–11.0 (10.5) 0.63 (2) 8.2–9.7 (8.9) 1.06
Porphyrio alleni (3) 35.1–41.7 (38.0) 3.36 (3) 5.3–6.6 (5.8) 0.70 (3) 3.6–5.2 (4.3) 0.80
Porphyrio hochstetteri (4) 81.2–85.0 (83.0) 2.05 (4) 14.6–16.5 (15.4) 0.80 (4) 11.7–17.0 (13.4) 2.42
Gallinula chloropus (9) 41.4–48.1 (45.5) 2.20 (9) 5.4–6.6 (5.9) 0.43 (9) 3.8–4.4 (4.1) 0.20
Gallinula mortierii (3) 56.9–59.0 (57.6) 1.15 (3) 9.4–10.0 (9.6) 0.32 (3) 5.9–6.3 (6.1) 0.20
Gallinula ventralis (3) 42.6–47.3 (45.4) 2.51 (3) 6.5–7.4 (6.9) 0.45 (3) 4.0–4.2 (4.1) 0.10
Gallinula comeri (1) 52.4 (53.4) (1) 6.8 (6.8) (1) 4.9 (4.9)
Fulica atra (9) 52.1–63.2 (57.3) 3.62 (9) 7.7–8.7 (8.2) 0.32 (9) 5.0–6.4 (5.7) 0.40
Fulica cristata (7) 48.0–58.9 (53.2) 3.86 (7) 6.7–8.7 (7.8) 0.76 (7) 5.0–5.7 (5.3) 0.28
Fulica americana (2) 52.3–54.1 (53.2) 1.27 (2) 6.9–7.1 (7.0) 0.14 (2) 4.9–5.6 (5.2) 0.49
Fulica cornuta (1) 74.0 (74.0) (1) 11.2 (11.2) (1) 8.5 (8.5)
†Fulica chathamensis (1) 67.1 (67.1) (1) 10.1 (10.1) (1) 8.2 (8.2)
HUME
Species RMaL (n) range RMaL (m) SD RMaW (n) range RMaW (m) RMaD (n) range RMaD (m) SD
†Erythromachus leguati (1) 25.8 (25.8) (1) 3.6 (3.6) (1) 2.9 (2.9)
Rallicula rubra (1) 9.9 (9.9) (1) 3.0 (3.0) (1) 1.9 (1.9)

Laterallus rogersi (2) 5.5–6.6 (6.0) 0.77 (2) 1.9 (1.9) (2) 1.5–1.6 (1.5) 0.07
Fulica atra (9) 7.3–11.8 (9.5) 1.56 (9) 3.2–5.1 (4.2) 0.67 (9) 3.0–4.9 (3.7) 0.64
Fulica cristata (7) 6.8–10.0 (8.2) 1.15 (7) 4.0–4.8 (4.4) 0.29 (7) 2.9–3.4 (3.2) 0.18
Fulica cornuta (1) 13.7 (13.7) (1) 7.3 (7.3) (1) 6.2 (6.2)

†Fulica chathamensis (1) 11.6 (11.6) (1) 6.2 (6.2) (1) 5.5 (5.5)
87
TABLE 4. Measurements (mm) of the scapula of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD PW (n) range PW (m) SD PD (n) range PD (m) SD
†Erythromachus leguati - - (1) 7.0 (7.0) (1) 3.2 (3.2)
Mentocrex kioloides (3) 35.8–36.7 (36.1) 0.47 (3) 5.9–6.7 (6.2) 0.43 (3) 2.6–2.9 (2.7) 0.15

Laterallus rogersi (3) 19.5–19.7 (19.6) 0.14 (3) 2.8 (2.8) (3) 1.1–1.2 (1.1) 0.07
†Laterallus podarces (1) 33.1
†Amaurolimnas concolor concolor (1) 35.1 (35.1) (1) 6.2 (6.2) (1) 2.3 (2.3)
Porphyrio porphyrio porphyrio (1) 63.8 (63.8) (1) 12.3 (12.3) (1) 5.5 (5.5)
Gallinula chloropus chloropus (8) 41.8–54.1 (48.6) 4.08 (8) 6.5–8.6 (7.4) 0.83 (8) 2.5–3.5 (3.0) 0.38
Gallinula chloropus pyrrhorrhoa (1) 43.5 (43.5) (1) 7.7 (7.7) (1) 2.8 (2.8)
Gallinula chloropus galeata (2) 47.2–49.5 (48.3) 1.62 (2) 7.9–8.0 (7.9) 0.07 (1) 2.8 (2.8)
Gallinula comeri (3) 47.6–49.9 (49.0) 1.27 (3) 8.5–9.3 (8.8) 0.41 (3) 3.5–4.3 (3.8) 0.43
Gallinula nesiotis (1) 44.8 (44.8) (1) 8.2 (8.2) (1) 3.0 (3.0)
Fulica atra (9) 60.0–68.7 (65.3) 3.00 (9) 9.2–11.0 (10.1) 0.60 (9) 3.2–4.6 (3.7) 0.46
Fulica cristata (7) 60.2–68.2 (64.0) 3.13 (7) 9.8–10.8 (10.1) 0.35 (7) 3.4–3.9 (3.6) 0.20
Fulica cornuta (1) 85.8 (85.8) (1) 15.2 (15.2) (1) 6.0 (6.0)
Fulica gigantea (1) 69.2 (69.2) (1) 12.4 (12.4) (1) 3.9 (3.9)
†Fulica newtonii (1) 64.9 (64.9) (2) 11.5–11.9 (11.7) (0.28) (2) 4.1–4.3 (4.2) (0.14)
†Fulica chathamensis (2) 71.0–71.1 (71.0) 0.07 (2) 11.7–11.8 (11.7) 0.07 (2) 3.5–4.5 (4.0) 0.70
HUME
TABLE 5. Measurements (mm) of the coracoid of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD SW (n) range SW (m) SD SD (n) range SD (m) SD DW (n) range DW (m)
vidual MU 923
†Erythromachus leguati (2) 23.5–25.0 (24.2) 1.06 (2) 3.3–3.5 (3.4) 0.14 (2) 2.1 (2.1) (2) 8.9 (8.9)
Mentocrex kioloides (3) 22.6–23.8 (23.0) 0.69 (3) 3.2–3.6 (3.3) 0.23 (3) 1.9–2.1 (2.0) 0.10 (3) 7.6–9.3 (8.4) 0.85

†Amaurolimnas concolor concolor (1) 20.6 (20.6) (1) 2.3 (2.3) (1) 1.9 (1.9) (1) 7.5 (7.5)
Porphyrio porphyrio porphyrio (1) 38.5 (38.5) (1) 5.0 (5.0) (1) 3.2 (3.2) (1) 14.3 (14.3)
Gallinula comeri (3) 26.4–26.7 (26.5) 0.15 (3) 2.9–3.2 (3.1) 0.17 (3) 2.4–2.5 (2.4) 0.05 (3) 11.3–12.4 (11.9) 0.55
†Gallinula nesiotis (1) 25.0 (25.0) (1) 2.7 (2.7) (1) 2.3 (2.3) (1) 8.4 (8.4)
Fulica atra (9) 32.7–37.0 (34.9) 1.51 (9) 4.1–5.0 (4.4) 0.29 (9) 2.6–3.5 (3.0) 0.25 (9) 13.8–16.8 (15.5) 1.14
Fulica cristata (7) 32.7–38.8 (35.6) 2.39 (7) 3.9–4.9 (4.4) 0.33 (7) 2.8–3.4 (3.0) 0.25 (7) 11.7–17.1 (15.3) 1.84
Fulica cornuta (1) 47.5 (47.5) (1) 6.7 (6.7) (1) 4.8 (4.8) (1) 17.6 (17.6)
Fulica gigantea (1) 43.0 (43.0) (1) 5.1 (5.1) (1) 3.7 (3.7) (1) 18.5 (18.5)

89
TABLE 6. Measurements (mm) of the sternum of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD GW (n) range GW (m) SD LW (n) range
†Aphanapteryx bonasia (1) 59.5 (59.5) (2) 26.6–29.0 (27.8) 1.69 (2) 16.8–18.0
†Aphanapteryx bonasia Associated individual MU 923 (1) 59.5 (59.5) (1) 26.6 (26.6) (1) 16.8
†Erythromachus leguati (4) 45.2–47.3 (46.1) 0.92 (5) 23.8–26.8 (25.1) 1.11 (5) 9.9–11.5
Mentocrex kioloides (3) 46.4–47.8 (46.9) 0.73 (2) 12.3–14.8 (13.5) 1.76 (2) 8.2–9.2
Gallirallus australis (4) 43.7–50.6 (46.6) 3.48 (4) 19.1–23.1 (21.4) 1.77 (4) 8.4–11.6
Hypotaenidia sylvestris (2) 39.6–41.2 (40.4) 1.13 (2) 17.1–18.1 (17.6) 0.70 (2) 9.9–10.4
Hypotaenidia owstoni (4) 31.9–37.0 (35.3) 2.36 (4) 11.3–12.8 (12.2) 0.68 (4) 7.0–7.6
Hypotaenidia rovianae (4) 45.8–48.8 (47.4) 1.41 (4) 21.8–25.4 (23.1) 1.63 (4) 9.1–11.5
†Hypotaenidia wakensis (3) 23.4–30.3 (26.8) 3.45 (3) 10.4–14.3 (12.1) 1.98 (3) 6.7–8.6
†Diaphorapteryx hawkinsi (1) 59.0 (59.0) (1) 30.1 (30.1) (1) 15.6
Rallus madagascariensis (1) 33.0 (33.0) (1) 9.2 (9.2) (1) 5.3
Dryolimnas cuvieri cuvieri (3) 44.2–51.6 (48.9) 4.08 (3) 13.1–14.9 (13.7) 1.03 (3) 7.2–8.1
Dryolimnas cuvieri aldabranus (16) 34.8–38.3 (36.7) 1.14 (14) 12.1–13.6 (12.7) 0.49 (15) 6.3–7.9

†Dryolimnas cuvieri abbotti (1) 43.7 (43.7) (1) 13.5 (13.5) (1) 7.7
Laterallus rogersi (3) 14.1–15.0 (14.6) 0.45 (3) 7.9–8.6 (8.1) 0.37 (3) 5.5–6.1
†Amaurolimnas concolor concolor (1) 39.1 (39.1) (1) 11.3 (11.3) (1) 8.0
†Zapornia palmeri (3) 17.1–19.1 (18.0) 1.00 (3) 8.4–9.6 (9.0) 0.61 (3) 5.9–6.3
Porphyrio porphyrio porphyrio (1) 67.9 (67.9) (1) 20.2 (20.2) (1) 15.3
Porphyrio porphyrio melanotus (2) 70.6–73.3 (71.9) 1.90 (2) 21.2–23.0 (22.1) 1.27 (2) 15.0–16.8
Porphyrio alleni (3) 35.6–44.7 (38.8) 5.11 (3) 11.3–14.3 (12.5) 1.55 (3) 7.5–9.4
Porphyrio hochstetteri (4) 65.5–67.8 (66.6) 0.97 (4) 34.0–37.3 (35.7) 1.69 (4) 23.4–25.2
Gallinula chloropus chloropus (8) 42.5–50.4 (47.2) 2.66 (8) 13.9–18.1 (16.6) 1.25 (8) 10.8–14.0
Gallinula chloropus pyrrhorrhoa (1) 48.0 (48.0) (1) 17.0 (17.0) (1) 13.8
Gallinula chloropus galeata (2) 48.6–48.7 (48.6) 0.07 (2) 17.6–19.0 (18.3) 0.98 (2) 13.7–14.1
Gallinula mortierii (3) 55.2–56.8 (56.2) 0.87 (3) 38.4–40.4 (39.3) 1.01 (3) 18.7–20.0
Gallinula ventralis (3) 48.8–59.2 (52.9) 5.53 (3) 23.7–24.2 (23.9) 0.25 (3) 13.0–13.5
Gallinula comeri (3) 44.1–45.2 (44.5) 0.60 (3) 19.1–20.5 (19.7) 0.70 (3) 16.2–16.3
†Gallinula nesiotis (1) 37.6 (37.6) (1) 15.8 (15.8) (1) 12.8
Fulica atra (10) 57.2–68.1 (63.4) 4.01 (10) 20.5–25.8 (23.5) 1.52 (10) 17.7–22.5
Fulica cristata (7) 58.6–69.9 (64.0) 4.14 (7) 23.2–27.0 (25.0) 1.31 (7) 19.5–22.4
Fulica americana (2) 54.6–59.3 (56.9) 3.32 (2) 20.6–23.6 (22.1) 2.12 (2) 16.0–18.3
Fulica cornuta (1) 83.9 (83.9) (1) 39.5 (39.5) (1) 29.0
Fulica gigantea (1) 75.7 (75.7) (1) 36.6 (36.6) (1) 24.6
†Fulica newtonii (1) 69.7 (69.7) (1) 36.9 (36.9) (1) 21.6
†Fulica chathamensis (4) 74.7–77.5 (76.5) 1.26 (3) 43.0–46.0 (44.0) 1.67 (6) 24.3–28.3
HUME
Species LW (m) SD LC (n) range LC (m) SD DK (n) range DK (m) SD
†Aphanapteryx bonasia (17.4) 0.84 (1) 48.0 (48.0) (2) 5.5–7.6 (6.5) 1.48
†Aphanapteryx bonasia Associated individual MU 923 (16.8) (1) 48.0 (48.0) (1) 7.6 (7.6)
†Erythromachus leguati (10.5) 0.63 (4) 33.8–40.1 (38.1) 2.92 (7) 6.5–8.4 (7.6) 0.65
Mentocrex kioloides (8.7) 0.70 (3) 41.4–43.4 (42.5) 1.01 (3) 10.7–12.8 (11.9) 1.11
Gallirallus australis (10.3) 1.48 (4) 33.3–37.4 (35.6) 1.78 (3) 6.8–7.8 (7.3) 0.46
Hypotaenidia sylvestris (10.1) 0.35 (2) 35.1–35.5 (35.3) 0.28 (2) 7.7 (7.7)
Hypotaenidia owstoni (7.2) 0.26 (7) 26.1–31.5 (29.3) 2.31 (4) 6.3–7.8 (7.1) 0.62
Hypotaenidia rovianae (10.0) 1.17 (4) 39.9–43.6 (42.2) 1.63 (4) 9.3–10.3 (9.6) 0.43
†Hypotaenidia wakensis (7.5) 0.96 (3) 18.9–25.9 (22.0) 3.54 (3) 4.7–5.6 (5.1) 0.45
†Diaphorapteryx hawkinsi (15.6) (1) 37.1 (37.1) (1) 4.4 (4.4)
Rallus madagascariensis (5.3) (1) 28.6 (28.6) (1) 9.3 (9.3)
Dryolimnas cuvieri cuvieri (7.5) 0.51 (3) 40.0–48.3 (44.9) 4.38 (3) 11.1–14.1 (12.8) 1.56
Dryolimnas cuvieri aldabranus (7.1) 0.53 (12) 28.6–31.7 (30.1) 0.97 (11) 7.0–8.9 (7.9) 0.61
†Dryolimnas cuvieri abbotti (7.7) (1) 41.4 (41.4) (1) 10.3 (10.3

Laterallus rogersi (5.8) 0.30 (3) 6.9–8.9 (8.1) 1.05 (3) 2.2–3.1 (2.5) 0.47
†Amaurolimnas concolor concolor (8.0) (1) 38.3 (38.3) (1) 12.4 (12.4)
†Zapornia palmeri (6.1) 0.20 (3) 10.1–12.2 (10.9) 1.09 (3) 3.4–3.7 (3.5) 0.15
Porphyrio porphyrio porphyrio (15.3) (1) 60.8 (60.8) (1) 17.1 (17.1)
Porphyrio porphyrio melanotus (15.9) 1.27 (2) 59.4–66.8 (63.1) 5.2 (2) 18.6–20.7 (19.6) 1.48
Porphyrio alleni (8.2) 1.02 (3) 30.3–40.0 (33.7) 5.46 (3) 10.3–13.5 (11.9) 1.60
Porphyrio hochstetteri (23.9) 0.86 (4) 50.6–52.7 (51.3) 0.94 (4) 8.1–9.2 (8.8) 0.48
Gallinula chloropus chloropus (12.7) 1.01 (8) 39.5–50.2 (45.8) 3.25 (8) 12.7–16.0 (14.1) 1.26
Gallinula chloropus pyrrhorrhoa (13.8) (1) 46.3 (46.3) (1) 13.2 (13.2)
Gallinula chloropus galeata (13.9) 0.28 (2) 49.0–50.7 (49.8) 1.20 (2) 14.5–16.5 (15.5) 1.41
Gallinula mortierii (19.3) 0.65 (3) 48.7–51.3 (50.2) 1.37 (3) 9.3–9.8 (9.6) 0.26
Gallinula ventralis (13.1) 0.28 (3) 47.6–58.1 (51.7) 5.61 (3) 13.7–15.5 (14.5) 0.90
Gallinula comeri (16.2) 0.05 (3) 38.9–39.5 (39.2) 0.30 (3) 7.3–8.3 (7.8) 0.50
†Gallinula nesiotis (12.8) (1) 31.8 (31.8) (1) 7.1 (7.1)
Fulica atra (20.1) 1.54 (56.6–66.0 (62.5) 3.65 (10) 15.1–19.0 (16.9) 1.18
Fulica cristata (20.4) 1.02 (7) 56.0–65.8 (61.7) 93.55 (7) 15.1–17.3 (16.0) 0.87
Fulica americana (17.1) 1.62 (2) 51.6–59.4 (55.5) 5.51 (2) 15.5–16.1 (15.8) 0.42
Fulica cornuta (29.0) (1) 82.4 (82.4) (1) 33.0 (33.0)
Fulica gigantea (24.6) (1) 73.6 (73.6) (1) 18.9 (18.9)
†Fulica newtonii (21.6) (1) 67.6 (67.6) (1) 16.9 (16.9)

†Fulica chathamensis (26.5) 1.30 (4) 62.6–68.0 (64.8) 2.33 (5) 11.8–14.7 (13.1) 1.14
91
TABLE 7. Measurements (mm) of the humerus of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD PW (n) range PW (m) SD PD (n) range PD (m) SD SW (n) range
†Aphanapteryx bonasia (12) 56.2–66.5 (61.2) 3.53 (15) 11.2–13.2 (12.0) 0.65 (13) (6.2–8.0) (7.1) 0.63 (20) 3.7–5.3
†Aphanapteryx bonasia Associated individual MU 923 (1) 61.0 (61.0) (1) 12.4 (12.4) (1) 6.3 (6.3) (1) 4.3
Rallicula rubra - - - - - - (1) 2.1
Mentocrex kioloides (1) 37.4 (37.4) (3) 8.4–8.7 (8.5) (0.17) (3) 4.9–5.2 (5.0) (0.15) (3) 3.2
Hypotaenidia insignis - - - - - - (1) 3.1
†Hypotaenidia wakensis (2) 30.8–34.0 (32.4) 2.26 (3) 6.0–6.9 (6.5) 1.09 (3) 3.3–4.1 (3.8) 0.93 (2) 1.8–2.0
†Hypotaenidia dieffenbachii (1) 44.1 (44.1) (1) 8.9 (8.9) (1) 5.6 (5.6) (1) 3.2
†Diaphorapteryx hawkinsi (1) 56.5 (56.5) (1) 11.7 (11.7) (1) 8.4 (8.4) (1) 4.9

†Dryolimnas chekei (1) 47.5 (47.5) (2) 8.8–9.5 (9.1) (0.49) - - (3) 2.9–3.2
†Dryolimnas augusti (1) 48.0 (48.0) (1) (10.0) (10.0) - - (1) 3.3
Eulabeornis castaneoventris - - - - - - (1) 2.1
Megacrex inepta - - - - - - -
Porphyrio porphyrio melanotus (2) 82.6–85.7 (84.1) 2.19 (2) 17.7–18.4 (18.0) 0.49 (2) 7.7–8.1 (7.9) 0.28 (2) 6.1
Gallinula mortierii (3) 64.6–65.3 (64.9) 0.35 (3) 13.1–13.5 (13.3) 0.2 (3) 8.4–9.1 (8.7) 0.35 (3) 4.4
†Gallinula nesiotis - - (1) 10.1 (10.1) (1) 5.5 (5.5) -
Fulica atra (9) 73.2–84.0 (79.5) 3.68 (9) 14.5–16.5 (15.6) 0.74 (9) 7.1–9.1 (7.7) 0.63 (9) 4.5–5.4
Fulica cristata (7) 73.1–86.5 (79.0) 5.22 (7) 14.5–17.1 (15.8) 1.04 (7) 6.1–8.8 (7.9) 0.91 (7) 4.5–5.3
Fulica cornuta (1) 110.9 (110.9) (1) 22.3 (22.3) (1) 11.6 (11.6) (1) 6.5
†Fulica newtonii (6) 75.8–89.2 (84.0) 4.64 (8) 14.7–18.6 (17.0) 1.19 (8) 6.8–9.1 (7.5) 0.92 (8) 5.2–6.3
Fulica chathamensis (13) 85.8–99.4 (93.8) 4.52 (13) 17.2–20.1 (18.8) 0.95 (13) 8.4–10.8 (9.8) 0.71 (13) 5.5–6.6
HUME
Species SW (m) SD SD (n) range SD (m) SD DW (n) ange DW (m) DD (n) range DD (m) SD
†Aphanapteryx bonasia Associated individual MU 923 (4.3) (1) 4.0 (4.0) (1) 10.0 (10.0) (1) 7.7 (7.7)
Rallicula rubra (2.1) (1) 1.7 (1.7) (1) 5.4 (5.4) (1) 3.0 (3.0)
Mentocrex kioloides (3.2) (3) 2.6–2.7 (2.6) (0.05) (1) 7.7 (7.7) (1) 4.0 (4.0)
Hypotaenidia insignis (3.1) (1) 3.1 (3.1) (1) 7.2 (7.2) (1) 4.3 (4.3)
†Hypotaenidia wakensis (1.9) 0.14 (2) 1.8–2.0 (1.9) 0.14 (2) 4.7–5.3 (5.0) 0.42 (2) 3.1–3.3 (3.2) 0.14
†Hypotaenidia dieffenbachii (3.2) (1) 3.0 (3.0) (1) 7.1 (7.1) (1) 4.3 (4.3)
†Diaphorapteryx hawkinsi (4.9) (1) 4.5 (4.5) (1) 10.0 (10.0) (1) 7.0 (7.0)
Dryolimnas cuvieri aldabranus (2.3) 0.40 (15) 1.8–2.5) (2.2) 0.18 (15) 5.7–6.8 (6.1) 0.46 (15) 3.4–3.9 (3.5) 0.32

†Dryolimnas chekei (3.0) (0.17) (3) 3.0–3.3 (3.1) (0.17) (1) 7.5 (7.5) (1) 4.3 (4.3)
†Dryolimnas augusti (3.3) (1) 3.3 (3.3) - - - -
Laterallus rogersi (1.0) 0.12 (3) 1.1 (1.1) (3) 2.8 (2.8) (2) 1.7–1.8 (1.7) 0.07
†Zapornia palmeri (1.2) 0.05 (3) 1.2 (1.2) (3) 3.0–3.1 (3.0) 0.05 (3) 1.7–1.9 (1.8) 0.11
Megacrex inepta - - - (1) 9.3 (9.3) (1) 5.6 (5.6)
Porphyrio porphyrio melanotus (6.1) (2) 5.1–5.2 (5.1) 0.07 (2) 12.6–13.1 (12.8) 0.35 (2) 7.3–7.8 (7.5) 0.35
Gallinula mortierii (4.4) (3) 4.1–4.5 (4.3) 0.2 (3) 9.7–10.4 (9.9) 0.40 (3) 6.3–6.6 (6.4) 0.15
†Gallinula nesiotis - - - - - - -
Fulica atra (5.0) 0.30 (9) 4.0–4.7 (4.3) 0.25 (9) 10.0–11.1 (10.6) 0.41 (9) 5.7–6.9 (6.3) 0.39
Fulica cristata (4.8) 0.27 (7) 4.1–4.8 (4.4) 0.26 (7) 10.2–11.6 (10.8) 0.55 (7) 6.3–7.1 (6.7) 0.28
Fulica cornuta (6.5) (1) 5.9 (5.9) (1) 15.6 (15.6) (1) 9.1 (9.1)
†Fulica newtonii (5.5) 0.57 (8) 4.1–5.4 (4.6) 0.47 (7) 10.0–13.2 (12.2) 1.09 (7) 5.8–7.7 (7.0) 0.62
Fulica chathamensis (6.1) 0.34 (13) 4.6–5.5 (5.1) 0.26 (13) 11.7–14.2 (13.3) 0.80 (13) 7.0–8.3 (7.8) 0.45

93
TABLE 8. Measurements (mm) of the radius of Mascarene rails compared with other Rallidae.
†Aphanapteryx bonasia Associated individual (1) 40.0 (40.0) (1) 4.5 (4.5) (1) 3.4 (3.4) (1) 2.6
MU 923
Rallicula rubra (1) 24.5 (24.5) (1) 1.9 (1.9) (1) 1.6 (1.6) (1) 0.9
Hypotaenidia sylvestris (2) 37.3–38.3 (37.8) 0.70 (2) 2.9 (2.9) (2) 2.8–3.2 (3.0) 0.28 (2) 1.4–1.5
Hypotaenidia woodfordi (1) 41.5 (41.5) (1) 4.1 (4.1) (1) 2.5 (2.5) (1) 1.6
Diaphorapteryx hawkinsi (1) 34.2 (34.2) (1) 3.9 (3.9) (1) 3.9 (3.9) (1) 3.0

†Dryolimnas cuvieri abbotti (1) 34.1 (34.1) (1) 3.0 (3.0) - - (1) 1.3
Laterallus rogersi (1) 13.4 (13.4) (1) 1.4 (1.4) (1) 1.2 (1.2) (1) 0.6
Fulica atra (9) 59.2–69.1 (65.7) 3.62 (9) 3.4–4.2 (3.9) 0.28 (9) 3.2–4.3 (3.8) 0.36 (9) 1.8–2.4
Fulica cristata (7) 55.6–73.0 (64.5) 6.29 (7) 3.7–4.5 (4.0) 0.34 (7) 3.5–4.4 (3.88) 0.30 (7) 1.8–2.2
Fulica cornuta (1) 96.2 (96.2) (1) 5.4 (5.4) (1) 5.8 (5.8) (1) 3.1
†Fulica newtonii (2) 74.5–75.1 (74.8) 0.42 (2) 4.7–5.3 (5.0) 0.42 (2) 4.9–5.2 (5.0) 0.21 (2) 3.1–3.4
†Fulica chathamensis (2) 72.7–74.6 (73.6) 1.34 (1) 4.6 (4.6) (2) 3.7–4.8 (4.2) 0.77 (2) 2.8–3.0
HUME
Species SW (m) SD SD (n) range SD (m) SD DW (n) range DW (m) DD (n) range DD (m) SD
†Aphanapteryx bonasia Associated individual (2.6) (1) 2.6 (2.6) (1) 3.7 (3.7) (1) 4.8 (4.8)
MU 923
Rallicula rubra (0.9) (1) 1.1 (1.1) (1) 2.2 (2.2) (1) 1.7 (1.7)
Hypotaenidia sylvestris (1.4) 0.07 (2) 1.4–1.5 (1.4) 0.07 (2) 3.9–4.1 (4.0) 0.14 (2) 2.8 (2.8)
Hypotaenidia woodfordi (1.6) (1) 1.6 (1.6) (1) 2.6 (2.6) (1) 3.1 (3.1)
Diaphorapteryx hawkinsi (3.0) (1) 2.7 (2.7) (1) 4.8 (4.8) (1) 3.9 (3.9)
Rallus madagascariensis (0.9) (1) 0.9 (0.9) (1) 2.5 (2.5 (1) 1.6 (1.6)

Dryolimnas cuvieri cuvieri (1.5) 0.14 (2) 1.2–1.5 (1.3) 0.21 (2) 3.6 (3.6) (2) 2.4–2.6 (2.5) 0.14
Laterallus rogersi (0.6) (1) 0.9 (0.9) (1) 2.0 (2.0) (1) 1.4 (1.4)
†Zapornia palmeri (0.5) 0.05 (3) 0.5–0.6 (0.5) 0.05 (3) 1.4–1.6 (1.4) 0.11 (3) 1.0–1.5 (1.2) 0.25
Porphyrio porphyrio melanotus (2.6) (2) 2.8 (2.8) (2) 6.4–6.5 (6.4) 0.07 (2) 3.6–4.0 (3.8) 0.28
Gallinula mortierii (2.4) 0.05 (3) 2.1 (2.1) (3) 4.8–5.0 (4.9) 0.10 (3) 3.2–3.6 (3.4) 0.20
Fulica atra (2.1) 0.21 (9) 2.1–2.5 (2.3) 0.13 (9) 4.4–5.1 (4.8) 0.28 (9) 3.0–3.6 (3.2) 0.24
Fulica cristata (2.1) 0.15 (7) 2.1–2.7 (2.5) 0.20 (7) 4.3–5.3 (4.9) 0.37 (7) 3.0–3.7 (3.2) 0.28
Fulica cornuta (3.1) (1) 3.3 (3.3) (1) 6.9 (6.9) (1) 4.7 (4.7)
†Fulica newtonii (3.2) 0.21 (2) 3.0–3.1 (3.0) 0.07 (2) 6.1–6.3 (6.2) 0.14 (2) 4.1–4.2 (4.1) 0.07
†Fulica chathamensis (2.9) 0.14 (2) 2.6–3.0 (2.8) 0.28 (2) 4.7–6.1 (5.4) 0.98 (2) 3.2–3.3 (3.2) 0.07

95
TABLE 9. Measurements (mm) of the ulna of Mascarene rails compared with other Rallidae.
†Aphanapteryx bonasia Associated individual MU 923 (1) 42.5 (42.5) (1) 5.9 (5.9) - -
†Erythromachus leguati (7) 38.1–40.5 (39.7) 0.82 (8) 4.4–6.0 (5.3) 0.54 (7) 3.6–5.9 (4.8) 0.77
Mentocrex kioloides - - (1) 4.8 (4.8) (1) 4.7 (4.7)
Rallicula rubra (1) 26.6 (26.6) (1) 3.1 (3.1) (1) 2.7 (2.7)
Hypotaenidia woodfordi (1) 45.8 (45.8) (1) 5.5 (5.5) (1) 5.7 (5.7)
Hypotaenidia insignis (1) 41.3 (41.3) (1) 4.8 (4.8) (1) 4.9 (4.9)

Rougetius rougetii (1) 41.9 (41.9) (1) 4.3 (4.3) (1) 4.3 (4.3)
Laterallus rogersi (1) 13.7 (13.7) (1) 2.4 (2.4) (1) 1.7 (1.7)
†Laterallus podarces - - (1) 6.2 (6.2) (1) 5.9 (5.9)
Megacrex inepta (1) 45.7 (45.7) (1) 6.0 (6.0) (1) 6.4 (6.4)
Porphyrio porphyrio melanotus (2) 53.5–53.8 (53.6) 0.21 (2) 10.0–10.8 (10.4) 0.56 (2) 5.0 (5.0)
Fulica atra (9) 39.5–45.6 (43.3) 2.17 (9) 7.3–8.9 (8.2) 0.62 (9) 3.8–4.6 (4.2) 0.26
Fulica cristata (7) 59.9–77.3 (68.7) 6.27 (7) 6.7–8.1 (7.5) 0.55 (7) 6.2–7.6 (6.8) 0.53
Fulica cornuta (1) 101.7 (101.7) (1) 9.4 (9.4) (1) 10.3 (10.3)
†Fulica newtonii (1) 73.2 (73.2) (1) 8.0 (8.0) (1) 8.2 (8.2)
†Fulica newtonii (juv) (1) 71.7 (71.7) (1) 6.8 (6.8) (1) 5.7 (5.7)
HUME
TABLE 9. (continued)
Species SW (n) range SW (m) SD SD (n) range SD (m) SD DD (n) range DD (m) SD
†Erythromachus leguati (8) 2.3–3.6 (3.2) 0.50 (8) 2.5–3.3 (2.9) 0.27 (6) 4.3–5.4 (4.8) 0.36
Rallicula rubra (1) 1.6 (1.6) (1) 1.7 (1.7) (1) 2.7 (2.7)
Hypotaenidia sylvestris (2) 2.5–2.6 (2.5) 0.07 (2) 2.9 (2.9) (2) 4.7–5.0 (4.8) 0.21
Hypotaenidia insignis (1) 2.4 (2.4) (1) 2.7 (2.7) (1) 4.1 (4.1)
†Hypotaenidia wakensis (2) 1.6–1.7 (1.6) 0.07 (2) 1.9 (1.9) (2) 3.0–3.2 (3.1) 0.35
Dryolimnas cuvieri cuvieri (2) 2.3–3.0 (2.6) 0.49 (2) 2.6–2.9 (2.7) 0.21 (2) 4.3 (4.3)

†Laterallus podarces (1) 3.6 (3.6) (1) 3.5 (3.5) - -
Megacrex inepta (1) 2.5 (2.5) (1) 3.3 (3.3) (1) 5.2 (5.2)
Fulica atra (9) 2.4–3.2 (2.9) 0.24 (9) 2.8–3.4 (3.1) 0.21 (9) 3.4–3.9 (3.5) 0.17
Fulica cristata (7) 2.9–4.2 (3.8) 0.43 (7) 3.8–4.5 (4.2) 0.26 (7) 6.3–7.2 (6.6) 0.34
Fulica cornuta (1) 5.3 (5.3) (1) 5.7 (5.7) (1) 9.7 (9.7)
†Fulica newtonii (1) 4.7 (4.7) (1) 4.9 (4.9) (1) 8.0 (8.0)
†Fulica newtonii (juv) (1) 4.7 (4.7) (1) 4.0 (4.0) (1) 6.5 (6.5)

97
TABLE 10. Measurements (mm) of the carpometacarpus of Mascarene rails compared with other Rallidae.
Dryolimnas cuvieri cuvieri (2) 28.1–29.6 (28.8) 1.06 (2) 6.0 (6.0) (2) 2.9–3.0 (2.9) 0.07

†Zapornia palmeri (3) 9.9–10.6 (10.2) (3) 2.4–2.7 (2.6) 0.17 (3) 1.3–1.6 (1.5) 0.17
0.36
Megacrex inepta (1) 29.5 (29.5) (1) 7.7 (7.7) (1) 3.4 (3.4)
Porphyrio porphyrio melanotus (2) 53–5–53.8 (53.6) 0.21 (2) 10.0–10.8 (10.4) 0.56 (2) 5.0 (5.0)
Gallinula chloropus (8) 29.0–34.6 (31.60 2.12 (8) 6.1–7.2 (6.6) 0.41 (8) 3.0–3.9 (3.5) 0.32
Fulica atra (9) 39.5–45.6 (43.3) 2.17 (9) 7.3–8.9 (8.2) 0.62 (9) 3.8–4.6 (4.2) 0.26
Fulica cristata (7) 37.2–48.6 (43.4) 3.76 (7) 7.4–9.2 (8.4) 0.67 (7) 4.1–4.8 (4.4) 0.26
Fulica cornuta (1) 62.7 (62.7) (1) 12.2 (12.2) (1) 6.4 (6.4)
†Fulica newtonii (1) 49.7 (49.7) (1) 10.0 (10.0) (1) 5.0 (5.0)
HUME
Species SW (n) range SW (m) SD SD (n) range SD (m) DD (n) range DD (m)
SD SD
Hypotaenidia sylvestris (2) 2.3–2.6 (2.4) 0.21 (2) 2.2–2.3 (2.2) 0.07 (2) 3.1 (3.1)

Megacrex inepta (1) 1.8 (1.8) (1) 2.1 (2.1) (1) 2.8 (2.8)
Fulica atra (9) 2.4–3.2 (2.9) 0.24 (9) 2.8–3.4 (3.1) 0.24 (9) 3.4–3.9 (3.5) 0.17
Fulica cristata (7) 2.6–3.2 (3.0) 0.19 (7) 2.9–3.7 (3.2) 0.26 (7) 3.1–3.8 (3.4) 0.27
Fulica cornuta (1) 3.8 (3.8) (1) 4.5 (4.5) (1) 4.7 (4.7)
†Fulica newtonii (1) 3.2 (3.2) (1) 3.9 (3.9) (1) 3.9 (3.9)

99
TABLE 11. Measurements (mm) of the pelvis of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD PW (n) range PW (m) SD PD (n) range PD (m) SD GW (n) range GW (m) SD
vidual MU 923
Mentocrex kioloides (3) 44.9–46.9 (45.9) 1.00 (3) 22.6–22.7 (22.6) 0.05 (3) 17.5–18.4 (18.0) 0.47 (3) 9.3–10.1 (9.8) 0.43
Hypotaenidia sylvestris (1) 59.7 (59.7) (2) 24.7–25.9 (25.3) (0.84) (1) 18.8 (18.8) (1) 14.0 (14.0)
†Hypotaenidia dieffenbachii (1) 53.6 (53.6) (1) 23.7 (23.7) (1) 16.7 (16.7) (1) 12.6 (12.6)
†Cabalus modestus (1) 28.4 (28.4) (1) 12.0 (12.0) (1) 9.4 (9.4) (1) 6.4 (6.4)

†Dryolimnas chekei - - (1) 15.2 (15.2) - - - -
†Laterallus podarces - - - - - - (1) 25.9 (25.9)
†Hovacrex roberti (1) 82.9 (82.9) (1) 39.7 (39.7) (1) 22.8 (22.8) - -
Gallinula comeri (3) 56.3–63.7 (60.2) 3.53 (4) 26.0–28.8 (27.8) 1.77 (3) 17.9–22.6 (20.7) 2.49 (3) 12.1–14.1 (13.2) 1.00
Fulica atra (9) 72.5–81.4 (78.0) 3.43 (9) 17.2–21.8 (20.2) 1.50 (9) 12.5–14.8 (13.7) 0.74 (9) 21.0–26.0 (24.2) 1.34
Fulica cristata (7) 69.6–88.2 (80.7) 6.56 (7) 22.9–26.9 (25.1) 1.52 (7) 17.3–21.8 (19.6) 1.59 (7) 13.4–16.3 (15.1) 0.95
Fulica cornuta (1) 103.4 (103.4) (1) 33.2 (33.2) (1) 26.8 (26.8) (1) 19.8 (19.8)
†Fulica newtonii (4) 67.8–88.8 (79.6) 8.90 (4) 22.4–31.7 (28.3) 4.80 (4) 14.7–23.6 (19.2) 3.63 (4) 17.2–27.1 (20.7) 4.37
HUME
TABLE 12. Measurements (mm) of the femur of Mascarene rails compared with other Rallidae.
†Aphanapteryx bonasia (7) 69.6–76.3 (73.2) 2.86 (9) 12.8–17.7 (15.6) 1.47 (8) 8.3–12.8 (10.2) 1.32 (14) (5.4–7.6)
†Aphanapteryx bonasia Associated individual (1) 74.7 (74.7) (1) 15.2 (15.2) (1) 13.1 (13.1) (1) 6.0
MU 923
Mentocrex kioloides (3) 43.6–45.6 (44.7) 1.01 (3) 8.7–8.9 (8.7) 0.11 (3) 5.1–5.8 (5.5) 0.37 (3) 3.7–4.3
Hypotaenidia sylvestris (2) 61.8–63.4 (62.6) 1.13 (2) 11.6 (11.6) (2) 8.2–8.4 (8.3) 0.14 (2) 4.4–4.7
†Hypotaenidia dieffenbachii (2) 55.5–59.3 (57.4) 2.68 (2) 10.0–11.0 (10.5) 0.70 (2) 7.1–7.8 (7.4) 0.49 (2) 4.3–4.4

Dryolimnas cuvieri aldabranus (18) 42.1–47.9 (44.7) 2.01 (18) 6.1–8.7 (7.9) 0.60 (18) 4.7–7.1 (5.7) 0.64) (18) 2.7–3.5
†Dryolimnas chekei (1) 50.0 (50.0) - - (1) 7.8 (7.8) (1) 3.8
†Laterallus podarces (2) 68.0–68.3 (68.1) 0.21 (2) 12.7–12.9 (12.8) 0.14 (2) 8.3–9.2 (8.7) 0.63 (2) 5.0
†Hovacrex roberti (1) 83.0 (83.0) (1) 14.1 (14.1) (1) 17.1 (17.1) (1) 6.9
Porphyrio porphyrio melanotus (2) 75.9–78.3 (77.1) 1.69 (2) 13.2–14.5 (13.8) 0.91 (2) 9.7–10.1 (9.9) 0.28 (2) 5.9–6.3
Porphyrio hochstetteri (4) 100.5– (102.1) 1.96 (4) 20.5–21.9 (21.0) 0.59 (4) 13.2–15.8 (14.1) 1.16 (4) 9.3–10.2
105.0
Gallinula comeri (3) 54.1–62.3 (58.5) 4.14 (4) 10.4–11.9 (11.4) 0.67 (4) 6.8–8.0 (7.5) 0.55 (4) 4.3–4.6
Fulica atra (10) 53.9–62.6 (59.0) 3.36 (10) 11.1–13.0 (11.9) 0.61 (10) 7.3–11.7 (8.9) 1.31 (10) 4.4–5.3
Fulica cristata (7) 52.0–66.3 (60.5) 4.98 (7) 11.1–13.3 (12.4) 0.81 (7) 7.8–9.7 (8.8) 0.64 (7) 4.4–5.5
Fulica cornuta (1) 79.4 (79.4) (1) 15.9 (15.9) (1) 13.8 (13.8) (1) 6.4
†Fulica newtonii (5) 70.8–77.6 (72.9) 2.71 (5) 13.9–16.8 (14.7) 1.18 (4) 9.4–13.0 (10.5) 1.65 (6) 6.3–7.3

†Fulica chathamensis (13) 80.5–88.5 (84.1) 2.37 (13) 17.6–20.0 (18.4) 0.66 (13) 11.3–15.2 (13.5) 1.34 (13) 6.8–8.0
101
Species SW (m) SD SD (n) range SD (m) DW (n) range DW (m) DD (n) range DD (m)
SD SD
†Aphanapteryx bonasia Associated individual (6.0) (1) 6.7 (6.7) (1) 15.6 (15.6) (1) 12.8 (12.8)
MU 923
†Erythromachus leguati (5.1) 0.32 (18) (5.1–7.2 (6.2) 0.55 (20) 11.5–14.2 (12.7) 0.83 (20) 9.0–10.9 (9.9) 0.55
Mentocrex kioloides (3.9) 0.30 (3) 3.7–3.9 (3.7) 0.11 (3) 7.9–8.3 (8.1) 0.20 (3) 6.1–6.7 (6.4) 0.30
†Hypotaenidia dieffenbachii (4.3) 0.07 (2) 4.5–4.6 (4.5) 0.07 (2) 9.9–10.6 (10.2) 0.49 (2) 8.4–8.8 (8.6) 0.49

†Dryolimnas chekei (3.8) (1) 3.8 (3.8) - - - -
Laterallus rogersi (1.6) 0.05 (3) 1.8 (1.8) (3) 4.1–4.2 (4.1) 0.05 (3) 3.2–3.5 (3.3) 0.17
†Laterallus podarces (5.0) (2) 5.7–5.9 (5.8) 0.14 (1) 13.3 (13.3) (2) 10.3–10.6 (10.4) 0.21
†Hovacrex roberti (6.9) (1) 7.4 (7.4) (1) 17.0 (17.0) (1) 13.8 (13.8)
Porphyrio porphyrio melanotus (6.1) 0.28 (2) 6.8 (6.8) (2) 12.9–13.9 (13.4) 0.70 (2) 11.1–12.0 (11.5) 0.63
Gallinula mortierii (7.3) 0.26 (3) 8.0–8.4 (8.1) 0.20 (3) 15.8–16.4 (16.0) 0.32 (3) 13.4–14.3 (13.8) 0.45
Gallinula comeri (4.4) 0.17 (4) 4.2–4.6 (4.4) 0.19 (3) 10.3–10.8 (10.5) 0.25 (4) 7.7–8.9 (8.4) 0.62
Fulica atra (4.9) 0.27 (10) 4.7–5.9 (5.2) 0.31 (10) 10.2–12.0 (11.1) 0.72 (10) 8.5–9.9 (9.1) 0.41
Fulica cristata (5.0) 0.41 (7) 5.6–5.7 (5.2) 0.38 (7) 11.0–12.6 (11.9) 0.62 (7) 8.9–10.9 (9.8) 0.88
Fulica cornuta (6.4) (1) 6.8 (6.8) (1) 15.2 (15.2) (1) 12.4 (12.4)
†Fulica newtonii (6.7) 0.33 (6) 7.0–7.6 (7.3) 0.25 (6) 12.3–16.4 (13.5) 1.51 (4) 11.2–13.1 (11.9) 0.74
HUME
TABLE 13. Measurements (mm) of the tibiotarsus of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) (SD) PW (n) range PW (m) (SD) PD (n) range PD (m) (SD) SW (n) range SW (m) (SD)
†Aphanapteryx bonasia (31) 100.2–121.3 (111.5) 6.08) (37) 12.5–17.8 (14.2) 1.39 (32) (15.2–21.6 (18.3) 1.65 (49) 5.2–7.2 (6.1) 0.51
†Aphanapteryx bonasia Associated (1) 110.2 (110.2) (1) 16.3 (16.3) (1) 17.4 (17.4) (1) 5.6 (5.6)
individual MU 923
Mentocrex kioloides (1) 67.1 (67.1) (1) 8.2 (8.2) (1) 9.4 (9.4) (1) 3.1 (3.1)
Rallicula rubra - - - - - - (1) 2.7 (2.7)
Hypotaenidia woodfordi (1) 95.0 (95.0) - - - - (1) 5.6 (5.6)
Hypotaenidia insignis - - - - - - (1) 4.7 (4.7)
Dryolimnas cuvieri cuvieri (2) 75.3–81.8 (78.5) 4.59 (2) 8.7–9.2 (8.9) 0.35 (2) 10.1–10.4 (10.2)0.21 (2) 3.3–3.8 (3.5) 0.35

†Dryolimnas chekei (2) 78.1–90.0 (84.0) 8.41 (3) 9.1–10.6 (9.6) 0.81 (2) 10.4–11.0 (10.7) 0.42 (4) 3.7–3.9 (3.7) 0.10
†Dryolimnas augusti - - (1) 9.7 (9.7) (1) 12.1 (12.1) - -
Rougetius rougetii (1) 77.0 (77.0) (1) 7.1 (7.1) (1) 7.3 (7.3) (1) 3.4 (3.4)
†Laterallus podarces (1) 108.2 (108.2) - - - - (1) 5.5 (5.5)
Amaurornis isabellina (1) 93.8 (93.8) (1) 8.1 (8.1) - - (1) 4.0 (4.0)
Eulabeornis castaneoventris - - - - - - (1) 5.1 (5.1)
Megacrex inepta (1) 121.0 (121.0) (1) 15.3 (15.3) (1) 17.8 (17.8) (1) 5.8 (5.8)
†Hovacrex roberti (1) 139.7 (139.7) (1) 14.0 (14.0) (1) 19.9 (19.9) (1) 7.3 (7.3)
Porphyrio hochstetteri (4) 144.5–151.2 (148.5) 2.98 (4) 18.1–18.8 (18.3)0.31 (4) 19.7–25.2 (23.3) 2.52 (4) 8.0–9.0 (8.6) 0.45
Gallinula comeri (1) 91.6 (91.6) (1) 9.6 (9.6) (1) 13.4 (13.4) (1) 4.1 (4.1)
Fulica atra (9) 90.9–108.4 (103.4) 5.95 (9) 8.9–11.5 (10.4) 0.91 (9) 12.2–14.4 (13.4) 0.76 (9) 4.5–5.7 (5.1) 0.76
Fulica cristata (7) 94.3–115.0 (105.3) 8.56 (7) 9.1–12.6 (11.0) 1.21 (7) 12.5–15.4 (13.8) 1.11 (7) 4.9–5.8 (5.4) 0.32
Fulica cornuta (1) 137.9 (137.9) (1) 13.4 (13.4) (1) 18.3 (18.3) (1) 7.5 (7.5)

†Fulica newtonii (26) 119.5–150.0 (127.4) 6.95 (24) 11.3–16.4 (13.5) 1.33 (24) 14.9–16.4 (16.8) 1.17 (30) 5.4–7.3 (6.6) 0.44
103
Species SD (n) range SD (m) (SD) DW (n) range DW (m) (SD) DD (n) range DD (m) (SD) WCC (n) range WCC (m) (SD)
†Aphanapteryx bonasia (49) 4.3–6.6 (5.7) 0.44 (44) 11.2–14.7 (12.9) 0.88 (42) 10.5–15.4 (13.1) 1.17 (28) (102.9–124.9 (112.9) 6.61
†Aphanapteryx bonasia Associated (1) 6.2 (6.2) (1) 12.9 (12.9) (1) 13.0 (13.0) (1) 115.2 (115.2)
individual MU 923
†Erythromachus leguati (22) 3.9–5.0 (4.3) 0.26 (23) (9.1–12.1) (10.0) 0.69 (23) 9.0–11.8 (10.3) 0.69 (16) (83.6–102.0 (90.7) 5.53
Mentocrex kioloides (1) 3.0 (3.0) (1) 6.3 (6.3) (1) 6.4 (6.4) - -
Rallicula rubra (1) 2.5 (2.5) (1) 5.2 (5.2) (1) 4.7 (4.7) - -
Gallirallus australis (4) 5.5–6.8 (6.1) 0.60 (4) 12.1–16.0 (14.0) 2.08 (4) 10.3–12.8 (11.6) 1.30 - -
Hypotaenidia owstoni (4) 2.8–3.4 (3.1) 0.25 (4) 6.4–6.7 (6.6) 0.14 (4) 6.3–7.0 (6.7) 0.29 - -
Hypotaenidia woodfordi (1) 4.0 (4.0) (1) 8.3 (8.3) (1) 9.1 (9.1) - -
Hypotaenidia insignis (1) 4.0 (4.0) (1) 8.6 (8.6) (1) 8.5 (8.5) - -
Rallus madagascariensis (1) 2.2 (2.2) (1) 5.0 (5.0) (1) 5.2 (5.2) - -
Dryolimnas cuvieri cuvieri (2) 3.1–3.4 (3.2) 0.21 (2) 6.6–6.7 (6.6) 0.07 (2) 6.8–6.9 (6.8) 0.07 - -

Dryolimnas cuvieri aldabranus (15) 2.4–3.0 (2.7) 0.16 (15) 5.5–6.8 (6.1) 0.39 (15) 5.8–6.8 (6.3) 0.30 - -
†Dryolimnas cuvieri abbotti (1) 3.0 (3.0) (1) 6.5 (6.5) (1) 7.0 (7.0) - -
†Dryolimnas chekei (4) 3.4–3.6 (3.4) 0.09 (2) 7.4 (7.4) (2) 7.3–8.0 (7.6) 0.49 - -
†Dryolimnas augusti - - (1) 7.8 (7.8) (1) 8.0 (8.0)
Rougetius rougetii (1) 2.8 (2.8) (1) 6.1 (6.1) (1) 6.4 (6.4) - -
Laterallus rogersi (2) 1.5–1.7 (1.6) 0.14 (2) 3.6–3.7 (3.6) 0.07 (2) 3.5–3.7 (3.6) 0.14 (1) 40.7 (40.7
†Laterallus podarces (1) 4.8 (4.8) (1) 10.2 (10.2) (1) 11.2 (11.2) - -
Amaurornis isabellina (1) 3.8 (3.8) (1) 7.5 (7.5) (1) 7.7 (7.7) - -
Eulabeornis castaneoventris (1) 4.2 (4.2) (1) 9.9 (9.9) (1) 10.1 (10.1) - -
Megacrex inepta (1) 5.8 (5.8) (1) 11.9 (11.9) (1) 12.7 (12.7) (1) 124.4 (124.4)
†Hovacrex roberti (1) 6.0 (6.0) (1) 12.6 (12.6) (1) 13.2 (13.2) (1) 142.9 (142.9)
Porphyrio porphyrio melanotus (2) 5.6 (5.6) (2) 11.1–11.2 (11.1) 0.07 (2) 11.3–11.8 (11.5) 0.35 (2) 132.9–137.3 (135.1) 3.11
†Gallinula nesiotis (1) 4.1 (4.1) (1) 10.2 (10.2) (1) 8.3 (8.3) - -
Gallinula comeri (1) 3.8 (3.8 (1) 9.1 (9.1) (1) 9.4 (9.4) (1) 95.3 (95.3)
Fulica atra (9) 3.8–4.5 (4.1) 0.27 (9) 8.5–10.0 (9.4) 0.59 (9) 8.2–9.8 (9.4) 0.58 (9) 97.1–115.2 (109.4) 6.20
Fulica cristata (7) 3.7–4.7 (4.3) 0.35 (7) 9.1–10.8 (10.0) 0.69 (7) 9.0–10.2 (9.5) 0.51 (7) 99.8–121.7 (110.4) 8.46
Fulica cornuta (1) 5.6 (5.6) (1) 14.1 (14.1) (1) 13.8 (13.8) (1) 144.5 (144.5)
†Fulica newtonii (30) 4.5–6.1 (5.1) 0.39 (31) 11.3–15.3 (12.4) 0.76 (30) 10.1–14.4 (11.5) 0.98 (18) 122.1–160.0 (134.4) 9.09
HUME
TABLE 14. Measurements (mm) of the fibula of Mascarene rails compared with other Rallidae.
Species TL (n) range TL (m) SD PW (n) range PW (m) SD PD(n) range PD (m) SD
†Aphanapteryx bonasia Asso- - - (1) 9.2 (9.2) (1) 3.2 (3.2)
ciated individual MU 923
†Erythromachus leguati (1) 45.3 (45.3) (2) 6.6-7.7 (7.1) 0.77 (2) 2.4 (2.4)
Dryolimnas cuvieri cuvieri (1) 74.7
Dryolimnas cuvieri aldabranus (1) 43.3 (43.3) (1) 4.5 (4.5) (1) 1.8 (1.8)
†Dryolimnas cuvieri abbotti (1) 74.7
†Hypotaenidia wakensis - - (1) 4.3 (4.3) (1) 1.4 (1.4)
TABLE 15. Measurements (mm) of the tarsometatarsus of Mascarene rails compared with other Rallidae.
†Aphanapteryx bonasia Associated individual MU 923 (1) 69.5 (69.5) (1) 13.7 (13.7) (1) 11.0 (11.0) (1) 5.7
Rallicula rubra (1) 38.8 (38.8) (1) 5.4 (5.4) (1) 5.1 (5.1) (1) 2.2
Hypotaenidia woodfordi (1) 60.5 (60.5) (1) 9.0 (9.0) (1) 8.7 (8.7) (1) 4.4
Hypotaenidia insignis (1) 63.3 (63.3) (1) 9.0 (9.0) (1) 9.0 (9.0) (1) 3.8

†Dryolimnas chekei (2) 45.3–52.5 (48.9) 5.09 (2) 7.4–7.9 (7.6) 0.35 (2) 7.0–8.0 (7.5) 0.70 (2) 3.6–4.0
†Dryolimnas augusti (2) 53.0–53.1 (53.0) 0.07 (2) 8.7–8.8 (8.7) 0.07 (2) 9.2–9.5 (9.3) 0.21 (2) 4.3–4.4
Laterallus rogersi (2) 24.6–26.0 (25.3) 0.98 (2) 3.7–4.2 (3.9) 0.35 (1) 3.6 (3.6) (2) 1.5–1.8
†Laterallus podarces (1) 72.5 (72.5) (1) 11.6 (11.6) - - (1) 5.1
Amaurornis isabellina (1) 66.1 (66.1) (1) 8.1 (8.1) (1) 8.5 (8.5) (1) 3.5
Megacrex inepta (1) 87.6 (87.6) (1) 12.8 (12.8) (1) 11.5 (11.5) (1) 5.7
Gallinula mortierii (3) 79.3–80.8 (80.1) 0.77 (3) 12.5–13.0 (12.8) 0.28 (3) 12.8–13.9 (13.4) 0.58 (3) 5.5
Gallinula nesiotis comeri (1) 56.1 (56.1) (1) 9.6 (9.6) (1) 8.8 (8.8) (1) 4.3
†Pareudiastes pacificus - - - - - - -
Fulica atra (9) 54.8–67.1 63.2) 4.50 (9) 9.1–10.2 (9.9) 0.58 (9) 8.1–10.2 (9.3) 0.80 (9) 3.4–4.5
Fulica cristata (7) 58.5–86.3 (71.8) 9.90 (7) 10.0–17.1 (12.9) 2.99 (7) 7.0–10.5 (8.9) 1.19 (7) 3.9–5.1
Fulica americana (2) 54.7–60.9 (57.8) 4.30 (2) 8.8–9.4 (9.1) 0.42 (2) 8.4 (8.4) (2) 3.3–3.9
Fulica cornuta (1) 88.1 (88.1) (1) 14.4 (14.4) (1) 14.6 (14.6) (1) 5.5
†Fulica newtonii (24) 75.1–91.2 (83.8) 3.92 (23) 12.1–14.5 (13.1) 0.68 (22) 10.2–13.5 (13.1) 0.70 (30) 5.3–6.4
†Fulica chathamensis (17) 86.6–96.9 (92.4) 2.89 (17) 14.0–17.0 (16.0) 0.80 (17) 12.9–15.0 (14.0) 0.59 (17) 6.2–7.6
HUME
Species SW (m) SD SD (n) range SD (m) DW (n) range DW (m) DD (n) range DD (m)
SD SD
†Aphanapteryx bonasia Associated individual MU 923 (5.7) (1) 5.3 (5.3) (1) 13.8 (13.8) (1) 12.3 (12.3)
Rallicula rubra (2.2) (1) 2.1 (2.1) (1) 5.8 (5.8) (1) 3.2 (3.2)
Hypotaenidia woodfordi (4.4) (1) 3.6 (3.6) (1) 9.1 (9.1) (1) 6.9 (6.9)
Hypotaenidia insignis (3.8) (1) 3.0 (3.0) (1) 9.2 (9.2) (1) 6.9 (6.9)

†Dryolimnas chekei (3.8) 0.28 (2) 3.6–4.0 (3.8) 0.28 (2) 3.2–3.4 (3.3) 0.14 (1) 8.7 (8.7)
†Dryolimnas augusti (4.3) 0.07 (2) 3.8–3.9 (3.8) 0.07 (1) 9.5 (9.5) (1) 7.2 (7.2)
Laterallus rogersi (1.6) 0.21 (2) 1.4–2.1 (1.7) 0.49 (2) 3.9–4.0 (3.9) 0.07 (2) 2.8–3.3 (3.0) 0.35
†Laterallus podarces (5.1) (1) 4.8 (4.8) (1) 11.8 (11.8) (1) 8.6 (8.6)
Amaurornis isabellina (3.5) (1) 3.1 (3.1) (1) 8.5 (8.5) (1) 6.0 (6.0)
Megacrex inepta (5.7) (1) 4.3 (4.3) (1) 13.5 (13.5) (1) 9.9 (9.9)
Porphyrio porphyrio melanotus (6.7) (2) 5.5 (5.5) (2) 12.5–12.8 (12.6) 0.21 (2) 9.1–9.7 (9.4) 0.42
Gallinula mortierii (5.5) (3) 4.9–5.1 (5.0) 0.11 (3) 12.9–13.9 (13.5) 0.55 (3) 10.2–11.0 (10.6) 0.41
Gallinula nesiotis comeri (4.3) (1) 3.9 (3.9) (1) 9.6 (9.6) (1) 7.9 (7.9)
†Pareudiastes pacificus - - - (1) 7.9 (7.9) (1) 5.3 (5.3)
Fulica atra (4.0) 0.36 (9) 3.7–4.5 (4.1) 0.25 (9) 8.8–10.8 (9.9) 0.71 (9) 7.0–9.2 (7.9) 0.66
Fulica cristata (4.5) 0.42 (7) 3.5–4.7 (4.5) 0.39 (7) 10.0–11.5 (10.5) 0.56 (7) 6.1–8.8 (7.2) 0.99
Fulica americana (3.6) 0.42 (2) 3.6 (3.6) (2) 8.6–9.1 (8.8) 0.35 (2) 6.4–6.9 (6.6) 0.35
Fulica cornuta (5.5) (1) 5.5 (5.5) (1) 15.3 (15.3) (1) 12.1 (12.1)
†Fulica newtonii (5.8) 0.28 (30) 4.2–5.4 (4.9) 0.30 (28) 12.8–14.6 (13.6) 0.58 (25) 9.1–11.6 (10.3) 0.67

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Zootaxa 4626 (1): 001–107 ISSN 1175-5326 (print edition)

Systematics, morphology and ecology of rails (Aves: Rallidae)

JULIAN PENDER HUME

Accepted by T. Worthy: 30 May 2019; published: 3 Jul. 2019

FIRST PUBLISHED IN 2019 BY

© 2019 Magnolia Press

ISSN 1175-5326 (Print edition)

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TABLE 1. Nomenclatural status of real and supposed endemic Mascarene rails.

Materials and methods

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Order Gruiformes Bonaparte, 1854

Family Rallidae Rafinesque, 1815

Genus Aphanapteryx Frauenfeld, 1868a

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†Mauritian Red Rail (Poule Rouge) Aphanapteryx bonasia (Sélys-Longchamps, 1848)

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Measurements: See Appendix 2.

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Diagnosis: As for genus.

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Genus Erythromachus Milne-Edwards, 1874

Erythromachus Milne-Edwards 1874, p.6, pls.11–12. Type by monotypy: E. leguati Milne-Edwards

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Holotype: None designated.

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Genus Hypotaenidia Reichenbach, 1853

Buff-banded Rail Hypotaenidia philippensis (Linnaeus, 1766)

Rallus philippensis: Linnaeus, 1766, p.263

Genus Dryolimnas Sharpe, 1893

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White-throated Rail Dryolimnas cuvieri (Pucheran, 1845)

Rallus cuvieri: Pucheran, 1845, p.279 (Mauritius)

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Measurements: See Appendix 2.

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TABLE 3. Invasive probable rail predators and their dates of introduction

†Cheke’s Wood Rail Dryolimnas chekei sp. nov

Dryolimnas cuvieri: Cowles, 1987, p.96; Cheke, 1987, p.37

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†Réunion Wood Rail Dryolimnas augusti Mourer-Chauviré et al. 1999

Holotype: Tarsometatarsi MHNR 13 (R), MHNR 14 (L), associated.

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Genus Porphyrio Brisson, 1760

†Réunion Gallinule (Oiseaux Bleu) ‘Porphyrio caerulescens’ (Sélys-Longchamps, 1848)

Holotype: None designated.

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African Swamphen Porphyrio porphyrio madagascariensis (Latham, 1801)

Gallinula porphyrio Latham, 1801, p.lxviii.

Allen’s Gallinule Porphyrio alleni Thomson, 1842

Porphyrio alleni Thomson, 1842, p.204; E. Newton, 1888, p.552

Genus Fulica Linnaeus, 1758