Article
An Extinct New Rail (Gallirallus, Aves: Rallidae) Species from
Rapa Island, French Polynesia †
Rodrigo B. Salvador 1 , Atholl Anderson 2
1
2
*
†
and Alan J. D. Tennyson 1, *
Museum of New Zealand Te Papa Tongarewa, Wellington 6011, New Zealand; salvador.rodrigo.b@gmail.com
Department of Archaeology and Natural History, Australian National University, Canberra 2600, Australia;
atholl.anderson@anu.edu.au
Correspondence: alant@tepapa.govt.nz
urn:lsid:zoobank.org:pub:B77575A7-DEFA-43A5-BAC9-A79BC76C301C.
Abstract: A new species of rail, Gallirallus astolfoi sp. nov., is described from Rapa Island (Rapa Iti),
French Polynesia. The holotype (and single known specimen) is a left tarsometatarsus recovered
from Tangarutu Cave. This rail species was apparently endemic to Rapa Iti and potentially flightless.
It became extinct after human colonisation of the island.
Keywords: endemic species; flightlessness; Gallirallus astolfoi sp. nov.; Holocene; Rapa Iti
Citation: Salvador, R.B.; Anderson,
A.; Tennyson, A.J.D. An Extinct New
Rail (Gallirallus, Aves: Rallidae)
Species from Rapa Island, French
Polynesia. Taxonomy 2021, 1, 448–457.
https://doi.org/10.3390/
taxonomy1040032
Academic Editor:
Mathias Harzhauser
Received: 24 November 2021
Accepted: 15 December 2021
Published: 20 December 2021
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1. Introduction
Rapa Island, also known as Rapa Iti, is a subtropical island in the South Pacific and
the southernmost inhabitable island of French Polynesia. It has an area of circa 38 km2 and
its origin as a volcanic caldera resulted in a jagged topography that rises abruptly from
the sea, with its highest point being Mont Perau at 650 m above sea level [1]. The first
Polynesian settlers most likely arrived on Rapa Iti between 1100 and 1200 CE and the first
Europeans in 1791 CE [2–4].
The island fauna was dominated by birds, with no native mammals or terrestrial
reptiles [5]. Among other species, Rapa Iti is home to the critically endangered Rapa fruitdove Ptilonopus huttoni Finsch, 1874, while the Rapa shearwater Puffinus myrtae Bourne,
1959, once present on the island, is now confined to offshore islets.
A number of bird bones from archaeological sites are also known from Rapa Iti, which
were studied by Tennyson & Anderson [5]. Among the species those authors identified
was a “Gallirallus-type rail”, which they hypothesized could be an endemic form and
commented that further analysis was necessary to determine its identity. The occurrence
of island-endemic species, particularly when reduced flight capability (or flight loss) is
in play, is a common occurrence in many genera of Rallidae worldwide [6–12]. This is a
particularly widespread phenomenon on the islands of the Caribbean and the South Pacific,
including several species known only from Quaternary fossil material [8,13–16].
In the present paper, we analyse the rail material of Tennyson & Anderson [5] from
Rapa Iti, comparing it to other known living and extinct Gallirallus spp. and describing it
as a new species.
2. Material & Methods
The material of the “Gallirallus-type rail” available from the study of Tennyson &
Anderson [5] is a single left tarsometatarsus, housed in the Museum of New Zealand Te
Papa Tongarewa (NMNZ, Wellington, New Zealand). It is part of a large series of animal
remains (mostly fish) recovered from archaeological sites along the coast of Rapa Iti. The
rail bone is very well preserved and was found in Tangarutu Cave (in Section S1, described
in [17]). For more details on the site and depositional environment, see [17].
4.0/).
Taxonomy 2021, 1, 448–457. https://doi.org/10.3390/taxonomy1040032
https://www.mdpi.com/journal/taxonomy
Taxonomy 2021, 1
449
The new tarsometatarsus was compared to specimens of living and extinct rails
described in the literature, with a focus in the South Pacific taxa of Gallirallus Lafresnaye,
1841, as well as specimens in the collection of the NMNZ (refer to Appendix A for a full
list). A digital calliper (0.01 mm precision, rounded to the nearest 0.1 mm) was used to
take the following measurements of the specimen of interest and comparative material
(Table 1): total length, distal width, proximal width, mid-shaft width, and mid-shaft depth.
Measurements of further species for comparison were taken from the literature (Table 1).
The osteological nomenclature used here follows [18–20].
The species classification in Gallirallus used here follows [21]. A recent phylogenetic
study [22] has split Gallirallus Lafresnaye, 1841 into four genera. For those authors, Gallirallus contains only two species: the weka G. australis and the New Caledonian rail G.
lafresnayanus. They classified the invisible rail G. wallacii into the monospecific genus
Habroptila G.R. Gray, 1861, and erected a further monospecific genus, Aptenorallus Kirchman, McInerney, Giarla, Olson, Slikas & Fleischer, 2021 (formally described in [23], a
corrigendum to that publication), solely for the Calayan rail G. calayanensis. All remaining
species were allocated to the genus Hypotaenidia Reichenbach, 1853 by those authors. While
the taxonomy of this group remains in flux, we retain all species in Gallirallus (Table 1).
Taxonomy 2021, 1
450
Table 1. Gallirallus spp., with their known distribution, status (extinct or living), flight capability, and tarsometatarsus measurements (given in mm). Sex: females (F) and males (M), when
known; Fossil material (†). Number of measured specimens indicated inside parentheses. References for the measurements are given in the respective column (see Appendix A for a list of
specimens measured for the present study).
Gallirallus spp.
G. astolfoi sp. nov.
Geographic
Distribution
Fench Polynesia
(Rapa Iti)
Flight
Status
Total Length
Distal Width
Proximal Width
Mid-Shaft
Depth
†: 2.1 (1)
Reference
†: 5.8 (1)
Mid-Shaft
Width
†: 2.7 (1)
flightless
extinct
†: 34.6 (1)
†: 5.8 (1)
this paper
G. australis
(Sparrman, 1786)
New Zealand
flightless
living
†: 60.8–70.7 (7)
†: 11.1–12.8 (7)
†: 9.7–11.9 (7)
†: 4.7–6.0 (7)
†: 4.1–5.0 (7)
this paper
G. calayanensis
Allen et al., 2004
Philippines
(Calayan Is.)
flightless
living
—
—
—
—
—
—
G. dieffenbachii
(G.R. Gray, 1843)
G. epulare Kirchman &
Steadman, 2007
New Zealand
(Chatham Is.)
French Polynesia
(Nuku Hiva)
flightless
extinct
†: 42.0–48.0 (10)
†: 7.8–8.8 (10)
†: 7.3–8.3 (10)
†: 3.4–4.3 (10)
†: 3.2–4.0 (10)
this paper
flightless
extinct
—
†: 6.5 (1)
—
†: 3.5 (1)
†: 2.4 (1)
[24]
G. ernstmayri Kirchman &
Steadman, 2006
G. gracilitibia Kirchman &
Steadman, 2007
Papua New Guinea
(New Ireland)
French Polynesia
(Ua Huka)
flightless
extinct
—
—
—
—
—
[8]
flightless
extinct
—
—
—
—
—
[8]
G. huiatua Steadman
et al., 2000
Niue
flightless
extinct
†: 39.2 (1)
†: 6.5 (1)
†: 5.4 (1)
†: 2.7 (1)
†: 2.5 (1)
this paper
G. insignis (P.L.
Sclater, 1880)
Papua New Guinea
(New Britain)
flightless
living
—
—
—
—
—
—
G. lafresnayanus Verreaux
& Des Murs, 1860
New Caledonia
flightless
extinct
—
—
—
—
—
—
G. modestus
(Hutton, 1872)
New Zealand
(Chatham Is.)
flightless
extinct
†: 26.4–32.1 (4)
†: 4.8–5.4 (4)
†: 4.4–5.0 (4)
†: 2.1–2.4 (4)
†: 1.7–2.1 (4)
this paper
G. okinawae (Yamashina &
Mano, 1981)
Japan (Okinawa)
flightless
living
—
—
—
—
—
—
G. owstoni
(Rothschild, 1895)
G. pacificus (Gmelin, 1789)
Guam, Mariana Islands
flightless
living
—
—
[13]
flightless?
extinct
F: 7.1 (1)
M: 7.1–8.2 (2)
—
—
French Polynesia
(Tahiti)
F: 49.6 (1)
M: 49.6–53.7 (2)
—
—
—
[12]
—
Taxonomy 2021, 1
451
Table 1. Cont.
Gallirallus spp.
G. pendiculentus
Kirchman &
Steadman, 2006
Geographic
Distribution
Northern Mariana Is.
(Tinian)
Flight
Status
Total Length
Distal Width
Proximal Width
Mid-Shaft
Depth
—
Reference
†: 5.6–7.3 (17)
Mid-Shaft
Width
†: 2.7–3.7 (17)
flightless
extinct
†: 38.4–46.5 (7)
†: 6.1–6.8 (14)
[8]
G. philippensis
(Linnaeus, 1766)
Oceania
volant
living
F: 40.8–45.2 (6)
M: 45.3–49.0 (6)
†: 36.7–43.9 (4)
F: 6.0–6.9 (6)
M: 6.7–7.5 (6)
†: 6.2–7.5 (4)
F: 5.8–6.6 (6)
M: 6.7–7.2 (6)
†: 5.6–7.1 (4)
F: 2.8–3.3 (6)
M: 3.1–3.6 (6)
†: 3.1–3.9 (4)
†: 2.5–3.3 (4)
F/M: [8]
†: this paper
G. pisonii Kirchman &
Steadman, 2006
Northern Mariana Is.
(Aguiguan)
flightless
extinct
—
†: 6.1 (1)
†: 5.6–5.7 (2)
†: 3.0–3.1 (2)
—
[8]
G. poecilopterus
(Hartlaub, 1866)
Fiji
flightless
extinct
†: 66.6–70.0 (2)
†: 10.0 (1)
†: 9.5 (1)
†: 4.0 (1)
—
[25]
G. ripleyi Steadman, 1986
G. roletti Kirchman &
Steadman, 2007
Cook Is. (Mangaia)
French Polynesia
(Tahuata)
flightless
flightless
extinct
extinct
†: 33.1 (1)
—
†: 6.3–6.5 (3)
†: 8.7 (1)
†: 5.7–5.9 (2)
†: 7.9 (1)
†: 2.8–3.1 (2)
†: 3.5–3.9 (2)
—
†: 2.5–2.8 (2)
[8,13]
[24]
G. rovianae
Diamond, 1991
G. sp. sensu Steadman &
Bollt, 2010
G. steadmani Worthy &
Bollt, 2011
G. storrsolsoni Kirchman
& Steadman, 2006
G. sylvestris (P.L.
Sclater, 1870)
Solomon Is.
flightless
living
—
—
—
—
—
—
Fench Polynesia
(Rurutu)
Fench Polynesia
(Tubuai)
Fench Polynesia
(Huahine)
Australia (Lord
Howe Is.)
flightless
extinct
—
—
—
—
—
[26]
flightless
extinct
†: 40.0 (1)
†: 6.2 (1)
†: 5.9–6.4 (3)
†: 2.9–3.1 (3)
—
[27]
flightless
extinct
—
—
†: 7.4 (1)
†: 4.1 (1)
†: 2.8 (1)
[28]
flightless
living
M: 49.6 (1)
M: 8.9 (1)
M: 8.7 (1)
M: 4.4 (1)
M: 3.4 (1)
this paper
G. temptatus Kirchman &
Steadman, 2006
Northern Mariana
Is. (Rota)
flightless
extinct
†: 47.4 (1)
†: 7.4 (1)
†: 6.4–6.8 (2)
†: 3.3–3.5 (2)
—
[8]
G. torquatus
(Linnaeus, 1766)
Indonesia, Papua New
Guinea, Philippines
volant
living
M: 54.2 (1)
—
M: 7.4 (1)
—
—
[13]
Taxonomy 2021, 1
452
Table 1. Cont.
Gallirallus spp.
Geographic
Distribution
Tonga (Vava’u)
Flight
Status
Total Length
Distal Width
Proximal Width
flightless
extinct
†: 70.9–73.0 (2)
†: 12.2–13.1 (2)
G. vekamatolu Kirchman &
Steadman, 2005
Tonga (’Eua)
flightless
extinct
†: 45.1 (1)
G. wakensis
(Rothschild, 1903)
USA (Wake Is.)
flightless
extinct
G. wallacii (G.R.
Gray, 1861)
Indonesia (Halmahera)
flightless
G. woodfordi
(Ogilvie-Grant, 1889)
Solomon Is.
flightless
G. vavauensis (Worthy &
Burley, 2020)
Mid-Shaft
Depth
†: 5.4–6.0 (6)
Reference
†: 12.9–13.2 (3)
Mid-Shaft
Width
†: 5.9–6.7 (6)
—
†: 7.7–8.6 (2)
—
—
[13]
F: 33.3 (1)
M: 35.6 (1)
F: 5.9 (1)
M: 6.2 (1)
F: 5.5 (1)
M: 6.0 (1)
F: 2.5 (1)
M: 2.9 (1)
—
[8]
living
—
—
—
—
—
—
living
F: 67.7–72.0 (3)
M: 69.8–72.3 (3)
F: 10.5–10.6 (3)
M: 10.7–11.0 (3)
F: 9.9–10.1 (3)
M: 10.2–10.7 (3)
F: 4.4–4.7 (3)
M: 4.8–5.0 (3)
—
[8]
[25]
Taxonomy 2021, 1
453
3. Systematics
The morphology of the Rapa Iti tarsometatarsus indicates that it could only belong
to the Order Gruiformes and family Rallidae, given the presence of two open tendinal
canals, one distal foramen, and the tendinal bridge [19]. More specifically, the Rapa Iti
fossil can be allocated to the genus Gallirallus due to the following diagnostic features
of the tarsometatarsus (as per [8]: p. 8): “corpus tarsometatarsi much wider than deep;
medial sulcus hypotarsi not enclosed; fossa parahypotarsalis medialis shallow in proximal
aspect; fossa metatarsi I short and shallow; crista plantaris mediana slopes gradually (not
steeply) to hypotarsus; distal end of trochlea metatarsi tertii sloped toward medial trochlea;
cotyla medialis is rectangular in proximal aspect with flat (not rounded) dorsal margin”.
According to our comparative analysis with tarsometatarsi of living and extinct species
of Gallirallus (see below), we are confident in assigning the present fossil to a new species,
apparently endemic to Rapa Iti.
Family Rallidae
Genus Gallirallus Lafresnaye, 1841
Gallirallus astolfoi sp. nov.
(Figure 1)
rail (cf. Gallirallus): Tennyson & Anderson, 2012: 108.
ZooBank reg. nr.: urn:lsid:zoobank.org:act:F606759B-4C48-491E-93F0-0E8C018B3B97.
Holotype: NMNZ S.044399 (left tarsometatarsus; A. Anderson col. 21/vii/2002).
Type locality: French Polynesia, Rapa Island (Rapa Iti), Tangarutu Cave, Section S1
(30–40 cm).
Etymology: The specific epithet honours Astolfo, one of Charlemagne’s fictional paladins.
In the epic Orlando Furioso, Astolfo becomes trapped on a remote island because of the
sorceress Alcina.
Diagnosis: Tarsometatarsus small (ca. 34.5 mm long), of delicate appearance, with narrow
and shallow shaft, and narrow trochleae (particularly the trochlea metatarsi II).
Differential diagnosis: The tarsometatarsus of Gallirallus astolfoi sp. nov. (Figure 1) is
considerably smaller than most other congeners (Table 1), with the exception of G.
ripleyi from the Cook Islands and G. wakensis from the Wake Island, both flightless and
extinct [8,24]. Its shaft, however, is proportionately much narrower and shallower, which
gives the bone a more delicate appearance. The Chatham Island rail G. modestus (previously
classified into its own genus, Cabalus) is much smaller and its tarsometatarsi have a more
compact appearance.
The single known tarsometatarsus of G. huiatua from Niue is similar to that of G.
astolfoi sp. nov. in its delicate appearance (i.e., shaft width and depth, and trochleae
width), but it is more elongate. The tarsometatarsus of G. pendiculentus from the Northern
Marianas and of G. steadmani from Tubuai, French Polynesia, are similar in shape to G.
astolfoi sp. nov. but are much larger (Table 1). Furthermore, G. steadmani apparently
has proportionately narrower trochlea (trochlea metatarsorum III and IV), though that
could have been overemphasized by the sub-optimal preservation of the specimens ([27]:
figure 4L).
In comparison with larger congeners, the tarsometatarsus of G. astolfoi sp. nov. is more
delicate, with its proportionately narrower and shallower mid and distal shaft, as well
as its narrower trochleae. Compared with G. philippensis and other species from French
Polynesia, namely G. epulare from Nuku Hiva, G. roletti from Tahuata, and G. storrsolsoni
from Huahine, the latter two, in particular, have much stouter tarsometatarsi, with a wider
mid-shaft ([24]: figure 6A,B). Furthermore, the trochlea of digit 2 (trochlea metatarsi II)
of G. astolfoi sp. nov. is less expanded than in G. philippensis and some island-endemic
congeners such as G. roletti and G. dieffenbachii.
Taxonomy 2021, 1
454
Figure 1. Left tarsometatarsus (holotype, NMNZ S.044399) of Gallirallus astolfoi sp. nov. in different
views: (A) anterior, (B) caudal, (C) lateral, (D) medial, (E) proximal, (F) distal.
There are two further extinct endemic flightless species from French Polynesia: G.
gracilitibia from Ua Huka and G. pacificus from Tahiti [12,24] (Table 1). No tarsometatarsi are
known for these taxa, which precludes comparison with G. astolfoi sp. nov. Nevertheless,
there is little chance that they could be conspecific as these islands are respectively 2100 km
and 1200 km away from Rapa Iti. The only other Rallidae species on Rapa Iti is the
widespread and much smaller spotless crake, Zapornia tabuensis (Gmelin, 1789) [5]. There
is a further undescribed fossil species, Gallirallus sp. of [26], that is endemic to Rurutu (and
extinct), though there is no published information or photographs available.
4. Discussion
Tennyson & Anderson [5] identified four extinct species on Rapa Iti, among which
is Gallirallus astolfoi sp. nov. On the tropical Pacific islands, extinction of the avifauna
is assigned to anthropogenic impact and the numbers of extinct land bird species are
dominated by rails [29,30]. More specifically in French Polynesia (albeit similar to many
other islands worldwide), the main causes of population crashes and extinctions are
predation of adults, chicks and eggs by mammals (particularly humans, Pacific rats Rattus
exulans, and feral cats), alongside land alteration and habitat destruction by humans and
feral goats [3,31,32]. While Pacific rats were introduced by Polynesians, cats and goats
(alongside cattle, pigs and dogs) were introduced by Europeans [5,32–34].
While we have little direct evidence to ascertain whether Gallirallus astolfoi sp. nov.
was hunted for food by humans, this seems probable. While most of the animals eaten
by the inhabitants of rock shelters like Tangarutu Cave, where the present rail bone was
found, seem to have been fish [2], birds were potentially consumed too. Seven species of
birds were reported from Section S1 of Tangarutu Cave alone (from a total of 15 bird taxa
from all archaeological sites on Rapa Iti) [5]. While there was no clear evidence of the bones
being deposited as part of a midden rather than by natural accumulation (i.e., no signs of
butchery or charring), the cultural context of the sites is clear [5]. Occupation of Tangarutu
Taxonomy 2021, 1
455
Cave likely dates back to between 1400 and 1600 CE [2]. Thus, the deposit happened after
human settlement; furthermore, in the case of Tangarutu Cave, this age is also supported
by the presence of rat bones in Section S1.
Given its smaller size in comparison to volant congeners and its presence on a remote
island, we hypothesize that Gallirallus astolfoi sp. nov. had reduced flight capacity or, potentially, it was flightless. As mentioned above, flightlessness is a common feature in Rallidae,
occurring independently in several genera and different species within the same genus.
Earlier workers [11,35] hypothesized that flightlessness in rails was a neotenic condition
involving little genetic modification and thus able to occur through rapid evolution. Later
works offered support to the heterochrony hypothesis, arguing that a change in timing
of development could lead to the observed allometric differences in rails (see [9] for a
review). Likewise, molecular clock estimates have supported the hypothesis that evolution
of flightless Gallirallus species in Oceania has been rapid after the appearance of the genus
in the Late Miocene [22,36–38]. Furthermore, phylogenetic studies offered evidence that
flightlessness can evolve prior to reproductive isolation [38].
Only two out of circa 30 species of Gallirallus are volant and widespread (Table 1); most
species are flightless and endemic to single islands (or multiple islands, in cases where they
were formerly connected due to the lower sea levels of the late Pleistocene [28]). Of the two
volant species, G. philippensis is the most widespread and it is hypothesized that breakaway
populations of it have colonized many islands throughout the Pacific, giving rise to the
endemic forms [8]. As such, it is notable that most of these endemic species evolved
the common pattern of flightlessness plus increased body size (e.g., G. roletti in French
Polynesia; Table 1). It has been argued that loss of flight was typically related to increased
body size, which could mean that such a combination has a greater selective advantage
and/or is ontogenetically more parsimonious [9]. Nevertheless, Gallirallus astolfoi sp. nov.
is one of the few species of the genus that apparently underwent a decrease in size (Table 1).
Therefore, despite the relatively similar areas and environments of these islands, it seems
that different selective pressures towards local optima were in play on each of them and
that Gallirallus defies generalizations of insular gigantism/dwarfism (cf. [9]).
5. Conclusions
Gallirallus astolfoi sp. nov. from Rapa Iti is the seventh extinct species in the genus
to be described from French Polynesia (Table 1), excluding the potential undescribed one
mentioned above. Excluding the volant G. philippensis, the species geographically closest
to G. astolfoi sp. nov. is G. steadmani from Tubuai, ca. 700 km NW from Rapa Iti. As new
specimens continue to be discovered and described, the scenario of a multitude of endemic
rail species across the Pacific Islands is becoming more evident, offering further evidence
in support of the above-mentioned hypothesis [8,30]. Likewise, the list of species extinct
after human contact during the past millennia is becoming more extensive, adding to the
corpus of data on the demise of insular faunas.
Author Contributions: Conceptualization, methodology, R.B.S. and A.J.D.T.; investigation, data
curation, writing—original draft preparation, R.B.S.; writing—review and editing, R.B.S., A.A. and
A.J.D.T.; supervision, A.A. and A.J.D.T. All authors have read and agreed to the published version of
the manuscript.
Funding: This research received no external funding.
Data Availability Statement: All data can be found within the article.
Acknowledgments: We are very grateful to Jean-Claude Stahl (NMNZ) for photographs of the
specimen; to Christopher Milensky (Smithsonian National Museum of Natural History, Washington,
DC, USA) for comparative photos of G. wakensis; to Douglas Kennett and Eric Conte for co-organizing
the fieldwork in which the present fossil was discovered; and to the two anonymous reviewers for
their helpful comments and suggestions.
Conflicts of Interest: The authors declare no conflict of interest.
Taxonomy 2021, 1
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Appendix A
The following specimens of Rallidae from the NMNZ collection were used for comparison. The list is arranged alphabetically by species name, with acronym (NMNZ) omitted
from registration numbers for brevity. Measured specimens (used in Table 1) are identified
by an asterisk (*).
Capellirallus karamu Falla, 1954: S.023241. Diaphorapteryx hawkinsi (Forbes, 1892):
OR.007997. Fulica atra australis Gould, 1845: OR.024566. Fulica chathamensis Forbes, 1892:
S.026519. Fulica prisca Hamilton, 1893: S.033721. Gallinula hodgenorum (Scarlett, 1955):
S.033724. Gallinula tenebrosa Gould, 1846: OR.024520. Gallinula ventralis Gould, 1837:
OR.022100. Gallirallus australis australis (Sparrman, 1786): OR.014994. Gallirallus australis
greyi (Buller, 1888): S.00573, S.023353*, S.023387*, S.027815*, S.034078*, S.042193*, S.045588*.
Gallirallus australis hectori (Hutton, 1873): OR.0025361. Gallirallus australis scotti (OglivieGrant, 1905): OR.018327. Gallirallus dieffenbachii (G.R. Gray, 1843): S.026625*, S.026924*,
S.026943*, S.027616*, S.027766*, S.030037, S.031762*, S.032013*, S.032092*, S.033198*, S.044038*.
Gallirallus huiatua Steadman et al., 2000: S.037708* (holotype). Gallirallus modestus (Hutton,
1872): S.026258*, S.026967*, S.027607*, S.031303*, S.031755. Gallirallus philippensis assimilis
(G.R. Gray, 1843): OR.023821. Gallirallus philippensis goodsoni (Mathews, 1911): S.035228*.
Gallirallus philippensis mellori (Mathews, 1912): S.045796*. Gallirallus philippensis sethsmithi
(Mathews, 1911): S.038224*, S.038383*. Gallirallus sylvestris (P.L. Sclater, 1870): S.027218a*.
Lewinia muelleri (Rothschild, 1893): OR.025556. Porphyrio mantelli (Owen, 1848): S.028440.
Porphyrio hochstetteri (A.B. Meyer, 1883): OR.024641. Porphyrio melanotus melanotus Temminck, 1820: OR.024252. Zapornia pusilla affinis (J.E. Gray, 1845): OR.024417. Zapornia
tabuensis tabuensis (Gmelin, 1789): OR.020984, S.044596.
References
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
Anderson, A.; Kennett, D.J.; Conte, E. Archaeological research on Rapa Island, French Polynesia. Terra Aust. 2012, 37, 7–23.
Anderson, A.; Kennett, D.J.; Conte, E. The prehistory of Rapa Island. Terra Aust. 2012, 37, 247–256.
Kennett, D.; Anderson, A.; Prebble, M.; Conte, E.; Southon, J. Prehistoric human impacts on Rapa, French Polynesia. Antiquity
2006, 80, 340–354. [CrossRef]
Richards, R. The earliest foreign visitors and their massive depopulation of Rapa-iti from 1824 to 1830. J. Société Océanistes 2004,
118, 3–10. [CrossRef]
Tennyson, A.; Anderson, A. Bird, reptile and mammal remains from archaeological sites on Rapa Island. Terra Aust. 2012, 37,
105–114.
Alcover, J.A.; Pieper, H.; Pereira, F.; Rando, J.C. Five new extinct species of rails (Aves: Gruiformes: Rallidae) from the
Macaronesian Islands (North Atlantic Ocean). Zootaxa 2015, 4057, 151–190. [CrossRef] [PubMed]
Bourne, W.R.P.; Ashmole, N.P.; Simmons, K.E.L. A new subfossil night heron and a new genus for the extinct rail from Ascension
Island, central tropical Atlantic Ocean. Ardea 2003, 91, 45–51.
Kirchman, J.J.; Steadman, D.W. Rails (Rallidae: Gallirallus) from prehistoric archaeological sites in Western Oceania. Zootaxa 2006,
1316, 1–31. [CrossRef]
Livezey, B.C. Evolution of flightlessness in rails (Gruiformes: Rallidae): Phylogenetic, ecomorphological, and ontogenetic
perspectives. Ornithol. Monogr. 2003, 53, 1–654. [CrossRef]
Matsuoka, H. The Late Pleistocene fossil birds of the central and southern Ryukyu Islands, and their zoogeographical implications
for the recent avifauna of the archipelago. Tropics 2000, 10, 165–188. [CrossRef]
Olson, S.L. A classification of the Rallidae. Wilson Bull. 1973, 85, 381–416.
Taylor, B.; van Perlo, B. Rails. A Guide to the Rails, Crakes, Gallinules and Coots of the World; Pica Press: Sussex, UK, 1998; 600p.
Kirchman, J.J.; Steadman, D.W. Rails (Aves: Rallidae: Gallirallus) from prehistoric sites in the Kingdom of Tonga, including a
description of a new species. Proc. Biol. Soc. Wash. 2005, 118, 465–477. [CrossRef]
Olson, S.L.; Wingate, D.B. Two new species of flightless rails (Aves: Rallidae) from the Middle Pleistocene “crane fauna” of
Bermuda. Proc. Biol. Soc. Wash. 2000, 113, 356–368.
Olson, S.L.; Wingate, D.B. A new species of large flightless rail of the Rallus longirostris/elegans complex (Aves: Rallidae) from the
Late Pleistocene of Bermuda. Proc. Biol. Soc. Wash. 2001, 114, 509–516.
Takano, O.M.; Steadman, D.W. Another new species of flightless rail (Aves: Rallidae: Rallus) from Abaco, The Bahamas. Zootaxa
2018, 4407, 376–382. [CrossRef] [PubMed]
Anderson, A. Archaeology of the coastal sites on Rapa Island. Terra Aust. 2012, 37, 47–76.
Baumel, J.J.; Witmer, L.M. Osteologia. In Handbook of Avian Anatomy: Nomina Anatomica Avium; Baumel, J.J., King, A.S., Breazile,
J.E., Evans, H.E., Vanden Berge, J.C., Eds.; Nuttall Ornithological Club: Cambridge, UK, 1993; pp. 45–132.
Gilbert, B.M.; Martin, L.D.; Savage, H.G. Avian Osteology; Modern Printing: Laramie, WY, USA, 1981; p. 252.
Taxonomy 2021, 1
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
457
Livezey, B.C.; Zusi, R.L. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy.
II. Analysis and discussion. Zool. J. Linn. Soc. 2007, 149, 1–95. [CrossRef]
Winkler, D.W.; Billerman, S.M.; Lovette, I.J. Rails, gallinules, and coots (Rallidae), version 1.0. In Birds of the World; Billerman,
S.M., Keeney, B.K., Rodewald, P.G., Schulenberg, T.S., Eds.; Cornell Lab of Ornithology: Ithaca, NY, USA, 2020. [CrossRef]
Kirchman, J.J.; McInerney, N.R.; Giarla, T.C.; Olson, S.L.; Slikas, E.; Fleischer, R.C. Phylogeny based on ultra-conserved elements
clarifies the evolution of rails and allies (Ralloidea) and is the basis for a revised classification. Ornithology 2021, 138, 1–21.
[CrossRef]
Kirchman, J.J.; McInerney, N.R.; Giarla, T.C.; Olson, S.L.; Slikas, E.; Fleischer, R.C. Corrigendum to: Phylogeny based on ultraconserved elements clarifies the evolution of rails and allies (Ralloidea) and is the basis for a revised classification. Ornithology
2021, 138, ukab065. [CrossRef]
Kirchman, J.J.; Steadman, D.W. New species of extinct rails (Aves: Rallidae) from archaeological sites in the Marquesas Islands,
French Polynesia. Pac. Sci. 2007, 61, 145–163. [CrossRef]
Worthy, T.H.; Burley, D.V. Prehistoric avifaunas from the Kingdom of Tonga. Zool. J. Linn. Soc. 2020, 189, 998–1045. [CrossRef]
Steadman, D.W.; Bollt, R. Prehistoric Birds from Rurutu, Austral Islands, East Polynesia. Pac. Sci. 2010, 64, 315–325. [CrossRef]
Worthy, T.H.; Bollt, R. Prehistoric birds and bats from the Atiahara Site, Tubuai, Austral Islands, East Polynesia. Pac. Sci. 2011, 65,
69–85. [CrossRef]
Kirchman, J.J.; Steadman, D.W. New species of rails (Aves: Rallidae) from an archaeological site on Huahine, Society Islands. Pac.
Sci. 2006, 60, 281–297. [CrossRef]
Steadman, D.W. Extinction of birds in Eastern Polynesia: A review of the record, and comparisons with other Pacific Island
groups. J. Archaeol. Sci. 1989, 16, 177–205. [CrossRef]
Steadman, D.W. Prehistoric extinctions of Pacific island birds: Biodiversity meets zooarchaeology. Science 1995, 267, 1123–1131.
[CrossRef] [PubMed]
Seitre, R.; Seitre, J. Causes of land-bird extinctions in French Polynesia. Oryx 1992, 26, 215–222.
Thibault, J.-C.; Varney, A. Numbers and habitat of the Rapa fruit-dove, Ptilinopus huttoni. Bird Conserv. Int. 1991, 1, 75–81.
[CrossRef]
Holyoak, D.T.; Thibault, J.-C. Contribution à L’étude des Oiseaux de Polynésie Orientale. Mémoires du Muséum National D’Histoire
Naturelle Série A Zoologie 1984, 127, 1–209.
Thibault, J.-C.; Varney, A. Breeding seabirds of Rapa (Polynesia): Numbers and changes during the 20th century. Bull. Br. Ornithol.
Club 1991, 111, 70–77.
Olson, S.L. Evolution of the rails of the South Atlantic islands (Aves: Rallidae). Smithson. Contrib. Zool. 1973, 152, 1–53. [CrossRef]
García-R, J.C.; Gibb, G.C.; Trewick, S.A. Deep global evolutionary radiation in birds: Diversification and trait evolution in the
cosmopolitan bird family Rallidae. Mol. Phylogenet. Evol. 2014, 81, 96–108. [CrossRef] [PubMed]
García-R, J.C.; Gibb, G.C.; Trewick, S.A. Eocene diversification of crown group rails (Aves: Gruiformes: Rallidae). PLoS ONE 2014,
9, e109635. [CrossRef] [PubMed]
Kirchman, J.J. Speciation of flightless rails on islands: A DNA-based phylogeny of the typical rails of the Pacific. Auk 2012, 129,
56–69.