Youngetaldodonomenclature

Zoological Journal of the Linnean Society, 2024, 201, zlae086
https://doi.org/10.1093/zoolinnean/zlae086
Advance access publication 16 August 2024
Original Article
Original Article
The systematics and nomenclature of the Dodo and the
Solitaire (Aves: Columbidae), and an overview of columbid
family-group nomina
Downloaded from https://academic.oup.com/zoolinnean/article/201/4/zlae086/7733394 by guest on 20 August 2024

Mark T. Young1,*, , Julian P. Hume2, , Michael O. Day3, Robert P. Douglas4,
Zoë M. Simmons4, , Judith White2, Markus O. Heller5,6,7, , Neil J. Gostling1,7,*,
1
School of Biological Sciences, Faculty of Environmental and Life Sciences, University of Southampton, Southampton, UK
2
Bird Group, Natural History Museum, Akeman Street, Tring HP23 6AP, UK
3
Fossil Reptiles, Amphibians, and Birds Section, Natural History Museum, Cromwell Road, London SW7 5BD, UK
4
Oxford University Museum of Natural History, Parks Road, Oxford OX1 3PW, UK
5
Bioengineering Research Group, Faculty of Engineering and Physical Sciences, University of Southampton, Southampton, UK
6
Centre for Sport, Exercise and Osteoarthritis Research Versus Arthritis, Southampton, UK
7
Institute for Life Sciences, University of Southampton, Southampton, UK
Corresponding authors. School of Biological Sciences, Faculty of Environmental and Life Sciences, University of Southampton, Southampton, UK. E-mail: mark.
*
young@soton.ac.uk and Institute for Life Sciences, University of Southampton, Southampton, UK. E-mail: n.j.gostling@soton.ac.uk
ABSTR ACT
The Dodo and its extinct sister species, the Solitaire, are iconic exemplars of the destructive capabilities of humanity. These secondarily terrestrial
columbids became extinct within a century of their first encounter with humanity. Their rapid extinction, with little material retained in natural
history collections, led 18th and some early 19th century naturalists to believe that these aberrant birds were mythological. This meant that the
nomenclatural publications in which their scientific nomina were established were based on accounts written before the species became extinct.
As such, no type specimens were designated for either the Dodo or the Solitaire. Our in-depth historical overview of both species and associated
family-group nomina found that the nominal authority of the Dodo-based family group is not what is reported in the literature. Moreover, our
detailed review of the family-group nomina based on columbid genera ensures that the current columbid family-group systematization is valid.
Changing nomenclatural norms between the 19th and 20th centuries had a profound impact on Dodo nomenclature; so much so that the Dodo
is an example of how pervasive nomenclatural ‘ripples’ can be and a warning for our current world of multiple nomenclatural codes.
Keywords: Columbidae; Dodo; family group; nomenclature; Pezophaps; Raphus; validity
INTRODUCTION many persons with the Griffin and the Phœnix of mythological
The Mauritian Dodo, †Raphus cucullatus (Linnaeus, 1758) antiquity’. The existence of the Solitaire, in particular, was long
(Fig. 1), and the Rodrigues Solitaire, †Pezophaps solitaria doubted, because for several decades it was known solely from
(Gmelin, 1789) (Fig. 2), are textbook examples of evolutionary the descriptions by Leguat (1708). Strickland (1844: 324) men-
transitions and of human-made extinctions. Their morphologies tioned that the Solitaire had been considered either ‘fictitious,
were so aberrant that for a time, during the 18th and early 19th or to be founded on an imperfect description of the true Dodo’.
centuries, they were considered mythological (Duncan 1828, de A series of key papers during the early 19th century ‘res-
Blainville 1835, Strickland 1844, 1848, Hume 2006; see Figs 1, urrected’ the Dodo and the Solitaire from the realm of the
2). As said by Strickland (1848: 4): ‘So rapid and so complete mythological to the material (Duncan 1828, de Blainville 1835,
was their extinction that the vague descriptions given of them Strickland 1844). The seminal work of Strickland (1848) and
[Dodo and Solitaire] by early navigators were long regarded as Melville (1848), in their shared volume, described in detail the
fabulous or exaggerated, and these birds, almost contemporaries anatomy of specimens still found in European collections at that
of our great-grandfathers, became associated in the minds of time, in addition to giving an authoritative account of the history
Received 5 April 2024; revised 31 May 2024; accepted 14 June 2024

[Version of Record, first published online 16 August 2024, with fixed content and layout in compliance with Art. 8.1.3.2 ICZN. http://zoobank.org/urn:lsid:zoobank.
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© 2024 The Linnean Society of London.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits
unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.
2 • Young et al.

Figure 1. Dodo (†Raphus cucullatus) mounted composite skeleton [NHMUK S/1988.50.1 (PV A 3302)]. A, cranial view. B, left lateral view.
of the two species. However, it was not until new expeditions understanding of these aberrant birds is being revolutionized.
to the islands of Mauritius and Rodrigues during the 1860s To ensure that this work is on a firm basis, we need to ensure
that new incomplete skeletons of both species were discovered. that the alpha and beta taxonomy (and accompanying nomen-
The skeletal remains discovered in the ‘Mare aux Songes’ marsh clature) of both species is stable. As we will show, there are no
during 1865 (Clark 1866, Hume et al. 2009) allowed the Dodo known type specimens for either species. Moreover, given that
to be described more fully (Owen 1866), and the Solitaire was the use of Dodo-based (i.e. †Raphus) family-group nomina is
described by Newton and Newton (1868, 1869) after the Jenner now accepted within columbid systematics, we need to ensure
excavations of 1865 discovered skeletal remains (Parish 2013: that these names are themselves valid, in order to maintain the
234; Hume et al. 2015). nomenclatural stability of extant pigeons and doves. To those
There has been renewed interest in the biology of the Dodo ends, we provide an in-depth historical overview of the Dodo,
and the Solitaire in the 21st century. Studies have explored the Solitaire, and the family-group nomina based upon them. We
Dodo body mass (Brassey et al. 2016, van Heteren et al. 2017) also establish a new nomen to unite both species: †Raphina.
and bone histology (Angst et al. 2017), and the endocranial
anatomy of both species has been reconstructed digitally from Terminology and nomenclatural background
computed tomography scans (Gold et al. 2016). New Dodo Before starting our historical overview, it is worth stating that the
material has been discovered from Mare aux Songes, and the current rules of zoological nomenclatural are ‘relatively’ recent
ecosystem of the Mare aux Songes Lagerstätte has been studied and have evolved from prior rules/suggestions made during the
(see Rijsdijk et al. 2009, 2015, Meijer et al. 2012). The remark- 19th century. We wish this to be clear from the outset, in order
able ‘Thirioux Dodos’ have been described in-depth, which in- that readers will not mistake our comments hereafter as undue
cludes the most complete Dodo skeleton known (Claessens criticisms of past workers. There have also been dramatic shifts in
and Hume 2015; Claessens et al. 2015). There have even been both systematics (the paradigms and methods used to hypothe-
attempts to reconstruct digitally how these animals would size clades) and nomenclature (the establishment of names for
have looked (Rodríguez-Pontes 2016). With each decade, our said clades, and the rules governing those names) between the
Dodo systematics and nomenclature • 3
Figure 2. Solitaire (†Pezophaps solitaria) mounted skeletons (on display at the Royal College of Surgeons, London, UK in 2023). A, female
individual. B, male individual. Note the difference in skeleton size and robusticity between the sexes.
18th and 21st centuries. During the 18th and 19th centuries, the Ornithologists’ Union 1886, Blanchard 1889, Bütschli et al.
rules and norms of zoological nomenclature were being devel- 1893), prior to their widespread formulation and promulga-
oped (e.g. Linnaeus 1758, Kirby 1815, Westwood 1836, 1837a, tion during the 20th century (ICZN 1905, 1961, 1964, 1985,
1837b, Strickland 1837, 1878, Strickland et al. 1843, Dall 1878, 1999). Moreover, the paradigms used to hypothesize taxa were
Société Zoologique de France 1881, Douvillé 1882, American distinctly different, with the transition from a pre-evolutionary
4 • Young et al.
paradigm to an acceptance of paraphyletic groupings and groups nomenclature (for further details, see Linsley and Usinger 1959,
united based on shared similarity, which then shifted to our cur- Bock 1994).
rent paradigm based on shared common ancestry and mono- Zoological nomenclature of the 18th and early 19th cen-
phyletic groups (for a general overview of thought, see Mayr turies did not adhere to the quasi-legal system in place today.
1942, 1965, 1982, Hennig 1966, Nelson 1973, de Queiroz 1988, The renaming of pre-existing genera and specific epithets was
Mishler 2009; and for some clade-specific examples, see Allard commonplace (particularly up to the 1840s–1850s). Therefore,
et al. 1999, Dornburg and Near 2021, Cotterill et al. 2014 and the readers should not be surprised that the principal of priority
references therein). with regard to nominal authority was not adhered to in Dodo
The current International Code of Zoological Nomenclature nomenclature during this time period or that the formulation of
(the Fourth Edition, ICZN 1999, 2003, 2012, 2016; ‘Zoological names does not meet the requirements on the Zoological Code
Code’ hereafter) is a direct descendent of ‘Blanchard’s as we understand it today (ICZN 1999). It is also worth noting
Code’ (Blanchard 1889) via the Règles Internationales de la that when the Dodo and Solitaire were first named (Linnaeus

Nomenclature Zoologique [International Rules of Zoological 1766, Gmelin 1789), the concept of nomenclatural types did
Nomenclature] (ICZN 1905). Raphaël Blanchard, the ‘father of not exist. Witteveen (2016: 156) credited Westwood (1837a) as
International Zoological Nomenclature’ (Bock 1994: 33), was the originator of this concept, which then became incorporated
the Chair of the nomenclatural committee of the International into ‘Strickland’s Code’ (and subsequent nomenclatural codes).
Congress of Zoology, the first President of the International As such, type specimens were not designated for the Dodo or
Committee on Zoological Nomenclature, and the Editor of the Solitaire.
the French edition of the Règles Internationales. For the first Before continuing, we also need to define the terminology we
International Congress of Zoology, he wrote an overview of zoo- will be using. We will follow the suggestions and recommenda-
logical nomenclature and outlined what he believed would be an tions of Dubois and Fitzhugh. Dubois (2021: 39) noted that,
acceptable set of rules for the international corpus of zoologists ‘the term taxonomy is traditionally used in two distinct senses,
(‘Blanchard’s Code’; Blanchard 1889). ‘Blanchard’s Code’ did to designate either a scientific discipline, or any scientific classi-
not exist in a vacuum, because a plethora of nomenclatural codes fication of organisms produced by this discipline and adopted
for zoology had been proposed during the 19th century, with as valid by taxonomists’. In order to distinguish between both
the earliest comprehensive code being proposed by the British meanings, Dubois (2005: 406) erected the term ergotaxonomy
Association for the Advancement of Science (‘Strickland’s for the latter (‘classification used by a given author in a given
Code’). work’, Dubois 2006: 250). To remove any ambiguity, we use the
‘Strickland’s Code’ (Strickland et al. 1843) was formulated by a term ergotaxonomy to refer to any ‘taxonomic framework’ con-
committee of British zoologists and palaeontologists (including sidered valid by their proposer.
famous individuals, such as Charles Darwin and Richard Owen, We will use the term ‘systematics’ rather than ‘taxonomy’
in addition to Hugh Strickland, who was pivotal in our under- throughout. There is disagreement within the field of evolu-
standing of the Dodo and the Solitaire), who set down many of tionary biology regarding whether taxonomy and systematics
the norms we recognize today; norms of the so-called ‘Linnean’ are different subfields (e.g. Simpson 1961, Wiley and Lieberman
system of nomenclature, although this is perhaps more accur- 2011), whether taxonomy is a subdiscipline within systematics
ately called ‘Linnean–Westwoodian–Stricklandian’ nomen- (e.g. Michener et al. 1970, Dubois 2006, Pavlinov 2013, Winsor
clature (sensu Dubois 2011: 4–5). However, there were some 2023 and the references therein), or whether systematics is a
important differences between ‘Strickland’s Code’ and the cur- subfield of taxonomy (e.g. Toepfer 2011). However, others, such
rent Zoological Code (ICZN 1999), such as the proposed ‘start as Mayr and Ashlock (1991) and Fitzhugh (2008), have pro-
date’ for zoological nomenclature, which in ‘Strickland’s Code’ posed that taxonomy is a synonym of systematics. We will follow
was 1766, beginning with the publication of the 12th edition Fitzhugh (2008: 54) and use the term ‘systematics’ throughout.
of Systema Naturæ (Linnaeus 1766). The ensuing controversy We also use the term ‘systematization’ in preference to ‘clas-
over the ‘start date’ for zoological nomenclature cost ‘Strickland’s sification’ following Fitzhugh (2008). Fitzhugh (2008: 54)
Code’ support amongst zoologists (Linsley and Usinger 1959: defined classification as the ‘segregation of objects into classes
41), with Dall (1878: 15) noting that the starting point used based on specified properties’, whereas systematization is ‘the or-
by the British Association had begun ‘admitting to recogni- ganization of observations into a system of concepts, in the form
tion some ichthyological works printed between the dates of of hypotheses, according to theory’ (the definitions of these
the tenth and twelfth editions [of Systema Naturæ]’. Other na- terms given by de Queiroz 1988: 241 was similar). We consider
tional societies began proposing their own nomenclatural codes, the latter to be the best description of systematics, because both
including the American Association for the Advancement of species and ‘higher-level’ clades are explanatory hypotheses ra-
Science (Dall 1878), the Société Zoologique de France (Société ther than objects (e.g. see Fitzhugh 2005, 2008, Mortimer et al.
Zoologique de France 1881), the American Ornithologists’ 2021).
Union (American Ornithologists’ Union 1886), and the Herein, we follow ornithological convention and capit-
Deutsche Zoologische Gesellschaft (Bütschli et al. 1893), as did alize English vernacular names of species (Parkes 1978; and
the Congrès international de géologie [International Congress of the International Ornithological Committee World Bird List
Geology] (Douvillé 1882). It was ‘Blanchard’s Code’ (Blanchard v.13.2; https://www.worldbirdnames.org/english-names/
1889) and the subsequent Règles Internationales (ICZN 1905) spelling-rules/). Moreover, we use the English vernacular
that would begin to bring international stability to zoological names for columbid species that appear in the International
Ornithological Committee World Bird List v.13.2 (https:// the Zoological Code, which states that the ‘starting point’ for
www.worldbirdnames.org/new/bow/pigeons/), but with the zoological nomenclature is on 1 January 1758, and two works
following exceptions: (i) Didunculus strigirostris ( Jardine, 1845) are deemed to have been published on that date, one of which
is referred to as the Samoan Tooth-billed Pigeon, because an- is that of Linnaeus (1758). Therefore, the ‘Raphus’ in Möhring
other species (†Didunculus placopedetes Steadman, 2006) was (1752: 58) and ‘Cygnus cuculatus’ in Nieremberg (1635: 231)
present throughout the islands that constitute the Kingdom are not valid under the Zoological Code (note that Nieremberg
of Tonga until ~2850 years ago (Steadman 2006; Worthy and 1635 spelt ‘cuculatus’ with one ‘l’, unlike later authors who used
Burley 2020) and was also present on Efate Island, Vanuatu two). The ‘Raphus’ that appears in the 1758 Dutch translation
(Worthy et al. 2015); and (ii) we generally refer to †Pezophaps of Möhring (1752), (Möhring 1758: 44), is also not considered
solitaria as the Solitaire rather than the Rodrigues Solitaire, in valid because that work was suppressed for nomenclatural
order that it is consistent with the use of ‘the Dodo’ for †Raphus purposes by the International Commission on Zoological
cucullatus (i.e. not using Mauritian Dodo). We follow Dubois Nomenclature (the ‘Zoological Commission’ hereafter) (see

(2000: 39) in using the term nomen (plural nomina) for any Opinion 801 of the ICZN 1967).
‘scientific name’ that is formulated in compliance with a nomen- Opinion 241 of the ICZN (1954) stated that the Dutch trans-
clatural code, which, in this case, is the Zoological Code (ICZN lation of Möhring (1752), (= Möhring 1758) was not valid
1999, 2003, 2012, 2016). under the Règles Internationales de la Nomenclature Zoologique
Our open nomenclature and synonymy lists follow the re- (ICZN 1905) and placed it on the Official Index of Rejected
commendations of Richter (1948) (see: Matthews 1973, who Works (note that this opinion was made prior to the publication
outlined them in English, and Becker 2001, who gave a recent of the first edition of the current Zoological Code; ICZN 1961).
overview in German), Sigovini et al. (2016), and Horton et al. A new submission was made to the ‘Zoological Commission’
(2021). Finally, we use the dagger (†) symbol in front of nomen- by The Standing Committee on Ornithological Nomenclature
clatural nomina that denote extinct taxa (except when they ap- (Salomonsen et al. 1964) to have Möhring (1758) suppressed
pear in quotations). for nomenclatural purposes because there was disagreement re-
garding whether Möhring (1758) was a re-edition of Möhring
M ATERIALS AND METHODS (1752) or a new work with its own pagination and additions.
This resulted in Opinion 801 of the ICZN (1967) and the sup-
Abbreviations pression of Möhring (1758).
NHMUK, Natural History Museum, London, UK; NMP,
National Museum, Prague, Czech Republic; OUMNH, Linnaeus (1766), with comments on synonymy and type specimens
Oxford University Museum of Natural History, University of In the 12th edition of Systema Naturæ, Linnaeus (1766: 267)
Oxford, UK; ZMUK, Royal Zoological Museum, Copenhagen, proposed a new genus and species for the Dodo: †Didus ineptus.
Denmark. In his synonymy list for the species, he listed Brisson’s (1760a,
b) genus Raphus, his previous binomial, †Struthio cucullatus
Nomenclatural history of the Dodo Linnaeus, 1758, and works that pre-date 1758 (including
Nominal authority ‘Cygnus cuculatus’ in Nieremberg 1635 and ‘Gallus gallinaceus
In the 10th edition of Systema Naturæ, Linnaeus (1758: peregrinus’ in Clusius 1605). Given the synonymy list in
155) lists the oldest binomial for the Dodo that is available Linnaeus (1766: 267), we consider the following:
under the Zoological Code: †Struthio cucullatus. ‘Raphus’ in
Möhring (1752: 58) and †Struthio cucullatus in Linnaeus (1758: (i) The nomenclatural species †Struthio cucullatus Linnaeus,
155) both have in their synonymy lists ‘Cygnus cuculatus’ 1758 and †Didus ineptus Linnaeus, 1766 are objective
Nieremberg 1635. Note that when we use ‘Raphus’ without synonyms.
italics we are citing a version of the nomen that is not available (ii) The genera †Raphus Brisson, 1760b and †Didus Linnaeus,
under the Zoological Code (i.e. either pre-Linnean, before 1758, 1766 are also objective synonyms. (This opinion matches
or from a text suppressed for nomenclatural purposes by the that of Allen 1910: 329, who considered both genera to
Commission). Therefore, the genus ‘Raphus’ in Möhring (1752) be synonymous ‘with the same type’.)
and the nominal species cucullatus in Linnaeus (1758) are based
on the same indications to earlier publications (see Duncan The synonymy lists of Linnaeus (1758: 155), Brisson (1760b: 15),
1828, de Blainville 1835 and Strickland 1848 for an overview), and Linnaeus (1766: 267) contained three identical bibliographic
and can therefore be treated as synonyms. Brisson (1760b: 14) references: ‘Gallus gallinaceus peregrinus’ in Clusius (1605: 99),
followed Möhring when using the genus †Raphus (in the Latin ‘Cygnus cuculatus’ in Nieremberg (1635: 231), and Dronte in
genitive case: Raphi), as he stated in a footnote: ‘Raphus, nomen Bontius (1658: 70). Although the work by Brisson (1760b) con-
à Moehringio huic generi inditum’ (‘Raphus, a name given to this tained more bibliographic references (including Möhring 1752),
genus by Moehringius’). Note that Brisson (1760a, b) used both the three bibliographic references that appear in the work by
Raphus and Raphi for the generic name. Linnaeus (1758) are all present in the work by Brisson (1760b);
It is clear that neither the genus Raphus nor the specific epi- therefore, we consider †Struthio cucullatus Linnaeus, 1758 and
thet cucullatus originated with Linnaeus or Brisson, although †Raphus raphus Brisson, 1760b to be objective synonyms.
under the Zoological Code both Linnaeus and Brisson are the It is worth noting that the ‘starting date’ of zoological no-
correct nominal authorities. This is attributable to Article 3 of menclature was contested during the 19th century, and that is
6 • Young et al.
why †Didus ineptus was used during the 19th century in pref- that NHMUK PV A 9040 (a composite of multiple individuals)
erence to genus †Raphus and the species †Struthio cucullatus. forms the type specimen does not constitute a valid neotype
‘Strickland’s Code’ (Strickland et al. 1843) proposed the 12th designation under Article 75 of the Zoological Code. As such,
edition of Systema Naturæ (Linnaeus 1766) as the ‘start date’ for the specimen later described by Owen (1872) is not the type
zoological nomenclature, i.e. 1766. This rendered both †Struthio specimen of the Dodo.
cucullatus Linnaeus, 1758 and †Raphus raphus Brisson, 1760a, b Syntypes were listed by Mlíkovský et al. (2011), Mlíkovský
as unavailable nomina under that nomenclatural code. The first (2012), and Parish (2015). Mlíkovský et al. (2011: 140) and
nomenclatural code to use the 10th edition of Systema Naturæ Mlíkovský (2012: 105–106) listed a cranial rostrum (‘upper
(Linnaeus 1758) as its ‘start date’ was that of the American beak’, NMP P6V-004389) as being a syntype of †Struthio
Ornithologists’ Union (American Ornithologists’ Union 1886). cucullatus. Mlíkovský et al. (2011) and Mlíkovský (2012) stated
The starting date of 1758 for zoological nomenclature was also that the bony cranial rostrum is apparently the last known
used by Blanchard (1889; ‘Blanchard’s Code’) and was incorpor- element of a Dodo that was kept by Emperor Rudolf II Habsburg

ated into the Règles Internationales (ICZN 1905). This explains (1552–1612), a live individual that was stuffed, possibly around
why there was a shift to using †Didus cucullatus (Rothschild 1609. Therefore, if NMP P6V-004389 is indeed the last re-
1907), †Raphus cucullatus (Poche 1904, Oudemans 1917), and maining part of Emperor Rudolf II’s Dodo, it is too old to be
synonymizing †Didus with †Raphus (Poche 1904, Allen 1910, one of the specimens forming the basis of Bontius’ (1658) de-
Ridgway 1916) during the early 20th century. scription. It also cannot be from the same individual as the par-
Article 72.5.6 of the Zoological Code states that: ‘In the case tial leg that Clusius (1605) mentioned and described. But could
of a nominal species-group taxon based on an illustration or de- Emperor Rudolf II’s Dodo be one of the unknown individuals
scription, or on a bibliographic reference to an illustration or de- that the Dutch sailors figured and mentioned and which helped
scription, the name-bearing type is the specimen or specimens to form the basis Clusius’ (1605) description? There is no direct
illustrated or described (and not the illustration or description evidence to support that contention, but it is possible.
itself)’. Therefore, the type series of †Struthio cucullatus, †Raphus Parish (2013: 170–171) considered the source of Emperor
raphus, and †Didus ineptus are those that appear in bibliographic Rudolf II’s Dodo most likely to be Hans Schuurman’s fleet that
references: Clusius (1605: 99), Nieremberg (1635: 231), and returned to Texel, The Netherlands in April 1603. There are
Bontius (1658: 70). The specimens illustrated and/or described no records of live Dodos reaching Europe from previous ex-
therein are therefore the type series of the three nomenclatural peditions, and the length of those journeys would suggest that
species and are the basis for the genera †Raphus and †Didus. The Dodos might not have survived (see Parish 2013: 170–171).
location of those specimens is unknown. The source of Clusius’ (1605) information were the journals of
The ‘Gallus gallinaceus peregrinus’ of Clusius (1605: 99–100) Van Neck (his first expedition) and the 1603 expedition that re-
is based on a drawing from a now lost journal that was executed turned under Hans Schuurman (Parish 2013: 108). Therefore:
in 1598 during the voyage of Jacob Cornelis van Neck to the East (i) the provenance of NMP P6V-004389 is unknown, but it
Indies, in addition to observations of live Dodos by the Dutch might be part of Emperor Rudolf II’s Dodo; (ii) Emperor Rudolf
sailors of the van Neck voyage, and Clusius’ own observations II’s Dodo might have been on the expedition that returned to
of a partial leg and foot of a Dodo brought back to Leiden and Europe in April 1603; (iii) Clusius’ (1605) information on
kept at the house of Petrus Pauwius (Hume 2006, Parish 2013). Mauritius was derived, in part, from information on the 1603 ex-
Alas, the whereabouts of the partial leg and foot has been un- pedition; thus (iv) Emperor Rudolf II’s Dodo might have been
known since the 17th century (see Parish 2013: 197). The part of the basis for Clusius’ (1605) description.
‘Cygnus cuculatus’ that appears in Nieremberg (1635: 231) is Unfortunately, Clusius (1605) made no mention of a live
largely copied from Clusius (1605) (see Parish 2013: 109). The Dodo returning to The Netherlands and, as pointed out by
Dronte of Bontius (1658: 70–71) is a description of the Dodo Parish (2013: 170), a live Dodo is not mentioned in Clusius’
based on observations from Dutch sailors of unknown Dodo posthumous Curae posteriores published in 1611. Complicating
specimens (interestingly, Parish 2013: 114 noted that the author matters is that the provenance of NMP P6V-004389 is unclear,
of the Dodo description could in fact be Gulielmus [William] although it appears definitively on the auction inventory of the
Piso). Therefore, the type series of the Dodo is composed of: (i) Prague Castle collection from 3 January 1782 (see Parish 2013:
the lost partial leg and foot mentioned by Clusius (1605: 100); 185). As such, it is impossible to state definitively that NMP
(ii) the unknown specimens that formed the basis of Bontius’ or P6V-004389 is, or is not, a syntype of the Dodo. What we can
Piso’s (1658: 70–71) description; and (iii) the unknown speci- say is that there is no direct evidence to support that conclusion,
mens seen by Dutch sailors on the van Neck voyage that helped and until there is, we cannot consider NMP P6V-004389 to be
to form the basis of Clusius’ (1605: 99–100) illustrations and a syntype. Moreover, there is no direct evidence to link any of
description. the oldest known Dodo specimens with a specific collector or
A subsequent type designation for †Didus ineptus Linnaeus, to determine their precise time of arrival in Europe (such as the
1766 comes from Gray (1870a: 24), who stated: ‘The skeleton ‘Oxford’ or ‘Tradescant’ Dodo; OUMNH ZC-11605; Figs 3, 4).
of the Dodo, forming the type specimen on which Professor Parish (2015: 215–218) also listed four syntypes (based
Owen has written and published an elaborate memoir, has been on communication from Jan Bolding Kristensen), the first of
restored’. Owen’s (1866) discussion of his ‘type concept’ is a which, ZMUK AVES-105484, is a complete pelvis discovered
metaphysical one and not as a taxonomic voucher specimen in the Mare aux Songes marsh. This specimen cannot be a syn-
(i.e. a nomenclatural type). Regardless, Gray (1870a) stating type, because it was not collected until 1865 (Parish 2015: 215).
Figure 3. The Oxford Dodo (†Raphus cucullatus) (OUMNH ZC-11605). A, preserved skin in left lateral view. B, skull in left lateral view. C,
preserved skin in right lateral view. Scale bar: 3 cm.
8 • Young et al.

Figure 4. Drawings of the Oxford Dodo (†Raphus cucullatus) (OUMNH ZC-11605). A, preserved skin in right lateral view. B, skull in left
lateral view. C, preserved skin in right lateral view. D, scleral rings. Artwork by Julian Pender Hume.
Also listed are a skull (ZMUK AVES-105485) and two casts of François Leguat was the commander of a group of French
the ‘Prague beak’ (ZMUK AVES-105487 and ZMUK AVES- Protestant refugees, who, in 1691, were the first humans to
10588). Even if the ‘Prague beak’ is a syntype, its casts would settle on the island of Rodrigues (Strickland 1848: 46). Leguat’s
not ‘inherit’ its type status. The origin of the Copenhagen skull observations are therefore of live birds in their natural habitat
(ZMUK AVES-105485) is unknown. It was rediscovered in 1840 (Leguat 1708; Fig. 5). Unfortunately, the exact individuals he
(see Reinhardt 1842, 1843), and its provenance is disputed (see observed are unknown, and thus, the type specimens of the spe-
Hume 2006: 80–81; Parish 2013: 189–194). However, there is cies are also unknown.
no evidence to suggest that it came to Europe in one of the two
Dutch expeditions that Clusius (1605) used as the basis for his The Nazarene Dodos
description. Therefore, none of the specimens Parish (2015) The Nazarene Dodo, †Didus nazarenus Gmelin, 1789, has a con-
listed as syntypes are, in fact, type specimens. This means that fusing history because the nomen has been applied to individ-
there is not a valid type specimen for the Dodo. uals from both Mauritius and Rodrigues (i.e. to both the Dodo
and the Solitaire). The phrase Nazarene Dodo was first used by
Nomenclatural history of the Solitaire Latham (1785: 4–5) to denote the animals François Cauche
Although the nomenclatural history of the Solitaire (†Pezophaps called ‘Oiseaux de Nazaret’ during his voyage to Mauritius
solitaria) is straightforward, unfortunately, like the Dodo, no (Cauche 1651) (see Parish 2012: 6, for a list of the pre-Latham
type specimen has been designated. The binomial †Didus names used for the Nazarene Dodo). Latham (1785: 1–3) re-
solitarius was established by Gmelin (1789: 728–729) for the ferred to †Didus ineptus as the Hooded Dodo. The primary dif-
Solitaire. The name was not based on physical specimens; inference between the two dodo species was that the Nazarene
stead, it was based on two bibliographic references: Leguat Dodo was said by Cauche to have three pes digits, whereas the
(1708) and Latham (1785). Latham (1785: 3–4) referred to Hooded Dodo had four. Latham (1785) stated that both spe-
the species as the ‘Solitary Dodo’, and his comments were based cies were from Mauritius, whereas the Solitary Dodo (named
solely on the descriptions given by Leguat (1708: 98–104). after Leguat’s 1708 ‘Le Solitaire’) was from Rodrigues. Gmelin
Therefore, as noted by Strickland (1848: 46), who established (1789: 728–729) created the binomials †Didus nazarenus for
the genus †Pezophaps for the species, ‘Leguat’s bird is the type of the Nazarene Dodo and †Didus solitarius for the Solitary Dodo.
the “Didus solitarius” of systematists’. Latham (1790: 662–663) used Gmelin’s binomials for the three
Figure 5. Artistic life reconstruction of the solitaire (†Pezophaps solitaria). This is the only known drawing of a solitaire based on living
specimens in their natural habitat. Artwork from Leguat (1708).
10 • Young et al.
‘species’ of Dodos, but otherwise the nomen †D. nazarenus was However, given that he had demonstrated earlier that the Dodo
largely unused thereafter. and Solitaire were pigeons (Strickland 1848), Strickland (1859:
Strickland (1848: 21–22) considered †D. nazarenus to be a 190) noted that in extant columbids ‘the males and females pre-
‘phantom-species’, based on Cauche’s conjectures and possible sent very nearly the same dimensions’. Therefore, he did not con-
confusion with the Cassowary, such as the presence of three sider the sexual dimorphism hypothesis likely.
toes and the hypothesized lack of a tongue (‘which latter char- This resulted in two distinct ergotaxonomies (= systematiza-
acter was at that time falsely attributed to the Cassowary’). In tions or ‘taxonomic classifications’) for the perceived ‘didine’
Strickland’s (1848: 21) opinion, ‘There can be no doubt that bird species of Rodrigues during the 1850s–1870s. The larger
the bird described by Cauche was the Dodo’. Unfortunately, and smaller species were either referred to as †Didus nazarenus
the nomen was resurrected by Bartlett (1854 [1851]) for a dif- and †D. solitarius, respectively (Bartlett 1854), or as †Pezophaps
ferent animal. (Note that, although written in 1851, the volume solitaria and †P. minor, respectively (Strickland 1859, Owen
in which Bartlett’s work was contained was not published until 1872). Note that Newton (1865) followed Bartlett, but used

1854, see Duncan 1937.) †Pezophaps solitaria instead of †D. solitarius. Unfortunately, this
Prior to the late 19th century, there was little material of the resulted in the nomen †Pezophaps solitaria being used in both
Dodo and the Solitaire in European collections (e.g. see Duncan ergotaxonomies and, confusingly, for both morphs.
1828, de Blainville 1835, Strickland 1844, 1848). As such, after It was not until the Jenner excavations of 1866, during which
the publication of ‘The Dodo and its Kindred’ (Melville 1848, complete skeletons of male and female Solitaires were dis-
Strickland 1848), there was confusion about the presence of two covered, that the confusion ended (Hume et al. 2015), after
distinct morphs from Rodrigues. Based on isolated limb bones, which, only one species of ‘didine’ bird from Rodrigues was rec-
two distinct morphologies of Rodrigues Solitaires were recog- ognized (Newton and Newton 1868, 1869). †Pezophaps solitaria
nized: one large and robust, and another smaller and more gra- was thereafter known to be a sexually dimorphic species, with
cile. Bartlett (1854: 284), however, considered there to be three adult male individuals being noticeably larger and more robust
species from Rodrigues: one ‘which was apparently identical than adult female individuals (Newton and Newton 1869, Owen
with the Dodo (Didus ineptus) of the Mauritius; a second, which 1878, Newton and Clark 1879, Lydekker 1891; see Fig. 2).
was well described under the name of Solitaire; and a third,
which was much larger than either of the above’. For the third, The White Dodo and Réunion Solitaire
much larger species, Bartlett (1854: 284) assigned Gmelin’s The Nazarene Dodos of Mauritius and Rodrigues were not the
nomen †D. nazarenus. The specimens Bartlett’s (1854: 283) re- only species subsequently shown not to exist; there was also
ferral was based on were figured in plate XV of Melville (1848) the White Dodo and Solitaire of Réunion Island. Hume and
and Strickland (1848), and these are, indeed, the large robust Cheke (2004) have already given a detailed overview of how
morph. Bartlett’s (1854: 281) belief that the true Dodo (†Didus an extinct species of quasi-flightless ibis became confused with
ineptus) was present on both Mauritius and Rodrigues comes the Dodo. In short, there was confusion about the identity of a
from his acceptance of Sir Thomas Herbert’s account that the white, large-bodied bird species, thought to be flightless, from
species was on both islands. the island of Bourbon (now called Réunion). Strickland (1848:
Strickland (1859: 191) came to believe that the two morphs 59) noted that ‘Bourbon [Réunion] was formerly inhabited by
represented distinct species. He considered the larger ro- a brevipennate bird called the solitaire, whose white or light
bust morph to be Leguat’s Solitaire (= †Didus solitarius and yellow plumage and woodcock-like beak proves it to have been
†Pezophaps solitaria), and Strickland provisionally established distinct from the dodo of Mauritius and from the so-called soli-
the nomen †Pezophaps minor for the smaller morph. Note taire of Rodrigues’. Hume and Cheke (2004: 59) noted that
that there has been some confusion about exactly when the during the 1850s, the life reconstructions of the ‘Réunion White
nomen †Pezophaps minor was established. The article in which Dodo’ were not actually of a Dodo, but of an animal resembling
Strickland provisionally erected †P. minor was published in an ibis or a stork. Such reconstructions came later, in the 1860s,
August 1859. However, the nomen had been known to mem- once the white bird of Réunion had been linked to the Dodo.
bers of the Zoological Society of London since 1851, because A scientific nomen for this mysterious species was established
it appears in a Society report dating August–December 1851 by de Sélys-Longchamps (1848: 293), who called it †Apterornis
(Mitchell 1852: 19). The gap between Strickland reading to solitarius (alongside two more extinct bird species from the
the Zoological Society and final publication explains why Mascarene Islands: †Apterornis cœrulescens and †Apterornis
†Pezophaps minor appears in synonymy lists from before 1859 bonasia). He placed †Apterornis solitarius within the ‘Famille
(e.g. Bonaparte 1854b in Bonaparte 1850–1865). Dididées’ alongside †Didus ineptus (the Dodo) and †Pezophaps
In the footnote accompanying †Pezophaps minor, Strickland solitaria (the Solitaire). Therefore, the confusion surrounding
discussed Bartlett (1854). Strickland (1859: 191) dismissed the the identity of this species continued into the scientific litera-
possibility of three species having been present on Rodrigues, ture. Unfortunately, the confusion became further ‘tangled’
stating: ‘I have, however, endeavoured to prove that they be- during the 1850s. Schlegel (1854: 244) referred to the species
long to two species only, neither of which can be referred to the as †Didus apterornis, and Bonaparte (1854b in 1850–1865: 2)
Mauritian D. ineptus, nor to the D. nazarenus, which is merely established the genus †Ornithaptera to replace †Apterornis, be-
a synonym of that bird, based on the erroneous description of cause it was already preoccupied. However, Bonaparte (1854b
Cauche’. Strickland (1859: 190) did consider the possibility in 1850–1865: 2) used the binomial †Ornithaptera borbonica.
that the two morphs were the result of sexual dimorphism. In his synonymy list for †Ornithaptera borbonica, Bonaparte
(1854b in 1850–1865: 2) included both †Apterornis solitarius de The Didus-based family group in the 1830s and 1840s
Sélys-Longchamps, 1848 and †Didus apterornis Schlegel, 1854. During the 1830s and 1840s, family-group nomina specifically
During the 19th and 20th centuries, different authors sug- for the Dodo began to appear in systematic ergotaxonomies
gested that the Réunion ‘didine’ was a species of Dodo or (= ‘taxonomic classifications’). These were always based on
Solitaire (see the historical overviews by Mourer-Chauviré et al. Linnaeus’ genus †Didus. The oldest occurrence we can find
1995, 1999, Hume and Cheke 2004). Hachisuka (1937), how- is by Swainson (1835: 239), with the use of †Didiadæ, and
ever, believed that there had been two species of ‘didine’ birds on the same page the Dodo is mentioned explicitly. This ori-
on the island of Réunion, one a ‘dodo’ and another a ‘solitaire’. ginal spelling is incorrect as per the Zoological Code (Article
He applied the previous name to his hypothesized Réunion 32.5.3.1), namely the suffix should be -idae not -iadæ. This
Solitaire (= †Ornithaptera solitaria) and included it in the family was corrected to †Dididæ by Swainson (1836: 286). Although
†Pezophapidae [sic] alongside †Pezophaps solitaria. In his †Didus was not was included explicitly in his †Dididæ, in the
†Raphidae, Hachisuka (1937: 71) placed †Raphus cucullatus and index Swainson (1836: 364) stated that ‘Dodo, affinities of ’ oc-

a new genus and species: †Victoriornis imperialis. †Victoriornis curred on page 286. Therefore, we consider that to be an indi-
imperalis Hachisuka, 1937 was his new nomen for the ‘White cation that the genus †Didus was indeed within †Dididæ, and
Dodo of Réunion’. Hachisuka’s (1937) systematization was therefore it fulfils Article 11.7.1.1 of the Zoological Code.
based not on specimens but on water colour paintings and the The subfamily †Didinæ, placed within Vulturidæ, was used by
accounts of explorers from 17th and 18th centuries. de la Fresnaye (1839: 193–194). The genus †Didus and †Didus
Mourer-Chauviré et al. (1995, 1999) placed †Apterornis ineptus were included explicitly in the subfamily. Lesson (1842:
solitarius in the genus Threskiornis and demonstrated that there column 1036) used a different nomen based on the genus
had never been a ‘Réunion dodo or solitaire’. They instead pos- †Didus: †Didusideae. The genus †Didus, and †Raphus attributed
ited that the bird described by explorers to Réunion had, in fact, to Möhring, were included explicitly, as was the species †Didus
been a species of ibis. ineptus.
The systematizations of the 1830s and early 1840s either
Cyanornis and the Dodo species of Schlegel followed Linnaeus (1758, 1766) in placing the Dodo in the
When de Sélys-Longchamps (1848: 293) established same group as the Ostrich (such as Gray 1840, Lesson 1842,
†Apterornis solitarius, as noted above, he also created two more Bonaparte 1842a, b) or allied the Dodo with vultures (such as
nomina for extinct bird species from the Mascarene Islands: †A. Swainson 1837, de la Fresnaye 1839). Both Gray (1840: 64) and
cœrulescens and †A. bonasia. They were also placed in the ‘Famille Bonaparte (1842a: 14, 89, 1842b: 65) included either †Dididae
Dididées’. Bonaparte (1854b in 1850–1865: 3) established the or †Didinae (family or subfamily rank) in their ergotaxonomies
genus †Cyanornis for these species and continued to place them to contain the Dodo, and placed it within the same order as the
alongside the Dodo and the Solitaire. However, †Apterornis Ostrich. Bonaparte’s earlier systematic frameworks lacked a
coerulescens de Sélys-Longchamps, 1848 is now considered to be family group specifically for the Dodo (e.g. Bonaparte 1841).
an extinct species of swamp hen of the extant genus Porphyrio In their seminal shared volume, Strickland (1848) and Melville
(Olson 1977: 365), whereas †A. bonasia de Sélys-Longchamps, (1848) used Didinæ to contain both the Dodo and the Solitaire,
1848 is an extinct species of rail of the genus †Aphanapteryx and placed it within the pigeon and dove family Columbidæ.
(Olson 1977: 357). The two Dodo species Schlegel (1854) es- This ergotaxonomy was followed by Gray (1848: 482–483, in
tablished, †Didus broeckei and †D. herbertii, are now considered Gray 1844–1849), and subsequently, the inclusion of the Dodo
to be subjective junior synonyms of the Red Rail †Aphanapteryx and the Solitaire alongside pigeons has become the prevailing
bonasia (Olson 1977: 357). systematic opinion.
Nomenclatural history of the Dodo family group The †Raphus-based family group
The first Didus-based family group Unfortunately, the nominal authority of the †Raphus-based
Illiger (1811: 243–244) established Columbidae (as Columbini) family group is confusing. The first to use the †Raphus-based
for the genus Columba, and Inepti on page 245 for the genus family group was Poche (1904: 500). Poche (1904: 500) also
†Didus. Illiger (1811: 245) used Linnaeus’ (1766) genus Didus demonstrated that the oldest available specific epithet for the
for the Dodo, although he did list Brisson’s (1760a, b) genus Dodo is cucullatus Linnaeus, 1758, and that should be used
†Raphus as a synonym. Interestingly, instead of using the generic instead of ineptus Linnaeus, 1766 (reflecting the shift in the
name as the stem for his family group, Illiger (1811) used the ‘starting date’ of zoological nomenclature from 1766 to 1758,
specific epithet of †Didus ineptus to form his family group for the as outlined above). Poche (1904) was therefore the first to rec-
Dodo. Under Article 11.7.1.1 of the Zoological Code, Inepti is ommend the binomial †Raphus cucullatus be used for the Dodo.
invalid, because the stem of the family group was based not on Poche (1904) based his †Raphidae on the 1758 Dutch trans-
a generic name but on a specific epithet. Therefore, the oldest lation of the work by Möhring (1752). At that time, Möhring
known family-group nomen for the Dodo is not valid under the (1758) had not been suppressed for nomenclatural purposes by
Zoological Code (also noted by Mlíkovský, 2000: 79). Use of the Zoological Commission, which did not occur until the pub-
Inepti as a family-group nomen did not seem to become wide- lication of Opinion 801 (see ICZN 1967).
spread in the literature, because we can find few instances of its The validity of †Raphidae Poche, 1904 has been chal-
usage. Notable exceptions include von Eichwald (1831: 257) lenged (Bock 1994), and that challenge has also been disputed
and Kaup (1836: 232–234). (Mlíkovský 2000). Bock (1994: 94–95) argued that Poche’s
12 • Young et al.
†Raphidae was invalid, owing to Poche (1904: 500) basing his Möhring (1758) and the names contained therein were not sup-
family group on †‘Raphus’ Möhring, 1752 not †Raphus Brisson, pressed in 1904. Therefore, an appeal to Article 11.5 does not
1760b. Given that the former is not an available genus (because apply in this situation.
it was established prior to 1758), the associated family group
would also not be valid (under Article 11.7.1.1 of the Zoological Pigeon family-group nomina
Code). However, Poche (1904: 495) discussed the 1758 Dutch The higher-level systematics and nomenclature of pigeons, doves,
translation of the work by Möhring (1752), not the 1752 work. the Dodo, and the Solitaire have long been in flux (see Sibley and
Article 11.7.1.5 of the Zoological Code states that family- Ahlquist 1990: 422–425, for an overview). Here, we will focus
group names when first published must ‘not be based on a on those systematizations that included the Dodo alongside ex-
genus-group name that has been suppressed by the Commission’. tant pigeons (also see Supporting Information, Appendix S1).
Möhring (1758) was suppressed by the Zoological Commission Note that we retain the ligature ‘æ’ from the original publications
(ICZN 1967), thereby making the genus †‘Raphus’ contained rather than correcting them to ‘ae’. This is done solely to preserve

therein unavailable of nomenclatural purposes. However, when original formatting.
Poche (1904) established †Raphidae, the genus †‘Raphus’ As noted above, some of the early ergotaxonomies (= system-
Möhring, 1758 was available. Because Poche (1904) treated the atizations or ‘taxonomic classifications’) united the Dodo with
genus and his family group as being valid, Mlíkovský (2000: 81) the struthionids/struthiones (following Linnaeus). Gray (1840:
contended that Poche’s †Raphidae is, in fact, valid under Article 64) provisionally included the subfamily †Didinæ in the family
11.5.2 of the Zoological Code. We disagree. Struthionidæ (in the order Cursores). Extant pigeons were in the
Mlíkovský’s (2000: 81) interpretation of Article 11.5.2 is that order Gyrantes, in the family Columbidæ (which contained four
because: (i) Poche (1904) used the genus †Raphus for a taxon subfamilies: Treroninæ, Columbinæ, Turturinæ, and Gourinæ)
he considered to be valid, and (ii) he attributed it to an earlier (Gray 1840: 57–59). Bonaparte (1842a: 14, 1842b: 65) had a
work with a clear bibliographic reference (Möhring 1758: 44), very similar ergotaxonomy to Gray (1840), where the family
Poche (1904) made the genus †Raphus available (with Poche †Dididae (with the subfamily †Didinae) was within the order
being the nominal authority). However, the genus †‘Raphus’ Struthiones. Extant pigeons were in the order Columbae, within
Möhring, 1758 was valid in 1904. It did not become invalid until the family Columbidae (which contained three subfamilies:
1967. Article 39 of the Zoological Code is clear that: ‘The name Columbinae, Treroninae, and Gourinae) (Bonaparte 1842a: 13,
of a family-group taxon is invalid if the name of its type genus 1842b: 64).
is a junior homonym or has been totally or partially suppressed Bonaparte’s (1854a: 139–140) later ergotaxonomies still had
by the Commission’. This means that when †‘Raphus’ Möhring, the Dodo and the Solitaire in a separate order from pigeons.
1758 was rendered invalid by Opinion 801 (ICZN 1967), The Dodo and the Solitaire were in the order †Inepti, with a
†Raphidae Poche, 1904 also became invalid. single family, †Dididæ. Within †Dididæ he had two subfamilies:
The oldest valid †Raphus-based family group is often quoted †Epyornithinæ (to contain the Réunion ‘dodo and solitaire’:
to be authored by Wetmore (1930: 5). However, Wetmore †Ornithaptera and †Cyanornis, alongside †Æpyornis Geoffroy
(1930) simply stated that †Raphidae included ‘Dodos, Saint-Hilaire, 1851a, b; note that the original paper was published
Solitaires’ and did not state whether the genus †Raphus Brisson, in two different journals, with one accompanied by an additional
1760b or †‘Raphus’ Möhring, (1752 or 1758) was stem for his note, Geoffroy Saint-Hilaire 1851c) and †Didinæ (containing
family group. Although, since Peters (1937), the genus †Raphus †Didus and †Pezophaps). Extant pigeons and doves were within
Brisson, 1760b has been considered the basis for Dodo nomen- the order Columbæ, within which there were five families. The
clature, Oudemans (1917) also used the binomen †Raphus first family was Didunculidæ, which had the sole subfamily
cucullatus and the family-group nomen †Raphidae. Oudemans Didunculinæ. The second family was Treronidæ, which had three
(1917: 6) stated: ‘Brisson 1760 vindt, dat de Dodo in een apart subfamilies: Treroninæ, Ptilopodinæ, and Alectraenadinæ. The
genus moet geplaast worden: Raphus2) (de oudste genusnaam!)’ third family, Columbidæ, had six subfamilies: Lopholaeminæ
[‘Brisson 1760 thinks, that the Dodo should be placed in a sep- [sic], Carpophaginæ, Columbinæ (with two tribes, Columbeæ
arate genus: Raphus2) (the oldest genus name!)’]. In footnote and Macropygieæ), Turturinæ, Zenaidinæ, and Phapinæ. The
2, which accompanies †Raphus, Oudemans (1917: 6) stated: fourth family, Calliænadidæ [sic], contained the sole subfamily
‘Ontleend aan Moehring 1752; dit werk heb ik niet kunnen Calliænadinae [sic]. The final family, Gouridæ, included the sub-
raadplegen’ [‘Taken from Moehring 1752; I have not been able family Gourinæ.
to consult this work’]. On page 101, Oudemans (1917) explicitly Bonaparte (1854b in Bonaparte 1850–1865) had a similar
used ‘Briss’. after †Raphus. As such, the †Raphus cucullatus and ergotaxonomy to his earlier publication. Again, the Dodo and
†Raphidae of Oudemans (1917) was based on Brisson (1760b), the Solitaire were united in the order †Inepti, with a single
and therefore he should be considered the nominal authority of family, †Dididæ. Within †Dididae he had two subfamilies:
the †Raphus-based †Raphidae, not Wetmore (1930). Epyornithinae (now containing only †Æpyornis) and †Didinae
Note that Mlíkovský (2000: 81) argued that, under Article (containing †Didus and †Pezophaps, in addition to the Réunion
39, precedence should be given to †Raphidae Wetmore, 1930 ra- ‘dodo and solitaire’ †Ornithaptera and †Cyanornis). Extant pi-
ther than †Raphidae Poche, 1904. This is owing to Mlíkovský’s geons and doves were again within the order Columbæ, with
opinion that †‘Raphus’ Poche, 1904 is a junior homonym of five families. The first family was Didunculidae, including the
†Raphus Brisson, 1760. However, as we outline above, Poche sole subfamily Didunculinae. The second family was Treronidae,
(1904) did not establish an available genus †Raphus, because which had four subfamilies: Treroninae, Ptilopodinae (with two
tribes: Ptilopodeae and Chrysaeneae), Alectraenadinae, and suffices. In his 1862 text, Reichenbach united dodos and pigeons
Carpophaginae. The third family, Columbidae, had five subfam- in Columbariæ, alongside taxa now included in Cracidae and
ilies: Lopholaeminae, Columbinae (with three tribes: Palumbeae, Megapodidae. The Dodo and Solitaire were united with modern
Columbeae, and Macropygieae), Turturinae, Zenaidinae, and cracids (in the family Alectorinae), while extant columbids were
Phapinae (with three tribes: Phapeae, Chalcophapeae, and placed in the families Columbinae and Peristerinae. The internal
Geopelieae). The fourth family, Caloenadidae, contained the ergotaxonomy of these families is complicated by Reichenbach’s
sole subfamily Calaenadinae [sic]. The final family, Gouridae, in- use of the -inae suffix at all ranks, thus, we summarize them in
cluded the subfamily Gourinae. Supporting Information, Appendix S1.
Gray’s latter ergotaxonomy (Gray, 1848 in Gray 1844–1849, Sundevall (1873 in Sundevall 1872–1873: 97–101) united
1870b) united dodos and pigeons in the order Columbæ. Within the Dodo and pigeons in the cohors Peristeroïdeæ. He had three
Columbidæ, he had five subfamilies: Columbinæ, Treroninæ, families: the first was Didinæ, which included the Dodo (†Didus)
Gourinæ, Didunculidæ, and †Didinæ. By the 1870s, Gray’s and the extant genus Didunculus Jardine, 1845 (for a discussion

ergotaxonomy had changed slightly (Gray 1870b), whereby he on the correct nominal authority of this genus, see Bruce and
raised Didunculidæ and †Didinæ to the family rank. This re- Bahr 2020). All other extant pigeons were placed in the families
sulted in three families: †Dididæ (which had a sole subfamily, Columbinæ and Megapeliinæ. Salvadori (1893) united dodos
†Didinæ, and a sole species, the Dodo), Columbidæ (which had and pigeons in the order Columbæ, within which Salvadori
three subfamilies: Columbinæ, Treroninæ, and Gourinæ), and had two suborders. The first, called †Didi, contained the family
Didunculidæ (with the subfamily Didunculinæ). The system- †Dididæ (for the Dodo and the Solitaire). The second suborder,
atization of Elliot (1885) was similar, with the Dodo, Solitaire, also called Columbæ, contained five families for extant pigeons
and extant pigeons united in the order Columbæ. Elliot had five and doves: Treronidæ (containing the subfamilies Treroninæ,
families: †Didiidæ (for the Dodo and the Solitaire), Columbidæ, Ptilopodinæ, and Carpophaginæ), Columbidæ (containing the
Carpophagidæ, Gouridæ, and Didunculidæ. subfamilies Columbinæ, Macropygiinæ, and †Ectopistinæ),
An important individual in columbid nomenclature and Peristeridæ (containing the subfamilies Zenaidinæ, Turturinæ,
systematization is Reichenbach. The publication dates for Geopeliinæ, Peristerinæ, Phabinæ, Geotrygoninæ, and
Reichenbach’s work are, in the words of Bock (1994: 21): ‘a Calœnadinæ), Gouridæ, and Didunculidæ. Sharpe’s (1899) sys-
major nomenclatural headache’. There were numerous, un- tematization was identical to that of Salvadori (1893), differing
dated short contributions, in addition to larger works that had only in Sharpe’s use of Columbiformes rather than Columbæ for
sections written at different times (even different decades, see the name of the order.
Meyer 1879, Bock 1994). The situation is complicated further Martin (1904) also united the Dodo, the Solitaire, and extant
by online versions of Reichenbach’s work often having the wrong pigeons in the order Columbiformes, with the same two sub-
publication date(s). It is also worth noting that: (i) Reichenbach orders: †Didi and Columbae. Interestingly, he had two families in
amended the spelling of several pigeon generic names (such as his suborder †Didi, †Dididae and †Pezophabidae, for the Dodo
Geopelia to Geopeleia), and (ii) he used the -inae suffix for nomina and the Solitaire, respectively. Extant pigeons were all placed in
at the family, subfamily, and below subfamily ranks (at what we the suborder Columbae, with five families within: Columbidae
today would call the tribe and subtribe ranks). The text and (with the subfamilies Columbinae and Caloenadinae),
plates within the work by Reichenbach (1849–1853) were com- Peristeridae (with the subfamilies Peristerinae, Phabinae, and
pleted at different times. According to Meyer (1879), the plates Ptilopodinae), Treronidae (with the subfamily Treroninae),
with columbid nomina were published in June 1850, whereas Carpophagidae (with the subfamilies Carpophaginae, Gourinae,
the text was published in October 1852. At the top of plate 22 and Otidiphabinae), and Didunculidae (with the subfamily
(XXII) there is Palumbinae and Treroninae, and at the top of Didunculinae). Peters (1937) simplified the systematics and no-
plate 23 (XXIII) is Peristerinae. It is not possible to determine menclature of pigeons, uniting dodos and pigeons in the order
what nomenclatural rank they were established at, but we can as- Columbiformes, within which he had two families, †Raphidae
sume that they were not meant to be at the family level, because (for the Dodo and Solitaire) and Columbidae (for all other pi-
none of them appears in the 1852 text. On plate 22, Palumbus geons and doves). Peters’ Columbidae had four subfamilies:
Kaup, 1829 does not appear, but Treron Vieillot, 1816 does, Treroninae, Columbinae, Goürinae [sic], and Didunculinae.
while Peristera Swainson, 1827 appears on plate 23. However, we Verheyen (1957), however, proposed a very different sys-
can assume Palumbinae was established for Palumbus, because tematization of pigeons. Within the order Columbiformes, he
the genus appears in the 1852 text. We can therefore consider had three families. The first was Caloenididae, within which
Reichenbach 1851 in Reichenbach (1849–1853) as the nom- there was three subfamilies: Caloenidinae, Goürinae [sic], and
inal authority of the Palumbus-based family group. Interestingly, †Raphinae (which had two tribes, †Raphini and †Pezophabini,
in his 1852 ergotaxonomy, Reichenbach united the Dodo with for the Dodo and Solitaire, respectively). The second pigeon
Goura Stephens, 1819, in the family Gourinae [sic]. Note that family was Duculidae, containing five subfamilies: Ptilinopinae
although both Temminck (1813: 377) and Vieillot (1816: 49) (containing the tribes Ptilinopini and Alectroenini),
used Goura prior to Stephens (1819), they did not use it as a Megaloprepiinae, Tréroninae [sic], Didunculinae, and
generic name. Duculinae (containing the tribes Duculini and Hemiphagini).
Reichenbach (1862: 13–160) later created an expansive The final pigeon family was Columbidae, which had eight
ergotaxonomy for columbids. Reichenbach (1862) also used, subfamilies: Columbinae (containing the tribes Columbini,
for what we today would call the subtribe rank, the -eae and -rae Macropygiini, Chalcophabini, and Oenini), Claravisinae [sic],
14 • Young et al.
Zenaidinae, Geotrygoninae, Starnoeninae, Turturinae (con- (e.g. Wetmore 1960, Sibley and Ahlquist 1990, Janoo 1996).
taining the tribes Turturini, Phabini, Geopeliini, Ocyphabini, Others, such as Cracraft (1981: 699), placed both species within
and Cosmopeliini), Gallicolumbinae (containing the tribes Columbidae, stating that ‘the systematic problem here is to iden-
Gallicolumbini, Geophabini, Leucosarciini, and Trugonini), tify their close relatives within the columbids’. However, the re-
and Otidiphabinae. In comparison to other pigeon and dove sults of modern phylogenetic analyses were not consistent with
ergotaxonomies, Verheyen’s (1957) was far more extensive. any previous ergotaxonomy (e.g. see Janoo 2000, Johnson and
It also placed the Dodo and the Solitaire directly within the Clayton 2000, Shapiro et al. 2002, Pereira et al. 2007, Fulton et al.
flighted columbiform radiation, uniting them with the Nicobar 2012, Heupink et al. 2014, Besnard et al. 2016, Nowak et al. 2019,
Pigeon Caloenas nicobarica (Linnaeus, 1758) and the crowned Chen et al. 2022, Boyd et al. 2022, Oliver et al. 2023). The ‘trad-
pigeons (the genus Goura). itional doves’ were never recovered as monophyletic; whereas
Although based on comparative osteology, Verheyen’s the Dodo, Solitaire, the crowned pigeons, Samoan Tooth-billed
ergotaxonomy was not broadly accepted. For Goodwin (1959: Pigeon, and Pheasant Pigeon were all recovered as members of

3), who prefaced his comments that he did not feel compe- the ‘fruit dove’ group (Treroninae sensu Baptista et al. 1997).
tent to criticize Verheyen’s work because it was based on oste- Since the work of Pereira et al. (2007), a three-clade subdivision
ology, some of Verheyen’s decisions were ‘so contrary to the of Columbidae has been recognized, with those clades referred
evidence from external taxonomic characters and behaviour- to as Columbinae, Peristerinae, and Raphinae by Dickinson and
patterns that I hesitate to follow his conclusions without Remsen Jr (2013: 25, 36). Dickinson and Raty (2015), however,
further evidence’. Johnston (1962: 69) was also hesitant to noted that Claravinae should be used instead of Peristerinae,
use Verheyen’s ergotaxonomy, noting that it was based on a which has been followed since (e.g. Boyd et al. 2022). As such,
sample size of only 20 species. Johnston (1962: 69) agreed since 2013, there has been an acceptance of using a Dodo-based
with Goodwin (1959) and concluded that, ‘I cannot accept family-group nomen for the clade that unites species that trad-
Verheyen’s taxonomy without a great deal of additional evi- itionally were placed in Gourinae, Otidiphabinae, Didunclinae,
dence’. Unfortunately, this meant that the hypothesis uniting and Treroninae (sensu Baptista et al. 1997). Moreover, the tribe
the Dodo and the Solitaire within the flighted columbiforms, Raphini has been used for the clade composed of the Dodo
and alongside the genera Caloenas Gray, 1840 and Goura spe- and closely related extant genera: Caloenas, Goura, Didunculus,
cifically, did not gain traction. This has proved to be unfortu- Otidiphaps Gould, 1870, and Trugon Gray, 1849 (see Chen et al.
nate, because that grouping has been supported consistently 2022, Oliver et al. 2023).
by molecular-based phylogenetic analyses (see Close relatives
of the Dodo and the Solitaire). Close relatives of the Dodo and the Solitaire
The ergotaxonomy of Brodkorb (1971) had two families within A close relationship between the Dodo and the Solitaire has
the suborder Columbae (within the order Columbiformes). long been suspected (e.g. Latham 1785, 1790, Gmelin 1789,
†Raphidae united the Dodo and the Solitaire, while all extant Melville 1848, Strickland 1848) and has been supported by
volant pigeons and doves were within Columbidae (which had morphology-based ( Janoo 2000, Livezey and Zusi 2006, 2007)
three subfamilies: Columbinae, Gourinae, and Ptilinopinae). In and molecular-based (Shapiro et al. 2002, Pereira et al. 2007,
contrast, the ergotaxonomy of Wolter (1975) had eight families Heupink et al. 2014, Besnard et al. 2016, Soares et al. 2016,
within the order Columbiformes. The first family was †Raphidae, Bruxaux et al. 2018, Oliver et al. 2023) phylogenetic analyses.
containing the subfamilies †Pezophabinae (for †Pezophaps See Livezey (1993: 279–281) for an overview of alternative hy-
solitaria) and †Raphinae (for both †Raphus cucullatus and the potheses, where the Dodo and the Solitaire are not closely re-
‘White Dodo’ †Raphus solitarius). The other seven families con- lated. However, their relationships with other pigeons have been
tained extant species: Caloenadidae, Gouridae, Otidiphabidae, more contentious, ever since Strickland (1848: 40–41) first hy-
Columbidae, Duculidae, Treronidae, and Didunculidae. pothesized that the Dodo was not simply ‘intermediate’ between
Janoo (1996: fig. 7) created a hypothetical branching diagram pigeons and ‘gallinaceous’ birds (sensu Reinhardt), but a ‘col-
of columbiform relationships. Columbiformes was composed ossal, brevipennate, frugivorous Pigeon’. Note that we have not
of a polytomy of Pteroclidae (the sandgrouses), †Raphidae been able to find any publication where Reinhardt discusses his
(containing the subfamilies †Raphinae and †Pezophabinae, hypotheses regarding the ‘position’ of the Dodo amongst birds
for the Dodo and the Solitaire, respectively), and Columbidae. (e.g. Reinhardt 1842, 1843); our only conduit to his thoughts
Columbidae was composed of a polytomy of five subgroups: come via Strickland (1848) and Owen (1866: 18).
Didunculidae, Treroninae, (Columbinae + Caloenadinae), (Pha Molecular-based phylogenetic analyses have consist-
binae + Peristerinae + Ptilopodinae), and (Otidiphabinae + Ca ently recovered the genus Caloenas as the sister taxon to the
rpophaginae + Gourinae). Dodo + Solitaire clade (Shapiro et al. 2002, Pereira et al. 2007,
Before the molecular phylogenetic revolution, the broadly ac- Heupink et al. 2014, Besnard et al. 2016, Soares et al. 2016,
cepted systematization of Columbidae (e.g. Baptista et al. 1997) Bruxaux et al. 2018, Oliver et al. 2023). This is true regard-
had five extant subfamilies: Columbinae (‘typical’ pigeons, less of whether only the Nicobar Pigeon (Caloenas nicobarica)
181 species), Otidiphabinae (containing only the Pheasant or the Nicobar Pigeon and the extinct Spotted Green Pigeon
Pigeon), Gourinae (crowned pigeons, composed of only the †Caloenas maculata (Gmelin, 1789) are included in said ana-
genus Goura), Didunculinae (Samoan Tooth-billed Pigeon, lyses. An early molecular-based phylogenetic analysis recovered
one species), and Treroninae (fruit doves, 123 species). The ex- only a Dodo + Solitaire + Caloenas subclade (although Goura
tinct Dodo and Solitaire were often placed in a separate family spp. and Didunculus strigirostris were found to be relatively close
(†Raphidae) alongside Columbidae within Columbiformes to this subclade in their topology) (Heupink et al. 2014). Six

Figure 6. Skulls, in right lateral view, of extant raphine columbids closely related to the Dodo and the Solitaire. A, Didunculus strigirostris
(NHMUK 1875.10.16.6). B, Caloenas nicobarica (NHMUK 1890.11.3.11). C, Goura cristata (NHMUK 1891.7.20.40). Scale bar: 3 cm.
studies found the Samoan Tooth-billed Pigeon (Didunculus island endemic pigeons (although its composition can vary, the
strigirostris), the crowned pigeons (Goura spp.), the Nicobar genera Caloenas, Goura, and Didunculus are the most consistent
Pigeon (Caloenas nicobarica), the Dodo and the Solitaire to close relatives; see Fig. 6).
form a clade of ground-dwelling island endemics (Shapiro et al.
Columbid family-group nomina
2002, Pereira et al. 2007, Besnard et al. 2016, Soares et al. 2016,
Bruxaux et al. 2018, Oliver et al. 2023). Chen et al. (2022) did Validity of columbid family-group nomina
not include the Solitaire (†Pezophaps solitaria) in their analysis, Given that Dodo-based family-group nomina are now accepted
but they still recovered a Dodo + Caloenas clade, with Goura and by columbid workers (see above), we need to ensure that the
Didunculus as successive sister taxa. †Raphus-based family group is indeed the oldest available
Interestingly, the only known morphology-based pigeon name for the subclades referred to as Raphinae and Raphini
phylogeny also recovered the Dodo + Solitaire subclade as (sensu Dickinson and Remsen Jr 2013, Chen et al. 2022, Oliver
being united with the genera Caloenas and Goura ( Janoo 2000). et al. 2023). To do so, we must consult Article 11.7.1.1 of the
However, that analysis found the crowned pigeons (Goura Zoological Code (see Dubois and Ohler 2015). Article 11.7.1.1
spp.) to be the sister taxon to the Dodo and Solitaire subclade, states that, to be available, a new zoological family group must:
with the Nicobar Pigeon (Caloenas nicobarica) being the basal- be a noun in the nominative plural formed from the stem of an
most member of the clade. The Samoan Tooth-billed Pigeon available generic name [Article 29] (indicated either by express
(Didunculus strigirostris) was not found to be closely related reference to the generic name or by inference from its stem, but
to the Goura, Caloenas, †Raphus, and †Pezophaps clade. The for family-group names proposed after 1999, see Article 16.2);
second morphology-based phylogeny that included a wide sam- and the generic name must be a name then used as valid in the
pling of pigeons is that of Livezey and Zusi (2006, 2007). They new family-group taxon [Articles 63 and 64] (use of the stem
found Didunculus and Goura to be successive sister taxa to the alone in forming the name is accepted as evidence that the au-
†Raphus + †Pezophaps subclade. The Nicobar Pigeon (Caloenas thor used the generic name as valid in the new family-group
nicobarica) was not included in their analysis. taxon unless there is evidence to the contrary). Therefore, to
Regardless of whether a morphology-based or molecular- be available, a new family-group nomen must be: (i) based on
based approach is taken, every phylogenetic analysis undertaken an available genus, and (ii) said genus must be within the new
has found †Raphus and †Pezophaps to be sister taxa. Moreover, family group. However, new family-group nomina established
all analyses agree that they form a subclade deep within after 1930 must also satisfy Article 13.1, being accompanied
Columbidae and are not outside of the columbid radiation. And with a description that includes characters that purport to dif-
finally, all analyses agree that the †Raphus + †Pezophaps subclade ferentiate the taxon, or a bibliographic refence to such a pub-
is a member of a wider clade composed of ground-dwelling lished statement.
16 • Young et al.
Most of the columbid family-group nomina were established This means that by the end of the 1830s there were six valid
before 1930, particularly between 1811 and 1862. Illiger (1811: family-group nomina: based on Columba (Illiger 1811), †Didus
243–244) established the first pigeon family-group nomen for (Swainson 1835), †Ectopistes and Ptilinopus (Selby 1835), and
the genus Columba Linnaeus, 1758, which he referred to as Goura and Vinago (Hodgson 1836). Gray (1840: 57–58) cre-
Columbini. Note that this was before Kirby (1815) introduced ated two new family-group nomina, all as subfamilies within
the -idae (-idæ) suffix into zoological nomenclature. Leach Columbidæ: Treroninæ (Gray 1840: 57), which explicitly
(1819: 66) is the oldest reference to Columbidæ we can find. contained the genus Treron, and Turturinæ (Gray 1840: 58),
Therefore, the Columba-based family-group nomen is available. which explicitly included the genus Turtur Boddaert, 1783.
As noted above, Illiger (1811: 245) created Inepti for †Didus Bonaparte (1840: 26) included the subfamily Ptilophyrinae
ineptus, but Inepti is not available because it was based on the within Columbidae; although Bonaparte did not list the con-
specific epithet, not the stem of an available generic name (i.e. tents of his columbid subfamilies, we can use the stem of the
Didus). The valid establishment of the †Didus-based family family group to consider it to be based on the genus Ptilophyrus

group comes from †Didiadæ of Swainson (1835: 239), who later Swainson, 1837. Gray, 1848 in Gray (1844–1849: 480–481) es-
emended it to †Dididæ (Swainson 1836: 286). tablished the subfamily Didunculinæ for the genus Didunculus.
Also established in 1835 were †Ectopistinæ, Peristerinæ, Gnathodon, the objective senior synonym of Didunculus (see
and Ptilinopinæ. They were created by Selby (1835) as sub- Bruce and Bahr 2020), was also the basis of a family-group
families within Columbidæ, alongside his use of the subfamily nomen: Gnathodontinæ, by Melville (1848: 119). The pigeon
Columbinæ. Note that under Article 36 of the Zoological Code genus Gnathodon Jardine, 1845 is a junior homonym of the bi-
(Principal of Coordination applied to family-group names), valve genus Gnathodon Sowerby, 1832 (and possibly other
when a valid family-group nomen is established, it is con- earlier genera, but we have been unable to confirm them our-
sidered to be established simultaneously at all ranks within the selves), which renders the pigeon Gnathodon-based family group
family group (therefore, the nominal authority of the subfamily invalid under Article 39 of the Zoological Code. Interestingly,
Columbinae is not Selby, 1835, but Illiger, 1811). Selby (1835) however, Melville (1848: 119) established his Gnathodontinæ
is the oldest instance of a subfamily rank within Columbidae we specifically for Didunculus and not Gnathodon. Therefore, it is de-
can find, because although Bonaparte (1831) is often quoted as batable whether Melville’s Gnathodontinæ is valid even under
using Columbinæ (i.e. Brodkorb 1971), Bonaparte (1831: 53) Article 11.7.1.1.
did not use any ranks below the rank of family for columbids. During the 1850s and 1860s, more than a dozen new
†Ectopistinæ was created explicitly for †Ectopistes Swainson, family-group nomina for extant columbids were established.
1827 (Selby 1835: 166–168), Peristerinæ contained the genus Reichenbach (1851 in Reichenbach 1849–1853) established
Peristera (Selby 1835: 191, 193, 213), and Ptilinopinæ con- the Palumbus-based family group (as we discuss above). Le
tained the genus Ptilinopus Swainson, 1825 (Selby 1835: Maout (1853) used the term Lophyriens as one of two ‘tribus’
119–120, 124–125, 128, 192). As such, the †Ectopistes-based within Colombiens (= Columbidae). Lophyriens included only
and Ptilinopus-based family groups are available; note that the genus Lophyrus Vieillot, 1816 (Le Maout 1853: 306). Under
Selby (1835) used the spelling Ptilinopinæ, not Ptilonopinæ Article 11.7.2 of the Zoological Code, Lophyriens can be con-
or Ptilinopodinæ (contra Brodkorb 1971, Bock 1994). Contra sidered a valid family group because Bock (1994) latinized it to
Bock (1994), Peristerinæ was used by Selby (1835: 191–193, Lophyrinae.
213); however, as noted by Oberholser (1899: 203), the pi- Bonaparte (1853) established family-group nomina explicitly
geon genus Peristera Swainson, 1827 is a junior homonym of for the genera Alectroenas Gray, 1840 (Alectrænadinæ [sic]),
the mollusc genus Peristera Rafinesque, 1815. Under Article 39 Caloenas (Calænadidæ and Calænadinæ [sic]), Carpophaga
of the Zoological Code, homonymy of the type genus renders Selby, 1835 (Carpophaginæ), Lopholaimus Gould, 1841
any family-group nomina based upon it invalid. Also, there does (Lopholaiminæ), Phaps Selby, 1835 (Phapinæ [sic]), Ptilopus
not appear to be a Carpophaginae in Selby (1835), contra Bock Strickland, 1841 (Ptilopodinæ), and Zenaida Bonaparte,
(1994). Selby (1835: 87) used the plural noun Carpophagæ 1838 (Zenaidinæ). The next year, Bonaparte (1854a) es-
(alongside the plural noun Ptilinopi for Ptilinopus), but we tablished Macropygieæ for the genus Macropygia Swainson,
cannot find any family-group nomen for the genus within the 1837. Finally, Bonaparte (1854b in Bonaparte 1850–1865)
text. Finally, as noted by Dickinson and Raty (2015: 188), there established family-group nomina for the genera Chalcophaps
have been several reprints of Selby (1835). The most commonly Gould, 1848 (Chalcophapeae), Chamaepelia Swainson, 1827
cited reprint is Selby (undated), an edition perhaps from the (Chamaepelieae), Chrysoena Bonaparte, 1854c (Chrysaeneae),
early 1850s, that is frequently available online. We believe we Geopelia Swainson, 1837 (Geopelieae), and Starnoenas
found one of the 1835 editions, but given how frequently Selby Bonaparte, 1838 (Starnoenadeae).
(1835) was reprinted we cannot be entirely sure. Reichenbach (1862) established seven new valid
Often, the family group based on the genus Goura is con- nomina: Ducula Hodgson, 1836 (Duculinae), Geotrygon
sidered to have been established by Gray (1840: 58). However, Gosse, 1847 (Geotrygoninae), Myristicivora Reichenbach,
Hodgson (1836: 160) had already used Gourinæ, alongside 1853 in Reichenbach (1851–1854) (Myristicivorae, not
Columbinæ and Vinaginæ, as subfamilies within Columbidæ. It Mystricivorinae as listed by Bock 1994: 139), Phapitreron
is clear from the text that he considered the genera Goura and Bonaparte, 1854c (Phapitreroninae), Leucotreron Bonaparte,
Vinago Cuvier, 1817 to be valid (Hodgson 1836: 159). As such, 1854c (Leucotreroneae), Osmotreron Bonaparte, 1854c
the nominal authority for both the Goura-based and Vinago- (Osmotreroneae), and Zonoenas Reichenbach, 1853 in
based family groups should instead be Hodgson, 1836. Reichenbach (1851–1854) (Zonoeninae). Reichenbach (1862)
also created a subfamily-ranked Chlorotreroneae; however, 1909), †Microgoura (Richmond, 1909), Claravis (Todd, 1913),
there was no generic name included within called ‘Chlorotreron’, and †Raphus (Oudemans, 1917).
hence the family group is not valid. The earliest usage of the After 1930, Article 13.2 of the Zoological Code adds the re-
genus Chlorotreron we can find is by Salvadori (1882). Note, quirement that all available family-group nomina be accom-
contra Bock (1994: 139), Reichenbach (1862: 13–160) did panied by a description or bibliographic reference to such a
not create family-group nomina for the genera Geophaps Gray, published statement. However, Article 13.2.1 adds a complica-
1842, Ocyphaps Gray, 1842, Phalacrotreron Bonaparte, 1854c, tion: ‘A family-group name first published after 1930 and before
or Sphenocercus Gray, 1840. Reichenbach (1862) listed both 1961 which does not satisfy the provisions of Article 13.1 is
Geophaps and Ocyphaps within Phapinae [sic], Phalacrotreron available from its original publication only if it was used as valid
was within an unnamed subgroup of Vinagineae called before 2000, and also was not rejected by an author who, after
‘Der Schnabel niedergedrückt’ (‘The beak depressed’), and 1960 and before 2000, expressly applied Article 13 of the then
Sphenocercus was within an unnamed subgroup of Vinagineae current editions of the Code’. Although Article 13.2.1 requires

called ‘Schwanz spitzfederig keilförmig’ (‘tail pointed-feathered, that all family-group nomina published between 1930 and 1961
wedge-shaped’). Sundevall (1873 in Sundevall 1872–1873: must be checked with the wider literature to determine whether
101) established the family Megapeliinae, in which he included they is valid or not, all family-group nomina established post-
the genus Megapelia Kaup, 1836. Note that Sundevall (1873 in 1961 must be accompanied by a description (including those
Sundevall 1872–1873) used the -inae suffix for his families. established herein).
In the early 20th century, more columbid family-group nomina Luckily, the only publication we can find that established new
were established. Martin (1904: 335, 336, 348) established family-group nomina post-1930 is by Verheyen (1957). As noted
family-group nomina for the genera Otidiphaps and †Pezophaps, above, Verheyen’s (1957) ergotaxonomy was extensive, including
Otidiphabinae and †Pezophabidae, respectively. McGregor numerous family-group nomina, of which he established nine
(1909: 42) established the subfamily Muscadivorinæ to include new nomina explicitly for the genera Cosmopelia Sundevall,
the genera Muscadivores Lesson, 1831 and Ptilocolpa Bonaparte, 1873 in Sundevall (1872–1873) (Cosmopeliini), Hemiphaga
1854c. Richmond (1909: 621) used ‘Microgouridæ?’ as the Bonaparte, 1854c (Hemiphagini), Gallicolumba Heck, 1849
family that contained the extinct genus †Microgoura Rothschild, (Gallicolumbinae), Geophaps (Geophabini), Leucosarcia Gould,
1904. Although Richmond was expressing uncertainty, it was 1848 (Leucosarciinae), Megaloprepia Reichenbach, 1853 in
used explicitly to contain the genus †Microgoura. We provision- Reichenbach (1851–1854) (Megaloprepiinae), Ocyphaps
ally list Richmond (1909) as the nominal authority, as we have (Ocyphabini), Oena Swainson, 1837 (Oenini), and Trugon
not been able to find an earlier instance of Microgouridae. We Gray, 1849 (Trugonini). Verheyen (1957: 30–36) provided
raise this issue because Richmond (1917) had been considered a list of characters that defined most of the nomina he used in
the nominal authority of Claraviinae, but as shown by Dickinson his ergotaxonomy, thus those nomina fulfil Article 13.2.1 of the
and Raty (2015), it was in fact Todd (1913). Todd (1913: 512) Zoological Code. The family-group nomina based on Cosmopelia,
established the family-group Claraviinae for the genus Claravis Gallicolumba, Geophaps, Leucosarcia, Ocyphaps, and Oena expli-
Oberholser, 1899, but Dickinson and Raty (2015) noted that citly had diagnostic characters listed, and characters that dis-
the spelling Claravinae should instead be used. tinguished Megaloprepiinae from Ptilinopinae were given by
This means that before 1930 the following columbid Verheyen (1957: 32), thus we consider the Megaloprepia-based
genera were the basis for valid family-group nomina: Columba family group available as well. However, no diagnostic characters
(Illiger, 1811), †Ectopistes (Selby, 1835), Ptilinopus (Selby, were provided for the Trugon-based family group (Trugonini),
1835), †Didus (Swainson, 1835), Goura (Hodgson, 1836), with Verheyen (1957: 35) stating: ‘anatomie inconnue’
Vinago (Hodgson, 1836), Treron (Gray, 1840), Turtur (‘anatomy unknown’). No diagnostic characters were listed for
(Gray, 1840), Didunculus (Gray 1848 in Gray 1844–1849), Hemiphagini either. As such, the Trugon- and Hemiphaga-based
Ptilophyrus (Bonaparte, 1840), Palumbus (Reichenbach, 1851 family groups do not fulfil Article 13.2.1 of the Zoological Code
in Reichenbach 1849–1853), Lophyrus (Le Maout, 1853), and are not available.
Alectroenas (Bonaparte, 1853), Caloenas (Bonaparte, 1853),
Carpophaga (Bonaparte, 1853), Lopholaimus (Bonaparte, The Claravis-based family group
1853), Phaps (Bonaparte, 1853), Ptilopus (Bonaparte, 1853), One of the consequences of our historical overview of columbid
Zenaida (Bonaparte, 1853), Macropygia (Bonaparte, 1854a), family-group nomina is the realization that within the clade
Chalcophaps (Bonaparte, 1854b in Bonaparte 1850–1865), Claravinae, the oldest valid family-group nomen is based on the
Chamaepelia (Bonaparte, 1854b in Bonaparte 1850– genus Chamaepelia (a subjective junior synonym of Columbina).
1865), Chrysoena (Bonaparte, 1854b in Bonaparte 1850– However, Article 35.5 of the Zoological Code allows us to retain
1865), Geopelia (Bonaparte, 1854b in Bonaparte 1850–1865), Claravinae instead of using Chamaepeliinae. Article 35.5 states
Starnoenas (Bonaparte, 1854b in Bonaparte 1850–1865), that if: ‘after 1999 a name in use for a family-group taxon (e.g. for
Ducula (Reichenbach, 1862), Geotrygon (Reichenbach, 1862), a subfamily) is found to be older than a name in prevailing usage
Myristicivora (Reichenbach, 1862) Phapitreron (Reichenbach, for a taxon at higher rank in the same family-group taxon (e.g.
1862), Leucotreron (Reichenbach, 1862), Osmotreron for the family within which the older name is the name of a sub-
(Reichenbach, 1862), Zonoenas (Reichenbach, 1862), Megapelia family) the older name is not to displace the younger name’. The
(Sundevall, 1873 in Sundevall 1872–1873), Otidiphaps (Martin, Chamaepelia-based family group was established by Bonaparte
1904), †Pezophaps (Martin, 1904), Muscadivores (McGregor, (1854b in Bonaparte 1850–1865) at a rank below the subfamily,
18 • Young et al.
Table 1. List of family-group nomina validly established under the Zoological Code within Columbinae Illiger, 1811 and Claravinae Todd,
1913 (sensu Dickinson and Remsen Jr 2013, Dickinson and Raty 2015). Nominal authors who first used a particular rank are given, and in
square brackets [] is how the nomen was first written. The oldest nomen within each family group, hence the nominal authority under the
Zoological Code, is in bold. The nomenclatural acts written with strikethrough are unavailable.
Stem Family Subfamily Below subfamilya
Columbinae: Columbini Illiger, 1811
Columba Illiger, 1811 Selby, 1835 Bonaparte, 1854a
[Columbini] [Columbinæ] [Columbeæ]
Palumbus Reichenbach, Bonaparte, 1854b
1851b [Palumbeae]
[Palumbinae]

Macropygia Salvadori, 1893 Bonaparte, 1854a
[Macropygiinæ] [Macropygieæ]
†Ectopistes Selby, 1835 Ridgway, 1916
[Ectopistinæ] [Ectopisteae]
Columbinae: Zenaidini Bonaparte, 1853
Zenaida Burmeister, 1856 Bonaparte, 1853 Bonaparte, 1854b
[Zenaididae] [Zenaidinæ] [Zenaideae]
Geotrygon Reichenbach,
1862b
[Geotrygoninae]
Claravinae Todd, 1913
Chamaepeliac Reichenbach, 1862b Bonaparte,
[Chamaepeleiinae] 1854b
[Chamaepelieae]
Claravis Richmond ,1917 Todd, 1913
[Claraviidae] [Claraviinae]
Peristerad Burmeister, 1856 Selby, 1835 Bonaparte, 1854b
[Peristerinae] [Peristerinæ] [Peristereae]
a
Bonaparte (1854b in Bonaparte 1850–1865) and Ridgway (1916) referred to the rank below subfamily as ‘series’. They used the suffix -eæ.
b
Reichenbach used the -inae suffix for nomina at the family, subfamily, and below subfamily ranks (at what we today would call the tribe and subtribe ranks). At what we today would
call the subtribe rank, he also used the -eae and -rae suffices.
c
Chamaepelia is a subjective junior synonym of Columbina. As we show, it does not displace the Claravis-based family group owing to Article 35.5 of the Zoological Code.
d
The nomina based on the genus Peristera are not available.
and the Claravis-based family group was established (and is cur- Treron-based family group has been in prevailing usage since the
rently used) at the subfamily rank. Therefore, Article 35.5 ap- 19th century (see the historical section above and Supporting
plies in this situation. Information, Appendix S1).
Following recommendation 40A, we cite the nominal au-
The Treron-based family group thority as: Treronini Gray, 1840 (1836). Under Article 40.2, the
Another consequence of our historical overview of columbid date of priority for the Treron-based family group is 1836. Under
family-group nomina is the realization that within the clade standard avian taxonomic practice, Treronini can be cited as:
Treronini, the oldest valid family-group nomen is based on the Treronini Gray.
genus Vinago (a subjective junior synonym of Treron). In ac-
cordance with Article 40.2 of the Zoological Code, we retain Ergotaxonomy of Columbidae
the Treron-based family group established by Gray (1840) ra- We can therefore confirm that the family-group nomina of the
ther than using the Vinago-based family group established by current systematization (Dickinson and Remsen Jr 2013, Chen
Hodgson (1836), even though the latter has priority. This is be- et al. 2022, Oliver et al. 2023) of pigeons is valid (see Tables 1 and
cause of the conditions set out in Article 40.2, i.e. was the name 2). Based on the consensus topology in the work by Oliver et al.
replaced before 1961, and is the replacement name in prevailing (2023), the current ergotaxonomy of Columbidae is as follows
usage? In this instance, both conditions are met, because since [also see Supporting Information, Appendix S2; genera they did
Hartert and Goodson (1918: 349) adopted Treron over Vinago, not recover as monophyletic are indicated with an asterisk (*)]:
the synonymy of these two genera has not been challenged.
Usage of the Vinago-based family group comes from the 19th Family Columbidae Illiger, 1811
century, being used by Hodgson (1836) for a subfamily-rank Subfamily Claravinae Todd, 1913
nomen and by Reichenbach (1862) for a below subfamily-rank Genera: Claravis, Paraclaravis Sangster et al.,
nomen. The nominal authority of the Vinago-based family group 2018, Columbina von Spix, 1825, Metriopelia
was often given as Reichenbach, 1862 (e.g. Bock 1994); how- Bonaparte, 1855, and Uropelia Bonaparte,
ever, as we have shown, it was first used by Hodgson (1836). The 1855.
Table 2. List of family-group nomina validly established under the Zoological Code within Raphinae (sensu Dickinson and Remsen Jr 2013,
Dickinson and Raty 2015). Nominal authors who first used a particular rank are given, and in square brackets [] is how the nomen was first
written. The oldest nomen within each family group, hence the nominal authority under the Zoological Code, is in bold. The nomenclatural
acts written with strikethrough are unavailable.
Raphini Oudemans, 1917 (1835)
†Didusb Swainson, 1835 de le Fresnaye, 1839
[Didiadæ] [Didinæ]
†Raphusb Oudemans, 1917 Verheyen, 1957 Verheyen, 1957
[Raphidae] [Raphinae] [Raphini]
†Pezophaps Martin, 1904 Wolters, 1975 Verheyen, 1957

[Pezophabidae] [Pezophabinae] [Pezophabini]
†Microgoura Richmond, 1909?
[Microgouridæ]
Goura Reichenbach, 1852c Hodgson, 1836
[Gourinae] [Gourinæ]
Ptilophyrus Bonaparte, 1840
[Ptilophyrinae]
Lophyrus Le Maout, 1853
[Lophyriens]
Megapelia Sundevall, 1873d
[Megapeliinæ]
Didunculus Bonaparte, 1850 Gray, 1848
[Didunculidæ] [Didunculinæ]
Gnathodone Melville, 1848
[Gnathodontinæ]
Caloenas Bonaparte, 1854a Bonaparte, 1853
[Calliænadidae] [Calænadinæ]
Otidiphaps Martin, 1904
[Otidiphabinae]
Trugonf Verheyen, 1957
[Trugonini]
Turturini Gray, 1840
Turtur Gray, 1840 Verheyen, 1957
[Turturinæ] [Turturini]
Oena Verheyen, 1957
[Oenini]
Chalcophaps Bonaparte, 1854b
[Chalcophapeae]
Treronini Gray, 1840 (1836)
Treron Bonaparte, 1853 Gray, 1840 Chen et al., 2022?
[Treronidæ] [Treroninæ] [Treronini]
Osmotreron Reichenbach, 1862c
[Osmotreroneae]
Vinago Hodgson, 1836 Reichenbach, 1862c
[Vinaginæ] [Vinagineae]
Ptilinopini Selby, 1835
Ptilinopus Selby, 1835 Verheyen, 1957?
[Ptilinopinæ] [Ptilinopini]
Chlorotreronf Reichenbach, 1862c
[Chlorotreroneae]
Leucotreron Reichenbach, 1862c
[Leucotreroneae]
Megaloprepia Verheyen, 1957
[Megaloprepiinae]
Ptilopus Bonaparte, 1853 Bonaparte, 1854b
[Ptilopodinæ] [Ptilopodeae]
20 • Young et al.
Table 2. Continued

Ducula Verheyen, 1957? Verheyen, 1957? Reichenbach, 1862c
[Duculidae] [Duculinae] [Duculinae]
Carpophaga Elliot, 1885 Bonaparte, 1853 Reichenbach, 1862c
[Carpophagidæ] [Carpophaginæ] [Carpophaginae]
Myristicivora Reichenbach, 1862c
[Myristicivorae]
Zonoenas Reichenbach, 1862c
[Zonoeninae]

Muscadivores McGregor, 1909
[Muscadivorinæ]
Hemiphagag Verheyen, 1957
[Hemiphagini]
Phapitreron Reichenbach, 1862c
[Phapitreroneae]
Lopholaimus Bonaparte, 1853 Reichenbach, 1862c
[Lopholaiminæ] [Lopholaiminae]
Alectroenas Bonaparte, 1853 Verheyen, 1957
[Alectrænadinæ] [Alectroenini]
Chrysoena Bonaparte, 1854b
[Chrysaeneae]
Phabini Bonaparte, 1853
Phaps Bonaparte, 1853 Bonaparte, 1854b
[Phapinæ] [Phapeae]
Cosmopelia Verheyen, 1957
[Cosmopeliini]
Gallicolumba Verheyen, 1957 Verheyen, 1957
[Gallicolumbinae] [Gallicolumbini]
Geopelia Reichenbach, 1862c Bonaparte, 1854b
[Geopeleiinae] [Geopelieae]
Geophaps Verheyen, 1957
[Geophabini]
Leucosarcia Verheyen, 1957
[Leucosarciini]
Ocyphaps Verheyen, 1957
[Ocyphabini]
Starnoenas Burmeister, 1856 Verheyen, 1957? Bonaparte, 1854b
[Starnoenidae] [Starnoeninae] [Starnoenadeae]
a
Bonaparte (1854b in Bonaparte 1850–1865) referred to the rank below subfamily as ‘series’. He used the suffix -eæ.
b
Didus is an objective synonym of Raphus.
c
Reichenbach used the -inae suffix for nomina at the family, subfamily and below subfamily ranks (at what we today would call the tribe and subtribe ranks). At what we today would
call the subtribe rank, he also used the -eae and -rae suffices.
d
Sundevall 1873 in Sundevall (1872–1873) also used the -inae suffix for nomina at the family rank.
e
The nomina based on the genus Gnathodon are not available.
f
Reichenbach (1862) used a ‘Chlorotreron’-based family-group nomen. However, he did not include a genus with that nomen. The earliest version of Chlorotreron we can find is
Salvadori (1882).
g
Nomen is not available as per Article 13.2.1 of the Zoological Code.
Subfamily Columbinae Illiger, 1811 Tribe Zenaidini Bonaparte, 1853

Genera: Geotrygon, Leptotrygon Banks et al.,
Tribe Columbini Illiger, 1811 2013, Leptotila Swainson, 1837, Zenaida, and
Genera: Columba, †Ectopistes, Macropygia, Zentrygon Banks et al., 2013.
Nesoenas, Patagioenas Reichenbach, 1853 Subfamily Raphinae Oudemans, 1917 (1835)
in Reichenbach 1851–1854, Reinwardtoena
Bonaparte, 1854c, Spilopelia Sundevall, 1873 Tribe Phabini Bonaparte, 1853
in Sundevall 1872–1873, Streptopelia, and Genera: Gallicolumba, Geopelia, Geophaps,
Turacoena Bonaparte, 1854c. Henicophaps Gray, 1862, Leucosarcia, Ocyphaps,
Pampusana Bonaparte, 1855, Petrophassa Gould, the genus or family-group spellings are preserved under Articles
1841, and Phaps. 29.5 and 33.2.3.1 or 33.3.1). The correct stem for the Raphium-
based family group is Raphi- not Raph- (as shown by the
Tribe Ptilinopini Selby, 1835 Rhaphium-based family-group nomen Rhaphiinae). Therefore,
Genera: Alectroenas, Drepanoptila Bonaparte, under Article 35.4.1 the Raphium-based family group should be
1855, Ducula, Gymnophaps Salvadori, 1874, corrected to Raphiinae, which removes the homonymy.
Hemiphaga, Lopholaimus, Phapitreron, and
Ptilinopus*. RESULTS
Tribe Raphini Oudemans, 1917 (1835) SYSTE M ATIC ZOOLOGY
Genera: Caloenas, Didunculus, Goura, Otidiphaps, Aves Linnaeus, 1758

†Pezophaps, †Raphus, and Trugon. Columbiformes Latham, 1790 (as Columbæ)
Columbidae Illiger, 1811 (as Columbini)
Tribe Treronini Gray, 1840 (1836)
Raphinae Oudemans, 1917 (1835)
Genus: Treron.
Type genus: †Raphus Brisson, 1760b.
Tribe Turturini Gray, 1840
Genera: Chalcophaps, Oena, and Turtur. Nomenclatural note
In accordance with Article 40.2 of the Zoological Code, we re-
Note that Oliver et al. (2023) did not include three extant genera tain the †Raphus-based family group established by Oudemans
considered valid by the International Ornithological Committee (1917) rather than use the †Didus-based family group estab-
World Bird List v.13.2 (https://www.worldbirdnames.org/new/ lished by Swainson (1835) even though the latter has priority.
bow/pigeons/): Alopecoenas Bonaparte, 1855, Cryptophaps This is owing to the conditions set out in Article 40.2, i.e. was
Salvadori, 1893, and Starnoenas, the last of which is currently the name replaced before 1961, and is the replacement name in
incertae subfamiliae. Many of the known fossil or extinct genera prevailing usage? In this instance, both conditions are met, be-
are likewise not included in this systematization. Based on the cause since Peters (1937: 10) adopted the use of †Raphus and
molecular-based supertree of Oliver et al. (2023, see their sup- Raphidae over †Didus and Dididae, the †Raphus-based family
plementary information for full details), the internal relation- group has been in prevailing usage ever since (e.g. Verheyen
ships of columbids can be summarized in parenthetical form 1957, Wetmore 1960, Brodkorb 1971, Wolters 1975, Sibley and
as: Claravinae + (Columbinae + ((Treronini + Turturini) + Ahquist 1990, Witmer and Cheke 1991, Bock 1994, Mourer-
(Phabini + (Raphini + Ptilinopini)))). Their results suggest Chauviré et al. 1995, Weber and Krell 1995, Janoo 1996, 2000,
that: (i) Claravinae is monophyletic and the sister taxon to an Mlíkovský 2000, Livezey and Zusi 2006, 2007, Mlíkovský et al.
unnamed Columbinae + Raphinae clade; (ii) Columbinae is 2011, Dickinson and Remsen Jr 2013, Parish 2013, Heupink
monophyletic and the sister taxon to Raphinae; and (iii) within et al. 2014, van Grouw 2014, Byrkjedal et al. 2016, Gold et al.
Raphinae there is a deep split between the Treronini + Turturini 2016, Rodríguez-Pontes 2016, Watanabe et al. 2018, Nowak et
subclade (unnamed) and the Phabini + Raphini + Ptilinopini al. 2019, Boyd et al. 2022, Chen et al. 2022, Oliver et al. 2023,
subclade (unnamed). The analyses of Oliver et al. (2023) sam- Szymański et al. 2023, Wyndham and Park 2023).
pled Columbidae more broadly (with 250 species included) Following recommendation 40A, we cite the nominal au-
and thus differed in some respects from earlier analyses (such thority as: Raphinae Oudemans 1917 (1835). Under Article
as Pereira et al. 2007, from which the three clade subdivision of 40.2, the date of priority for the †Raphus-based family group is
Columbidae originates). 1835, thus it has priority over most other family groups within
Columbidae (see above). As we noted above, the correct nom-
Homonymy problem inal authority of the †Raphus-based family group is Oudemans
There is a homonymy problem with Raphinae/Raphidae. A (1917), not Poche (1904) or Wetmore (1930). Under standard
Raphium-based Raphinae appeared in the work of Karsh (1883: avian taxonomic practice, Raphinae can be cited as: Raphinae
336), prior to the †Raphus-based Raphidae (based on Brisson Oudemans.
1760a, b) that first appeared in work of Oudemans (1917:
100–101). Note that under the Zoological Code, the Principal Composition
of Coordination (Article 36.1) results in the nominal authority Following Dickinson and Remsen Jr (2013: 25, 36), the
of a family group spanning all ‘ranks’ of the group (in this case, †Raphus-based family group has been used to provide nomina
Raphidae and Raphinae). Raphium Latreille, 1829 is a junior for major columbid subclades. Raphinae is currently used to
synonym of Rhaphium Meigen, 1803 (see O’Hara et al. 2011); unite the ‘traditional’ subfamilies Gourinae, Otidiphabinae,
the latter is still used in the dipteran literature and has its own Didunculinae, and Treroninae (sensu Baptista et al. 1997).
family-group nomen, Rhaphiinae Bigot, 1852.
Under Article 35.4.1 of the Zoological Code, if a family-group Raphini Oudemans, 1917 (1835)
nomen is established upon an unjustified emendation or incor-
rect spelling of the type genus it must be corrected (unless either Type genus: †Raphus Brisson, 1760b.
22 • Young et al.
Nomenclatural note ZooBank registration number [urn:lsid: zoobank.org

As with Raphinae above, we follow recommendation 40A of the :pub:A79C5ED9-6E44-4DE7-AC70-CD77267FC655].
Zoological Code and cite the nominal authority of Raphini as:
Raphini Oudemans 1917 (1835). Article 36 of the Zoological Type genus: †Raphus Brisson, 1760b.
Code (Principal of Coordination applied to family-group
Nomenclatural note
nomina) states that when a name (= nomen) is established at
one rank, it is simultaneously established at all other ranks in As with Raphinae and Raphini above, we follow recommenda-
said family group. All the nomina at these various ranks retain tion 40A of the Zoological Code and cite the nominal authority
the same nominal authority (authorship and date). Following of †Raphina as: †Raphina Oudemans, 1917 (1835). Note that
Article 40.2, the date of priority is 1835. Under standard avian although we formally established the new subtribe †Raphina,
taxonomic practice, Raphini can be cited as: Raphini Oudemans. the correct nominal authority of †Raphina is Oudemans, 1917.
This is owing to Article 36 of the Zoological Code (Principal

Composition of Coordination applied to family-group nomina). Following
Raphini is currently used to unite the ‘traditional’ subfamilies Article 40.2, the date of priority is 1835. Under standard
Gourinae, Otidiphabinae, and Didunculinae (sensu Baptista et al. avian taxonomic practice, †Raphina can be cited as: †Raphina
1997). This clade is composed of the extinct Dodo and Solitaire, Oudemans.
alongside the extant genera Caloenas, Goura, Didunculus,
Otidiphaps, and Trugon (see Chen et al. 2022, Oliver et al. 2023). Geographical range
Note that this clade is more expansive than the subclade char- The islands of Mauritius and Rodrigues, Mascarene Islands.
acterized as a ‘ground-dwelling island endemics’ radiation (i.e.
†Raphus, †Pezophaps, Caloenas, Goura, and Didunculus). It is Diagnosis
possible that the extinct genera †Microgoura and †Natunaornis Columbids with the following unique combination of charac-
Worthy, 2001 also belong to this subclade. ters: large body size; sexual dimorphism of the skeleton is ex-
treme in the Solitaire and noticeable in the Dodo; basipterygoid
†Raphina subtribus nova processes of the parasphenoid are reduced; pectoral girdle is
small relative to volant columbids; pelvic girdle is large relative
• 1811 Inepti fam. nov.—Illiger, p. 245. [Invalid—stem
to volant columbids; scapulocoracoidal angle is ~100°–110°;
based on a specific epithet.]
scapula–coracoid articulation is typically not fused; modal
• 1835 Didiadæ fam. nov. [sic]—Swainson, p. 239.
number of sternal ribs is four to five; ulnae have conspicuous pa-
• 1836 Dididæ emend. (Swainson)—Swainson, p. 286.
pillae remigiales caudales; sternum basin is broad and dorsally
• 1837 Dididæ (Swainson)—Swainson, p. 200.
concave; sternal carina is ventrally truncated and round, with the
• 1839 Didinæ subfam. nov. (Swainson)—de la Fresnaye,
cranial margin forming an obtuse angle with the base; the pila
p. 193–194.
carinae, sulcus carinae, and crista lateralis are greatly reduced;
• 1840 Didinæ (Swainson)—Gray, p. 64.
sternal trabecula lateralis are distally reduced and essentially
• 1842a Dididæ (Swainson)—Bonaparte, p. 14, 89.
straight; sternal trabecula intermedia are absent; sternal fenestra
• 1842a Didinæ (Swainson)—Bonaparte, p. 14.
medialis is absent; sternal rostrum spinae externa and interna
• 1842b Dididæ (Swainson)—Bonaparte, p. 65.
are greatly reduced; lineae intermusculares of the sternum con-
• 1842b Didinæ (Swainson)—Bonaparte, p. 65.
verge noticeably cranially to the margo caudalis; the cranial por-
• 1842 Didusideae fam. nov. [sic]—Lesson, column 1036.
tions of the ilia are typically bowed dorsally, meeting medially
• 1854a Inepti order nov.—Bonaparte [also contained the
in a curved arch dorsal to the crista dorsalis of the synsacrum.
Réunion ‘dodo and solitaire’].
(The preceding characters were modified from table 2 of Livezey
• 1854a Dididæ (Swainson)—Bonaparte [also contained
1993, comparing †Raphus cucullatus and †Pezophaps solitaria
the Réunion ‘dodo and solitaire’].
with volant members of Columbidae.) Enlargement of the olfac-
• 1862 Didinae (Swainson)—Reichenbach, p. 124.
tory bulbs and Wulst. [Based on the endocranial reconstructions
• 1870b Dididæ (Swainson)—Gray, p. 248.
by Gold et al. (2016), comparing the Dodo and the Solitaire with
• 1870b Didinæ (Swainson)—Gray, p. 248.
volant columbids.]
• 1904 Raphidae fam. nov.—Poche, p. 500. [Based
on Raphus Möhring, 1758—genus is contained in a
Composition
work suppressed for nomenclatural purposes by the
Commission. Unavailable.] †Raphus and †Pezophaps.
• 1916 Raphi subord. nov.—Ridgway, p. 275–277.
• 1917 Raphidae fam. nov.—Oudemans, p. VI, 98, 100, Comments
101. †Raphina is a new nomen for the clade uniting †Raphus and
• 1971 Raphidae (Oudemans)—Brodkorb, p. 199. †Pezophaps, which historically has been referred to as Dididae,
• 1991 Raphidae (Oudemans)—Witmer and Cheke, p. Didinae, Raphidae, Raphinae, or Raphini (see historical section
134. above). We chose to establish a new family-group nomen rather
• 1995 Raphidae (Oudemans)—Weber and Krell, p. 171. than synonymize †Pezophaps with †Raphus, a possibility that
[Nominal authority cited was Wetmore 1930.] Livezey (1993) and Janoo (2000) previously suggested. Our
• 2013 Raphini (Oudemans)—Parish, p. xi. rationale for this is 2-fold: (ii) both genera are morphologically
• 2014 Raphinae (Oudemans)—van Grouw, p. 300. distinct and can be distinguished readily from one another (see
the diagnoses for both taxa below; the figures in the studies by the Dodo. Poche (1904: 500) was the first to use the binomen
Meville 1848, Strickland 1848, Livezey 1993, and the discussion †Raphus cucullatus, but as outlined above, the genus †‘Raphus’
on the differences between both taxa by Newton and Newton used by Poche is no longer valid. This is owing to Poche (1904)
1868, 1869, Owen 1872, 1878, Newton and Clark 1879); and using †‘Raphus’ Möhring, 1758, a name that appears in a sup-
(ii) the lack of a fossil record. At present, we cannot distinguish pressed work (see Opinion 801 in ICZN 1967), and not †Raphus
adequately between the following two competing hypotheses: Brisson, 1760b.
(i) the common ancestor of the Dodo and the Solitaire was vo- Following recommendation 51G of the Zoological Code, the
lant, with two independent transitions from powered flight to type species can be referred to as †Raphus cucullatus (Linnaeus,
terrestrial cursoriality; or (ii) their common ancestor was also 1758) Oudemans, 1917. It can also be written more simply as
secondarily flightless. Should fossil evidence support hypoth- †Raphus cucullatus (Linnaeus, 1758).
esis (i), then retaining †Raphus and †Pezophaps as distinct
genera would reflect parallel evolution of a secondarily terres- Etymology

trial morphotype. Molecular-based divergence estimates place According to ( Jobling 2010: 331): raphos was the name given
the separation of †Raphus and †Pezophaps between 17.6 and to the Great Bustard by Galenus (190 AD) and was ‘probably a
35.9 Mya, which is far older than the estimates for emergence misreading of outis or ōtis’. Jobling (2010: 331) also mentioned
of the islands of Mauritius and Rodrigues, both of which are that rhaphos is a ‘dubious bird name mentioned by Hesychius in
<10 Mya (Shapiro et al. 2002). This would support hypothesis the plural form rhaphoi’. Parish (2012: 6, 2013: 140–141) out-
(i). However, as noted by Shapiro et al. (2002), other ridges of lined a possible etymology for Raphus, based on Hesychius, that
the Mascarene Plateau would have been above sea level during derives from Rhamphios/Rhamphos, by the way of Ramphos,
the divergence estimation period of the Dodo and the Solitaire. possibly referring to the pelican of Egypt, and in particular its
Without fossil evidence, we cannot determine raphinan modes large beak. The origin of the nomen Raphus remains unclear.
of dispersal during this time span or when the transition(s) to
secondary cursoriality occurred. However, if new fossil discov- Diagnosis
eries support hypothesis (ii), then it will be up to future workers Same as the only species (see below).
to decide whether the genus †Pezophaps and our new nomen,
†Raphina, are required. †Raphus cucullatus (Linnaeus, 1758)
We propose the English vernacular term ‘Mascarene giant (Figs 1, 3, 4)
ground doves’ for raphinans (i.e. the Dodo and the Solitaire).
Our phraseology is based on that of previous workers, such as • 1605 Gallus gallinaceus peregrinus (pre-Linnean)—
Owen (1872: 514, 515, 518), who referred to them as ‘Ground- Clusius, p. 99–101, figured on p. 100.
Doves’, Verheyen (1957: 30), who referred to them as ‘pigeons • 1635 Cygnus cuculatus (pre-Linnean)—Nieremberg, p.
géants’ [giant pigeons], and Worthy (2001: 791), who used the 231, figured on p. 231.
phrase ‘Mascarene giant pigeons’. Referring to both species as • 1656 Dodar (pre-Linnean)—Tradescant, p. 4.
dodos would be confusing, because the vernacular name ‘dodo’ • 1658 Dronte (pre-Linnean)—Bontius, p. 70–71, figured
has become engrained with †Raphus cucullatus in popular cul- on p. 70.
ture. • 1752 Raphus (pre-Linnean)—Möhring, p. 58.
• 1758 Raphus (work suppressed for nomenclatural pur-
†Raphus Brisson, 1760b (as both Raphus and Raphi) poses by the Commission)—Möhring, p. 44.
• 1758 Struthio cucullatus sp. nov.—Linnaeus, p. 155.
Type species: †Raphus raphus Brisson, 1760b. Type species by • 1760b Raphi raphus gen. et sp. nov.—Brisson, p.
monotypy. 14–15 (generic name had genitive suffix when used
The use of †Raphus raphus (as Raphi raphus, where the genus as a binomial).
had the genitive Latin suffix when appearing next to the spe- • 1766 Didus ineptus gen. et sp. nov.—Linnaeus, p. 267.
cific epithet) first appeared in Brisson (1760b: index table). • 1785 Hooded Dodo—Latham, p. 1–3, plate LXX.
Rothschild (1907: 172–173) synonymized †Didus ineptus • 1785 Nazarene Dodo—Latham, p. 4–5.
with †Struthio cucullatus, creating the new combination †Didus • 1789 Didus ineptus Linnaeus—Gmelin, p. 728.
cucullatus. Ridgway (1916: 277) synonymized †Didus with • 1789 Didus nazarenus sp. nov.—Gmelin, p. 729.
†Raphus, creating the new combination †Raphus cucullatus, but • 1790 Didus ineptus Linnaeus—Latham, p. 662.
Ridgway did not mention whether Brisson or Möhring was the • 1790 Didus nazarenus Gmelin—Latham, p. 663.
source of the †Raphus he used. The first to use †Raphus Brisson, • 1811 Didus ineptus Linnaeus—Illiger, p. 245.
1760b explicitly was Oudemans (1917: 14, 101), who also • 1828 Didus ineptus Linnaeus—Duncan, p. 554–566, figs
used the combination †Raphus cucullatus. Peters (1937: 10) 1–3.
synonymized †Raphus raphus with †Struthio cucullatus, using the • 1835 Didus ineptus Linnaeus—de Blainville, p. 1–36,
combination †Raphus cucullatus. Note that Allen (1910: 329) plates 1–4.
had previously synonymized the genera †Raphus and †Didus, • 1839 Didus ineptus Linnaeus—de la Fresnaye, p. 194.
and stated that †Raphus should be used instead of †Didus, but • 1840 Didus ineptus Linnaeus—Gray, p. 64.
he did not mention specific epithets. Allen (1910) is, we believe, • 1842 Didus ineptus Linnaeus—Bonaparte, p. 89.
the first to use †Raphus Brisson, 1760b as the generic name for • 1842 Didus ineptus Linnaeus—Lesson, column 1036.
24 • Young et al.
• 1844 Didus ineptus Linnaeus—Strickland, p. 324. • 2001 Raphus cucullatus (Linnaeus)—Worthy, p. 766.
• 1848 Didus ineptus Linnaeus—Strickland, p. 7–45, plates • 2002 Raphus cucullatus (Linnaeus)—Shapiro et al., p.
1, 5, 6, 8, 9, 11, 12. 1683.
• 1848 Didus ineptus Linnaeus—Melville, p. 71–112, • 2003 Raphus cucullatus (Linnaeus)—Roberts and Solow,
plates 1, 5, 6, 8, 9, 11, 12. p. 245, fig. 1.
• 1854 Didus ineptus Linnaeus—Bartlett, p. 280–284, plate • 2005 Raphus cucullatus (Linnaeus)—Janoo, p. 167–178,
XLV (fig. 3). [Published in 1854.] fig. 2.
• 1854b Didus ineptus Linnaeus—Bonaparte, p. 2. • 2006 Raphus cucullatus (Linnaeus)—Cheke, p. 155–157.
• 1854b Didus nazarenus Gmelin—Bonaparte, p. 2–3. • 2007 Raphus cucullatus (Linnaeus)—Pereira et al., p.
• 1859 Didus ineptus Linnaeus—Strickland, p. 187, 195– 656–658, 660, 662, 663, 665, 667, 669.
196. • 2008 Raphus cucullatus (Linnaeus)—Cheke and Hume,
• 1862 Didus ineptus Linnaeus—Reichenbach, p. 125– p. 26, 50, 162, 400, plate.

128. • 2009 Raphus cucullatus (Linnaeus)—Rijsdijk et al., p. 14,
• 1862 Didus nazarenus Gmelin—Reichenbach, p. 128. fig. 3G.
• 1868 Didus ineptus Linnaeus—Newton and Newton, p. • 2011 Struthio cucullatus Linnaeus—Mlíkovský, p. 140.
428–431. • 2012 Raphus cucullatus (Linnaeus)—Meijer et al., p. 177,
• 1869 Didus ineptus Linnaeus—Newton and Newton, p. figs 4–6.
327, 331–350. • 2012 Raphus cucullatus (Linnaeus)—Mlíkovský, p. 105–
• 1870b Didus ineptus Linnaeus—Gray, p. 248. 106, fig. 10.
• 1878 Didus ineptus Linnaeus—Owen, p. 87–96. • 2012 Raphus cucullatus (Linnaeus)—Roberts, p. 1478–
• 1872 Didus ineptus Linnaeus—Owen, p. 513–523, plates 1480.
LXIV, LXV, LXVI (figs 8–11, 14–17). • 2013 Raphus cucullatus (Linnaeus)—Parish, [numerous
• 1891 Didus ineptus Linnaeus—Lydekker, p. 129–130, fig. occurrences].
31. • 2014 Raphus cucullatus (Linnaeus)—Heupink et al., p. 1,
• 1893 Didus ineptus Linnaeus—Sharpe, p. 632–635. 3.
• 1904 ‘Raphus’ cucullatus (Linnaeus) comb. nov.— • 2014 Raphus cucullatus (Linnaeus)—van Grouw, p. 291,
Poche, p. 500. [‘Raphus’ Möhring 1758—a work 300.
that was later suppressed by the Commission in • 2015 Raphus cucullatus (Linnaeus)—Claesson and
1967.] Hume, p. 21, figs 2, 4–10.
• 1907 Didus cucullatus (Linnaeus) comb. nov.— • 2015 Raphus cucullatus (Linnaeus)—Claessons et al., p.
Rothschild, p. 172–173, plate 24. 29, figs 1–9, 11–65.
• 1916 Raphus cucullatus (Linnaeus)—Ridgway, p. 277. • 2015 Raphus cucullatus (Linnaeus)—Rijsdijk et al., p. 4,
[Does not state which Raphus was used.] figs 1, 3, 6–7.
• 1917 Raphus cucullatus (Linnaeus) comb. nov.— • 2016 Raphus cucullatus (Linnaeus)—Brassey et al., p.
Oudemans, p. 14, 101. [Raphus Brisson, 1760b was 1–3, 5, 7–14, fig. 3.
used.] • 2016 Raphus cucullatus (Linnaeus)—Besnard et al., p.
• 1937 Raphus cucullatus (Linnaeus)—Peters, p. 10. 76–78.
[Raphus Brisson, 1760b was used.] • 2016 Raphus cucullatus (Linnaeus)—Byrkjedal et al., p.
• 1957 Raphus cucullatus (Linnaeus)—Verheyen, p. 3, 6, 199.
17, 19, 21, 24, 30. • 2016 Raphus cucullatus (Linnaeus)—Gold et al., p. 950–
• 1971 Raphus cucullatus (Linnaeus)—Brodkorb, p. 199– 961, figs 2A, 3A, 4A, 5, 6A, 7A, 8A.
200. • 2016 Raphus cucullatus (Linnaeus)—Rodríguez-Pontes,
• 1975 Raphus cucullatus (Linnaeus)—Wolters, p. 43. p. 398, fig. 8.
• 1989 Raphus cucullatus (Linnaeus)—Brom and Prins, • 2016 Raphus cucullatus (Linnaeus)—Soares et al., p. 1–3,
p. 233–245, plates 1–5. 5.
• 1990 Raphus cucullatus (Linnaeus)—Sibley and Ahquist, • 2017 Raphus cucullatus (Linnaeus)—Angst et al., p. 1–8,
p. 421. figs 1–5.
• 1991 Raphus cucullatus (Linnaeus)—Witmer and Cheke, • 2017 Raphus cucullatus (Linnaeus)—van Heteren et al.,
p. 134. p. 1–15, fig. 1.
• 1993 Raphus cucullatus (Linnaeus)—Kitchner, p. 279– • 2018 Raphus cucullatus (Linnaeus)—Bruxaux et al., p.
299, figs 1–3, 8–10. 250, 254.
• 1993 Raphus cucullatus (Linnaeus)—Livezey, p. 247– • 2020 Raphus cucullatus (Linnaeus)—Cheke and Parish,
282, plate 1 (fig. 2a). p. 1–12.
• 1996 Raphus cucullatus (Linnaeus)—Janoo, p. 57–75, • 2021 Raphus cucullatus (Linnaeus)—Warnett et al.,
figs 1–5, 7. p. 2247–2255, figs 1–6.
• 2000 Raphus cucullatus (Linnaeus)—Janoo, p. 323, 325– • 2022 Raphus cucullatus (Linnaeus)—Chen et al., p. 1005,
327, fig. 1. 1011.
Type specimen Diagnosis

No type specimen has been designated formally for †Raphus Columbids with the following unique combination of char-
cucullatus (the Mauritian Dodo, or simply the Dodo). Several acters: modest sexual dimorphism of the skeleton; the frontal
syntypes have been postulated (see Mlíkovský et al. 2011, processes (= processus frontales) of the premaxilla meet prox-
Mlíkovský 2012, Parish 2015), but, as discussed above, there is imally without any space along their medial margins; distally,
no direct evidence that any of those specimens form part of the the premaxillary frontal processes are enlarged, gradually
original type series. Alas, we cannot locate any members of the tapering, fusing well proximally to their distal terminus; an-
original type series, which, as we stated above, is based upon: terior border of the nares (= apertura nasalis ossea) is slit-like;
(i) the lost partial leg and foot brought to Europe (Clusius the body of the premaxilla (corpus ossis premaxillare) forms
1605: 100); (ii) the unknown individuals that Bontius’ (1658: the large, porous, bulbous tip of the upper bill; the maxillary
70–71) description was based upon; and (iii) the unknown in- process (= processus maxillaris) of the premaxilla is vertically
dividuals that Dutch sailors saw during the van Neck voyage, oriented distally, converging with the nasal process to form a

which, in part, formed the basis of Clusius’ (1605: 99–100) slit-like nares (= apertura nasalis ossea); the distal ventral sur-
illustrations and description. As such, we cannot designate a faces of the premaxilla are apneumatic; the distal end of the
lectotype. lower jaw is comparatively deep and strong; the proximal end
At present, we cannot formally designate a neotype for of the lower jaw is comparatively shallow and caudally exten-
†Raphus cucullatus. In order to be fully compliant with Article sive, with obtuse posterior margins of the rami; the quadrate–
75 of the Zoological Code, a neotype designation cannot be lower jaw articulation is situated comparatively posteriorly on
‘an end in itself, or a matter of curatorial routine, and any such the cranium; the lacrimal processes of the frontal are greatly
neotype designation is invalid’ (Article 75.2), and ‘there is an reduced; the frontal is dorsally convex and dome-shaped;
exceptional need’ (Article 75.3). There is no exceptional need the frontals extend anteromedially to the frontonasal hinge,
for a neotype for the Dodo. As we have shown above, only covering the posteromedial portions of the nasals and frontal
one raphinan species is known from the Island of Mauritius, processes of the premaxillae; the orbits are relatively small; the
and there is no nomenclatural or systematic confusion for ventral surface of the endocranial cavity forms an acute angle
said species. Should fossil raphinan remains be discovered in with the basioccipital plane; the palatines are deeply ventrally
Mauritius, it might be worth revisiting the typification of the curved; the cervical vertebrae are comparatively large relative
Dodo. to volant columbids; the modal number of synsacral verte-
brae is 17; the modal number of caudal vertebrae (excluding
Locality the pygostyle) is six; the pectoral girdle and limbs are reduced
relative to volant columbids; the sternum sulcus articularis
Island of Mauritius, Mascarene Islands. Janoo (2005: 172) stated
coracoideus is comparatively shallow and laterally located;
that the type locality was ‘Type locality: Mauritius, Plaisance,
the coracoid processus lateralis is well developed, curved, and
Grand Port (Lydekker, 1891); Mare aux Songes (Lambrecht,
pointed; the scapula blade is typically curved, especially dis-
1933)’. However, no further information is presented, nor was a
tally; the scapula has a distinct tubercle on the cranioventral
type specimen mentioned.
edge of the blade; the furcula is U-shaped distally; the prox-
imal end of the humerus is displaced ventrally relative to the
Etymology axis of the shaft; the humerus incisura capitis is comparatively
‘Hooded Dodo’. The specific epithet, cucullatus, is from the Late short, broad and rounded; the humerus crista deltopectoralis
Latin for hooded ( Jobling 2010: 124). The first usage of cucullatus is variable in morphology; the distal ends of the radii and ulnae
for the Dodo, was as ‘Cygnus cuculatus’ by Nieremberg (1635), lack exostosis; the carpometacarpus processus extensorius
in pre-Linnean literature (hence why the binomial is not written is more block-like than in extant (volant) columbids; the
in italics). Note, Nieremberg (1635) used one ‘l’ (= cuculatus) carpometacarpus and the metacarpals majus and minus are
not two (= cucullatus) as later authors did. comparatively long, slender, and cylindrical, although in
some specimens the metacarpal minus is somewhat flattened;
carpometacarpus distal end is not deflected cranially relative
Geographical range
to the long axes of the metacarpals; the ilia are comparatively
Island of Mauritius, Mascarene Islands. Extinct since the 17th broad, especially caudal to the acetabula; the hindlimb bones,
century, although precisely when is still subject to debate (see relative to the shaft widths, are comparatively short and thick;
Roberts and Sollow 2003, Hume et al. 2004, Mlíkovský 2004, the femur crista trochanteris has a pneumatic foramen on the
Cheke 2006, Cheke and Hume 2008, Roberts 2013, Cheke and craniomedial edge; the femur linea intermuscularis caudalis is
Parish 2020). The IUCN Red List of Threatened Species has distinct but not conspicuously raised; and the tarsometatarsus
1662 as the year last seen, with individuals considered to have crista medialis hypotarsi is comparatively large. (The preceding
been the last of the species killed on the islet of île d’Ambre in characters were modified from table 3 of Livezey 1993, which
1662 (BirdLife International 2016a). compares the Dodo with the Solitaire.) The horizontal semi-
circular canal has a pronounced concavity at the mid-point of
Referred specimens
the canal. (Based on the endocranial reconstructions in Gold et
See Parish (2015) for a list of specimens in museum collections. al. 2016, comparing the Dodo with volant columbids, note that
Note that the specimens sent to institutions in Australia and the inner ears of the Solitaire are unknown.)
New Zealand are not listed.
26 • Young et al.
†Pezophaps Strickland, 1848 • 1872 Pezophaps minor Strickland—Owen, p. 519, plate

Type species LXVI (figs 5–7, 12).
• 1872 Pezophaps solitaria (Gmelin)—Owen, p. 519, plate
†Didus solitarius Gmelin, 1789. Type species by monotypy
LXVI (figs 1–4, 13).
(contra Peters 1937: 10, who stated that the species was the type
• 1878 Pezophaps solitaria (Gmelin)—Owen, p. 87–97,
by original designation; Strickland did not state that †Didus
plates VII and VIII.
solitarius was the type species of his new genus). Strickland
• 1879 Pezophaps solitaria (Gmelin)—Newton and Clark,
(1848: 69) stated: ‘Leguat’s bird is the type of the “Didus
p. 438–451, plates XLIV–L.
solitarius” of systematists’; there was no mention of generic type
• 1891 Pezophaps solitaria (Gmelin)—Lydekker, p. 128.
species.
• 1893 Pezophaps solitarius [sic] (Gmelin)—Sharpe,
Following recommendation 51G of the Zoological Code, the
p. 629–632.
type species can be referred to as †Pezophaps solitaria (Gmelin,
• 1907 Pezophaps solitarius [sic] (Gmelin)—Rothschild, p.

1789) Strickland, 1848. It can also be written more simply as
177–178, plates 23 and 25a (figs 1–3).
†Pezophaps solitaria (Gmelin, 1789).
• 1916 Pezophaps solitarius [sic] (Gmelin)—Ridgway, p.
Etymology 277.
• 1937 Pezophaps solitaria (Gmelin)—Peters, p. 10.
‘Pedestrian pigeon’, based on the Greek pezós (πεζός, ‘ped-
• 1957 Pezophaps solitaria (Gmelin)—Verheyen, p. 9, 18,
estrian’) and phaps (φάψ, ‘a pigeon’) (from Strickland 1848:
21, 27, 30.
54). The genus is of feminine grammatical gender, hence why
• 1971 Pezophaps solitaria (Gmelin)—Brodkorb, p. 200.
Strickland (1848: 46) declined the suffix of the specific epithet
• 1975 Pezophaps solitarius [sic] (Gmelin)—Wolters, p. 43.
solitarius (-us) to solitaria (-a).
• 1989 Pezophaps solitaria (Gmelin)—Brom and Prins,
p. 233, 236.
Diagnosis • 1990 Pezophaps solitaria (Gmelin)—Sibley and Ahquist,
Same as the only species (see below). p. 421.
†Pezophaps solitaria (Gmelin, 1789) • 1991 Pezophaps solitaria (Gmelin)—Witmer and Cheke,
p. 134.
(Figs 2, 5, 7–9) • 1993 Pezophaps solitaria (Gmelin)—Livezey, p. 247–
282, fig. 2b.
• 1708 le Solitaire (pre-Linnean)—Leguat p. 98–104. • 1996 Pezophaps solitaria (Gmelin)—Janoo, p. 57, 58, 67,
Figured on the page adjacent to p. 98. 69, 72, fig. 7.
• 1785 Solitary Dodo—Latham, p. 3–4. • 2000 Pezophaps solitaria (Gmelin)—Janoo, p. 323, 325–
• 1789 Didus solitarius sp. nov.—Gmelin, p. 728–729. 327, fig. 1.
• 1790 Didus solitarius Gmelin—Latham, p. 662–663. • 2001 Pezophaps solitaria (Gmelin)—Worthy, p. 766.
• 1844 Didus solitarius Gmelin—Strickland, p. 324–327. • 2002 Pezophaps solitaria (Gmelin)—Shapiro et al., p.
• 1848 Pezophaps solitaria (Gmelin) gen. nov. et just. 1683.
emend.—Strickland, p. 46–56, plates XIII–XV. • 2005 Pezophaps solitaria (Gmelin)—Janoo, p. 177.
• 1848 Pezophaps solitaria (Gmelin)—Melville, p. 113– • 2006 Pezophaps solitaria (Gmelin)—Cheke, p. 156.
119, plates XIII–XV. • 2007 Pezophaps solitaria (Gmelin)—Pereira et al., p.
• 1852 Pezophaps solitaria (Gmelin)—Mitchell, p. 19. 656–659, 662, 663, 665, 667, 669.
• 1852 Pezophaps minor [nomen nudum] Strickland— • 2008 Pezophaps solitaria (Gmelin)—Cheke and Hume,
Mitchell, p. 19. p. 30, 50, 167, 398, plate.
• 1854 Didus nazarenus Gmelin—Bartlett, p. 280–284, • 2013 Pezophaps solitaria (Gmelin)—Parish, [numerous
plate XLV (fig. 1). [Published in 1854.] occurrences].
• 1854 Didus solitarius Gmelin—Bartlett, p. 280–284, • 2014 Pezophaps solitaria (Gmelin)—Heupink et al., p. 1,
plate XLV (fig. 2). [Published in 1854.] 3.
• 1854b Pezophaps solitaria (Gmelin)—Bonaparte, p. 3. • 2014 Pezophaps solitaria (Gmelin)—van Grouw, p.
• 1859 Pezophaps solitaria (Gmelin)—Strickland, p. 187– 300.
196, plate LV (figs 1–4). • 2015 Pezophaps solitaria (Gmelin)—Claessons et al., p.
• 1859 Pezophaps minor sp. nov.—Strickland, p. 187–196, 31, 36, 184, 185–187.
plate LV (figs 5–7). • 2016 Pezophaps solitaria (Gmelin)—Besnard et al., p.
• 1862 Pezophaps solitaria (Gmelin)—Reichenbach, p. 76–78.
128. • 2016 Pezophaps solitaria (Gmelin)—Byrkjedal et al., p.
• 1865 Didus nazarenus Gmelin—Newton, p. 199–201, 199.
plate VIII. • 2016 Pezophaps solitaria (Gmelin)—Gold et al., p. 951–
• 1868 Pezophaps solitaria (Gmelin)—Newton and 957, 959–961, figs 2B, 3B, 4B, 7B, 8B.
Newton, p. 428–433. • 2016 Pezophaps solitaria (Gmelin)—Rodríguez-Pontes,
• 1869 Pezophaps solitaria (Gmelin)—Newton and p. 398–413, figs 1–11.
Newton, p. 327 – 359, plates 15–22, 24. • 2016 Pezophaps solitaria (Gmelin)—Soares et al., p. 1–3, 5.

Figure 7. Solitaire (†Pezophaps solitaria) referred specimen (NHMUK PV A 3505). Skull in left lateral view (A), occipital view (B), right
lateral view (C), dorsal view (D), and ventral view (E). Note that this is a male individual. Scale bar: 3 cm.
• 2017 Pezophaps solitaria (Gmelin)—Angst et al., p. 1, 4. of Réunion (also part of the Mascarene Islands). This species, the
• 2018 Pezophaps solitaria (Gmelin)—Bruxaux et al., p. Réunion or White Dodo, is, in fact, the extinct quasi-flightless
250, 254. ibis †Threskiornis solitarius (de Sélys-Longchamps, 1848) (see
Hume and Cheke 2004).
Type specimen
No type specimen has been designated for †Pezophaps solitaria Geographical range
(the Rodrigues Solitaire, or simply the Solitaire). As noted by Island of Rodrigues, Mascarene Islands. Extinct since the mid-
Strickland (1848: 56) the type series of †Pezophaps solitaria are 18th century (see Cheke and Hume 2008). The IUCN Red List
the unknown individuals that François Leguat observed while of Threatened Species has 1770 as the year last seen, and states
on the island of Rodrigues. Given that they were live animals ob- that the species was reported in 1761 but was extinct by 1778
served in their natural habitat, it is impossible to ascertain what (BirdLife International 2016b). Pingré, who visited Rodrigues in
specimens they were and whether they were ever collected. As 1761, did not see any Solitaires but was informed that they were
such, we cannot locate the original syntypes, and this precludes still alive (Parish 2013: 65).
us from designating a lectotype.
At present, we cannot formally designate a neotype for Referred specimens
†Pezophaps solitaria. In order to be fully compliant with Article See Parish (2015) for a list of specimens in museum collections.
75 of the Zoological Code, a neotype designation cannot be Note that the specimens sent to institutions in Australia and
‘an end in itself, or a matter of curatorial routine, and any such New Zealand are not listed.
neotype designation is invalid’ (Article 75.2), and ‘there is an
exceptional need’ (Article 75.3). There is no exceptional need Diagnosis
for a neotype for the Solitaire. As we have shown above, only Columbids with the following unique combination of charac-
one raphinan species is known from the Island of Rodrigues, ters: pronounced sexual dimorphism of the skeleton; at their
and there is no nomenclatural or systematic confusion for proximal end, the frontal processes (= processus frontales) of
said species. Should fossil raphinan remains be discovered in the premaxilla are separated by a narrow space of variable size;
Rodrigues, it might be worth revisiting the typification of the distally, the premaxillary frontal processes are not enlarged,
Solitaire. fusing more distally; the caudal margin of the corpus ossis
premaxillare has plate-like lateral extensions that result in the
Locality nares (= apertura nasalis ossea) being subtriangular; the upper
Island of Rodrigues, Mascarene Islands. bill tip is typically hooked; the maxillary process (= processus
maxillaris) of the premaxilla tapers distally to slender splints;
Etymology the premaxilla distal ventral surfaces typically have numerous
‘The solitary pedestrian pigeon’. The specific epithet is based neurovascular foramina in distinct depressions; the distal end of
on François Leguat’s name, ‘le Solitaire’. As noted by Strickland the lower jaw is comparatively shallow and weak; the proximal
(1848: 46), Leguat re-used the name ‘Solitaire’, which had ori- end of the lower jaw is comparatively deep and caudally abbre-
ginally been used to denote a different bird species on the island viated, with perpendicular posterior margins of the rami; the
28 • Young et al.

Figure 8. Solitaire (†Pezophaps solitaria) referred specimen (NHMUK PV A 3506). Skull in left lateral view (A), occipital view (B), right
lateral view (C), dorsal view (D), and ventral view (E). Note that this is a female individual. Scale bar: 3 cm.
Figure 9. Artistic life reconstructions of the solitaire (†Pezophaps solitaria). A, a solitary male. B, two males engaged in intraspecific conflict.
Artwork by Julian Pender Hume.
quadrate–lower jaw articulation is not situated comparatively Oliver et al. 2023), we undertook a historical review of columbid
posteriorly on the cranium; the lacrimal processes of the frontal family-group nomina. Although most of the columbid family-
are comparatively large; the frontal is dorsally flat; the frontals group nomina are valid under the Zoological Code, we found
are truncated anteromedially, leaving the posteromedial por- that the Trugon- and Hemiphaga-based nomina do not fulfil
tions of the nasals and frontal processes of the premaxillae ex- Article 13.2.1 of the Zoological Code and are therefore not
posed dorsally; the orbits are relatively large; the ventral surface available. Given that most of the family-group nomina within
of the endocranial cavity is nearly parallel with the basioccipital Columbidae were established prior to 1930 and that most of the
plane; the palatines are moderately ventrally curved; the cer- nomina in use are compliant with the Principal of Coordination
vical vertebrae are comparatively small; the modal number of for the family group (Article 36), they are not required to be ac-
synsacral vertebrae is 17 or 18; the modal number of caudal companied by a list of characters that purport to differentiate it
vertebrae (excluding the pygostyle) is five; the pectoral girdle from other taxa (Article 13.1.1) or by a bibliographic reference
and limbs are reduced relative to volant columbids; the sternum to such a published statement (Article 13.1.2). However, if a

sulcus articularis coracoideus is comparatively deep and medi- new family group were to be established, any new nomina would
ally located; the coracoid processus lateralis is reduced, with the need to be accompanied by a list of diagnostic characters.
lateral margin truncated; the scapula blade is typically straight; Our review also found that Oudemans (1917), not Poche
the scapula lacks a distinct tubercle on the cranioventral edge (1904) or Wetmore (1930), is the correct nominal authority of
of the blade; the furcula is essentially parabolic distally; the hu- the †Raphus-based family group. We cannot preclude the pos-
merus proximal end is not displaced ventrally relative to the sibility that an older undiscovered publication might alter the
axis of the shaft; the humerus incisura capitis is comparatively nominal authority again; this is owing to the early 20th cen-
long, with a narrow groove; the humerus crista deltopectoralis tury shift from using the genus †Didus and its family group, and
is rounded without a dorsal point; in males, the distal ends of the specific epithet †ineptus for the Dodo, to using the genus
the radii and ulnae variably are exostotic; presence of an out- †Raphus and specific epithet †cucullatus instead (Poche 1904,
growth of the processus extensorius of the carpometacarpus Rothschild 1907, Allen 1910, Ridgway 1916, Oudemans 1917).
(carpal knob sensu Hume and Steel 2013), and these exo- This shift and the nomenclatural impact it had are intriguing.
stoses can become greatly enlarged in males but can also be The cause is a simple one: the change in the ‘starting date’ of zoo-
present in females, albeit in a much more reduced manner; logical nomenclature. As we outlined above, during the 19th
the carpometacarpus and metacarpals majus and minus, are century different nomenclatural codes used different editions of
comparatively short, flat, and rectangular in cross-section; the Systema Naturæ as the ‘start’ of zoological nomenclature, with
carpometacarpus distal end is deflected cranially relative to the some codes preferring the 12th edition (e.g. Strickland et al.
long axes of the metacarpals; the ilia are comparatively narrow, 1843) and others the 10th edition (American Ornithologists’
especially caudal to the acetabula; the hindlimb bones, rela- Union 1886, Blanchard 1889). By the early 20th century, the
tive to the shaft widths, are comparatively long and slender; international community of zoologists (ICZN 1905) had settled
the femur crista trochanteris lacks a pneumatic foramen on the on using the 10th edition of Systema Naturæ (Linnaeus 1758)
craniomedial edge; the femur linea intermuscularis caudalis is and a date of 1758. It was this act that impacted Dodo nomen-
comparatively raised and more prominent; the tarsometatarsus clature so profoundly. By accepting the 10th edition of Systema
crista medialis hypotarsi is comparatively small. (The preceding Naturæ (Linnaeus 1758) as available for zoological nomencla-
characters were taken from table 3 of Livezey 1993, which com- ture, and a ‘starting date’ of 1758, it rendered both †Struthio
pares the Solitaire with the Dodo.) cucullatus Linnaeus, 1758 and †Raphus raphus Brisson, 1760a, b
as available nomina.
DISCUSSION This situation is an example of what we call herein nomen-
clatural ‘ripples’. A seemingly small change to a nomenclatural
Nomenclatural ‘ripples’ code, or a change in which nomenclatural code is in prevailing
Our overview of the nomenclatural and systematic history of the use, can ‘ripple’ outwards, impacting the validity of nomina (spe-
Dodo and Solitaire has shown that even iconic species can lack cific, generic, and family group) and which subclade names are
type specimens. As we have demonstrated, this ‘oversight’ is not available to systematists when overarching analyses of the in-
as surprising at it might first appear. Both species were extinct ternal relationships of a clade are undertaken. The fourth edition
by the time their scientific nomina were established (Linneaus of the Zoological Code (ICZN 1999) does have many caveats
1758, Gmelin 1789), and their nomina were not based on phys- in place to prevent threats to nomenclatural stability, such as
ical specimens but on written accounts of unknown live indi- in the Dodo example herein. But this example should be used
viduals and lost specimens (Clusius 1605, Nieremberg 1635, as a cautionary tale for systematists and those formulating/re-
Bontius 1658, Leguat 1708). Both species had begun to fall into vising nomenclatural codes. Although the impact we described
myth by the 18th century (Duncan 1828, de Blainville 1835, herein is restricted to an extinct species, how easily could future
Strickland 1844, 1848, Hume 2006), and the concept of the changes inadvertently impact the nomenclature of an endan-
nomenclatural type did not exist until the early 19th century gered species and thereby detrimentally affect conservation or
(Westwood 1837a, Witteveen 2016). law-enforcement efforts?
Given the recent use of nomina based on the †Raphus Alas, too many zoologists/systematists who are not ‘taxo-
family group by extant columbid workers for subclades within nomically’ or ‘nomenclaturally’ inclined dismiss the impact that
Columbidae (Dickinson and Remsen Jr 2013, Chen et al. 2022, even relatively minor changes to nomenclatural codes can have.
30 • Young et al.
For example, although well intentioned, the idea we can sweep of a secondarily terrestrial morphotype evolving from island-
away problematic nomina established during the age of coloni- dwelling raphinans: the †Raphus and †Pezophaps clade (which
alism (see the discussion and wider references in the works by: is closely related to the extant genus Caloenas) and †Natunaornis
Bae et al. 2023, Ceríaco et al. 2023, Cheng et al. 2023, Guedes (which might be closely related to the extant genus Goura).
et al. 2023, Jost et al. 2023, Orr et al. 2023, Pethiyagoda 2023, Another large-bodied columbid presumed to be secondarily
Raposo et al. 2023) without causing nomenclatural instability is terrestrial is the Saint Helena Dove †Dysmoropelia dekarchiskos
naïve. Although upsetting nomenclatural stability is not an ex- Olson, 1975, from the island of Saint Helena, in the South
cuse to do nothing, we must strive: (i) not to introduce unneces- Atlantic Ocean. Lewis (2008) dated Sugar Loaf site 1, from which
sary instability to our quasi-legal nomenclatural codes, because †Dysmoropelia dekarchiskos is known, to ~14 000 before present
they underpin both national and international wildlife laws and (i.e. late Pleistocene). The species is known from forelimb and
regulations (especially for endangered species); and (ii) to have hindlimb elements and the coracoid. Similar to †Natunaornis,
a reasonable idea about the potential impact that any proposed the forelimb and pectoral elements of †Dysmoropelia Olson,

changes to the Zoological Code could have. That is the only way 1975 are extremely reduced in size relative to the hindlimb
we can even attempt to ameliorate adverse effects (e.g. as the bones, with Olson (1975: 30) noting: ‘No other known
ICZN did with their amendments to the fourth edition of the columbiform bird besides the Dodos and Solitaires has evolved
Zoological Code to make nomenclatural acts published online anything like an equivalent condition’. Olson (1975: 31) posited
available and valid, ICZN 2012). a Streptopelia-like ancestor for †Dysmoropelia dekarchiskos but
Dear reader, if you have made it this far through our contri- stressed that ‘the differences between the two genera are great
bution, you know that addressing nomenclatural ‘ripples’ and and any relationship between them must be considered only
other ‘taxonomic tangles’ is neither quick nor easy; as you can hypothetical’. If †Dysmoropelia is closely related to Streptopelia,
imagine, they are time intensive (not only examining specimens then it is evidence that secondarily terrestrial morphotypes
first hand but reading through >400 years of relevant papers, evolved outside of Raphinae.
in our instance). If, as a community, we are going to address Within the raphine tribe Phabini, there are extinct species
problematic nomina, taxonomic vandalism, ‘nomenclatural referred to the genus Pampusana that were either larger bodied
harvesting’ (sensu Denzer and Kaiser 2023), and other instances or, possibly, secondarily terrestrial. These include †Pampusana
of using nomenclature as a weapon, there is a need for more longitarsus (Balouet & Olson, 1989) from New Caledonia and
trained taxonomists/systematists and more funding to support the possibly flightless species †Pampusana leonpascoi (Worthy
descriptive biology (see Boero 2010, Britz et al. 2020, Engel et & Wragg, 2003). Within the subfamily Columbinae, the ex-
al. 2021). Before we can elucidate proximate and ultimate causal tinct Mauritian Turtle Dove, †Nesoenas cicur Hume, 2011,
relationships, we must first make observation statements on is considered to be a terrestrial granivore (Hume 2011).
semaphoronts (which, in the 21st century, includes data gleaned Unfortunately, the lack of total-evidence phylogenetic analyses
from computed tomography datasets and DNA sequence align- hampers our understanding of the evolutionary relationships
ments), formulate hypotheses based on our observations (e.g. of extinct island-dwelling columbids. It also precludes us from
are the semaphoronts in front of me with ‘X’ and ‘Y’ unusual determining whether the flightless columbids †Natunaornis
characteristics a new species?), and if necessary, attach labels/ gigoura, †Dysmoropelia dekarchiskos, and †Pampusana longitarsus
nomina to those hypotheses (zoological nomenclature). To are closely related to †Raphina or are instances of convergent
quote Mayr (1982: 70): ‘It is sometimes overlooked how essen- evolution. It is our intention to incorporate osteology and soft-
tial a component in the methodology of evolutionary biology tissue characters alongside molecular data in future studies in
the underlying descriptive work is’. order that the relationships of subfossil, and older fossil, species
can be elucidated.
Island-dwelling and terrestrial columbids
The Mascarene giant ground pigeons (†Raphina subtribus
nova) are not the only island-dwelling columbids that evolved CONCLUSION
to become secondarily terrestrial. The Fiji Giant Ground Pigeon The Dodo (†Raphus cucullatus) is an icon of conservation and
†Natunaornis gigoura Worthy, 2001 is another notable ex- a reminder of the destructive potential of humanity towards
ample. †Natunaornis is known from cave deposits on the island the natural world. Along with its sister species, the Solitaire
of Viti Levu of unknown geological age, but the specimens (†Pezophaps solitaria), these aberrant columbids were named
are presumed to be from the Holocene (Worthy 2001: 767). after their extinction but before the development of the ‘no-
The species is known from forelimb and hindlimb elements, menclatural type’ (i.e. establishment of voucher specimens).
the coracoid and scapula, and possibly, an isolated premaxilla. Therefore, no type specimens have been designated previously
Although incompletely known, †Natunaornis is the third largest for these iconic species. Given that there is no exceptional need
known columbid, behind †Raphus and †Pezophaps, and, based for neotype designations for the Dodo or the Solitaire, we refrain
on the extreme size difference between the fore- and hindlimbs, from making them. We did, however, establish a new family-
it is thought to have been flightless. Based on the osteological group nomen, †Raphina subtribus nova, to unite them.
similarities between †Natunaornis gigoura and the extant genus Our historical overview of the Dodo and the Solitaire, and
Goura, Worthy (2001) hypothesized that they were closely re- the family-group nomina within Columbidae, has shown that
lated. However, this hypothesis has never been subjected to changing nomenclatural codes have had a profound impact
phylogenetic analysis. If supported, there would be two instances on Dodo nomenclature. During the 19th century, when the
first nomenclatural codes were being developed, the proposed AU THOR CON TIBU TIONS
‘starting date’ of zoological nomenclature was based on the 12th
NJG and MOH obtained funding, MTY led the project, conceived the
edition of Systema Naturæ (Linnaeus 1766), using 1766 as the
study and wrote the manuscript, JPH and NJG helped develop the pro-
cut-off date for valid nomenclatural acts. This is why the genus ject, all authors helped edit drafts and approved the final version to be
†Didus was consistently preferred over †Raphus, why the specific published.
epithet †ineptus was used over †cucullatus, and why the family
group was †Dididae/Didinae. With the development and even-
tual publication of the Règles Internationales (ICZN 1905), the FUNDING
corpus of international zoologists adopted the 10th edition of We thank the Institute for Life Sciences, University of Southampton for
Systema Naturæ (Linnaeus 1758) as the ‘starting date’ of zoo- funding this work. The funders had no role in study design, data collec-
logical nomenclature, thereby shifting the date back to 1758. tion and analysis, decision to publish, or preparation of the manuscript.
This explains the shift in Dodo nomenclature during early 20th

century (e.g. Poche 1904, Rothschild 1907, Allen 1910, Ridgway DATA AVAIL ABILIT Y STATE MEN T
1916, Oudemans 1917).
We are again entering a period of multiple active nomencla- All data are incorporated into the article and its online supplementary
tural codes in zoology, with the development of the fifth edition material.
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Zoological Journal of the Linnean Society, 2024, 201, zlae086

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Received 5 April 2024; revised 31 May 2024; accepted 14 June 2024

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Stem Family Subfamily Below subfamilya

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Subfamily Columbinae Illiger, 1811 Tribe Zenaidini Bonaparte, 1853

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Nomenclatural note ZooBank registration number [urn:lsid: zoobank.org

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Type specimen Diagnosis

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†Pezophaps Strickland, 1848 • 1872 Pezophaps minor Strickland—Owen, p. 519, plate

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https://archive.org/details/returnbritishmus56brit/page/n479/ ICZN [International Commission on Zoological Nomenclature].

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