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A LArge Fruit Pigeon (CoLumbidAe) From the eArLy
mioCene oF new ZeALAnd
R.
—Describimos un nuevo género y especie de paloma (Columbiformes) de un único coracoides de la fauna St Bathans de
Nueva Zelandia (– millones de años antes del presente). Es la primera especie de paloma descrita de depósitos pre-Pliocénicos en
Australasia. Dos apomorfías identifican el fósil como perteneciente al grupo ptilinopino de las palomas fruteras, entre las cuales es más
parecida a Hemiphaga, la gran paloma frutera actualmente endémica del área biogeográfica de Nueva Zelandia. Esto revela que el linaje
de Hemiphaga ha estado en Nueva Zelandia desde el Mioceno Temprano, lo cual avala estimados recientes de fechas de divergencia para
Hemiphaga y su taxón hermano moderno (Lopholaimus) basados en datos moleculares.
—
649
—
e Auk 126(3):649−656, 2009
e American Ornithologists’ Union, 2009.
Printed in USA.
e Auk, Vol. , Number , pages −. ISSN -, electronic ISSN -. by e American Ornithologists’ Union. All rights reserved. Please direct
all requests for permission to photocopy or reproduce article content through the University of California Press’s Rights and Permissions website, http://www.ucpressjournals.
com/reprintInfo.asp. DOI: ./auk..
Una Gran Paloma Frutera (Columbidae) del Mioceno Temprano de Nueva Zelandia
Tr e v o r H. Wo r T H y ,
1,2,5
Su z a n n e J. Ha n d ,
1
Je n n i f e r P. Wo r T H y ,
1
a
l a n J. d. Te n n y S o n ,
3
a n d r. Pa u l Sc o f i e l d
4
1
School of Biological, Earth and Environmental Sciences, University of New South Wales, New South Wales 2052, Australia;
2
Department of Earth and Environmental Sciences, University of Adelaide, Darling Building, DP 418, Adelaide, South Australia 5005, Australia;
3
Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand; and
4
Canterbury Museum, Rolleston Avenue, Christchurch 8001, New Zealand
5
E-mail: t.worthy@unsw.edu.au
A.—We describe a new genus and species of pigeon (Columbiformes) from a single coracoid from the St Bathans Fauna
of New Zealand (– mya). It is the first columbid species described from pre-Pliocene deposits in Australasia. Two apomorphies
identify the fossil as belonging to the ptilinopine group of fruit pigeons, among which it is most similar to Hemiphaga, the large fruit
pigeon currently endemic to the New Zealand biogeographic area. is reveals that the Hemiphaga lineage has been in New Zealand
since the Early Miocene, which supports recent divergence-date estimates for Hemiphaga and its modern sister taxon (Lopholaimus)
based on molecular data. Received November , accepted March .
Key words:
Columbidae, Hemiphaga, Miocene fossil coracoid, New Zealand, Rupephaps.
T
pigeons (Aves: Columbidae) in Australasia
is considerable, with ~ indigenous species (including three
recently extinct) in genera (Higgins and Davies , Hold-
away et al. , Christidis and Boles ). In New Zealand,
columbids are represented by the single genus Hemiphaga with
allopatric species on the Chatham Islands, Norfolk Island, and
mainland New Zealand (Turbott , Higgins and Davies ,
Holdaway et al. , Checklist Committee [O.S.N.Z.] ).
e fossil history of Hemiphaga is restricted to Late Pleisto-
cene and Holocene deposits in New Zealand (Worthy and Hold-
away ). e single exception is a distal ulna reported from
the Pliocene Whenuataru Tuff in the Chatham Islands by Eagle
et al. (). It is identical to Hemiphaga chathamensis, and its
preservation is very different (no infilled sediment, no wear, no
encrusting marine calcareous structures) from that of the other
bone described from the same deposit. Its preservation is typical
of Holocene bones from other sites on the islands (T. H. Worthy
and A. J. D. Tennyson pers. obs.), and it seems likely that this
specimen was an unrecognized Holocene intrusion in the Plio-
cene deposit.
e pre-Quaternary fossil record of columbiforms is mea-
ger compared with the fossil records of many birds (Olson ,
Becker and Brodkorb ). ere is no Paleogene record of spe-
cies referable to Columbidae in Europe (Mayr ). e putative
columbid Microena goodwini from the Early Eocene of England
(Harrison and Walker ) was relegated to Aves incertae sedis
(Mlíkovský ). e order Columbiformes is represented, how-
ever, by four species in two genera (Archaeoganga and Leptoganga)
of sandgrouse (Pteroclidae) from the Late Eocene–Oligocene and
Early Miocene of France (Olson , Mlíkovský ). Because
Columba calcaria Milne-Edwards, –, from the Lower
Miocene at Saint-Gérand-le-Puy in France, is now also considered
a sandgrouse, as Gerandia calcaria (Mlíkovský ), there is no
pre-Pliocene columbid record in Europe.
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Globally, the oldest described pigeons are known from abun-
dant fossils from the Early Miocene (– mya) of Florida (Olson
) referred to a modern dove genus, as Columbina prattae, by
Becker and Brodkorb (). However, Steadman () erected
a new genus, Arenicolumba, for this taxon, noting a greater phe-
netic similarity to the Old World Oena and Turtur than to any
New World taxa, but generic comparisons were limited.
Olson () suggested that columbids may have originated in
the Southern Hemisphere, but as yet there is little fossil evidence
to support this proposal. Pigeons and doves have a fossil record in
Australia extending back to the Late Oligocene–Early Miocene,
from lacustrine deposits at Lake Palankarinna (Etadunna For-
mation) and Lake Pinpa (Namba Formation) of central Australia
(Vickers-Rich , Boles ), but the material is meager and
undescribed. Similarly, the New Zealand St Bathans Fauna has re-
vealed a single fossil of a columbid, a distal right ulna of a bird
about the size of the Wompoo Fruit-dove (Ptilinopus magnificus;
Worthy et al. ).
e St Bathans Fauna is derived from sediments deposited
in a shallow freshwater lake ~, km
in area and is of late
Early Miocene age (– mya; Worthy et al. ). e faunal-
bearing sediments outcrop as the lower Bannockburn Formation
of the Manuherikia Group, in Otago, South Island, New Zea-
land. At the time of deposition, the climate was warm and the
surrounding vegetation included casuarinas (Casuarinaceae),
Eucalyptus (Myrtaceae), and palms (Arecaceae), in addition to
the typical “New Zealand” podocarps (Podocarpaceae), Nothofa-
gus, and araucarias (Araucariaceae) (Pole and Douglas , Pole
et al. ). is fauna provides the only data on the terrestrial
vertebrates that existed in New Zealand during the Tertiary and,
so far, has revealed ~ avian taxa. e avifauna of the St Ba-
thans Fauna is dominated by anseriforms, with a minimum of
eight taxa in five genera, but has a diverse fauna of other birds,
including procellariiforms, accipitriforms, gruiforms, a palael-
odid, charadriiforms, columbiforms, psittaciforms, apodiforms
including both swiftlets and owlet nightjars, and passeriforms
including a possible cracticid (Worthy et al. , ; T. H.
Worthy et al. unpubl. data). Here, we add to this diversity from
the St Bathans Fauna and, on the basis of a coracoid, describe the
first pre-Quaternary pigeon species from Australasia. We draw
phylogenetic inferences of its relationships and discuss its bio-
geographic implications.
Me t h o d s
Several molecular analyses have examined phylogenetic relation-
ships within Columbidae (e.g., Johnson and Clayton , Shapiro
et al. ), but the most comprehensive is that of Pereira et al.
(), which used >, base pairs from multiple nuclear and
mitochondrial genes from pigeon genera. We used this phylo-
genetic hypothesis and the distribution of morphological charac-
ters to assess the relationship of the fossil described here.
e osteology of the columbid coracoid was the subject of a
detailed description (Martin ), but this work outlined few in-
tergeneric distinguishing features. More recently, Worthy ()
detailed coracoid characters useful for generic identification, and
these were expanded by Worthy and Wragg (, ) and by
Steadman (). Our comparisons were mainly restricted to taxa
in the Australasian–Pacific region; because of geographic separa-
tion, taxa outside this region were considered unlikely relatives.
Abbreviations.—AM = Australian Museum, Sydney, Austra-
lia; AMNH = American Museum of Natural History, New York;
CM = Canterbury Museum, Christchurch, New Zealand; FM =
Fiji Museum, Suva, Fiji; LACM = Natural History Museum, Los
Angeles County, California; MVZ = Museum of Vertebrate Zool-
ogy, University of California, Berkeley; NMNZ = Museum of New
Zealand Te Papa Tongarewa, Wellington; SAM = South Australian
Museum, Adelaide, South Australia; USNM = U.S. National Mu-
seum, Smithsonian Institution, Washington, D.C.; and UWBM =
Burke Museum, University of Washington, Seattle.
Anatomical nomenclature.—Names for specific bone land-
marks follow Baumel and Witmer (). Some common terms
are abbreviated: artic. = articularis, m. = musculus, and proc. =
processus.
Measurements were made with Tesa dial calipers and rounded
to . mm. e fossil was compared with the following taxa and
specimens, with an emphasis on Australasian–Southwest Pacific
taxa. All material is from modern skeletons unless stated other-
wise; extinct taxa are indicated by a cross. †Bountyphaps obsoleta,
MNZ S. (fossil bones). Caloenas nicobarica, SAM B.,
SAM B., SAM B.. Otidiphaps nobilis, USNM .
Gallicolumba stairi, NMNZ S. ( fossil bones, individ-
uals). G. beccarii, AMNH , USNM , UWBM .
G. rubescens, MVZ . G. luzonica, MVZ , LACM ,
LACM . G. rufigula, LACM . G. criniger, LACM
. G. jobiensis, AM O., AMNH . Chalcophaps in-
dica, SAM B., SAM B., SAM B.. Columba leu-
comela, SAM B., AM O.. C. livia, SAM B., SAM
B., SAM B.. C. vitiensis, AM O., FM . Didun-
culus strigirostris, AM O.. Ducula bicolor, AM O., SAM
B. (as D. spilorrhoa). D. latrans, FM . D. pacifica, AM
O.. Geopelia cuneata, SAM B.. G. striata, SAM B..
Geophaps plumifera, SAM B., SAM B.. G. scripta, SAM
B.. G. smithii, SAM B.. Goura cristata, SAM B.,
SAM B., SAM B., SAM B.. Hemiphaga novaesee-
landiae, AM O.. Leucosarcia picata, SAM B. , SAM
B.. Lopholaimus antarcticus, AM O., AM O..
Macropygia amboinensis phasianella, SAM B., SAM
B . . Ocyphaps lophotes, SAM B. , SAM B., AM
B.. Phaps chalcoptera, SAM B., SAM B.. P. el-
egans, SAM B., SAM B.. Ptilinopus magnificus, SAM
B., SAM B.. P. magnificus keri, NMNZ OR..
P. regina, SAM B.. Streptopelia chinensis, SAM B.,
SAM B..
sy s t e M a t i c Pa l e o n t o l o g y
Columbiformes Latham,
Columbidae Illiger,
e fossil coracoid is assigned to Columbidae because it has the
following combination of features found only in this family: it is
elongate with a long shaft region; the proc. acrocoracoideus is large
and inflated, with a greater dorsoventral depth than that through
the facies artic. humeralis; the cotyla scapularis is rounded and
shallow, not a deep cup-like sulcus; the proc. procoracoideus lacks
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a foramen; and the impressio m. sternocoracoidei contains pneu-
matic foramina.
Rupephaps, gen. nov.
Type species.—Rupephaps taketake, sp. nov.
Diagnosis.—As for species.
Etymology.—Etymology: after Rupe, an honorific term for
the Kererū, or New Zealand Pigeon (H. novaeseelandiae) when
personified in Māori myths and a name widely used throughout
Polynesia for pigeons (Tregear , Best ), and phaps (Greek,
feminine noun for “wild pigeon”).
Rupephaps taketake, sp. nov.
Holotype.—NMNZ S., left coracoid (Fig. ), collected
January by the University of New South Wales–University of
Adelaide, South Australia–CM–NMNZ expedition.
Diagnosis.—A pigeon distinguished from all other taxa by
the following unique combination of characters. () e facies ar-
tic. clavicularis is planar, with a shallow notch at mid-depth, and
the ventral lobe extends farther sternally than the dorsal lobe.
() e ventromedial facies of the proc. acrocoracoideus, or that
ventrally of the facies artic. clavicularis, has two pits, interpreted
here as the insertion points for ligamentum acrocoraco-clavic-
ulare superficiale, the most ventral of which opens via a groove
to the ventral facies. () e facies artic. clavicularis extends as
a rounded lobe medially of the medial margin, rather than being
pointed and hooked sternally. And () the ridge extending crani-
ally along the medial margin above the angulus medialis does not
reach the facies artic. sternalis ventralis, being separated from it
and the medial angle by a rounded facies. e latter is an autapo-
morphy for the species.
Taxonomic remarks.—e fossil shares character with
Ptilinopus, Ducula, Lopholaimus, and Hemiphaga: the dorsal
and ventral lobes of the facies artic. clavicularis have about even
sternal extent and are separated by a marked notch in all other
compared taxa, except in Geophaps, in which the facies artic.
clavicularis is a medially convex structure without an obvious
dorsal lobe. e structure of the pit or pits for the insertion for
ligamentum acrocoraco-claviculare superficiale (figure . in
Baumel and Raikow ) on the ventromedial tip of the proc.
acrocoracoideus (character ) varies significantly among colum-
bids. e fossil shares with Ducula, Ptilinopus, Hemiphaga, and
Lopholaimus the conformation of two distinct pits separated by
a ridge, with the ventral-most pit opening to the ventral facies.
In many taxa (e.g., Columba, Leucosarcia, Streptopelia, Chal-
cophaps, Macropygia, Geophaps, Geopelia, and Caloenas), the
two pits are merged as a single elongate pit that opens ventrally,
whereas in Gallicolumba, Otidiphaps, Bountyphaps, Diduncu-
lus, Phaps, Ocyphaps, and Goura a single elongate pit is sepa-
rated by a ridge from the ventral facies. Both characters and
are consistent in allying Rupephaps with Ptilinopus, Ducula,
Lopholaimus, and Hemiphaga, to the exclusion of other taxa.
ese taxa, with the addition of Drepanoptila and Gymnophaps,
formed a strongly supported clade (Pereira et al. ), termed
the “fruit dove group” (Christidis and Boles ), for which
the name Ptilinopinae Selby, is available. We consider
characters and (as described above for Rupephaps) apomor-
phies for this clade.
Character , where the facies artic. clavicularis is a rounded
lobe extending medially of the sulcus m. supracoracoidei mar-
gin rather than being pointed and hooked sternally, was observed
only in Hemiphaga, Lopholaimus, and Natunaornis. We consider
its occurrence in the latter taxon to be non-homologous to the
state in Hemiphaga, a convergence that is probably the result of
reduction in pectoral structure associated with flightlessness in
this large species (Worthy ). Hemiphaga and Lopholaimus
are strong flyers. erefore, we suggest that this character is an
apomorphy for the well-established Hemiphaga + Lopholaimus +
Gymnophaps clade (Goodwin , Pereira et al. ). As such, it
associates Rupephaps with these large fruit pigeons to the exclu-
sion of other ptilinopines.
Character is an autapomorphy for the fossil taxon; in all
other taxa, the crest extends from the ventral sternal facet up the
medial margin.
Rupephaps is larger than all species in the genera Ducula and
Ptilinopus. It is slightly larger than Lopholaimus (– g) and
smaller than Hemiphaga (– g) and so is considerably larger
than the well-known Rock Pigeon (Columba livia; ~ g; Higgins
and Davies ).
In Rupephaps taketake, the impressio m. sternocoracoi-
dei extends to the medial margin of the bone and the facies ar-
tic. sternalis dorsalis has even breadth over the coracoid width,
which distinguishes it from Lopholaimus, in which the impres-
sio m. sternocoracoidei is separated from the medial margin
and the dorsal sternal facet is distinctly broader medially. Ru-
pephaps is distinguished from Hemiphaga novaeseelandiae by
two features apart from smaller size: a flat lateral facies above
the proc. lateralis and the form of the crest above the angulus
medialis.
Etymology.—From taketake, M
‒
aori for “long established, an-
cient, original” (Williams ), chosen to reflect the longevity of
the pigeon lineage in New Zealand. It is pronounced “ta-kay-ta-
kay.”
Type locality.—Bed HHa, a sand and mud cobble layer about
– cm thick with abundant bone and stromatolite fragments,
which is overlain by fine sand up to cm thick and is .–. m
above the base of the Bannockburn Formation. Home Hills Sta-
tion, true left side of Manuherikia River, Otago. °′.″S,
°′.′′E. New Zealand Fossil Record File Number
H/f.
Stratigraphy–age–fauna.—Bannockburn Formation, Manuher-
ikia Group, Early Miocene (Altonian); – mya; St Bathans Fauna.
Measurements of holotype.—Length from angulus media-
lis to tip of proc. acrocoracoideus = . mm; length from cotyla
scapularis to tip of proc. acrocoracoideus = . mm; shaft width =
. mm.
Description.—e fossil is stained black and is missing the
tip of the proc. procoracoideus, the proc. lateralis, a fragment of
the impressio m. sternocoracoidei, and a small section of the me-
dial margin of the shaft above the angulus medialis. It is slightly
crushed through the proc. acrocoracoideus in a zone extending
from above the facies artic. humeralis to below the facies artic.
clavicularis. In addition to the characters described above, the fos-
sil has the following features.
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Fig . 1. Left coracoids in (A) dorsal, (B) medial, (C) medial × 2, and ( D) ventral views of Rupephaps taketake NMNZ S. 51259 (top row) and Hemiphaga
novaeseelandiae AM O.65739 (bottom row). Abbreviations: ac = proc. acrocoracoideus, am = angulus medialis, cs = cotyla scapularis, fac = facies
artic. clavicularis, fah = facies artic. humeralis, fasv = facies artic. sternalis ventralis, pf = pneumatic foramina, pl = proc. lateralis, pr = proc. procora-
coideus, si = impressio m. sternocoracoidei, and ss = sulcus m. supracoracoidei. Scale bar = 1 cm.
e presence and size of pneumatic foramina in the sulcus
m. supracoracoidei below the dorsal lobe of the facies artic. clavic-
ularis differ among columbid taxa. e slight crushing through
the proc. acrocoracoideus precludes determining whether a small
pneumatic foramen was present under the dorsal lobe of the clav-
icle facet, as seen in Hemiphaga and Lopholaimus, but the area is
sufficiently well preserved to exclude the possibility that a large
pneumatic fossa as seen in Ptilinopus was present.
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e dorsal lobe of the facies artic. clavicularis is rounded and
protuberant over the sulcus m. supracoracoidei, but not so much
as to be prominent over the cotyla scapularis in lateral view. ere-
fore, it differs from Gallicolumba, Otidiphaps, Columba, Phaps,
Chalcophaps, Goura, Caloenas, Macropygia, Streptopelia, Leu-
cosarcia, Ocyphaps, Geophaps, and Geopelia, in which the dorsal
lobe of the facies artic. clavicularis markedly overhangs the cotyla
scapularis in lateral view.
e lateral surface distad of the facies artic. humeralis is
flattened to slightly convex and without a sulcus in the fossil, as
in Ptilinopus, Ducula, Columba, Hemiphaga, Lopholaimus, and
some species in the genera Geophaps and Geopelia, whereas a
marked sulcus is present in all other compared taxa.
e medial border of the proc. procoracoideus merges with
the shaft over a short length, and its cranial margin slopes ster-
nally from the cotyla scapularis, so that although the absolute tip
is not preserved, it clearly did not extend toward the proc. acro-
coracoideus. A short merging of the proc. procoracoideus with
the shaft is shared with most taxa, but Goura, Didunculus, and,
to some extent, Caloenas, are distinguished by a very elongate
merging of the process to the shaft. In most taxa, the tip of the
proc. procoracoideus extends toward the proc. acrocoracoideus
and extends craniad to the cotyla scapularis, but the fossil is like
Hemiphaga, Lopholaimus, Otidiphaps, and some Gallicolumba
species (G. stairi, G. beccarii, G. rubescens, G. leonpascoi, and G.
jobiensis, but not G. luzonica, G. criniger, or G. rufigula), in that
the tip does not extend cranially past the cotyla scapularis.
In the fossil, the impressio m. sternocoracoidei is well defined
but not deep, extends to the medial edge of bone, and has promi-
nent muscular striae, and the main area of pneumatic foramina
is located close to the angulus medialis. In Didunculus, Bountyp-
haps, Goura, Caloenas, Ocyphaps, and Geophaps, the main area
of pneumatic foramina is located more centrally in the extremitas
sternalis coracoidei. In most taxa, the impression is distinctly sep-
arated from the medial margin of the bone above the medial angle,
but Ptilinopus, Ducula, Hemiphaga, Streptopelia, Leucosarcia,
some Gallicolumba (e.g., G. stairi), Geophaps, and Geopelia are
similar to the fossil, in that the impression is confluent with the
medial edge of the bone. Ocyphaps and Otidiphaps differ mark-
edly in having a deeper, more extensively pneumatic impression,
and in Caloenas, Macropygia, and Geophaps the impression ex-
tends farther cranially, into the proximal half of the bone. In the
fossil, the impression does not reach the mid-length of the cora-
coid. e fossil has well-developed sternal articular facets, with
the dorsal one wider and of even width across the lateromedial
breadth of the coracoid, as in Hemiphaga. In most taxa, Lopholai-
mus included, the medial part of the facies artic. sternalis dorsalis
is distinctly wider than its lateral part.
Although the proc. lateralis is missing, the preserved lateral
margin of the impressio m. sternocoracoidei slopes steeply ven-
trally just above the end of the facies artic. sternalis dorsalis, in
a fashion that suggests that a pneumatic foramen was probably
present slightly farther laterad of this point, as in most other pi-
geons but with the notable exception of Didunculus and Galli-
columba jobiensis.
Many of the above features support inclusion of Rupephaps
as a member of the ptilinopine clade, and characters and are
apomorphies for this clade. Character is shared by Rupephaps,
Hemiphaga, and Lopholaimus, exclusive of other taxa. Rupephaps
is more similar to Hemiphaga and differs from Lopholaimus by
the impressio m. sternocoracoidei extending to the medial mar-
gin of the bone and the facies artic. sternalis dorsalis having even
breadth over the coracoid width. However, it differs from both in
that the crest extending along the medial margin above the an-
gulus medialis does not reach the facies artic. sternalis ventralis.
In addition, the lateral margin of the fossil immediately above the
missing angulus lateralis is distinctly flattened and is defined dor-
sally by a crest along the lateral margin of the impressio m. ster-
nocoracoidei and ventrally by a crest leading toward the angulus
lateralis. By contrast, this facies is rounded in Hemiphaga and
slightly flattened in Lopholaimus.
di s c u s s i o n
Rupephaps taketake is described on the basis of a single bone, but
it is significant as the first fossil columbid to be described from the
Australasian region that is older than the Pliocene. At – mya,
it provides the first indication of the evolutionary history for
columbids in Australasia, which, with species (Christidis and
Boles , Checklist Committee [O.S.N.Z.] ), has one of the
most diverse columbid faunas globally. Rupephaps taketake is
diagnosed as a ptilinopine fruit pigeon by two synapomorphies
and is only slightly smaller than the world’s largest fruit pigeon,
Hemiphaga novaeseelandiae, which at about – g is about
twice the weight of the Rock Pigeon. e fossil is more similar to
Hemiphaga than to any other Australasian pigeon, including the
extant sister-group species Lopholaimus antarcticus. Rupephaps
is distinguished from Hemiphaga by two features of the coracoid,
apart from size: a flat lateral facies above the proc. lateralis and the
form of the crest above the angulus medialis. e latter is an auta-
pomorphy for the fossil taxon and suggests that the coracoids in
Rupephaps had a slightly different relationship with the sternal ar-
ticulation and associated ligaments attached to the carina sterni.
Similarly, the flat lateral side to the coracoid suggests a different
relationship to the proc. craniolateralis of the sternum from that
in Hemiphaga.
Although our comparative studies were concerned only with
the coracoid, it is significant that we have identified two apo-
morphies to support the monophyly of the clade of Australasian
fruit-doves, or Ptilinopinae (Ptilinopus, Drepanoptila, Ducula,
Lopholaimus, Gymnophaps, and Hemiphaga): () the more ster-
nally elongate ventral lobe of the facies artic. clavicularis; and
() that the ligament attachment sites on the ventromedial re-
gion of the proc. acrocoracoideus form two distinct pits, with one
opening via a groove to the ventral facies, rather than a single pit
or groove. ese taxa form a well-supported clade based on ro-
bust and diverse molecular data (Pereira et al. ). ese apo-
morphies augment the identification of a characteristic sternal
morphology (Boles ); however, that character is ambiguous,
because a similar state is shared with Treron.
Steadman () described columbids as the most taxonomi-
cally diverse family of land birds in the Pacific Ocean region, but
until now the antiquity of such radiations in the region has been
unknown. ree closely related species in the genus Hemiphaga,
all of which inhabit forest, are currently recognized: H. novaesee-
landiae from mainland New Zealand and, formerly, the Kermadec
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Islands; the extinct H. spadicea on Norfolk Island, , km north-
west of Auckland; and H. chathamensis on the Chatham Islands,
km east-southeast of Wellington, New Zealand (Checklist
Committee [O.S.N.Z.] ). e sister-group taxa to Hemiphaga
are Lopholaimus of eastern Australian forests and Gymnophaps
from subtropical–tropical moist montane forests in Indonesia,
Papua New Guinea, and the Solomon Islands (Goodwin ,
Pereira et al. ). Lopholaimus and Hemiphaga are strong fli-
ers and often aggregate in large flocks, which predisposes them
to successful dispersal, as evidenced by the recent colonization of
Norfolk, Kermadec, and Chatham islands by Hemiphaga.
Our finding that R. taketake occurred on Zealandia (which
became modern-day New Zealand) in the Early–Middle Mio-
cene (– mya) and was closely related to both Lopholaimus and
Hemiphaga has significant biogeographic implications. It strongly
suggests that the Hemiphaga lineage has been present in New Zea-
land since at least mya. is fits with molecular evidence that
suggests that the most recent common ancestor of Hemiphaga
and Lopholaimus existed ~ mya (.–. mya) and a hypothe-
sized dispersal event from Australia ~ mya (Pereira et al. ).
If the ancestor of Rupephaps colonized Zealandia in the Early
Miocene, which would be consistent with the genetic data and the
age of the fossil, it would have found a land re-emerging from a
maximal submergence during the latest Oligocene–earliest Mio-
cene (Cooper and Millener , Landis et al. ) and so in-
creasing in area, geographic complexity, and ecological diversity.
Such conditions then probably predisposed Zealandia to coloni-
zation by birds. Other colonizing taxa in the Early Miocene proba-
bly included a Palaelodus sp. (T. H. Worthy et al. unpubl. data), but
other components of the St Bathans Fauna, such as New Zealand
wrens (Acanthisittidae; T. H. Worthy et al. unpubl. data) and sev-
eral waterfowl taxa (Worthy et al. , Worthy and Lee ), re-
veal an avifauna already then quite distinct from that of Australia.
us, even in the Early Miocene, New Zealand had a composite
biota with some long-established taxa and others that are recent
arrivals, much like today.
Rupephaps is not, however, the only pigeon taxon in the St Ba-
thans Fauna. e distal ulna NMNZ S. described by Worthy
et al. () from Bed HHa is from a smaller species, and there is a
third columbid fossil in the collections from the St Bathans Fauna.
NMNZ S. is a pedal phalanx digit I., also from Bed HHa.
It is much smaller than that of H. novaeseelandiae and similar in
size to that of Ptilinopus magnificus keri (NMNZ OR.); thus,
it conforms in size with the distal ulna but is not as flattened as in
these taxa. ese fossils are insufficient, however, to identify an
affinity within columbids, but minimally attest to a diversity of at
least two pigeon species in the St Bathans Fauna.
Given this diversity among pigeons of the Early Miocene and
the long tenure of the Hemiphaga lineage in mainland New Zea-
land, it is notable that there is no evidence for any modern diver-
sification within the genus. Two observations may explain this
apparent anomaly: () this lineage has a strong association with
forests, especially in warmer regions; and () Hemiphaga prefer-
entially eats fruit but otherwise eats leaf material (Clout et al. ,
Higgins and Davies ). Both fruit and leaves would have been
a very limited resource in New Zealand during the cold glacial pe-
riods, the most drastic effects of which peaked during the last and
penultimate glacials (McGlone , Newnham et al. ). In
those periods, forest was constrained to latitudinal regions <°S
(essentially to the Northland peninsula of the North Island; Mc-
Glone ) and, given the limited diversity of fruiting trees in the
flora, would have provided limited food for pigeons. We have no
knowledge of the avian biota during the period between deposi-
tion of the St Bathans Fauna and Recent times, but it seems likely
that the insular nature of New Zealand combined with the Pleis-
tocene glaciations may have led to the extinction of one or more
taxa, leaving just one surviving to the present on mainland New
Zealand.
Lastly, Rupephaps provides the first internal fossil-calibra-
tion point within the columbid phylogeny, with a minimum age
of – mya for the Lopholaimus–Hemiphaga split. Such cali-
bration points are few in the avian record (Ericson et al. ).
Because Ptilinopinae is a deeply nested clade within Columbidae,
Rupephaps confirms that modern generic diversification had al-
ready been attained in Australasia at least by ~ mya. e pres-
ent high diversity of columbids in Australia, most of which are
endemic, suggests that a diverse fossil record of pigeons is yet to
be revealed there. Undescribed Late Oligocene–Early Miocene
columbid fossils are present in the Riversleigh World Heritage
Area deposits of northwestern Queensland (T. H. Worthy pers.
obs.) and in lacustrine deposits at Lake Palankarinna (Etadunna
Formation) and Lake Pinpa (Namba Formation) in central Austra-
lia (Vickers-Rich , Boles ). Ultimately, we predict that a
diverse fauna of Australasian Oligocene–Miocene columbids will
be revealed in support of Olson’s () hypothesized southern or-
igin for the family.
ac k n o w l e d g M e n t s
We specifically thank the landowners A. and E. Johnstone, who
not only graciously allowed us access to their land but actively con-
tributed to the excavations by working a digger. Further, we ben-
efited greatly from the unstinting efforts of many field assistants
in the St Bathans Fauna project, especially M. Archer, J. Wood,
J. Nyugen, J. Louys, H. Godthelp, P. Creaser, and the other vol-
unteers for . is work has been supported by Australian
Research Council grant DP for collaborative work by Uni-
versity of New South Wales (UNSW) and New Zealand research-
ers, and by postdoctoral research support to T.H.W. through the
UNSW Strategic Initiative Funding Scheme. T.H.W. thanks W.
Boles of the Australian Museum and P. Horton of the South Aus-
tralian Museum for access to comparative material. We thank J.
Parish for images of Otidiphaps.
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