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Journal of Systematic Palaeontology
ISSN: 1477-2019 (Print) 1478-0941 (Online) Journal homepage: http://www.tandfonline.com/loi/tjsp20
Wading a lost southern connection: Miocene
fossils from New Zealand reveal a new lineage of
shorebirds (Charadriiformes) linking Gondwanan
avifaunas
Vanesa L. De Pietri, R. Paul Scofield, Alan J. D. Tennyson, Suzanne J. Hand &
Trevor H. Worthy
To cite this article: Vanesa L. De Pietri, R. Paul Scofield, Alan J. D. Tennyson, Suzanne J. Hand
& Trevor H. Worthy (2015): Wading a lost southern connection: Miocene fossils from New
Zealand reveal a new lineage of shorebirds (Charadriiformes) linking Gondwanan avifaunas,
Journal of Systematic Palaeontology, DOI: 10.1080/14772019.2015.1087064
To link to this article: http://dx.doi.org/10.1080/14772019.2015.1087064
Published online: 13 Oct 2015.
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Wading a lost southern connection: Miocene fossils from New Zealand reveal a
new lineage of shorebirds (Charadriiformes) linking Gondwanan avifaunas
Vanesa L. De Pietri
a,b
*,R.PaulScofield
b
, Alan J. D. Tennyson
c
, Suzanne J. Hand
d
and Trevor H. Worthy
a
a
School of Biological Sciences, Flinders University of South Australia, GPO 2100, Adelaide 5001, South Australia, Australia;
b
Canterbury Museum, Rolleston Avenue, Christchurch 8013, New Zealand;
c
Museum of New Zealand Te Papa Tongarewa,
Wellington 6140, New Zealand;
d
School of Biological, Earth and Environmental Sciences, University of New South Wales,
New South Wales 2052, Australia
(Received 28 April 2015; accepted 22 July 2015)
An endemic and previously unknown lineage of shorebirds (Charadriiformes: Scolopaci) is described from early Miocene
(1916 Ma) deposits of New Zealand. Hakawai melvillei gen. et sp. nov. represents the first pre-Quaternary record of the
clade in New Zealand and offers the earliest evidence of Australasian breeding for any member of the Scolopaci. Hakawai
melvillei was a representative of the clade that comprises the South American seedsnipes (Thinocoridae) and the
Australian Plains-wanderer (Pedionomidae), and presumed derived features of its postcranial skeleton indicate a sister
taxon relationship to Australian pedionomids. Our findings reinforce that terrestrial adaptations in seedsnipes and the
Plains-wanderer are convergent as previously proposed, and support an ancestral wading ecology for the clade. Although
vicariance events may have contributed to the split between pedionomids and H. melvillei, the proposed sister taxon
relationship between these taxa indicates that the split of this lineage from thinocorids must have occurred independently
from Australia and Zealandia’s separation from the rest of Gondwana.
http://zoobank.org/urn:lsid:zoobank.org:pub:FD3E50A9-EE95-4660-880A-A60B0DE2CEF4
Keywords: Scolopaci; Pedionomidae; Miocene; fossil birds; St Bathans Fauna
Introduction
New Zealand (NZ) has long been renowned for having a
distinctive terrestrial vertebrate faunal assemblage, char-
acterized by endemic lineages and dominated by birds
(Worthy & Holdaway 2002). Early-diverging crown-
group representatives of some of the most diverse lineages
within extant avian orders, such as songbirds and parrots
for which biogeographical patterns and sequence data
support a Gondwanan origin (Ericson et al.2002; de Kloet
& de Kloet 2005)have been a part of the avifauna of
NZ for more than 20 Ma (Worthy et al.2010,2011). The
survival of these lineages in NZ can be explained by its
geographical isolation following separation from the rest
of Gondwana. The approximate date when isolation
would have prevented taxa with low dispersal capabilities
from colonizing NZ from the rest of Gondwana is now
thought to be between 55 and 52 Ma (Schellart et al.
2006; Worthy et al.2010). Insights into the evolution,
taxonomic diversity and longevity of endemic lineages of
terrestrial vertebrates in NZ are hampered by a poor ter-
restrial fossil record, with the only pre-Quaternary terres-
trial fossils being those from the 1916 Ma St Bathans
Fauna (South Island, Central Otago region; Worthy et al.
2007,2009a, b,2010,2011; Jones et al.2009; Lee et al.
2009; Scofield et al.2010; Tennyson et al.2010; Hand et
al.2013).
A small wading bird was briefly reported from the early
Miocene St Bathans Fauna (Worthy et al.2007), represent-
ing the only pre-Quaternary record of the charadriiform
clade Scolopaci in NZ, and only the second record for Aus-
tralasia and the Pacific (De Pietri et al.2015). The suborder
Scolopaci is one of the three major clades in the species-
rich and ecologically diverse avian order Charadriiformes
(waders, gulls, auks and allies; Ericson et al.2003; Paton
et al.2003; Baker et al.2007). The Scolopaci contains five
family-level taxa, which include the painted-snipes (Ros-
tratulidae), jacanas (Jacanidae), seedsnipes (Thinocoridae),
the monospecific Plains-wanderer (Pedionomidae) and the
diverse radiation of sandpipers, snipes and allies (Scolopa-
cidae). Interrelationships amongst these lineages are well
resolved (Ericson et al.2003; Paton et al.2003; Baker
et al.2007; Gibson & Baker 2012), with a basal divergence
supporting a seedsnipe/Plains-wanderer clade and a jacana/
painted-snipe clade on the one hand, and the large scolopa-
cid radiation on the other.
*Corresponding author. Email: vdepietri@canterburymuseum.com
ÓThe Trustees of the Natural History Museum, London 2015. All Rights Reserved.
Journal of Systematic Palaeontology, 2015
http://dx.doi.org/10.1080/14772019.2015.1087064
Journal of Systematic Palaeontology
While the majority of members of the clade containing
jacanas, painted-snipes, seedsnipes and the Plains-wan-
derer currently breed in tropical to temperate areas of the
Southern Hemisphere, most species of Scolopacidae breed
in boreal, subarctic and arctic environments in the North-
ern Hemisphere (Piersma 1996). The explosive radiation
of scolopacids is probably linked to climate cooling dur-
ing the Neogene, and the evolution of grasslands, steppes
and the tundra (e.g. Kraaijeveld & Nieboer 2000; Ball-
mann 2004). Scolopacids that do breed south of the north-
ern temperate zone include both tropical and temperate
residents, such as some species of woodcock (Scolopax)
and snipes (Gallinago and Coenocorypha), as well as the
Tuamotu Sandpiper (Prosobonia parvirostris). The
endemic austral snipes (three living species of the genus
Coenocorypha), now distributed across the outlying
islands of NZ, are the only members of the Scolopaci that
currently breed in NZ, and otherwise only long-distance
migratory scolopacids that spend the boreal winter in NZ
are part of the extant avifauna. Together with some snipes
of the genus Gallinago, austral snipes are, however, likely
to have evolved from migratory ancestors (Gibson 2010;
Gibson & Baker 2012).
The pre-Quaternary Gondwanan fossil record of repre-
sentatives of the clade Scolopaci is extremely poor, with
most fossil representatives known from the Northern
Hemisphere (Mlikovsky 2002; Mayr 2009; De Pietri &
Mayr 2012). The only record for the Gondwanan lineage
represented by the South American seedsnipes and the
Australian Plains-wanderer is a pedionomid from the Oli-
gocene of South Australia (De Pietri et al.2015). It is also
worth noting that one of the longest branches in extant
charadriiform phylogeny separates the Plains-wanderer
from seedsnipes (Baker et al.2007; Gibson & Baker
2012), indicating lineage loss over time (e.g. Slack et al.
2007).
Here we describe a species of wading bird representing a
hitherto unknown lineage from the early Miocene of NZ,
belonging to the Scolopaci and showing close affinities
with the Plains-wanderer and seedsnipes, of which no rep-
resentatives are known from NZ. The fossils here described
also offer the earliest unequivocal evidence of Australasian
breeding for any member of the Scolopaci, likely indicat-
ing that this new taxon was another NZ endemic. This
record fills a significant gap in the Gondwanan fossil record
of the Scolopaci, and indeed Charadriiformes, and has
major implications for understanding Gondwanan phylo-
geography and the different evolutionary trajectories of
Scolopaci in the Northern and Southern hemispheres.
Material and methods
Our working hypothesis of relationships between and
within the different lineages of the suborder Scolopaci
follows Gibson & Baker (2012). Taxonomy and nomen-
clature is after Dickinson & Remsen (2013), and anatomi-
cal terminology follows Baumel & Witmer (1993). All
measurements are in mm and were rounded to the nearest
0.1. Commonly used abbreviations: proc, processus; R,
right; L, left; d, distal; p, proximal; ML, maximum
length; PW, proximal width; DW, distal width; Min.SW,
minimal shaft width. Much of the material is fragmentary
and therefore measurements are provided for type mate-
rial only. The specimen repositories are Canterbury
Museum, Christchurch, NZ (CM) and the Museum of
New Zealand Te Papa Tongarewa, Wellington, NZ
(NMNZ).
Comparative material
Comparative material was sampled from NMNZ;
Museum Victoria, Melbourne, Australia; Natural History
Museum Basel, Switzerland; South Australian Museum,
Adelaide, Australia; and Senckenberg Research Institute,
Frankfurt, Germany. Apart from other family-level taxa
within Charadriiformes (e.g. Glareolidae, Charadriidae),
comparisons were made with the following representa-
tives of extant species. Scolopaci: Scolopacidae: Actitis
macularius,Arenaria interpres,Bartramia longicauda,
Calidris acuminata,C. alba,C. alpina,C. canutus,C. fal-
cinellus,C. ferruginea,C. himantopus,C. minutilla,C.
pusilla,C. ruficollis,C. subruficollis,C. tenuirostris,Coe-
nocorypha aucklandica,Gallinago gallinago,G. hard-
wickii,Limnodromus griseus,L. scolopaceus,L.
semipalmatus,Limosa limosa,L. lapponica,Numenius
arquata,N. madagascariensis,N. phaeopus,N. tahitien-
sis,Phalaropus fulicarius,P. lobatus,Steganopus tri-
color,Scolopax minor,S. rusticola,Tringa incana,T.
glareola,T. nebularia,T. ochropus,Xenus cinereus. Ros-
tratulidae: Rostratula benghalensis,R. australis. Within
Jacanidae: Irediparra gallinacea,Jacana spinosa. Pedio-
nomidae: Pedionomus torquatus. Thinocoridae: Attagis
gayi,Thinocorus orbignyianus,T. rumicivorus.
Additionally, fossil material attributed to an unde-
scribed species of the extinct genus-level taxon Prosobo-
nia (Polynesian sandpipers, Scolopacidae) from
Henderson Island (Wragg 1995), and the extinct Coeno-
corypha neocaledonica (New Caledonian Snipe; Worthy
et al.2013), were examined.
Systematic palaeontology
Class Aves Linnaeus, 1758
Order Charadriiformes Huxley, 1867
Suborder Scolopaci Strauch, 1978 sensu Paton et al.
(2003)
2 V. L. De Pietri et al.
Journal of Systematic Palaeontology
Remarks. Referral to this clade is based on the following
combination of features. Coracoid (Figs 1CE, 2D, E, H)
with (1) facies articularis clavicularis ventrodorsally
broad; (2) proc. acrocoracoideus ventromedially protrud-
ing; (3) foramen nervi supracoracoidei absent. Humerus
(Figs 1IL, 2I, K, N, R) with (4) transverse ridge at inci-
sura capitis; (5) tuberculum dorsale proximodistally elon-
gated (except species of Numenius); (6) caudal surface of
crista deltopectoralis convex; (7) fossa pneumotricipitalis
lacking pneumatic foramina. Tarsometatarsus
(Figs 1OS, 2D0,G
0,J
0) with (8) tendon of musculus
flexor digitorum longus enclosed in bony canal; (9) canal
for tendon of musculus flexor digitorum longus situated
directly dorsal of sulcus for tendon of m. perforans et per-
foratus digiti 2 (some scolopacids are an exception; e.g.
Arenaria interpres, some species of Tringa and Calidris);
(10) fossa metatarsi I present. Features (1), (2) and (7) are
present in most Charadriiformes; (3), (4), (5), (6), (9) and
(10) are characteristic of all Scolopaci (see Mayr 2011;
De Pietri & Mayr 2012); features (8), (9) and (10) differ-
entiate Scolopaci from most representatives of Charadrii
(plovers and allies), and (3; except Cursorius), (5), (6)
and (8) differentiate them from Glareolidae (pratincoles
and coursers).
Family incertae sedis
Genus Hakawai gen. nov.
Type species. Hakawai melvillei sp. nov.
Derivation of name. The generic name refers to an
unseen and enigmatic ‘mystery bird’ in Maori mythology;
gender is feminine. Although recent research shows that
Maori from Rakiura (Stewart Island) considered the Coe-
nocorypha snipe’s nocturnal aerial calls to be those of the
Hakawai (Miskelly 1987), our choice alludes to a link
between the mythical Hakawai and waders of the suborder
Scolopaci.
Description. See species description below.
Hakawai melvillei sp. nov.
(Figs 1,2A, D, E, H, I, K, N, R, U, X, B0D0,
G0,J
0,L
0)
Holotype. NMNZ S.50806, right tarsometatarsus (ML D
26.3 mm, PW D3.8 mm, DW D4.0 mm, Min.SW D
1.5 mm).
Type locality, stratigraphy and age. Bed HH4,
Manuherikia River, near St Bathans, Central Otago, South
Island, NZ (44.907861S, 169.857233E). Located
25.6325.83 m above the base of lacustrine Bannockburn
Formation, Manuherikia Group; Altonian local stage,
early Miocene, 1916 Ma (Schwarzhans et al.2012). NZ
Fossil Record File Number H41/f0095.
Derivation of name. The specific name honours NZ-
based ornithologist and ecologist David Melville, for his
efforts in implementing conservation measures for shore-
birds, locally and globally.
Paratypes and measurements. The sites the elements
were recovered from are shown in parentheses (see
Schwarzhans et al.2012 for details). NMNZ S.53360, R
cranial extremity of scapula; cranial width D3.5 mm, W
of corpus scapulae D1.2 mm (HH4). NMNZ S.50490,
omal extremity of L coracoid; facies articularis humeralis
to proc. acrocoracoideus D3.6 mm, cotyla scapularis to
proc. acrocoracoideus D4.3 mm, W facies articularis
clavicularis D2.1 mm (HH1a). CM 2013.18.785, R cora-
coid; ML D13.3 mm (HH4). NMNZ S 43153, pR
humerus (HH1a). NMNZ S.50716, dR humerus; DW D
4.5 mm, dDepth D2.8 mm (HH4). CM 2013.18.195, pR
ulna; PW D3.4 mm, Min.SW D1.8 mm (HH1a); NMNZ
S.53375, dL ulna; DW D2.8 mm, Min.SW D1.7 mm
(HH4); NMNZ S.53399, L carpometacarpus; ML D
17.7 mm, PW including proc. extensorius D3.9 mm,
DW D2.3 mm, width of os metacarpale majus D1.2 mm
(HH4).
Additional referred material. Scapulae: CM
2013.18.1054, R cranial extremity (HH1a); NMNZ
S.53448, L cranial extremity, immature (HH4); NMNZ
S.52801, L cranial extremity (HH1b); NMNZ S.52755, R
cranial extremity (HH1b). Coracoids: CM 2013.18.781,
R, immature (HH4); NMNZ S.50711, R, immature
(HH2c); CM 2013.18.284, L omal end (HH1a); NMNZ
S.52160, R omal end (HH1a); NMNZ S.43975, L lacking
sternal end and part of omal end (HH1a); NMNZ
S.57903, R omal end (HH1a); CM 2013.18.114, R omal
end (HH1a); NMNZ S.51871, L omal end (HH1a);
NMNZ S.52771, L omal end, with procoracoid (HH1b);
NMNZ S.43975, L lacking sternal end (HH1a). Humeri:
NMNZ S.44064, dL, immature (HH1a); NMNZ S.42416,
pR, (Croc site); NMNZ S.50720, dR (HH4); NMNZ
S.52235, dL (HH1a); NMNZ S.51604, dL (HH1a);
NMNZ S.52548, dR (HH1a); NMNZ S.52321, dR (HH4).
Ulnae: NMNZ S.52461, pR (HH1a); NMNZ S.53080, dR,
immature (HH1b); NMNZ S.52629, dR (HH1a); CM
2013.18.1053, dR (HH1a); CM 2013.18.1055, dR
(HH1a); CM 2013.18.511, dL (HH0). Carpometacarpi:
NMNZ S.50808, L (HH4); NMNZ S. 50805, pR and dR
(HH4); CM 2013.18.527, pR (HH0); NMNZ S. 52108, pR
(HH1a); NMNZ S.51971, pL (HH1a); NMNZ S.53267,
pL (Mata Creek 5); NMNZ S.51488, pL (HH1a); NMNZ
S.50436, pR (HH2c); NMNZ S.51371, pR (HH1a);
NMNZ S.53365, pL (HH4); NMNZ S.53398, dL, dR
(HH4). Tibiotarsus: NMNZ S.50906, dL (HH4); NMNZ
S.52773, dL, immature (HH1b). Tarsometatarsi: MNZ
S.52846, pL (HH1b); MNZ S.50803, pL (HH4); MNZ
S.52414, pR (HH1a).
New Miocene shorebird from New Zealand 3
Journal of Systematic Palaeontology
4 V. L. De Pietri et al.
Journal of Systematic Palaeontology
Locality remarks. Full details of the site locations on
Home Hills Station (HH), Central Otago, South Island,
New Zealand, and their stratigraphical relationships are
given in Schwarzhans et al.(2012), except one. Bed HH0
is equivalent to FF1 of Lindqvist (1994) and is a fossil
stromatolite bed exposed at 44.90359S, 169.85840E,
Manuherikia River. The fossils were in clay surrounding
the stromatolites, New Zealand Fossil Record File (NZ
FRF) system administered by the Geoscience Society of
New Zealand and GNS Science H41/f059. Two sites on
nearby Dunstanburn Station also yielded fossils: Croc
Site, Layer 1, about 10-cm-thick sand and cobble layer,
3.5 m above the base of the Bannockburn Formation, in 3-
m north-facing cliff on a small hill on the west side of
Mata Creek, Otago, (44.889500S, 169.837833E), NZ
FRF H41/f84; and Mata Creek, Site 5, 44.88387S,
169.84113E, exposure on east side of Mata Creek with
stromatolite layer about 30 cm thick with a 10-cm-thick
conglomerate at its base containing fish and bird fossils;
NZ FRF H41/f0118.
The material attributed to Hakawai melvillei was col-
lected from the following sites (see Schwarzhans et al.
2012): HH4 (type locality), HH1a and HH1b, with few,
mostly single, elements coming from HH0, HH2c, Croc
site and Mata Creek 5. Despite stemming from different
sites, this material can be attributed to a single taxon by:
(1) size, as H. melvillei has smaller bones than all other
charadriiforms recovered from these localities (i.e. plov-
ers and gulls; Worthy et al.2007), and all elements fall
within the range expected for a single species (e.g. Ball-
mann 2004); and (2) because HH1a, HH1b and HH4 have
yielded comparable elements identical in morphology,
attesting to the presence of the same taxon in all these
sites.
Diagnosis. Small, long-legged wader, displaying the fol-
lowing combination of features (other than those noted
above). Scapula with: (1) round facies articularis humera-
lis, i.e. nearly equal craniocaudal and dorsoventral width
(Fig. 1A); (2) prominent and distinctly globular
tuberculum coracoideum, with pronounced notch separat-
ing it from facies articularis humeralis (Fig. 1B); (3) acro-
mion with weak cranial projection (Fig. 1A, B); (4) facies
articularis clavicularis nearly entirely on dorsal surface of
bone (Fig 1A). Coracoid with: (5) proc. acrocoracoideus
elongated along main axis (Fig. 1C); (6) marked recess
below facies articularis clavicularis absent (Fig. 1C, E);
(7) cotyla scapularis deep and distinctly round with maxi-
mum diameter about half the length of the facies articula-
ris humeralis (Fig. 1D); (8) shaft elongated in relation to
the omalsternal length of the omal extremity and medio-
lateral width of the facies articularis sternalis (Fig. 2H;
see also De Pietri et al.2015). Humerus with: (9) well-
developed dorsal fossa pneumotricipitalis (Fig. 1I); (10)
impressio coracobrachialis small and narrow, slightly kid-
ney-shaped (Fig. 1J); (11) condylus ventralis with cranial
surface dorsoventrally wide and proximodistally narrow,
with reduced distal and cranial projection (Fig. 1K); (12)
proc. flexorius projecting markedly ventrally (Fig. 1K, L);
(13) proc. supracondylaris dorsalis slightly prominent,
subtriangular in shape (Figs 1K, 2R); (14) fossa musculi
brachialis well developed (Fig. 1K). Ulna with: (15) well-
defined impressio scapulotricipitis (Fig. 1G); (16) condy-
lus dorsalis ulnaris with caudal margin flaring markedly
caudally and with ventral surface very broad craniocau-
dally (Fig. 1H). Carpometacarpus with: (17) proc. exten-
sorius proportionally narrow and upturned cranially and
proximally (Fig. 1M); (18) fovea carpalis caudalis deep
(Fig. 1N); (19) ventral facies of os metacarpale minus
with well-developed tuberosity proximally (Fig. 1N); (20)
os metacarpale minus dorsoventrally broad immediately
distal to the fovea carpalis caudalis (i.e. distal to synosto-
sis). Tarsometatarsus with: (21) trochlea metatarsi II dis-
tinctly proximal to other trochleae (Fig. 1P); (22) in
plantar aspect, medial rim of trochlea metatarsi III shorter
than lateral one (Figs 1Q, 2H0); (23) plantar projection on
trochlea metatarsi II with notch separating it from the
medial surface of the trochlea (Fig. 1S); (24) deep and
well-defined pit on proximal and dorsal surface of troch-
lea metatarsi III (Fig. 1O); (25) fossa metatarsi I well
Figure 1. Type material of Hakawai melvillei gen. et sp. nov. A, B, NMNZ S.53360, right scapula; A, lateral view; B, medial view.
CE, NMNZ S.50490, left coracoid; C, dorsal view; D, dorsolateral view; E, ventral view. F, G, CM 2013.18.195, right ulna; F, ventral
view; G, dorsal view. H, NMNZ S.53375, left ulna, dorsal view. I, J, NMNZ S.43153, right humerus; I, caudal view; J, cranial view. K,
L, NMNZ S.50716, distal right humerus; K, cranial view; L, caudal view. M, N, NMNZ S.53399, left carpometacarpus; M, ventral
view; N, caudal view. OS, NMNZ S.50806 (holotype), right tarsometatarsus; O, dorsal view; P, medial view, Q, plantar view; R, prox-
imal view; S, distal view. Abbreviations: acr, acromion; cdp, crista deltopectoralis; cdu, condylus dorsalis ulnaris; cdv, condylus ventra-
lis; cih, cristae intermediae hypotarsi; cmh, crista medialis hypotarsi; csc, cotyla scapularis; ctd, cotyla dorsalis; ctv, cotyla ventralis;
dfp, dorsal fossa pneumotricipitalis; epv, epicondylus ventralis; fac, facies articularis clavicularis; fah, facies articularis humeralis; fcc,
fovea carpalis caudalis; fdl, canal for tendon of musculus flexor digitorum longus; fdm, facies articularis digitalis major; fic, fossa infra-
cotylaris dorsalis; fmI, fossa metatarsi; fmb, fossa m. brachialis; fos, fossa; fpm, fossa parahypotarsalis medialis; icb, impressio coraco-
brachialis; fvd, foramen vasculare distale; ist, impressio scapulotricipitis; m&l tr. met III, medial and lateral rims of trochlea metatarsi
III; pex, proc. extensorius; pp2, sulcus for tendon of m. perforans et perforatus digiti 2; ppr, plantar projection; prf, proc. flexorius; psd,
proc. supracondylaris dorsalis; sht, sulcus humerotricipitalis; tbc, tuberculum coracoideum; tbd, tuberculum dorsale; tcp, tuberculum
carpale; tmII, trochlea metatarsi II; tmIII; trochlea metatarsi III; trd, transverse ridge; tub, tubercle. Scale bars D2 mm, except
OQD5 mm.
J
New Miocene shorebird from New Zealand 5
Journal of Systematic Palaeontology
6 V. L. De Pietri et al.
Journal of Systematic Palaeontology
marked (Fig. 1Q); (26) foramen vasculare distale large
(Fig. 1O, Q).
Differs from all examined species of Scolopacidae in
characters (2), (4), (16), (18), (20), (23) and (26) and from
most scolopacids in (1), (3), (8), (12), (13), (14), (17) and
(21). Characters (9) and (16) were present in members of
Thinocoridae; characters (2), (4), (6), (8), (24) and (26)
were found in Pedionomus torquatus; and (1), (7), (11),
(19), (20), (22) and (23) in both thinocorids and P. torqua-
tus (see Fig. 3). Character (7) is also present in Rostratuli-
dae, and (16), (19), (21) and (22) in Rostratulidae and
Jacanidae. Hakawai melvillei differs from members of
Rostratulidae and Jacanidae in characters (5), (6), (8), (9),
(10), (12), (13), (14), (23) and (24); character (26) is also
present in this clade, but the foramen is much larger and
situated in a very deep and wide groove, which is a
derived trait for species in Jacanidae and Rostratulidae
(see also Mayr 2011).
Description and comparisons. Although the scapula and
coracoid are about the size of those of the Red-necked
Stint Calidris ruficollis, one of the smallest scolopacids,
overall proportions across different postcranial elements
suggest Hakawai melvillei was larger than previously
reported (Worthy et al.2007), and about the size of a
male Plains-wanderer (Table 1). Pedionomus torquatus
displays pronounced sexual dimorphism, with females
being larger than males (Baker-Gabb 1996). All elements
except the tarsometatarsus (Fig. 3RW) agree in size and
proportions with those of Pedionomus torquatus, whereas
those of scolopacids and thinocorids are markedly differ-
ent (Table 1). Elements of the pectoral girdle and wing
display features that most closely match the condition in
the corresponding elements of P. torquatus and thinocor-
ids (Fig. 2).
The facies articularis humeralis is round and distinctly
offset, ventrolaterally, from the cranial extremity of the
scapula (Fig. 1A, B). The tuberculum coracoideum is
prominent and globular. Other than in P. torquatus
(Fig. 2B) and seedsnipes, this feature is also present in
species of Gallinago,Scolopax and Coenocorypha, but
unlike these taxa, the tuberculum coracoideum is dis-
tinctly separated from the facies articularis humeralis by a
notch (Fig. 2A). Unlike in several of the examined scolo-
pacids (Fig. 2C), the acromion is rounded and lacks the
marked dorsocranial projection. The facies articularis
clavicularis is robust and more dorsally rather than lat-
erally positioned compared to that of the examined scolo-
pacids and thinocorids, but resembling P. torquatus
(Fig. 2B). The facies articularis clavicularis of the cora-
coid has nearly the same breadth throughout its dorsoven-
tral extent (Fig. 2D), as in P. torquatus. In other examined
taxa, the dorsal section is more elongated in the omal
sternal direction than the ventral section. Also as in P. tor-
quatus, a well-developed recess under the facies articula-
ris clavicularis is absent (Fig. 1C; De Pietri et al.2015).
Unlike in pedionomids, the sulcus m. supracoracoidei is
more deeply excavated. The cotyla scapularis is round
and deep, with a maximum diameter of about half the
length of the facies articularis humeralis (Fig. 1D).
At the proximal end of the humerus, a dorsal fossa
pneumotricipitalis is well developed (Figs 1I, 2K). This
fossa is absent in P. torquatus, whereas the condition is
variable within Scolopacidae (Mayr 2011;Fig. 2M). In H.
melvillei (Fig. 2K), this fossa is much deeper and dorso-
ventrally wide, approaching the condition in thinocorids
(Fig. 2L). The shape of the caput humeri (Figs 1J, 2I)
resembles that of thinocorids (Fig. 2J), but much of its
caudal surface is worn in one of the specimens (NMNZ
S.42416; Fig. 2K). A markedly proximally protruding
caput humeri was considered a derived similarity of thino-
corids and P. torquatus (Olson & Steadman 1981), but it
occurs to some degree in certain scolopacids, and is as
such not a distinguishing character. The distal end closely
matches that of Thinocorus rumicivorus (Fig. 2N). The
fossa musculi brachialis is as deep as that of T. orbignyia-
nus. Similar to seedsnipes (except T. orbignyianus) and P.
torquatus, the cranial surface of the condylus ventralis is
Figure 2. Selected elements of Hakawai melvillei gen. et sp. nov. (A, D, E, H, I, K, N, R, U, X, B0,C
0,D
0,G
0,J
0,L
0) in comparison
with extant taxa of Scolopaci, illustrating differences in the features discussed. AC, right scapulae of A, H. melvillei, NMNZ S.53360;
B, Pedionomus torquatus;C, Calidris acuminata.DG, left coracoids of D, E, H. melvillei NMNZ S.50490; F, Thinocorus rumicivo-
rus;G, Calidris canutus.H, right coracoid of H. melvillei CM 2013.18.785, immature individual. IM, right proximal humeri of I, K,
H. melvillei NMNZ S.42416; J, L, T. rumicivorus;M, Gallinago hardwickii.NT, distal right humeri of N, R, H. melvillei NMNZ
S.50716, O, S, T. rumicivorus;P, T, G. hardwickii;Q, Calidris acuminata.UW, distal left ulnae of U, H. melvillei NMNZ S.53375;
V, T. rumicivorus;W, Limosa lapponica.XB0,left carpometacarpi of X, B0,H. melvillei NMNZ S.53399; Y, A0,T. rumicivorus;Z,
C. acuminata.C0,distal left tibiotarsus, H. melvillei NMNZ S.50906. D0N0,right tarsometatarsi of D0,G
0,J
0,L
0,
H. melvillei NMNZ S.50806 (holotype); E0,M
0,Calidris canutus;E0,H
0,N
0,T. rumicivorus;I0,G. hardwickii;K0,Limosa lapponica.
Abbreviations: acr, acromion; cdl, condyles lateralis; cdu, condylus dorsalis ulnaris; cdv, condylus ventralis; cph, caput humeri; csc,
cotyla scapularis; dfp, dorsal fossa pneumotricipitalis; dom, dorsal flare at os metacarpale minus; fac, facies articularis clavicularis; fah,
facies articularis humeralis; fdl, canal for tendon of musculus flexor digitorum longus; fdm, facies articularis digitalis major; fmb, fossa
m. brachialis; m&l tr. met III, medial and lateral rims of trochlea metatarsi III; not, notch; pex, proc. extensorius; ppr, plantar projection;
prf, proc. flexorius; psd, proc. supracondylaris dorsalis; pst, pons supratendineus; tbc, tuberculum coracoideum; tcp, tuberculum carpale;
tmII, trochlea metatarsi II; tre, tuberositas retinaculum extensoris; tub, tubercle. Scale bars D2 mm, except M, P, Q, T, WZ, E0D
5 mm.
J
New Miocene shorebird from New Zealand 7
Journal of Systematic Palaeontology
8 V. L. De Pietri et al.
Journal of Systematic Palaeontology
not markedly globular and distally protruding compared
to that of scolopacids (Figs 1K, 2O), being wider ventro-
dorsally and narrower proximodistally (Figs 1K, 3I). The
cranialmost surface of the proc. supracondylaris dorsalis
becomes gradually wider proximally, and is subtriangular
in shape (Figs 1L, 2R).
The ulna is similar to that of T. rumicivorus and P. tor-
quatus, differing primarily in the presence of a better-
defined impressio scapulotricipitis at the proximal end
(Fig. 1G). The tuberculum carpale is less cranially pro-
truding compared to that of P. torquatus and most scolo-
pacids (Fig. 2UW). As in thinocorids, the caudal margin
of the condylus dorsalis ulnaris flares markedly caudally,
and the ventral surface is craniocaudally very broad
(Figs 1H, 2U, V). As in thinocorids (Fig. 2Y), the proc.
extensorius of the carpometacarpus (Fig. 2X) is propor-
tionally narrow and upturned cranially and proximally.
The fovea carpalis caudalis (Figs 1N, 2B0) is deep but less
so than in P. torquatus. At the proximal end of the os
metacarpale minus there is a well-developed tuberosity
ventrally for ligamental attachment (Figs 1N, 2B0). This
tubercle is also present in thinocorids and P. torquatus,
albeit less pronounced (Fig. 2A0). In H. melvillei,P. tor-
quatus and thinocorids, the os metacarpale minus, proxi-
mally and immediately distal to the fovea carpalis
caudalis, is noticeably broad dorventrally, whereas scolo-
pacids lack the pronounced dorsal flare (Fig. 2A0). At the
distal end, the cranial projection of the dorsal facet of the
facies articularis digitalis major (Fig. 1M) is globular in
shape, and identical to that of P. torquatus.
The distal end of the tibiotarsus is only partially
preserved (Fig. 2C0). The tubercle lateral to the pons
supratendineus for the attachment of lig. meniscoti-
biale intertarsi is better developed than in most of the
other examined scolopacids, being absent in thinocor-
ids and very well developed in P. torquatus.The
pons supratendineus is proximodistally wide and the
tuberositas retinaculum extensoris is elongated and
ridge-like. In P. torquatus, this tuberositas is medio-
laterally broader and slightly longer proximodistally.
The condylus lateralis is not as laterally projecting as
that of T. rumicivorus. Both the epicondylus lateralis
and the depressio epicondylaris lateralis are well
developed.
In size the tarsometatarsus resembles that of P. torqua-
tus and is very much unlike the proportionally shorter and
stouter tarsometatarsus of thinocorids (Fig. 2F0). As in
nearly all members of Scolopaci, the tendon of musculus
flexor digitorum longus is enclosed in a bony canal, which
is medially positioned in the hypotarsus. The crista latera-
lis hypotarsi is dorsoplantarly short and projects markedly
laterally (Fig. 2L0). In plantar view (Fig. 1Q), the
Figure 3. Postcranial elements of Hakawai melvillei gen. et sp. nov. (A, C, E, H, K, N, P, R, T, V, X) in comparison with Pedionomus
torquatus (B, D, F, I, L, O, Q, S, U, W, Y) and Thinocorus rumicivorus (G, J, M). Illustrated characters (see Diagnosis for character
description) (1), (7), (11), (20), (23) and (24) are shared with members of Thinocoridae and P. torquatus. Characters (2), (4), (6), (8),
(25) and (27) are interpreted as presumably derived for P. torquatus and H. melvillei. Note the proportionally more elongate tarsometa-
tarsus of H. melvillei compared to the Plains-wanderer. Abbreviations: cdv, condylus ventralis; csc, cotyla scapularis; dom, dorsal flare
at os metacarpale minus; fac, facies articularis clavicularis; fah, facies articularis humeralis; m&l tr. met III, medial and lateral rims of
trochlea metatarsi III; pit, rounded depression on dorsal surface of trochlea metatarsi III; ppr, plantar projection; rec, recess below facies
articularis clavicularis (absent); shf, shaft; tbc, tuberculum coracoideum; tmII, trochlea metatarsi II. Not to scale.
Table 1. Comparative measurements (in mm) of selected post-
cranial elements of the Plains-wanderer Pedionomus torquatus,
Hakawai melvillei gen. et sp. nov., the Least Seedsnipe Thinoco-
rus rumicivorus, and the Sharp-tailed Sandpiper Calidris acumi-
nata. Both female (left) and male (right) individuals of
P. torquatus were measured. Measurements of H. melvillei were
obtained from type material only. Abbreviations: csc, cotyla
scapularis; fah, facies articularis humeralis; DW, distal width;
ML, maximum length; MSW, minimum shaft width; pac, proc-
essus acrocoracoideus; PW, proximal width. All measurements
are in mm.
Pedionomus
torquatus
yHakawai
melvillei
Thinocorus
rumicivorus
Calidris
acuminata
Coracoid
ML 17.2/15.6 13.3
17.1 17.5
cscpac 4.6/4.5 4.3 5.8 6.1
fahpac 3.8/3.7 3.6 4.8 5.6
MSW 1.7/1.5 1.3
(1.5) 1.8 1.9
Scapula
cranial width 3.7/3.2 3.5 5.1 5.3
Humerus
ML 34.8/30.7 33.2 31.1
DW 4.6/4.4 4.5 5.3 5.6
Ulna
ML 37.4/33.2 36.8 39.7
PW 4.0/3.7 3.5 4.4 4.4
DW 2.9/2.8 2.8 3.8 4.0
Carpometacarpus
ML 18.3/16.7 17.7 23.1 22.7
PW 4.4/3.9 3.9 5.0 5.4
Tarsometatarsus
ML 26.9/24.1 26.3 19.9 31.0
PW 4.7/3.9 3.8 4.0 4.0
DW 4.2/3.9 4.0 3.6 3.9
Coracoid CM 2013.18.785 is a juvenile (see Results section) with omal
and sternal ends not fully defined. The maximum length of an adult indi-
vidual is therefore likely to be greater. Shaft width of an adult individual
(MNZ S. 52771) is also noted. Proportions (Fig. 2H) indicate the cora-
coid of H. melvillei is elongated in the omalsternal direction, like that
of P. torquatus and unlike that of thinocorids and scolopacids
(De Pietri et al. 2015).
J
New Miocene shorebird from New Zealand 9
Journal of Systematic Palaeontology
hypotarsus closely resembles that of thinocorids, P. tor-
quatus, and species of Numenius in being proximodistally
shorter than that of most scolopacids. The proximal end of
the hypotarsus differs from that of species of Gallinago,
Coenocorypha,Limnodromus and Scolopax in that in
these, the crista medialis hypotarsi almost completely enc-
loses the sulcus for the tendon of m. perforans et perfo-
rates digiti 2, and there is more than one bony canal for
the tendons of the extensor muscles of the foot. The troch-
lea metatarsi II (Fig. 1P) at the distal end is more proxi-
mally situated than that of most scolopacids, resembling
that of thinocorids (Fig. 2F0) and P. torquatus, albeit this
condition is also present in species of Limnodromus and
Gallinago.InH. melvillei, the medial rim of the trochlea
metatarsi III is, plantarly, very short compared to the lat-
eral one (Figs 1Q, 2G0), which it does not reach proxi-
mally (Fig. 2I0). This asymmetry was absent in all
examined scolopacids but, as in H. melvillei, is very pro-
nounced in thinocorids (Fig. 2H0). The plantar projecting
flange on trochlea metatarsi I0(in distal view) makes a
notch with the medial surface of the trochlea (Fig. 1S)
this condition is present in thinocorids and P. torquatus
and is absent in the examined scolopacids (Fig. 2K0). A
fossa metatarsi I is well marked (Fig. 1Q). The round pit
on the proximal and dorsal surface of trochlea metatarsi
III resembles that of P. torquatus, and is much deeper
than in all other examined taxa (Fig. 1O). The foramen
vasculare distale is larger than that found in scolopacids
and seedsnipes (Fig. 1Q), and resembles that of P.
torquatus.
Ontogenetic stage of osteologically immature
individuals
Bone surface texture is a useful indicator of skeletal matu-
rity in modern and fossil birds, provided the studied taxa
undergo similar growth patterns (Tumarkin-Deratzian
et al.2006). Most (but not all) neornithine birds are con-
sidered to obtain postcranial skeletal maturity within the
first year of life (Padian et al.2001); they can, however,
reach adult size as early as the time of fledging, i.e. before
becoming osteologically mature (Tumarkin-Deratzian
et al.2006; Watanabe & Matsuoka 2013). Although few
skeletal ontogenetic studies have been conducted for neo-
gnathous birds, congruence in patterns of surface texture
and age classes across diverse taxa (ducks, geese, cranes,
herons, gulls; Gotfredsen 1997; Serjeantson 1998; Tumar-
kin-Deratzian et al.2006; Watanabe & Matsuoka 2013)
suggests results may be largely applicable to taxa within
Neognathae (Tumarkin-Deratzian et al.2006).
Two coracoids (CM 2013.18.781; NMNZ S.50711) of
Hakawai melvillei (Fig. 2H) show a striated, rough sur-
face, with longitudinal ridges, deep grooves and trans-
verse struts (Fig. 2H). This texture is consistent with an
early juvenile or late chick stage, i.e. before fledging and
before functionally mature plumage is obtained (Wata-
nabe & Matsuoka 2013). Striations in the osseous surface
of bones tend to be absent in immature but adult-sized
(i.e. sub-adult) individuals (Tumarkin-Deratzian et al.
2006). Osteological detail of structures and muscle attach-
ment scars not observable in detail at the chick stage
become progressively evident from fledging to the adult
stage. Because of this, much detail is lacking in the sternal
end of both juvenile coracoids of H. melvillei, and
although some features of the omal end are recognizable
(cotyla scapularis, facies articularis humeralis), the facies
articularis clavicularis, the proc. acrocoracoideus and the
medial surface of the bone are poorly developed
(Fig. 2H). Scapula NMNZ S.53448 (Fig. 2A) and ulna
NMNZ S.53080 are of adult size but are not yet fully skel-
etally mature (sub-adult sensu Tumarkin-Deratzian et al.
2006). These elements still show some porosity in the tex-
ture of the shaft, and well-defined longitudinal lines still
remain (Fig. 2A). All these observations provide unequiv-
ocal evidence that H. melvillei was breeding in the vicin-
ity of palaeolake Manuherikia in the early Miocene.
Discussion
Hakawai melvillei gen. et. sp. nov. represents a unique and
distinct lineage within Scolopaci, with close affinities to
the clade encompassing thinocorids and pedionomids.
The overall similarity in morphology to members of this
clade is evident in all examined elements of the postcra-
nial skeleton, particularly the scapula, coracoid, humerus
and tarsometatarsus. While several distinctive postcranial
features (characters 1, 7, 11, 19, 20, 22 and 23 in Diagno-
sis) are present in both thinocorids and pedionomids,
some (2, 4, 6, 8, 24 and 26) appear to be shared only by H.
melvillei and Pedionomus torquatus. Despite there being
a relatively high proportion of characters that exhibit
homoplasy within Scolopaci, especially in Scolopacidae,
the unique combination of features present in seedsnipes,
the Plains-wanderer and H. melvillei (Fig. 3) is not repli-
cated in any lineage within Scolopacidae, and represents
strong evidence for a link between these taxa. Despite the
overall derived morphology of the Plains-wanderer com-
pared to other Scolopaci (Bock & McEvey 1969; Olson &
Steadman 1981), we propose a sister taxon relationship
between H. melvillei and P. torquatus based on presum-
ably derived features retained in both lineages (Fig. 3).
The absence of several traits that are possibly derived
for scolopacids indicates that H. melvillei was not a repre-
sentative of this lineage, contrary to previous reports
(Worthy et al.2007), and therefore precludes it from
being closely related to the endemic austral snipes (spe-
cies of Coenocorypha). Late Oligoceneearly Miocene
(c.2420.5 Ma) scolopacids from Europe already
10 V. L. De Pietri et al.
Journal of Systematic Palaeontology
possessed a combination of features characteristic of
extant scolopacids (De Pietri & Mayr 2012), and resem-
bled early-diverging taxa such as Numenius and Limosa.
Given the biogeographical, phylogenetic and boreal
breeding patterns of extant scolopacids (Piersma 1996;
Gibson 2010; Gibson & Baker 2012), it is likely that
members of the crown group did not arrive in Australasia
until after shifts in non-breeding ranges (in response to
climate cooling during the Neogene) prompted the onset
of long-distance migration (Louchart 2008). Response to
geographical and environmental changes from the early to
mid-Cenozoic has therefore contributed to the different
evolutionary trajectories for Scolopaci in the Northern
and Southern hemispheres. High endemism resulted from
geographical isolation in Australia and NZ, while crown-
group scolopacids breeding in the Northern Hemisphere
formed a Neogene radiation characterized by migratory
behaviour.
In the absence of terrestrial predators, several endemic
NZ birds have become flightless (Tennyson & Martinson
2010), but there is still little indication that the St Bathans
Fauna possessed a high proportion of flightless birds
(Worthy et al.2007; Worthy et al.2009b; Tennyson et al.
2010). Morphology and limb proportions do not suggest
that Hakawai melvillei was flightless or possessed dimin-
ished flight capabilities. The morphology of the humerus,
carpometacarpus and ulna, which closely matches that of
seedsnipes (Fig. 3), may indicate that H. melvillei was a
better flier than the extant Plains-wanderer, which is a
notoriously poor flier (e.g. De Pietri et al.2015). How-
ever, the presence of osteologically immature individuals
at chick or near fledging stage demonstrates that H. mel-
villei bred in Zealandia, which supports it having been an
insular, endemic taxon nonetheless.
The proportionally elongated and gracile tarsometatar-
sus, which resembles that of wading scolopacids (also in
the arrangement of the hypotarsal canals), is unlike the
stouter tarsometatarsi of seedsnipes, the Plains-wanderer
and species of Coenocorypha (austral snipes), all of which
inhabit terrestrial environments. This likely indicates that
H. melvillei was a littoral zone feeder, which is supported
also by its abundance in these lacustrine fossil sites. The
terrestrial adaptations of P. torquatus and seedsnipes were
proposed to be convergent for both taxa (De Pietri et al.
2015), which is further substantiated by the wading habits
of H. melvillei. We propose that wading was likely to
have been the ancestral ecology of the clade, as suggested
by the wading habits of other Scolopaci (Scolopacidae,
Rostratulidae and Jacanidae).
Combined knowledge of present and extinct NZ groups
points to long-term conservatism in its distinctive and
endemic fauna, at least from the mid-Cenozoic onwards.
Nevertheless, because of the scarcity of terrestrial faunas
pre-dating the Quaternary in NZ, it is not possible to
determine when H. melvillei went extinct or for how long
after the early Miocene its lineage was a part of the fauna
of NZ. Vulnerability after the Oligocene bottleneck, i.e.
the reduction of land area as a result of eustatic sea-level
Figure 4. Proposed Gondwanan diversification of the lineage from which the South American seedsnipes (Thinocoridae), the Australian
Plains-wanderer (Pedionomidae) and the New Zealand (NZ) Hakawai melvillei gen. et sp. nov. originated. Continents are shown in their
current position but the splits are likely to have taken place before 40 Ma (see main text). The possibility of trans-Tasman dispersal
between Australia and NZ is indicated by a dotted arrow and a question mark. However, there is no evidence of a common ancestor of
H. mellvillei and pedionomids in Australia, and Oligocene pedionomids already shared derived traits with Pedionomus torquatus (De
Pietri et al.2015) that are absent in H. melvillei. We propose instead that both lineages have common ancestry in East Gondwana before
its final fragmentation, and became independently isolated following complete separation of NZ and Australia from the rest of Gond-
wana. Insets: A, phylogeny of Scolopaci (Gibson & Baker 2012), showing the proposed sister taxon relationship between H. melvillei
and pedionomids; B, position of continents at 55 Ma (Lawver & Gahagan 2003). Bird images reproduced with permission of HBW Alive
(Baker-Gabb 1996; Fjeldsa
1996).
New Miocene shorebird from New Zealand 11
Journal of Systematic Palaeontology
rise and associated niche diversity loss during the late Oli-
gocene and earliest Miocene, and the climatic fluctuations
that reduced opportunity for refugia after the middle Mio-
cene, have been listed as contributing factors to floral
overturn and faunal changes during the Cenozoic (but see
Tennyson 2010), leading to the disappearance of many
taxa, including crocodilians and tropical birds (Cooper &
Cooper 1995; Worthy et al.2007; Pole 2008,2014; Jones
et al.2009; Reichgelt et al.2015).
Pedionomids and Hakawai melvillei were part of the
ancestral radiation of a lineage that, similar to sphenodon-
tids (Jones et al.2009), may have been more widespread
in the past, but only managed to survive locally in Austral-
asian landmasses (Olson & Steadman 1981). Hakawai
melvillei fills an important gap in the fossil record of the
Charadriiformes in the Southern Hemisphere, and demon-
strates that, as proposed for passerines and parrots, the
splits between seedsnipes, pedionomids and H. melvillei
are to some degree likely to be associated with the
breakup of Gondwana and the resulting isolation of land-
masses that prevented gene flow amongst the ancestors of
these lineages during the period 5535 Ma (Fig. 4; Schel-
lart et al.2006; Selvatti et al.2015). Nevertheless, con-
trary to biogeographical models proposed for passerines
(e.g. Ericson et al.2003; Selvatti et al.2015), sequential
vicariance alone probably does not explain these patterns,
as the proposed sister taxon relationship between pediono-
mids and H. melvillei indicates that the split of this lineage
from the seedsnipe lineage must have occurred indepen-
dently from Zealandia’s separation from the rest of Gond-
wana (Fig. 4). Due to the incomplete nature of the fossil
record, this phylogeographical pattern could as well
reflect differential extinctions in post-Gondwana land-
masses. We note that this less parsimonious scenario is
not supported by the available data. Likewise, the role of
dispersal across open water, albeit over relatively short
distances (Fig. 4), from the remaining East Gondwana
landmass to Australia and NZ is at present difficult to
assess. A parallel amongst other vertebrate groups might
be found in the divergence of Australasia’s endemic mys-
tacinid bats (the extinct Icarops lineage in Australia and
surviving Mystacina lineage in NZ (Hand et al.2013,
2015) from ancestral noctilionoids 5141 Ma (Teeling et
al.2005) in East Gondwana before its final fragmentation
(Gunnell et al.2014).
Acknowledgements
We are grateful to K. Roberts, P. Horton and G. Mayr for
access to comparative material. We thank Lynx Edicions
for permission to use the images in Figure 4. We are grate-
ful to the landowners A. and E. Johnstone of Home Hills
Station and J. and T. Enright, Dunstanburn Station, South-
ern Lakes Holdings Ltd, for access to sites, and to J.
Worthy for sorting the material. We also thank G. Gully
and B. Choo for their assistance with preparing figures.
This work was funded by grants from the Australian
Research Council DP0770660 to S. Hand et al. and
DP120100486 to T. H. Worthy et al. We are thankful for
comments by A. Louchart and one other, anonymous,
reviewer which helped improve this manuscript.
ORCID
Vanesa L. De Pietri http://orcid.org/0000-0002-3786-9741
R. Paul Scofield http://orcid.org/0000-0002-7510-6980
Trevor H. Worthy http://orcid.org/0000-0001-7047-4680
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