Journal of Vertebrate Paleontology

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A new species of crested pterosaur

(Pterodactyloidea, Anhangueridae) from the Lower
Cretaceous (upper Albian) of Richmond, North
West Queensland, Australia

Timothy M. Richards, Paul E. Stumkat & Steven W. Salisbury

To cite this article: Timothy M. Richards, Paul E. Stumkat & Steven W. Salisbury (2021): A new
species of crested pterosaur (Pterodactyloidea, Anhangueridae) from the Lower Cretaceous (upper
Albian) of Richmond, North West Queensland, Australia, Journal of Vertebrate Paleontology, DOI:
10.1080/02724634.2021.1946068

To link to this article: https://doi.org/10.1080/02724634.2021.1946068

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Published online: 09 Aug 2021.

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Journal of Vertebrate Paleontology e1946068 (14 pages)
© by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2021.1946068

ARTICLE

A NEW SPECIES OF CRESTED PTEROSAUR (PTERODACTYLOIDEA, ANHANGUERIDAE)

FROM THE LOWER CRETACEOUS (UPPER ALBIAN) OF RICHMOND, NORTH WEST
QUEENSLAND, AUSTRALIA

TIMOTHY M. RICHARDS, *,1 PAUL E. STUMKAT,2 and STEVEN W. SALISBURY 1

1
School of Biological Sciences, The University of Queensland, St. Lucia, Queensland, 4072, Australia, t.richards@uqconnect.edu.au;
s.salisbury@uq.edu.au;
2
P.O. BOX 165, Killarney, Queensland, 4373, Australia, paul@stumkatstudios.com.au

ABSTRACT—Pterosaur fossils from Australia are exceptionally rare. Since the discovery of the continent’s first pterosaur
some 40 years ago, fewer than 20 specimens have been described. The Lower Cretaceous (upper Albian) Toolebuc
Formation of North West Queensland is the most productive horizon for Australian pterosaurs. Herein, we describe a new
species of pterosaur, Thapunngaka shawi gen. et sp. nov., from the Toolebuc Formation, near Richmond, North West
Queensland. The specimen (KKF494) comprises the rostral portion of a crested mandible and represents the largest
pterosaur yet described from Australia. The new species presents features that indicate an affinity with Anhangueridae,
which is consistent with their reported cosmopolitan distribution during this period. Thapunngaka shawi can be
distinguished from other anhanguerids through the possession of a mandible with a smooth dorsal surface medially and
uniquely sized alveoli that are positioned laterally along the jaw. Phylogenetic analysis reveals a close relationship among
all Australian anhanguerids and points to an endemic Australian radiation within Anhangueridae. Thapunngaka shawi has
the largest mandibular crest of any anhanguerian worldwide, and provides further evidence for the existence of an
increasingly diverse range of large crested pterosaurs in the Australian part of eastern Gondwana during the Cretaceous.

http://zoobank.org./urn:lsid:zoobank.org:pub:B0281A27-DDDC-4AD9-AFBF-50C7A4D8540D

SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Citation for this article: Richards, T. M., P. E. Stumkat, and S. W. Salisbury. 2021. A new species of crested pterosaur
(Pterodactyloidea, Anhangueridae) from the Lower Cretaceous (upper Albian) of Richmond, North West Queensland,
Australia. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2021.1946068

INTRODUCTION Archaeopterodactyloidea (sensu Kellner, 2003). However, dis-

agreements over the original interpretation of overall skull
Pterosaur fossils from Australia are exceptionally rare, com-
length, corresponding tooth positions, and estimated wing span
prising fragmentary and predominately isolated bones from the
led Fletcher and Salisbury (2010) and Brougham et al. (2017)
Cretaceous of Queensland, New South Wales, Victoria and
to suggest that Mythunga should be reclassified as a pteranodon-
Western Australia (Fig. 1). Since the discovery of the first ptero-
toid (sensu Kellner et al., 2010), with probable affinities to
saur fossils almost 40 years ago, fewer than 20 specimens have
Anhangueria (sensu Andres et al., 2014). Recent phylogenetic
been described. From these, only three taxa have been named:
assessments have consistently supported the placement of
Mythunga camara Molnar and Thulborn, 2007; Aussiedraco
Mythunga camara as an anhanguerian and as a close relative of
molnari Kellner, Rodrigues, and Costa, 2011; and Ferrodraco
Ferrodraco lentoni (Pentland and Poropat, 2019; Pentland
lentoni Pentland, Poropat, Tischler, Sloan, Elliott, Elliott,
et al., 2019; Holgado and Pêgas, 2020).
Elliott, and Elliott, 2019. All three taxa come from the mid-Cre-
Aussiedraco molnari is based on the rostral part of a mandible
taceous rocks of the Rolling Downs Group of the Eromanga
(QMF10613), found in the Toolebuc Formation near the Hamilton
Basin of North West and Central West Queensland.
River, approximately 70 km east of Boulia, western Central West
Mythunga camara (QMF18896) was Australia’s first named
Queensland (Fig. 1). Molnar and Thulborn (1980) initially con-
pterosaur (Molnar and Thulborn, 2007). It is based on a partial
sidered the mandible very similar to material previously assigned
rostrum comprising part of the maxilla and the corresponding
to species of Ornithocheirus from the Cenomanian Cambridge
portion of the mandible, recovered from the Toolebuc For-
Greensand of England and assigned the mandible to Ornithocheir-
mation, near Hughenden, North West Queensland (Fig. 1).
oidea (sensu Unwin, 2003) aff. Ornithocheirus. However, based on
Noting a unique suite of characters, relating mainly to the denti-
a combination of features pertaining to the dentition, the mor-
tion, Molnar and Thulborn (2007) assigned the skull to a new
phology of the symphysis and the lack of mandibular crest,
genus and species, and provisionally placed it within
Kellner et al. (2011) argued that it cannot be referred to any pre-
viously recognized genera and erected Aussiedraco molnari,
regarding it as being closely related to (but not a member of) the
*Corresponding author
Anhangueridae. Pêgas et al. (2019) recently assigned Aussiedraco
Color versions of one or more of the figures in the article can be found to the newly erected clade Targaryendraconia, which is the sister-
online at www.tandfonline.com/ujvp. clade to Anhangueria (sensu Holgado and Pêgas, 2020).

Published online 09 Aug 2021

 Richards et al.—New anhanguerid from Australia (e1946068-2)
Australia’s most complete pterosaur, Ferrodraco lentoni
(AODF876), derives from the Cenomanian–lower Turonian
portion of the Winton Formation, north-east of Winton,
Central West Queensland (Fig. 1) (Pentland et al., 2019). The
holotype is represented by parts of the skull (partial premaxillae,
maxillae and dentaries, the left frontal and the articular region of
the left mandibular ramus) and associated postcranial elements
(five cervical vertebrae, the right scapulocoracoid, the left ulna
and radius, the left proximal and distal carpals, and the left meta-
carpal IV). Using a modified dataset of Andres et al. (2014),
Ferrodraco was assigned to Anhangueria (as the sister taxon to
Mythunga) by Pentland et al. (2019) due to the presence of pre-
maxillary and mandibular crests and ‘spike-shaped’ teeth with an
oval base. Ferrodraco lentoni can be diagnosed by two autapo-
morphies relating to its dentition: the first pair of the teeth in
the premaxillae and mandible being smaller than the other
rostral teeth; and the fourth to seventh teeth being smaller
than the third and eighth teeth (Pentland et al., 2019). Supporting
the close relationship between the Australian taxa proposed by
Pentland et al. (2019), Holgado and Pêgas (2020) recently
assigned Ferrodraco lentoni and Mythunga camara to
Tropeognathinae, a cosmopolitan anhanguerid clade that also
includes Tropeognathus mesembrinus and Siroccopteryx
moroccensis.
Fletcher and Salisbury (2010) described four pterosaur speci-
mens from various localities throughout North West and
Central West Queensland (Fig. 1): an incomplete mandible
FIGURE 1. Maps of: A, Australia; B, Queens-
land, showing locations (circles) at which pre-
viously described pterosaur specimens were
found and the extent of outcrop from the
Rolling Downs Group. The star represents the
location where the new pterosaur described in
this study was found.
(QMF44423) discovered in a Toolebuc age-equivalent formation
east of Hughenden, assigned to an indeterminate ornithocheirid
(sensu Unwin, 2003); the proximal end of a left metacarpal IV
(QMF44321) from near Boulia, and the distal portion of a wing
phalanx (QMF44312) from between Richmond and Hughenden,
both from the Toolebuc Formation and assigned to cf.
Anhanguera; also described was the proximal end of a left
humerus (QMF42739) from south-west of Hughenden in the
slightly younger Mackunda Formation (Fig. 1), and tentatively
identified as an indeterminate ctenochasmatoid, which remains
the only described specimen of non-ornithocheiroid pterosaur
(sensu Andres et al., 2014) from Australia. Other pterosaur
fossils from the Toolebuc Formation include a complete right
metacarpal IV (NMV P197962) and a cervical vertebra (SAM
P41968) lacking most of the neural arch (Kellner et al., 2010).
Both specimens were found in isolation on Slashers Creek
Station east of Boulia and likely represent non-azhdarchid
ornithocheiroids, possibly anhanguerids.
Two isolated teeth (LRF 759 and LRF 3142) from the middle
Albian Griman Creek Formation of Lightning Ridge (Fig. 1)
assigned to Anhangueria indet. represent the only formally
described pterosaur material from New South Wales (Brougham
et al., 2017). Western Australian specimens represent the only
Upper Cretaceous pterosaur occurrences in Australia. A jaw
fragment (WAM 68.5.11) from the Cenomanian–Coniacian
Molecap Greensand near Gingin in south-western Western
Australia (Fig. 1), which Kear et al. (2010) tentatively assigned
 Richards et al.—New anhanguerid from Australia (e1946068-3)
to Ornithocheiridae, extended the known range of this clade in
the southern hemisphere through to the Upper Cretaceous.
Interestingly, an incomplete right proximal ulna (WAM 60.57)
from the upper Maastrichtian Miria Formation of north-
western Western Australia (Fig. 1) is unique in that it represents
the only known azhdarchid remains from Australia (Bennett and
Long, 1991). Other pterosaur specimens have been described
from Australia (Molnar and Thulborn, 1980), however their frag-
mentary nature renders them largely indeterminate.
Here we describe the crested mandible of a new anhanguerian
pterosaur from the Lower Cretaceous (upper Albian) Toolebuc
Formation of Richmond, North West Queensland. It is the
fourth named pterosaur species from Australia and, based on
crest size, represents the largest pterosaur currently known
from Australia, the largest mandibular crest known from any
anhanguerian, and the third largest anhanguerian pterosaur
known worldwide.
Institutional Abbreviations—AOD, Australian Age of Dino-
saurs, Winton, Australia; GLGMV, Guilin Longshan Geological
Museum, Guangxi, China; KK, Kronosaurus Korner, Richmond,
Australia; LRF, Australian Opal Centre, Lightning Ridge, Aus-
tralia; MN, Museu Nacional, Universidade Federal do Rio de
Janeiro, Rio de Janeiro, Brazil; MNHS, Museum of Natural
History Sintra, Sintra, Portugal; NMV, National Museum of Vic-
toria, Melbourne, Australia; QM, Queensland Museum, Bris-
bane, Australia; SAM, South Australia Museum, Adelaide,
Australia; SAO, Sammlung Oberli, St. Gallen, Switzerland;
SMNK, Staatliches Museum für Naturkunde Karlsruhe,
Germany; SMNS, Staatliches Museum für Naturkunde Stuttgart,
Stuttgart, Germany; SNSB-BSPG, Staatliche Naturwissenschaf-
tliche Sammlungen Bayerns-Bayerische Staatssammlung für
Paläontologie und Geologie, Munich, Germany; WAM,
Western Australia Museum, Perth, Australia.
GEOLOGICAL SETTING
Local fossicker Len Shaw discovered KKF494 in June 2011 at a
site known as the ‘water pond’ at the ‘8 mile pits’ (now known as
the ‘Free Fossil Hunting Site 1’), approximately 12 km NW of the
town of Richmond, North West Queensland (Fig. 1). Originally
developed as a quarry for road dressing, this site exposes a
heavily weathered 4–5 meter thick sequence of marls and coqui-
nas of the Toolebuc Formation. The Toolebuc Formation is part
of the Rolling Downs Group, a stratigraphic unit made up of a
continuous sequence of strata deposited within the Eromanga
Basin during the ‘mid’ Cretaceous, spanning the upper Aptian
to lower Turonian (Tucker et al., 2013) (Fig. 2). It records a
period of extensive marine inundation when shallow seas
covered approximately 60% of the Australian continent
(Frakes et al., 1987; Campbell and Haig, 1999).
The Toolebuc Formation crops out extensively within the
northern and central Eromanga Basin, covering an area of
approximately 300,000 km2 (Jiang et al., 2018). It is easily recog-
nized on the basis of a strong gamma-ray anomaly in wireline-
logging (Exon and Senior, 1976). Conformably overlain by the
Allaru Mudstone and overlying the Wallumbilla Formation
(Fig. 2), the Toolebuc Formation has an average thickness of 15
meters but reaches a maximum thickness of over 60 meters
near the center of the basin (Senior et al., 1975; Jell, 2013;
Jiang et al., 2018). Compared with other units within the
Rolling Downs Group, the Toolebuc Formation is considerably
thinner, which is interpreted as being the result of reduced volca-
nic activity (Jell, 2013). Comprising oil and kerigenous shales,
coquinitic and nodular limestones and minor labile sandstone,
the formation represents maximum flooding marine conditions
within an epeiric basin system (Henderson, 2004; Jell, 2013).
Based on palynological evidence, the Toolebuc Formation
correlates with the Coptospora paradoxa Zone and
Phimopolshawites pannosus Zone (Burger, 1982) and the lower
part of the Endoceratium ludbrookiae dinocyst Zone (McMinn
and Burger, 1986), indicative of a lowermost upper Albian age.
Further support for this age is given by Bralower et al. (1993),
who determined the Toolebuc Formation to contain the lower-
most upper Albian Axopodorhabus albianus NC 9B nannofossil
Zone.
Indicative of a productive high latitude marine ecosystem, the
Toolebuc Formation is richly fossiliferous and is one of Austra-
lia’s most prolific sources of Mesozoic marine fossils. It has
yielded enormous quantities of invertebrate remains including
molluscs, pelecypods, gastropods, ammonites and belemnites
(Day, 1969; Exon and Senior, 1976; Henderson and Kennedy,
2002). Numerous teleosts have also been found, including the
ichthyodectiforms Cooyoo australis (Lees and Bartholomai,
1987) and Dugaldia emmilta (Cavin and Berrell, 2019), along
with other predatory fish such as the aspidorhynchid
Richmondichthys sweeti (Bartholomai, 2004), and the pachycor-
mid Australopachycormus hurleyi (Kear, 2007a). The shark
fauna includes the cretoxyrhinids Archaeolamna kopingengis
and Cretalamna appendiculata (McHenry, 2009). Undoubtedly,
the best-documented finds include marine tetrapods such as
the endemic ichthyosaur Platypterygius australis (Zammit,
2010), the elasmosaurid plesiosaur Eromangasaurus australis
(Kear, 2007b), and the pliosaurid Kronosaurus queenslandicus
(McHenry, 2009). Various protostegid sea turtles including
the massive Cratochelone berneyi (Longman, 1915), and
Bouliachelys suteri (Kear and Lee, 2006), as well as the ubiqui-
tous Notochelone costata (Owen, 1882), which Molnar (1991)
described as the most common reptile in the Toolebuc For-
mation, are also frequently reported. Despite the marine deposi-
tional setting, several non-marine tetrapod fossils have also been
discovered within the Toolebuc Formation and likely represent
‘bloat and float’ carcasses from terrestrial habitats that existed
adjacent to the Eromanga Sea. These include remains attributed
to the ornithopod Muttaburrasaurus sp. (Molnar, 1996), an inde-
terminate titanosauriform (Molnar, 1991; Molnar and Salisbury,
2005; Poropat et al., 2017), along with several fragments thought
to pertain to indeterminate dinosaurs (McHenry, 2009).
In the Richmond area, the Toolebuc Formation is dominated
by extensive coquinas formed by fragments of the characteristi-
cally prismatic shells of the large oyster-like Inoceramus, along
with the smaller Aucellina hughendenensis (Day, 1969). These
coquinitic layers are interspersed with thin layers of either fine
labile sandstone or dark muddy shales, suggestive of dysoxic
and anoxic benthic conditions, respectively (Henderson, 2004).
Additionally, extensive beds of fish ‘mash’ consisting of teeth,
bones and scales from small osteichthyan fish are also common
(T.M.R. and P.E.S., pers. obs.).
MATERIALS AND METHODS
KKF494 is housed at Kronosaurus Korner Regional Museum
(an accredited Queensland State Government repository for sig-
nificant natural history and cultural artifacts), in Richmond,
North West Queensland, Australia. It was prepared by one of
the authors (P.E.S.) using mechanical methods. Where necessary,
cyanoacrylate and acetone-soluble acrylic resin (ethyl-methacry-
late copolymer) were used to consolidate and seal the preserved
bone.
Systematic Framework—Dispute over the taxonomic content
of Ornithocheiroidea (Kellner, 2003; Unwin, 2003) and the
appropriate use of Ornithocheiridae or Anhangueridae is con-
siderable and ongoing. The systematic and nomenclatural
issues at the core of the debate are well documented in the litera-
ture (e.g., Veldmeijer et al., 2006; Rodrigues and Kellner, 2013,
Andres et al., 2014; Vidovic and Martill, 2017; Jacobs et al.
2019; Pêgas et al., 2019; Holgado and Pêgas, 2020) and, as such,
 Richards et al.—New anhanguerid from Australia (e1946068-4)
a detailed treatment will not be presented here. This study
follows the phylogenetic framework of Holgado and Pêgas
(2020), which is presently the most recent and comprehensive
framework regarding the phylogenetic relationships of anhan-
guerians. This framework differs most notably from alternative
pterosaur phylogenies (e.g., Unwin, 2003; Andres et al., 2014)
in recovering Ornithocheirus simus in a basal position within
Ornithocheirae, and in nesting Tropeognathus mesembrinus
and Coloborhynchus clavirostris within Anhangueridae. As
such, Anhangueria consists of two clades: Hamipteridae and
Anhangueridae. The Hamipteridae, comprising Hamipterus
tianshanensis and Iberodactylus andreui, share two synapomor-
phies: well-defined parallel and forward curved striae and sulci
on the rostral region of the premaxillary crest and a rounded ros-
trodorsal expansion of the rostral margin of the premaxillary
crest (Holgado et al., 2019). Anhangueridae comprises three
main clades: Anhanguerinae, Coloborhynchinae and Tropeog-
nathinae. Anhanguerinae comprises the genus Anhanguera,
along with Maaradactylus kellneri, Cearadactylus atrox,
Liaoningopterus gui, Ludodactylus sibbicki, Guidraco venator
and Caulkicephalus trimicrodon, and is regarded as a sister-
group to Coloborhynchinae, with species within both clades
sharing the possession of a medially displaced upper fifth tooth
position. The clade Coloborhynchinae comprises Aerodraco
sedgwickii, Coloborhynchus clavirostris, Nicorhynchus spp. and
Uktenadactylus spp. and is supported by five synapomorphies:
lateral margins of the premaxilla and anterior expansion straight
and subparallel; a dorsal deflection of the palate forming an angle
of 90o; third pair of upper teeth twice the size of the fourth pair;
first pair of upper teeth slender, elongated and elliptical in cross-
section. Within this framework, Coloborhynchus clavirostris is
considered less closely related to Tropeognathus mesembrinus
and Ornithocheirus simus than to members of Anhanguerinae
(contra Andres et al., 2014; Longrich et al., 2018; Jacobs et al.,
2019). Tropeognathinae comprises Tropeognathus mesembrinus,
Siroccopteryx moroccensis, and the Australian taxa Ferrodraco
FIGURE 2. Stratigraphy of the Eromanga
Basin, western Queensland (after Syme et al.,
2016). The exact position of Thapunngaka
shawi gen. et sp. nov. KKF494 within the Toole-
buc Formation is presently unclear. Abbrevi-
ation: P/N/Q, Paleogene/Neogene/Quaternary.
lentoni and Mythunga camara. This clade is supported by three
synapomorphies: reduced lateral expansion of the premaxilla/
dentary, short teeth and premaxillary crests continuing to the
rostral tip. With regard to the present study, it is worth noting
that Ornithocheiridae (sensu Unwin, 2003) and Anhangueria
(sensu Andres et al., 2014) are both approximately equivalent
to Ornithocheirae (sensu Pêgas et al., 2019).
SYSTEMATIC PALEONTOLOGY
PTEROSAURIA Kaup, 1834
PTERODACTYLOIDEA Plieninger, 1901
ORNITHOCHEIROIDEA Seeley, 1870
ANHANGUERIA after Rodrigues and Kellner, 2013
ANHANGUERIDAE Campos and Kellner, 1985 (sensu
Holgado and Pêgas, 2020)
TROPEOGNATHINAE Holgado and Pêgas, 2020
THAPUNNGAKA SHAWI gen. et sp. nov.
(Fig. 3)
Holotype—The rostral portion of a mandible, KKF494.
Locality—The holotype was found at ‘Free Fossil Hunting Site
1’ (20°38′49.31′′S, 143°5′58.34′′E), approximately 12 km north-
west of Richmond, on Wanamara Country, North West Queens-
land, Australia.
Horizon—Toolebuc Formation, Rolling Downs Group, Lower
Cretaceous, upper Albian (Boreham and Powell, 1987).
Etymology—The genus names incorporate words from
Wanamara [wun-a-mah-ra], one of the Mayi languages
spoken by the people of the Wanamara Nation, on whose
Country the holotype was found. Wanamara Country encom-
passes the headwaters of the Flinders River east to Richmond,
south to Kynuna, west to the Williams River near Cloncurry
and north to Cambridge Downs and Dalgonally (Breen,
1981). The generic name incorporates thapun [ta-boon] and
 Richards et al.—New anhanguerid from Australia (e1946068-5)

FIGURE 3. Thapunngaka shawi gen. et sp. nov., KKF494, the rostral portion of a pterosaur mandible from the upper Albian Toolebuc Formation near
Richmond, north-west Queensland, Australia. Photo and schematic interpretation in: A and B, right lateral view; C and D, left lateral view; E and F,
dorsal view; G and H, rostral view. Hatched shading in D indicates surface breaks. Cranial direction is indicated by an arrow in A–F, and numbers
correspond to alveolar positions. Abbreviations: crd, crista dentalis; sf, shell fragment.

ngaka [nga-ga], the Wanamara words for ‘spear’ and ‘mouth’, Description
respectively. The specific epithet shawi honors Mr. Len Shaw,
Preservation—The specimen is an incomplete pterosaur man-
discoverer of the holotype. The name therefore means
dibular rostrum, comprising the rostral, symphysial portions of
‘Shaw’s spear mouth’.
the left and right mandibular rami (Fig. 3). Alveoli 1–10 are pre-
Diagnosis—Anhanguerian pterodactyloid with the following
served on the left dentary and 1–13 on the right. The specimen
apomorphies: dorsal surface of mandibular rostrum flat medially,
has suffered considerable ‘egg-shell’ fracturing due to lateral
lacking a sagittal groove; third and sixth pair of dentary alveoli
compaction, a common condition observed in pterosaur fossils.
largest in the mandibular toothrow, and subequal in size;
The ventral portion of the mandibular crest has undergone
dentary alveoli 6 through to 13 positioned laterally on the mand-
minor lateral deformation, as indicated by a slight bend to the
ible; dentary alveolar diastema increase gradually in size up to the
right. There is no evidence of further plastic deformation to
sixth alveolus; large increase in diastema between seventh and
other parts of the specimen. No in situ teeth were preserved.
eighth dentary alveoli (almost double the length of preceding dia-
The third and sixth alveoli on both sides are slightly crushed,
stema between sixth and seventh dentary alveoli). Thapunngaka
with the dorsal margin of their bony collars having partially col-
shawi can be further distinguished from other anhanguerians by
lapsed. The remaining alveoli show no signs of distortion and are
the following combination of characters: deep, ‘blade-like’ asym-
generally well preserved, though most contain varying amounts
metrical sagittal crest on the mandible with a gently concave
of matrix.
rostral margin (shared with Anhanguera robustus); mandibular
Where preserved, the cortical bone is thin (0.5–1.7 mm) and
crest beginning at the rostral tip (shared with Tropeognathus
brownish grey in color. When viewed laterally, both sides of
mesembrinus and Ferrodraco lentoni); very slight lateral expansion
the specimen show damage to the central portion of the mandib-
of rostral part of the mandibular rostrum (shared with tropeog-
ular crest. The cortical bone in this area is missing and, in some
nathines); rostral portion of the mandibular rostrum directed dor-
places, where bone is completely absent, small void spaces are
sally in lateral aspect (shared with Anhanguera spp.,
present. Several of these voids are partially filled with matrix.
Brasileodactylus araripensis and Guidraco venator).
 Richards et al.—New anhanguerid from Australia (e1946068-6)

A fracture near the ventral margin of the crest has been repaired TABLE 1. Measurements of preserved anterior portion of mandible of
with acrylic resin. Despite the mandibular crest missing a small Thapunngaka shawi gen. et sp. nov., in millimeters.
fragment of the ventral margin, the general outline is still
inferable. Length as preserved—right dentary 303
Although lacking some cortical bone, the rostral-most part of Length as preserved—left dentary 201
Maximum height of dentary crest 124
the mandible is well preserved and undistorted. Caudal to the Length of dentary crest >195
third pair of alveoli, the occlusal surface is fractured, presumably Width at:
due to compaction. Beginning between the third and fourth pair 1st pair of alveoli 16.6
of alveoli, a small fracture tapering in width from 1–0.4 mm 2nd pair of alveoli 14.4
extends caudally along the occlusal surface to the end of the pre- 3rd pair of alveoli 14.9
served portion of the mandible (Fig. 3E, F). Caudal to the tenth 4th pair of alveoli 12.3
5th pair of alveoli 8.2
alveolus, the left dentary has broken off, revealing the symphysial 6th pair of alveoli 7.6*
surface of the right dentary. Unfortunately, this area has suffered 7th pair of alveoli 6.7
considerable damage rendering it otherwise uninformative. 8th pair of alveoli 7.2
Immediately ventral to the fifth right alveolus, a small crescent- 9th pair of alveoli 7.5
shaped fragment of shell measuring 6.3 mm in length is pre- 10th pair of alveoli 8.1
Right alveoli (mesiodistal length × labiolingual width)
served. Based on its size, degree of curvature and the presence 1 8.3 × 7.9
of faint concentric undulations, it is interpreted as representing 2 10.0 × 10.2*
the broken margin of a molluscan valve, possibly belonging to 3 11.8 × 9.0
Aucellina hughendenensis, a buchiid bivalve frequently found 4 9.4 × 8.4
in this horizon (Day, 1969). 5 8.9 × 7.8
Osteology—The preserved portion of the mandible is extre- 6 10.8 × 9.4*
7 8.8 × 7.3
mely narrow mediolaterally relative to its dorsoventral depth. 8 10.3 × 8.8*
The right dentary, being the more complete, is 303 mm long, 9 8.6 × 6.8
whilst the left is 201 mm (Table 1). In dorsal aspect, the mandible 10 8.7 × 6.7
is 12.3 mm wide at a point level with the fourth pair of alveoli, 11 7.6 × 6.5
narrowing to 8.2 mm at the fifth pair of alveoli. Rostral to the 12 7.8 × 5.2
fourth pair of alveoli, the mandible slightly expands laterally, 13 7.6 × 5.0
Left alveoli (mesiodistal length × labiolingual width)
reaching a maximum mediolateral width of 16.6 mm between 1 8.3 × 7.5
the first pair of alveoli. 2 8.9 × 6.1
The left and right dentaries appear completely fused and the 3 11.8 × 9.4
mandibular symphysis cannot be discerned. In lateral aspect, 4 10.6 × 8.1
the caudal two-thirds of the dorsal margin of the mandible is 5 8.1 × 7.4
6 12.1 × 9.9*
straight, but the rostral third, beginning near the fifth pair of 7 10.4 × 7.4
alveoli, is deflected rostrodorsally at an angle of 20° relative to 8 10.5 × 7.1
the plane defined by the caudal occlusal surface. Rostral to the 9 9.7 × 7.5
mesial margin of the second pair of alveoli, the dorsal outline 10 8.6 × 5.9
of the occlusal surface becomes rostrodorsally convex. In Right dentary—diastemae distance
rostral aspect (Fig. 3G, H), the mandible is subtriangular in 1–2 5.5
2–3 8.4
shape, with the first pair of alveoli located on its dorsolateral ver- 3–4 7.3
tices. Despite a minor amount of fracturing on the occlusal 4–5 6.6
surface of the mandibular rostrum, there is no evidence of a sagit- 5–6 12.8
tal groove and the dorsal surface is flat. 6–7 12.2
The most striking feature of this specimen is the presence of a 7–8 26.1
large sagittal mandibular crest (see Fig. 3). The crest is rostro- 8–9 20.7
9–10 20.1
caudally asymmetrical in outline, with the rostral margin much 10–11 19.5
steeper relative to the caudal margin. In lateral aspect, the rostro- 11–12 17.9
ventral margin of the crest is gently concave, falling steeply from 12–13 10.1
a point immediately caudal to the tip of the rostrum, ventral to Left dentary—diastemae distance
the first and second pairs of alveoli, reaching a maximum depth 1–2 6.1
2–3 7.9
of 12.4 cm immediately ventral to the sixth pair of alveoli. The 3–4 4.1
caudoventral margin remains convex caudally until the ninth 4–5 6.9
pair of alveoli, where it is broken off. Immediately caudal to 5–6 9.8
this break is a relatively smooth, 20.5 mm long, section of cortical 6–7 12.7
bone that runs subparallel to the occlusal surface of the mandible 7–8 22.2
and widens dorsocaudally. The caudal end of this bone measures 8–9 20.4
9–10 18.1
4 mm in width and is deflected dorsally. This region is tentatively
interpreted as being the part of the dentary immediately adjacent *: approximate.
to the caudal end of the mandibular crest. If interpreted cor-
rectly, and taking the curvature of the preserved ventral margin
into consideration, the mandibular crest terminated at a point
just level with the ninth or tenth pair of dentary alveoli. mandibles with a similarly elongated mandibular symphysis
The preserved portion of mandible has 13 alveoli on the right and comparable patterns of alveolar size and spacing, such as
side and 10 on the left, indicating a minimum of 26 teeth in the some specimens assigned to Anhanguera (see Kellner and
lower jaw. As both dentaries are incomplete, it is difficult to Tomida, 2000; Veldmeijer, 2003), we conservatively estimate
verify the exact number of teeth that were originally present that KKF494 may have had 30–40 teeth in the mandible (15–20
and how far they extended caudally. However, when compared teeth in each dentary). All preserved alveoli have a mesiodistally
with presumably closely related taxa that preserve complete oval outline (Table 1). The first pair of alveoli is located slightly
 Richards et al.—New anhanguerid from Australia (e1946068-7)
caudal to the tip of the rostrum and is directed rostrodorsally.
The second through to the sixth alveoli are positioned laterodor-
sally on the dentary. Caudally, the remaining alveoli are posi-
tioned on the lateral surface of the dentary. The third pair of
alveoli is the largest, followed by the sixth pair. Together, the
sixth and third pairs of alveoli form the distinctive ‘double-
peak’ dentition observed in most ornithocheirids (sensu Unwin,
2003). The second, fourth, seventh and eighth alveoli are all sub-
equal in size, though smaller than the third and sixth alveoli, and
considerably larger than the first and fifth, which are also com-
parable in size. The first and fifth alveoli are approximately
only two-thirds the size of the third and sixth. Caudally, from
the ninth alveoli, mesiodistal alveolar length diminishes gradu-
ally. The interalveolar distance gradually increases in size caud-
ally until the sixth alveolus. The diastema between the seventh
and eighth alveoli is almost twice the length of the preceding dia-
stema (Table 1). Beginning at the fourth alveolus and continuing
caudally, the alveoli on the right side are positioned slightly more
caudally with regards to the corresponding alveoli on the left
side. With the exception of the first and second alveoli, alveoli
3–13 are surrounded by robust bony collars. Between the
second and seventh alveoli, the lateral dentigerous surface of
each dentary is deeply scalloped and pierced by several small for-
amina. The angular and splenial are not visible in KKF494.
Comparisons—The lightly built mandibular rostrum and the
extremely thin cortical bone with a smooth external surface
identify KKF494 as a pterosaur. Dentaries with teeth that have
an oval or elliptical cross section are typical for pterosaurs and
the strong resemblance of the specimen to the mandible of
anhanguerians such as Anhanguera robustus SMNK 2302 PAL
(Fastnacht, 2001) and Anhanguera sp. SAO 200602 (Veldmeijer
et al., 2005) verifies the pterosaurian identification. Among
toothed pterosaurs, the presence of a large mandibular crest
and a mandibular rostrum with triangular cross section are
restricted to some, but not all, anhanguerians. This, the relatively
slender proportions of the mandible, including the degree of
lateral expansion, and the size, spacing and orientation of the
alveoli are all diagnostic features of Anhangueria (sensu
Holgado and Pêgas, 2020). The anhanguerian affinities of the
specimen are also in accord with its Lower Cretaceous age.
KKF494 was therefore compared with a range of anhanguerians
and related taxa represented by mandibular material.
Regarding pterosaur material from Australia, most specimens
are isolated postcranial elements that are not comparable with
KKF494 (see Introduction for examples). Of the remaining
material, Mythunga camara (QMF18896; Molnar and Thulborn,
2007) is known from a partial skull comprising maxillary and
mandibular elements. Based upon a combination of characters,
Molnar and Thulborn (2007) assigned M. camara to Archaeop-
terodactyloidea (sensu Kellner, 2003). Critical to this assignment
was their interpretation that the first premaxillary teeth sat in
confluent alveoli, thus delineating the tip of the maxillary
rostrum and implying that the rostral end was nearly complete.
Noting that the dentition of M. camara was inconsistent with
any archaeopterodactyloid reported thus far, Fletcher and Salis-
bury (2010) and Kellner et al. (2010) concluded that the rostral
end of the specimen was incomplete (as opposed to nearly com-
plete). Fletcher and Salisbury (2010) suggested that the observed
alveolar pattern, characterized by larger teeth mesially and sets
of smaller and then larger sets of teeth distally, was more consist-
ent with Ornithocheiridae (sensu Unwin, 2003). A similar con-
clusion was reached by Pentland and Poropat (2019) and
further supported by Holgado and Pêgas (2020). We agree that
Mythunga camara belongs within Anhangueria (sensu Holgado
and Pêgas, 2020), and, as such, a comparison with KKF494 is war-
ranted. Unfortunately, the rostral end of the mandible is missing
on QMF18896, as are the ventral margins of each mandibular
ramus, such that comparisons are limited to the mandibular
dentition and alveoli. The base of the first lower tooth preserved
on M. camara is similar in size to the eighth right dentary alveolus
preserved on KKF494 (10.5 mm and 10.3 mm, respectively).
Caudally, the relative sizes of the alveoli in both specimens are
also comparable (see Molnar and Thulborn, 2007 for dental
measurements). Both specimens present alveoli that are labiolin-
gually compressed, set in a dentary that is deeply scalloped
between the alveoli for the reception of the occluding maxillary
teeth. The corresponding diastemae between alveoli is slightly
smaller in KKF494 than M. camara, though a gradual decrease
in interalveolar distance caudally is observed in both specimens.
If the first pair of lower teeth preserved on M. camara is inter-
preted as being the eighth or ninth pair, then the alveolar pat-
terns in both specimens are strikingly similar. However,
KKF494 is distinguished from Mythunga camara by having
bony collars present on all alveoli caudal to the second pair,
which, in the latter, is present only on the first preserved pair
on the maxilla. In KKF494, all alveoli caudal to the sixth pair
are positioned almost laterally on the dentary, contrasting with
the more dorsolaterally positioned alveoli on Mythunga
camara. Given the fragmentary nature of both specimens and
the subsequent lack of overlapping diagnostic features, it is not
possible to make further meaningful comparisons. Although
there is a general similarity between the alveolar patterns
observed in KKF494 and Mythunga camara, this is only diagnos-
tic at the level of Anhangueria. Until more complete material is
found that shows otherwise, we regard KKF494 to be distinct
from Mythunga camara.
A jaw fragment (WAM 68.5.11) from the Upper Cretaceous
(Cenomanian–Coniacian) Molecap Greensand of Western
Australia was described by Kear et al. (2010). The specimen is
extremely fragmentary and it is difficult to determine both its
precise orientation in the tooth row and whether it belongs to
the maxilla or dentary. Nevertheless, WAM 68.5.11 preserves
two adjacent alveoli which are ovate, labiolingually compressed
and relatively widely spaced. Based on these characters, and
despite its relative incompleteness, the authors tentatively
assigned WAM 68.5.11 to Ornithocheiridae (sensu Unwin,
2003). This combination of characters, along with alveoli that
are surrounded by robust bony collars, is also observed in
KKF494. However, the relative degree of labiolingual com-
pression with respect to mesiodistal length is much greater in
the alveoli of the Western Australian specimen. The characters
shared by WAM 68.5.11 and KKF494 indicate that they are
both referable to Anhangueria. However, until additional
material is discovered, further comparisons are purely speculat-
ive. We therefore do not currently regard these specimens as
conspecific.
KKF494 differs from Aussiedraco molnari (QMF10613;
Kellner et al., 2011) based on the presence of a large mandibular
crest and slight lateral expansion of the rostral end of the mand-
ible, both of which are lacking in the latter. Furthermore, the
dorsal margin of the mandibular rostrum in KKF494 is straight,
contrasting the markedly dorsally convex margin observed in
A. molnari, a character that Kellner et al. (2011) considered auta-
pomorphic. KKF494 also lacks the deep mandibular groove that
is evident on the occlusal surface of the mandibular rostrum of
Aussiedraco.
KKF494 and Ferrodraco lentoni are similar in that both possess
a relatively large mandibular crest and raised alveolar collars;
however, KKF494 differs from F. lentoni in that it does not
possess a mandibular groove and the rostral-most portion of
the mandible is slightly expanded laterally. By contrast, in
F. lentoni, the rostral-most portion of the mandible is less later-
ally expanded and CT scan data reveal a mandibular groove
that gradually deepens caudally, terminating at the caudal
margin of the mandibular rostrum (Pentland et al., 2019:fig.
4C–I). Furthermore, the rostral portion of the mandible is
 Richards et al.—New anhanguerid from Australia (e1946068-8)
dorsally deflected in KKF494, a feature also observed in Anhan-
guera spielbergi (Veldmeijer, 2003:fig. 4), “Anhanguera” robustus
(SNSB-BSPG 1987 I 47) (Pinheiro and Rodrigues, 2017:fig. 6H)
and Anhanguera piscator (Kellner and Tomida, 2000:figs. 4, 5),
which presumably allows for tight occlusion with the upper jaw,
which is similarly dorsally deflected. This contrasts with
F. lentoni in which the dorsal surface of the mandible is straight
in lateral aspect.
KKF494 can further be distinguished from F. lentoni based on
its alveolar morphology. In F. lentoni, the first pair of alveoli are
situated on the rostral-most margin of the mandible and, caud-
ally, the remaining alveoli face dorsally. By contrast, the first
pair of alveoli on KKF494 are situated more rostrolaterally,
with all the proceeding more caudal alveoli positioned almost lat-
erally along the dentary. Interalveolar distance increases caud-
ally in F. lentoni, unlike KKF494 where it increases from the
first to third pair of alveoli, then decreases from the third to
sixth pair, increasing again to the 13th (the last pair preserved
on the specimen). Considered autapomorhic to Ferrodraco
lentoni, the rostral-most alveoli of the mandible (and premaxilla)
are smaller than all the other rostral alveoli (Pentland et al.,
2019). This also differs from the condition seen in KKF494
where the first and fifth alveoli are subequal in size.
Finally, despite the incomplete preservation of the mandible
on both specimens, it is clear that the holotype individual of
F. lentoni was considerably smaller than KKF494. The mandibu-
lar symphysis of F. lentoni is 180 mm long, significantly shorter
than the minimum length of 303 mm observed on KKF494.
This difference may in fact be far greater given that the caudal
extent of the mandibular symphysis is not preserved on
KKF494. Similarly, the rostrocaudal length of the mandibular
crest in F. lentoni (max. 126 mm) is also much shorter than
KKF494 (min. 195 mm). Given the F. lentoni holotype is con-
sidered an ontogenetically mature individual based on the
fusion of several postcranial elements (e.g., scapulocoracoid
and wing phalanx IV–1) (Pentland et al., 2019), we consider it
unlikely that this difference is related to ontogeny. However,
we do convey some caution regarding the diagnostic utility of
size differences, as conspecific taxa can show considerable size
variability throughout ontogenetic stages (Prondvai et al., 2014).
In comparison with Chinese anhanguerians (or anhanguerian-
like pterosaurs), KKF494 is clearly distinct from boreopterines in
terms of its mandibular morphology. Contrasting to KKF494,
Boreopterus cuiae (Junchang and Qiang, 2005) possesses small
dentary alveoli that are closely spaced. Boreopterus cuiae also
lacks a diastema between the fourth and fifth pair of dentary
alveoli. Similarly, Feilongus youngi (Wang et al., 2005) bears a
greater number of alveoli, possessing 20 pairs in the rostral-
most 100 mm of the mandible (compared with five pairs in
KKF494). Zhenyuanopterus longirostris (Lu, 2010) and the pter-
odactyloid Haopterus gracilis (Wang and Lü, 2001) also differ
from KKF494 in that the orientation of their rostral-most pairs
of alveoli are almost perpendicular to the occlusal surface of
the dentary, contrasting with the dorsolateral orientation dis-
played by the corresponding pairs on KKF494. The rostralmost
part of the mandible of Liaoningopterus gui, a crested anhan-
guerian from the Jiufotang Formation (Wang and Zhou, 2003),
is more robust in comparison with that of KKF494. Despite
minor loss of the cortical surface due to poor preservation
(Rodrigues et al., 2015), this bulbous appearance is a result of
the rostral outline of the mandibular crest being convex; in con-
trast, the rostral outline of the mandibular crest of KKF494 is
concave in lateral aspect. Guidraco venator (Wang et al., 2012),
also from the Jiufotang Formation, shares with KKF494 (and
most anhanguerians) large rostral teeth in the lower jaw.
However, the teeth are lacking robust alveolar borders and
display a different alveolar pattern. In Guidraco venator, the
alveoli decrease in size caudally with the exception of the sixth
alveolus, which is much smaller than the fifth and seventh
(Wang et al., 2012). This is in stark contrast to KKF494 where
the first three alveoli increase in size caudally and the sixth alveo-
lus is larger than the fifth and seventh. It is also noted that the
entire mandible of Guidraco venator measures 330 mm in
length, which is only slightly larger than the preserved symphy-
sial length of KKF494, measuring 303 mm. However, given that
the holotype is the only known specimen of Guidraco and the
authors consider it a subadult animal, we acknowledge that this
difference may be due to ontogenetic variation and must be
viewed with caution.
The anhanguerian Hamipterus tianshanensis from the Turpan-
Hami Basin, north-western China, is known from dozens of
specimens, seven with complete lower jaws (Wang et al., 2014).
KKF494 clearly differs from all known lower jaws of Hamipterus
tianshanensis by lacking a well-developed mandibular groove, a
prominent rostral lateral expansion (similar to the condition
observed in anhanguerines), and a ‘hook-shaped’ bony process
on the tip of the dentary. KKF494 also has a minimum of 13
teeth along the length of the mandibular symphysis, whereas
Hamipterus constantly maintains 11 teeth throughout ontogeny
(Wang et al., 2014).
Possessing larger teeth (alveoli) rostrally and a large mandib-
ular crest, KKF494 is superficially similar to many crested anhan-
guerians from South America and comparisons with specimens
where the mandible is preserved are therefore warranted.
However, as the diagnostic utility of characters relating to the
morphology of crests remains debated (Bantim et al., 2015; Pin-
heiro and Rodrigues, 2017), comparisons were also made with
crestless anhanguerians from South America.
The rostral end of the mandible of KKF494 and Tropeognathus
mesembrinus (SNSB-BSPG 1987 I 46) are similar in that they are
not mediolaterally expanded or ‘spoon-shaped’ as in species of
Anhanguera. However, when viewed laterally, the rostral
outline of the crest in T. mesembrinus is convex, making the
overall crest appear symmetrical, differing from KKF494 where
the rostral outline of the mandibular crest is concave causing
an overall asymmetrical appearance. Furthermore, the mandibu-
lar crest of KKF494 is significantly dorsoventrally deeper (124 to
51 mm), and rostrocaudally longer (min. 195 to 126 mm) than
that of T. mesembrinus. The mandibular symphysis of KKF494
is also much longer rostrocaudally and contains a significantly
greater number of alveoli (minimum of 13 versus 8). The
alveoli are also positioned more laterally in KKF494 and, ros-
trally, show greater variation in size. The median dorsal surface
of the mandibular symphysis in KKF494 is smooth, and lacking
a mandibular groove, further distinguishing it from
T. mesembrinus where a deep mandibular groove is observed.
All species of Anhanguera possess a mandibular groove on the
median dorsal surface of the symphysis. This is in stark contrast to
KKF494 where the corresponding area is smooth and a mandibu-
lar groove is absent. Although the diagnostic utility of the mandib-
ular groove has been questioned by some authors (e.g., Elgin and
Frey, 2011), most of the ambiguity relates to subtle morphological
variations, and the extent to which the mandibular groove extends
caudally or rostrally (e.g., Veldmeijer, 2003), not its absence or
presence. Therefore, we consider the flat dorsal surface of the
mandible (i.e., absence of a mandibular groove) on KKF494 to
be a diagnostic character. However, it is noted that the presence
(or absence) of a mandibular groove in Anhanguera piscator
(Kellner and Tomida, 2000) cannot be assessed as the dorsal
surface of the mandible is still embedded in matrix.
Regardless of the diagnostic ambiguity surrounding the dorsal
surface of the mandible noted above, KKF494 further differs
from all species of Anhanguera with regards to its alveoli
pattern. The recently revised diagnosis of Anhanguera provided
by Pinheiro and Rodrigues (2017:16) includes the synapomorphy
“5th and 6th upper dental alveoli smaller than the 4th and 7th
 Richards et al.—New anhanguerid from Australia (e1946068-9)
ones”. Although this character clearly applies to the upper jaw,
Kellner and Tomida (2000:27) note that “this same variation in
size is also seen in the teeth of the lower jaw”. This variation is
clearly different to the pattern exhibited in KKF494 where the
third and sixth alveoli are larger than the fourth and seventh.
The alveoli pattern of KKF494 also differs to that observed on
members of Targaryendraconidae (i.e., Barbosania gracilirostris,
Aussiedraco molnari and Targaryendraco wiedenrothi) in which
the first mandibular alveolus is the largest and the diameter of
subsequent alveoli decreases caudally (Pêgas et al., 2019). Fur-
thermore, members of this clade are united through the posses-
sion of a relatively deep mandibular groove with lateral ridges
raised above the alveolar margin, extending to the level of the
first tooth pair. These features are not observed on KKF494.
Brasileodactylus araripensis (MN 4804-V), a crestless anhanguer-
ian, also clearly differs from KKF494 in possessing a mandibular
groove and having the rostral-most pair of alveoli positioned ros-
trally (Kellner, 1984).
Aetodactylus halli (SMU 76383) is the only cimoliopterid rep-
resented by mandibular material (Myers, 2010) and clearly
differs from KKF494 by exhibiting a relatively short (158 mm
long) mandibular symphysis that is strongly compressed dorso-
ventrally. Moreover, caudal to the fourth alveoli pair, the alveo-
lar spacing in Aetodactylus halli remains constant, whereas the
alveolar spacing in KKF494 greatly increases in this portion of
the mandible. Typical of all cimoliopterids (Pêgas et al., 2019),
the first three pairs of alveoli on Aetodactylus halli are closely
spaced, with the first two alveoli pairs being the largest. This con-
trasts with the alveolar pattern seen in KKF494 where the first
five pairs of alveoli are evenly spaced, with the third and sixth
teeth being the largest. KKF494 also lacks a mandibular groove
which, in Aetodactylus halli, extends caudally from the second
pair of alveoli to the caudal end of the dentary symphysis.
In light of these observations, we therefore regard KKF494 as
a distinct anhanguerian, with a unique mandibular morphology
not seen in other taxa described to date. Despite the fragmentary
nature of the specimen, we feel justified in erecting a new genus
and species, Thapunngaka shawi.
Phylogenetic Analysis—In order to assess the phylogenetic
relationships of Thapunngaka shawi within Pterosauria, we scored
it using two datasets. The primary phylogenetic analysis used the
dataset of Holgado and Pêgas (2020) (Supplemental Data S1); the
secondary analysis used a modified dataset of Andres et al. (2014)
(Supplemental Data S2). Both analyses were conducted using the
software TNT v. 1.5 (Goloboff and Catalano, 2016), following the
methodology outlined by the original authors (see Supplemental
Data S3 and Supplemental Data S4, respectively).
Using the dataset of Holgado and Pêgas (2020), the analysis
resulted in nine most parsimonious trees with a length of 416
steps (CI = 0.630, RI = 0.867) (see Fig. S1 for bootstrap
values). The strict consensus tree recovered Thapunngaka
shawi as an anhanguerid within Tropeognathinae, nested in
a polytomy with the Australian anhanguerids Mythunga
camara and Ferrodraco lentoni (Figs. 4, S2), supporting the
recent phylogenetic proposals of Pentland et al. (2019) and
Holgado and Pêgas (2020). The Australian anhanguerids are
united by a single synapomorphy: strongly raised alveoli
borders, a feature that is unique to the Australian taxa
among anhanguerians. Besides the inclusion of Thapunngaka
shawi, the remaining topology of our strict consensus tree
was nearly identical to that of the original tree presented by
Holgado and Pêgas (2020), with the exception that Anhan-
guerinae was slightly better resolved (Anhanguera piscator +
AMNH 2255 recovered as the sister group of Anhanguera
blittersdorffi; and Ludodactylus sibbicki + Guidraco venator
recovered as the sister group of Caulkicephalus trimicrodon).
However, in our analysis Boreopteridae was recovered as an
unresolved polytomy.
Tropeognathinae is supported by three synapomorphies: short
teeth (crown height under 3× diameter); premaxillary crest
reaching rostrum tip; and a reduced premaxillary/dentary expan-
sion width (under 130% post-rosette width) (Holgado and Pêgas,
2020). Because the holotype of Thapunngaka shawi comprises
only the rostral portion of a mandible (with no preserved
teeth), the first two synapomorphies cannot be assessed.
Although, if the mandible of Thapunngaka shawi presumably
reflects the condition seen in the upper jaw it could be argued
that the second synapomorphy (premaxillary crest reaching
rostral tip) is seen in the new Australian taxon. Regarding the
third synapomorphy, Thapunngaka shawi does share with
Ferrodraco lentoni and Tropeognathus mesembrinus a reduced
lateral expansion of the dentary, whereby the lateral margins
gently curve continuously to meet the post-rosette constriction,
thus supporting its position within Tropeognathinae.
Using the dataset of Andres et al. (2014), the analysis resulted
in a single most parsimonious tree with a length of 873.047 steps
(CI = 0.358, RI = 0.793) (see Fig. S3 for bootstrap values).
Despite being based on a diverging phylogenetic proposal (a dis-
cussion of which is beyond the scope of this paper), the secondary
analysis similarly recovered Thapunngaka shawi in a clade that
includes Mythunga camara and Ferrodraco lentoni within
Anhangueria (Figs. 4, S4; see Supplemental Data S5 for a brief
analysis of results).
DISCUSSION
Paleobiogeographic Implications
In both phylogenetic analyses, Thapunngaka shawi was con-
sistently recovered as an anhanguerian within a clade that also
includes Mythunga camara and Ferrodraco lentoni. These con-
vergent results support the anhanguerian affinity of
Thapunngaka shawi and the close relationship between
Australian anhanguerians. As the Australian specimens preserve
very little morphological overlap between each other, this
relationship is based solely on the shared presence of raised
alveolar borders. With regards to the results obtained from the
primary phylogenetic analysis using the dataset of Holgado and
Pêgas (2020), the Australian pterosaurs form a sister-group
with Tropeognathus mesembrinus from the Romualdo Formation
of Brazil and Siroccopteryx moroccensis from the Kem Kem
Group of Morocco. Given the cosmopolitan distribution of
anhanguerians during the Early Cretaceous (Upchurch et al.,
2015), the close relationship of the Australian taxa with anhan-
guerians from South America and North Africa is not surprising.
The erection of Thapunngaka now brings the total of named
Australian pterosaurs to four, and helps frame the earlier ideas
about the nature of the Australian pterosaur fauna. The place-
ment of Mythunga, Ferrodraco and Thapunngaka within Anhan-
gueridae suggests that anhanguerids were the dominant group of
Australian pterosaurs during the Early Cretaceous. Their close
relationship also hints at the presence of an endemic radiation
of Australian anhanguerians during this time. With its abundant
food sources and vast expanse, the Eromanga Sea would have
been an ideal geographic point of diversification.
Aussiedraco molnari is the only non-anhanguerian pterosaur
recognized from Australia. However, the ctenochasmatoid
humeral fragment (Fletcher and Salisbury, 2010) and the tenta-
tively assigned azhdarchid ulna from the upper Maastrichtian
of Western Australia (Bennett and Long, 1991) also possibly rep-
resent distinct non-anhanguerian Australian pterosaur taxa.
If these latter assignments are correct it is conceivable that at
least six pterosaur taxa, representing four distinct clades, were
present in Australia during the Cretaceous.
Given the taxonomic diversity of pterosaur assemblages seen
on other continents during the Early Cretaceous (Upchurch
 Richards et al.—New anhanguerid from Australia (e1946068-10)

FIGURE 4. Time-calibrated phylogenetic trees showing the relationship of Thapunngaka shawi gen. et sp. nov. within Lanceodontia. Taxon ranges,
denoted by the box adjacent to each taxon, include both the true stratigraphic range and uncertainty whereas the color of the box, in the online
version, reflects the paleocontinental origin of each taxon. Hatched box shows uncertain temporal range. A, based on the data matrix of Holgado
and Pêgas (2020), with Thapunngaka shawi gen. et sp. nov. included. B, based on the data matrix of Andres et al. (2014), with Thapunngaka shawi
gen. et sp. nov., Mythunga camara and Ferrodraco lentoni included. Note that outgroup relationships are not shown (see Figs. S2 and S4 for
details). New species proposed in this paper marked in bold. Abbreviations: Al, Albian; Ap, Aptian; Ba, Barremian; Be, Berriasian; Ca, Campanian;
Ce, Cenomanian; Co, Coniacian; Ha, Hauterivian; Ki, Kimmeridgian; Ma, Maastrichtian; Ox, Oxfordian; Sa, Santonian; Ti, Tithonian; Tu, Turonian;
Va, Valanginian.
 Richards et al.—New anhanguerid from Australia (e1946068-11)

TABLE 2. Estimated wingspan of Thapunngaka shawi (KKF494), based respectively on the mandibular symphysis length and wingspan ratios of
Tropeognathus mesembrinus (SNSB-BSPG 1987 I 46), Anhanguera piscator (NSM-PV 19892), Anhanguera araripensis (SNSB-BSPG 1982 I 89),
Anhanguera spielbergi (RGM 401 880) and Anhanguera robustus (SNSB-BSPG 1987 I 47). All measurements are given in meters. Abbreviations:
msl, mandibular symphysis length; ws, wingspan.

Ratio Estimated ws of Thapunngaka shawi

Specimen msl ws (msl/ws) (msl = 0.303) Reference
Tropeognathus mesembrinus 0.18* 5.6* 0.032 9.47 Wellnhofer, 1987; Kellner et al., 2013.
Anhanguera piscator 0.259 5.0 0.052 5.83 Kellner and Tomida, 2000.
Anhanguera araripensis 0.235* 4.7 0.05 6.06 Wellnhofer, 1985.
Anhanguera spielbergi 0.251* 5.9 0.043 7.05 Veldmeijer, 2003.
Anhanguera robustus 0.284* 6.2* 0.045 6.73 Wellnhofer, 1987; Pinheiro and Rodrigues, 2017.

*approximate.

FIGURE 5. Hypothetical outlines of Australian

pterosaurs showing relative wingspan sizes.
Human standing 1.8 m tall is used for scale. A,
Thapunngaka shawi gen. et sp. nov., KKF494;
B, Mythunga camara, QMF18896; C, Ferrodraco
lentoni, AODF876. Anhangueria silhouette
modified from Claessens et al. (2009:fig. 3d).
Human silhouette modified from Ristevski
et al. (2020:fig. 39).

FIGURE 6. Reconstruction of the skull of Tha-

punngaka shawi gen. et sp. nov. (KKF494).
 Richards et al.—New anhanguerid from Australia (e1946068-12)
et al., 2015) it is curious that representatives of other pterosaur
clades, such as Azhdarchoidea, are not well represented in
Australia. Conceivably, given the relative scarceness of the
Australian pterosaur record, and the fact that the vast majority
of Australian pterosaur specimens are found in rocks associated
with the Eromanga Basin, the lack of azhdarchoid material may
not be a true paleobiogeographic signal. Different habitat use,
preservational or sampling bias, or some other unknown cause
may account for the current paucity of azhdarchoid material in
the Australian fossil record.
Wingspan Estimation
Given that Thapunngaka shawi is known only from the rostral-
most portion of a mandible, any estimation regarding the wing-
span of this taxon must remain speculative until further material
is found. Nevertheless, a general estimation of wingspan can be
made by comparing a near-complete element of Thapunngaka
shawi (in this case, minimum mandibular symphysis length;
303 mm) to closely related pterosaurs with known symphysial
lengths and wingspans and scaling accordingly. The mandibular
symphysis lengths and wingspan estimates of anhanguerian pter-
osaurs were gathered from the literature (see Table 2). Results
show that there is a weak correlation between mandibular sym-
physis length and wingspan, with the ratio ranging from 0.032
in Tropeognathus mesembrinus (SNSB-BSPG 1987 I 46) to
0.052 in Anhanguera piscator (NSM-PV 19892). Using these
ratios, the estimated wingspan of Thapunngaka shawi ranges
from 5.83 m to 9.47 m, based on A. piscator and
T. mesembrinus, respectively. However, it is noted that the use
of Tropeognathus mesembrinus as a proxy for the symphysis/
wingspan ratio of Thapunngaka shawi may be problematic.
Tropeognathus mesembrinus has a very low symphysis/wingspan
ratio because it has a short mandibular symphysis (approxi-
mately one third of the mandibular length, with eight alveoli;
see Wellnhofer, 1987). Anhanguera piscator has a higher symphy-
sis/wingspan ratio because it has a longer symphysis (approxi-
mately half of the mandibular length, with 14 alveoli; see
references in Table 2). As Thapunngaka shawi has a similarly
elongated symphysis with at least 13 alveoli, we consider
Anhanguera piscator the more suitable proxy for symphysis/
wingspan ratio for estimating the wingspan of the new Australian
pterosaur. Therefore, disregarding the symphysis/wingspan ratio
obtained from Tropeognathus mesembrinus and given that the
mandibular symphysis of Thapunngaka shawi is incomplete, we
conservatively estimate that the wingspan of Thapunngaka
shawi was between 6 and 7 m, making it the largest known pter-
osaur from Australia (Fig. 5), and the third largest nominal
species of anhanguerian, after Tropeognathus mesembrinus
(approximately 8.5 m; Kellner et al., 2013) and Nicorhynchus
capito (approximately 7 m; Martill and Unwin, 2012), ever
reported.
CONCLUSION
Whilst the Australian pterosaur record remains comparatively
poor, it is improving. The erection of Thapunngaka shawi
(Fig. 6), Australia’s largest pterosaur and the world’s third
largest anhanguerian, contributes greatly to our understanding
of Australian pterosaur diversity and their paleobiogeography.
It confirms that large-crested pterosaurs, rivaling contempora-
neous South American, North African and European species in
size, were present in Australia during the Early Cretaceous.
Importantly, the similar anatomical characteristics shared
between Australian pterosaurs are indicative of a close relation-
ship between taxa and support the idea that there might have
been a radiation of Early Cretaceous pterosaurs that was
endemic to the Eromanga Basin. Finally, given the more
diverse pterosaur assemblages found in South America and
China, this new discovery continues to provide optimism that
additional Australian species will be found in the future.
ACKNOWLEDGMENTS
The authors thank M. Johnston and the staff at K.K. for allow-
ing T.M.R. access to the Thapunngaka shawi holotype. We also
thank Q.M. staff K. Spring, S. Hocknull and A. Rozefelds for
allowing T.M.R. access to the Mythunga camara and Aussiedraco
molnari holotype specimens. T.M.R. would like to personally
thank B. Andres, M. Habib, T. Rodrigues, D. Hone,
J. Ristevski, J. Nair, A. Faith, L. Mitchell, A. Romilio, V. Weis-
becker and A. Jannel for useful discussions and suggestions.
Special thanks to R. Molnar for providing unpublished material
and support. The Willi Hennig Society is thanked for making
TNT freely available. This research was supported by the Austra-
lian Government’s Australian Biological Resources Study
(ABRS) – National Taxonomy Research Grant Programme
(NTRGP). We express our gratitude to D. Hone and R. Pêgas
for their comments and suggestions that greatly improved this
manuscript. We also acknowledge the people of the Wanamara
Nation and their custodianship of the lands on which the holo-
type was found, and pay our respects to their Ancestors and
their descendants, who continue cultural and spiritual connec-
tions to Country. We recognize their valuable contributions to
Australian and global society.
ORCID
Timothy M. Richards http://orcid.org/0000-0002-0969-2635
Steven W. Salisbury http://orcid.org/0000-0003-4097-8567
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34:281–292.
Submitted January 4, 2021; revisions received May 31, 2021;
accepted June 14, 2021.
Handling Editor: Elizabeth Martin-Silverstone.