Historical Biology

An International Journal of Paleobiology

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Skull morphology of the enigmatic Genyornis

newtoni Stirling and Zeitz, 1896 (Aves,
Dromornithidae), with implications for functional
morphology, ecology, and evolution in the context
of Galloanserae

Phoebe L. McInerney, Jacob C. Blokland & Trevor H. Worthy

To cite this article: Phoebe L. McInerney, Jacob C. Blokland & Trevor H. Worthy (2024) Skull
morphology of the enigmatic Genyornis newtoni Stirling and Zeitz, 1896 (Aves, Dromornithidae),
with implications for functional morphology, ecology, and evolution in the context of
Galloanserae, Historical Biology, 36:6, 1093-1165, DOI: 10.1080/08912963.2024.2308212

To link to this article: https://doi.org/10.1080/08912963.2024.2308212

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HISTORICAL BIOLOGY
2024, VOL. 36, NO. 6, 1093–1165
https://doi.org/10.1080/08912963.2024.2308212

Skull morphology of the enigmatic Genyornis newtoni Stirling and Zeitz, 1896 (Aves,
Dromornithidae), with implications for functional morphology, ecology, and evolution
in the context of Galloanserae
Phoebe L. McInerney , Jacob C. Blokland and Trevor H. Worthy
College of Science and Engineering, Flinders University, Adelaide, SA, Australia

ABSTRACT ARTICLE HISTORY

The presence of Dromornithidae in the Australian Cenozoic fossil record was first reported in 1872, yet Received 20 November 2023
although eight species and hundreds of specimens are known, key information on their morphology Accepted 14 January 2024
remains elusive. This is especially so for their skulls, which contributes to a lack of resolution regarding KEYWORDS
their relationships within Galloanserae. The skull of the Pleistocene dromornithid, Genyornis newtoni, Skull morphology;
was initially described in 1913. Additional fossils of this species have since been discovered and Homology; Galloanserae;
understanding of avian skull osteology, arthrology, and myological correlates has greatly advanced. Musculus adductor
Here we present a complete redescription of the skull of Genyornis newtoni, updating knowledge on its mandibulae externus;
morphology, soft-tissue correlates, and palaeobiology. We explore the diversity within Dromornithidae Functional morphology
and make comprehensive comparisons to fossil and extant galloanserans. Furthermore, we expand on
the homologies of skull muscles, especially regarding the jaw adductors and address the conflicting
and unstable placement of dromornithids within Galloanserae. Findings support generic distinction of
Genyornis newtoni, and do not support the close association of Dromornithidae and Gastornithidae. We
thus recommend removal of the dromornithids from the Gastornithiformes. Considering character
polarities, the results of our phylogenetic analyses, and palaeogeography, our findings instead support
the alternative hypotheses, of dromornithids within, or close to, the Suborder Anhimae with
Anseriformes.

Introduction articulated fossils to be uncovered (see Stirling 1894 for origi­

The fossil evidence for an endemic Australian avian radiation
of evolutionarily distinct giant birds, the Dromornithidae,
shows this family had a long temporal range, extending from
at least the Eocene (Vickers-Rich and Molnar 1996) to the late
Pleistocene (Rich 1979; Wells and Tedford 1995; Murray and
Vickers-Rich 2004; Miller et al. 2005; Worthy et al. 2016a;
McInerney et al. 2022). Named taxa span a shorter interval,
however, and include late Oligocene-early Miocene
Barawertornis tedfordi Rich 1979; late Oligocene Dromornis
murrayi Worthy, Handley, Archer and Hand 2016a; middle
Miocene D. planei (Rich 1979); middle to late Miocene species
Ilbandornis woodburnei Rich 1979 and I. lawsoni Rich 1979;
late Miocene D. stirtoni Rich 1979; as well as possibly Pliocene
D. australis Owen 1872 (see Murray and Vickers-Rich 2004).
The youngest representative is Genyornis newtoni Stirling and
Zietz 1896, the only dromornithid species currently known to
have survived into the late Pleistocene; a range of dated and
undated fossil localities support a Pleistocene occurrence of
G. newtoni (e.g. as reviewed in Saltré et al. 2016). The most
recent and robustly dated site, Lake Callabonna shows
G. newtoni survived minimally 48 – ~45 thousand years ago
(Ka) in the north-eastern region of South Australia
(McInerney et al. 2022). Lake Callabonna is a unique fossil
locality in Australia and spans many square kilometres
wherein thousands of animals (mostly large mammals) were
mired. The unusual deposition has allowed for near-complete,
nal descriptions of Lake Callabonna). It is the type locality for
G. newtoni, and the source of the specimens described in this
paper.
Dromornithids are represented primarily by an abundance of
postcranial material (Rich 1979). Historically, cranial fossils were
rare, extremely fragmented, and/or inadequate for a comprehensive
morphological analysis (Murray and Megirian 1998; Worthy et al.
2016a). This is evidenced by the first described dromornithid skull
material: that of G. newtoni was reported by Stirling (1913; see
Appendix One: Figure A1), redescribed by Murray and Megirian
(1998), and again by Murray and Vickers-Rich (2004), despite
recognition of the specimen being ‘a friable mass of broken frag­
ments . . . [that] had suffered such great distortion from pressure
affecting it laterally that anything approaching a satisfactory recon­
struction was quite impossible’ (Stirling 1913, p. 111). Following
numerous attempts at stabilising and reconstructing the skull,
Stirling concluded that the specimen failed to yield important
information on shape and structure, although other described ele­
ments, i.e. quadrates and a mandible, were better preserved and
remained relatively informative. Regardless, the 1913 description of
this skull has formed the basis of all reconstructions of G. newtoni,
in association with interpretations that were misled by an early
consensus that the Dromornithidae had ratite affinities (see Owen
1874; Stirling and Zietz 1896, 1900, 1905; Stirling 1913; Lambrecht
1933; Rich 1979: fig. 1; Rich 1980; Murray and Vickers-Rich 2004;
Nguyen et al. 2010).

CONTACT Phoebe L. McInerney phoebe.mcinerney@flinders.edu.au College of Science and Engineering, Flinders University, Adelaide, SA, Australia
Supplemental data for this article can be accessed online at https://doi.org/10.1080/08912963.2024.2308212.
© 2024 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution,
and reproduction in any medium, provided the original work is properly cited. The terms on which this article has been published allow the posting of the Accepted Manuscript in a repository by
the author(s) or with their consent.
Published online 03 Jun 2024
1094 P. L. MCINERNEY ET AL.
Rich (1979) described five additional dromornithid species in
three genera, yet did not attribute cranial material to any taxa, nor
discuss Stirling’s (1913) skull material of G. newtoni, as noted by
Olson (1985). In 1998, the first skull materials for species of
Dromornis and Ilbandornis was described by Murray and
Megirian (1998), and those of G. newtoni redescribed despite the
authors not directly examining the fossils themselves (p. 74). They
concluded this latter material contributed important proportional
information for dromornithid morphology, identifying elements
which aligned the dromornithids with the Galloanserae. Crown
group galloanserans form two diverse orders of birds, that from
most to least basal, include the Megapodiidae, Cracidae,
Numididae, Odontophoridae, and Phasianidae – galliforms – and
the Anhimidae, Anseranatidae, and Anatidae – anseriforms. They
further suggested that G. newtoni closely resembled a goose (Anser,
Anatidae) or screamer (Chauna, Anhimidae) and concluded anseri­
form affinities; subsequent publications, therefore, treated the dro­
mornithids as anseriforms (see Nguyen et al. 2010; Angst and
Buffetaut 2017). Analysis of phylogenetic relationships among
extinct Galloanserae by Worthy et al. (2016b; 2017b) instead
resolved dromornithids as sister to the ordinal-level lineages,
Anseriformes and Galliformes.
Murray and Vickers-Rich (2004) discussed the skull mor­
phology of the dromornithids, and in 2016 new skull material
for species of Ilbandornis, Dromornis, and Barawertornis were
subsequently described (Worthy et al. 2016a). Additional key
dromornithid cranial characters were identified, and Worthy
et al. (2016a) reiterated the lack of informative skull material
available for G. newtoni. This is supported by the exclusion of
Genyornis newtoni from a study on dromornithid endocranial
anatomy by Handley and Worthy (2021) and recent observa­
tions of the original G. newtoni skull material (now SAMA
P10838). Although the ratite affinities of the Dromornithidae
have since been dismissed (Olson 1985; Murray and Megirian
1998; Murray and Vickers-Rich 2004; Mayr 2011, 2022a;
Worthy et al. 2017b), the lack of new skull material for
G. newtoni over the past 109 years has led to misconceptions
on morphology, derived from photos and descriptions of the
original skull (e.g. the shape of the upper bill), that have only
been partially resolved through inferences from other dromor­
nithids. As a result, despite the species being known for more
than one hundred years and one of the longest known and
best represented dromornithids – much of the leg, sternum,
pelvis, and vertebral column have been described (Stirling and
Zietz 1896, 1900, 1905; Stirling 1913; Murray and Vickers-
Rich 2004) – the morphology of the skull of Genyornis new­
toni requires a review. This is now possible due to discoveries
made during a series of expeditions to Lake Callabonna from
2013–2019 wherein multiple skull elements were recovered.
It is evident that an analysis of the skull of G. newtoni
based on modern knowledge of dromornithids and updated
literature on the skull of Aves is necessary, and so we present
detailed descriptions of the skull of Genyornis newtoni based
on the first new material discovered since 1913. These speci­
mens provide novel information on the morphology of the
braincase, upper bill, palate, quadrate, and lower bill. We
additionally use these skeletal elements as a framework of
osteological correlates for soft tissue structures which enable
justified inferences on kinetic capabilities, and skull myology
and syndesmology. This most especially relates to the adduc­
tor chamber – homological understanding of which has grown
substantially for Aves since the turn of the century, and has
been facilitated by digital dissections (e.g. Zusi and Livezey
2000; Holliday and Witmer 2007; Lautenschlager et al. 2014;
Jones et al. 2019; Smith-Paredes and Bhullar 2019). The
results of this study are used to refine the systematic place­
ment of dromornithids, make inferences on dromornithid
evolution, and infer the ecology of G. newtoni.
Materials and methods
Institutional Abbreviations. AMNH, American Museum of
Natural History, New York, U.S.A.; ANSTO, Australian Nuclear
Science and Technology Organisation, Sydney, New South Wales,
Australia; FU, Flinders University, Adelaide, Bedford Park, South
Australia; FUR, Flinders University Vertebrate Collection,
Palaeontology Laboratory, Flinders University, Adelaide, South
Australia; MHNT, Muséum d’Histoire naturelle de Toulouse,
France; NMV, Museums Victoria, Melbourne, Victoria; NTM,
Museum and Art Gallery of the Northern Territory, Alice Springs,
Northern Territory; QM, Queensland Museum, Brisbane,
Queensland; QVM, Queen Victoria Museum and Art Gallery,
Launceston, Tasmania; SAHMRI, South Australian Medical and
Health Research Institute, Adelaide, South Australia; SAMA, South
Australian Museum, Adelaide, South Australia; USNM, United
States National Museum, Washington D.C., U.S.A..
Specimen collection, preparation, and morphological analysis
We studied the morphology of the skull including the braincase,
upper bill, palate, parts of the jugal arches, hyoid skeleton,
lower bill, and quadrates of Genyornis newtoni from six speci­
mens (see Table 1), all from Lake Callabonna Fossil Reserve,
Pirlatapa and Adnyamathanha Country, South Australia,
Australia. For additional specimen photos, see Appendix One.
Specimens were collected during expeditions in 2013, 2014,
2018, and 2019, authorised by permits from the Department
of Environment, Water, and National Resources, Government
of South Australia (U26313–1, U26313–2, and in 2018 and 2019
File ref. DEWNRF-26892) and the South Australian Museum,
and were prepared at FU; fossils were mechanically cleaned of
matrix using both dental and PalaeoTools pneumatic Micro
Jack® tools and stabilised by infusion with ParaloidTM B-72
dissolved in acetone. Additional material of G. newtoni, stored
in FU and SAMA collections, were also examined, including
those available from the 1913 descriptions (see Stirling 1913).
Descriptions are based off morphological observations and com­
parisons with other dromornithids and galloanserans.
Measurements, made using Erskine Oral Care Dentagauge 2
callipers with results rounded to the nearest 0.1 mm, are given
in Appendix Two (Table A1).
Specimens SAMA P59516 and SAMA P59521 were scanned
(neutron beam scanning at ANSTO, Sydney by Joseph Bevitt
and µCT scanning at SAHMRI, Adelaide, respectively) to assess
internal structures although in neither case could the bone be
reliably distinguished from the infilling matrix (a mix of kaolin
and smectite clays, calcite concretions, and paraloid, as ascer­
tained by X-ray diffraction analysis, authors’ unpublished data
for SAMA P53833, field ID: Geny10) in subsequent reconstruc­
tions in Materialise Mimics (Materialise’s Interactive Medical
Image Control System) Innovation Suite (versions 18.0–22.0).
Assignment of skull material to taxa follows Murray and
Megirian (1998) for Dromornis planei, D. stirtoni, Ilbandornis
species, and Worthy et al. (2016a) for D. murrayi, Barawertornis

Table 1. Preserved skull elements, damage and taphonomic distortion in recently recovered Genyornis newtoni skull material from Lake Callabonna. Note that all of these
specimens are also associated with post-cranial material not described here.
Specimen
Number Preserved element(s)
SAMA Articulated skull: cranium, rostrum, mandible, quadrates,
P59516 pterygoids, paraglossum, basihyal, ceratobranchials, atlas,
axis
NMV Cranium, quadrate, mandible
P256893
SAMA Cranium, quadrate
P53830
SAMA Rostrum, mandible
P59517
SAMA Rostrum
P59521
SAMA Cranium, quadrate, mandible
P59520
and Ilbandornis ?woodburnei. All Genyornis newtoni skull mate­
rial is identified by its association with partial skeletons includ­
ing the diagnostic leg bones, comparison with material
described by Stirling (1913) and furthermore, G. newtoni is
the only dromornithid species known from the Pleistocene.
A three-dimensionally modelled Ilbandornis woodburnei cra­
nium was produced via segmentation of CT data in Materialise
Mimics (Materialise’s Interactive Medical Image Control System)
Innovation Suite 22.0 (for scan data, see Handley & Worthy, 2021),
digitally altered and reconstructed to be more complete using
Blender v. 2.93.2, and images of perspectives subsequently adapted
using Adobe Photoshop v. 24.
Comparative material
Dromornithid and galloanseran specimens were loaned to THW at FU
from SAMA, NTM, QM, and QVM. A range of comparative speci­
mens were also accessed using the Flinders University Vertebrate
Collection. Where specimens were unavailable for direct observation,
e.g. gastornithids and other fossil galloanserans, comparisons were
made using the available literature, photographs supplied by THW
(e.g. Gastornis giganteus, Presbyornis pervetus), and a scan of
Presbyornis pervetus (supplied by L. M. Witmer). All comparative
material used can be found listed in SI 1. All comparative specimens
including some with dried musculature intact, and targeted dissections
of single Gallus gallus, Anas superciliosa and Chenonetta jubata speci­
mens (unregistered), were used to support interpretations on the
myology which was derived from the published literature.
HISTORICAL BIOLOGY 1095
Brief description
Extensive deformation to the skull: some lateral compression, left side displaced caudally
relative to right; the mandible and quadrates are shifted and rotated caudally relative
to the cranium, but the quadrates remain articulated in the quadratic fossa; the
cranium is tilted rostroventrally, underlapping the caudally displaced rostrum; the
right mandibular ramus is heavily damaged on the lateral side; fragmentary
pterygoids are disarticulated caudally on either side of the cranium; the paraglossum
is displaced rostrally and crushed between the palatines and the medial side of the
right mandibular rami; the basihyal and ceratobranchials are disarticulated; and the
atlas and axis are together articulated but disarticulated from the skull.
Skull fragments include a mediolaterally compressed partial left caudal skull, separate
right quadrate, parts of a left ceratobranchial and mandible. The articulated left
quadrate is fragmented; the disarticulated right quadrate lacks the processus orbitalis
and has sustained damage to the lateral side and the medial apex of the condylus
mandibularis lateralis. A fragment of the rostral mandible preserves part of the left
ramus mandibulae and pars symphysialis, the fragmented articulated caudal
mandible preserves the processus medialis mandibulae and associated cotyla
medialis.
Only preserves the fragmented partial condylus occipitalis (occipital condyle) and partial
right quadrate. Of the quadrate, only the pars otica is without significant damage,
although a fragment of the orbital region is also retained.
Disarticulated elements. Near-complete rostrum but mediolaterally flattened, the height
of the rostrum and key features are still assessable, e.g. right palatinum largely intact.
There are two mandible fragments which represent the right and left ramus
mandibulae, partes intermedialis et caudalis. The right ramus has suffered much
damage with the processus medialis mandibulae and cotyla medialis poorly
preserved. The opposite ramus is completely eroded dorsally. Two fragments of shaft
from one ceratobranchial, no ends preserved.
Disarticulated and disassociated rostrum. Near complete specimen with only slight
bilateral compression mid-way along the flattened lateral plates of the praemaxillaria,
and thus, forms the main basis of the G. newtoni rostrum descriptions herein.
Fragmented section of the left side of the caudal skull. Preserves the osseus meatus
acusticus externus and processus paroccipitalis, and a partial left quadrate remains in
articulation; the pars otica is well preserved, pars mandibularis is heavily abraded, and
the articulatory region of the mandible, is crushed. A fragment of the right ramus
mandibulae, pars symphysialis, is also preserved.
Nomenclature
Higher taxonomic nomenclature follows Worthy et al (2017b;
2017a; Sun et al. 2017) except we use Galloanserae due to taxo­
nomic priority and thus, galloanseran, as the adjectival form and to
refer to members of Galloanserae. In addition, the Superfamily
Anatoidea, and its distinction with respect to the Suborder
Anseres, is used in the sense of Livezey (1997), as advocated by
Field et al. (2020). For anatomical nomenclature, we follow Baumel
and Raikow (1993) unless indicated otherwise. Selected updated
and synonymised terminology, i.e. for the bony palate proposed by
Zusi and Livezey (2006), the maxillary bone by Mayr (2018a), the
quadrate by Elzanowski et al. (2001) and Elzanowski and Stidham
(2010), and the myology of the adductor chamber by Holliday and
Witmer (2007), has been used when necessary. The prefix os is
omitted from the names of skull bones and the term cranium is
used to refer specifically to the neurocranium or braincase as per
Baumel and Witmer (1993: annot. 8a, p. 68), not to be mistaken for
the term skull, which is used here to refer to the entire head skeleton
including the cranium, jaws, quadrates, hyoid apparatus, and other
associated bones of the combined neurocranium and splanchno­
cranium (see Zusi 1993). While maxilla refers to the total structure
comprising the upper jaw (Baumel and Witmer 1993: annot. 93;
Zusi 1993, p. 394–395), we opt to use the term ‘rostrum’ herein
when referring to this complex (as per Mourer-Chauviré and
Balouet 2005), to avoid confusion with the maxillary bones (ossa
maxillaria) that participate in its composition (however, see Clarke
1993: annot 12, regarding the proper anatomical use of the term
rostrum). As per Mayr (2018a), we follow the correct Latin spelling
1096 P. L. MCINERNEY ET AL.
for the bilaterally paired upper jaw bones, the praemaxilla, rather
than ‘premaxilla’. For myological homologies, see Appendix Three
(Table A2), additional skeletal homologies are present in SI 2.
A notable subject of contention is the use of osteological terms
associated with the ‘fossa temporalis’ or ‘temporal fossa’, resulting
from imprecisions in referring to it through interpretation of the
associated myological homology in different clades (see Zusi and
Livezey 2000, pp. 165–166). In general, the temporal fossa accom­
modates adductor muscle origins in various configurations across
different groups of Neornithes, including musculus pseudotempor­
alis superficialis, bellies associated with musculus adductor mandi­
bulae externus (AME) profundus, and musculus AME superficialis
(using terminology of Holliday and Witmer 2007), the latter of
which being the most superficial of the external adductor muscu­
lature and attaching to the caudolateral region of the cranium (e.g.
see Goodman and Fisher 1962; Elzanowski 1987; Baumel and
Witmer 1993; Vanden Berge and Zweers 1993; Weber 1996; Zusi
and Livezey 2000; Holliday and Witmer 2007). Use of this term has
also included association with musculus depressor mandibulae (van
Gennip 1986, p. 5; Baumel and Witmer 1993: fig. 4.1; see below). To
address this issue, Zusi and Livezey (2000) advocated the use of
‘fossa muscularis temporalium’ as well as terms that specifically
relate to the musculature occupying each region (e.g. impressio
musculi adductor mandibulae externus (AME) profundus, pars
coronoideus, using applied nomenclature of Holliday and Witmer
2007), which we support. However, when referring to the complete
region of attachment for the musculus AME on the lateral squa­
mosum, we opt to use the generalised topographic term ‘temporal
fossa’ (Holliday and Witmer 2007). Additionally, following
Holliday and Witmer (2007), directional terms are used to refer to
specific structural regions within this (i.e. ‘dorsotemporal fossa’ and
‘caudotemporal fossa’). Hereon, in names for certain muscles and
nerves, ‘musculus’ (plural ‘musculi’) is abbreviated to ‘m.’ (‘mm.’)
and ‘nervus’ to ‘n.’ respectively.
The term ‘fossa subtemporalis’ (impressio temporalis, sensu
van Gennip 1986) is similarly ambiguous. Following Zusi and
Livezey (2000, p. 166), we refer to this depression with
regards to the associated musculature, although note that
here (in the context of the Dromornithidae) it is instead
solely related to the origin of m. depressor mandibulae (i.e.
impressio musculi depressor mandibulae).
Phylogenetic analyses
To qualitatively assess some of the hypotheses present herein,
and to precede future, more extensive analyses assessing the
phylogeny of Dromornithidae and their relationships to other
galloanseran families, we only present preliminary analyses
using a limited character and taxon sample. Our ingroup
includes 22 modern species from across the Galloanserae radia­
tion and 12 relevant fossil taxa that are hypothesised to have
galloanseran or near-galloanseran affinities, while the outgroup
consists of three palaeognaths (Eudromia elegans, Dromaius
novaehollandiae and Casuarius casuarius) and four neoavians
(Antigone rubicunda, Fulmarus glacialoides, Accipiter fasciatus
and Colius striatus). We have reassessed and added to the
characters of Worthy et al. (2017b; see Appendix Four) and
produced a character matrix comprising characters of the skull
only. These 100 morphological/standard discrete characters
address variation across the complete skull (3), cranium (28),
lacrimal (3), quadrate (22), mandible (29), and rostrum (15). Of
these characters, 51 are ordered. Missing data was coded as ‘?’,
and no gaps (‘-’) were coded. See SI 3 and 4 for the correspond­
ing NEXUS files that were used in phylogenetic analyses. These
analyses are summarised below. In consideration of the likely
extensive convergence in postcranial material and the modifica­
tions derived from the giant, flightless nature of the dromor­
nithids, a reassessment of dromornithid postcranial morphology
is required to inform on the characters and methods used in a
more complete analysis. This is beyond the scope of this study
which specifically focuses on testing the phylogenetic informa­
tion pertaining to the skull.
Currently, a lack of directly comparable skull elements for
some dromornithid species (resulting in missing data) and our
exclusion of postcranial material from these analyses, restrict
quantitative assessment of intrafamilial dromornithid relation­
ships. These phylogenetic analyses are thus focused on testing
the interfamilial relationships in the context of Galloanserae.
All analyses in the present study were constrained to mole­
cular-based topological relationships for modern taxa follow­
ing a combination of Wang et al. (2013), Burleigh et al.
(2015), Prum et al. (2015), and Kimball et al. (2019), and
appropriated to the taxon-selection herein at specific or gen­
eric levels. Molecular-based constraints for Anatidae were
relaxed following Worthy et al. (2022).
Parsimony analyses were conducted using PAUP*
v. 4.0a169 (Swofford 2003). The search for optimal trees
involved a heuristic approach, with 10,000 replicates of ran­
dom stepwise taxon addition using the tree-bisection-
reconnection (TBR) branch-swapping algorithm, holding 10
trees at each step, and saving no more than 100 trees of
a length greater than or equal to 1 in each replicate.
Subsequent bootstrapping involved 10,000 bootstrap replicates
with the following parameters: 100 random-addition sequence
replicates per bootstrap replicate; no more than 1,000 trees at
a score equal to or greater than 1 saved per bootstrap repli­
cate; holding 10 trees each step; and TBR branch swapping
implemented. Multistate characters were treated as
uncertainty.
An undated Bayesian analysis of the morphological data
was also performed using MrBayes v. 3.2.7 (Ronquist and
Huelsenbeck 2003), appropriating the same character and
taxon selection, ordering settings, and molecular-based topo­
logical constraints as the parsimony analysis. The Mk model
(Lewis 2001) was used to apply maximum likelihood phylo­
geny inference to the variable, discrete morphological dataset
(coding = variable). Evolutionary rate variability was distribu­
ted according to gamma parameter (rates = gamma). Four
independent analyses were simultaneously run for a total of
50,000,000 generations, and sampled every 5,000 generations,
to confirm convergence. The heating parameter was set as 0.1,
and four chains per analysis, one cold and three incrementally
heated, were used to better explore the tree topology space.
The first 20% of sampled trees from all runs were discarded
as relative burn-in, and the remaining samples combined to
produce a consensus tree.
 HISTORICAL BIOLOGY 1097

Figure 1. The near-complete, articulated skull of Genyornis newtoni (SAMA P59516): A. Left lateral view; B. Left lateral view outlined with major parts distinguished; C. Right
lateral view; D. Right lateral view outlined with major parts distinguished; E. Dorsal view; F. Caudal view; G. Ventral view; H. Rostral view. Annotations: cb., ceratobranchial;
cr., cranium; ma., mandible; or., orbit; pt., pterygoid; ro., rostrum; qu., quadrate; ve., articulated vertebrae. Scale bars: 50 mm, E. to G. all to same scale.
1098 P. L. MCINERNEY ET AL.
Systematic palaeontology
AVES Linnaeus, 1758
NEORNITHES Gadow, 1892
NEOGNATHAE Pycraft, 1900
GALLOANSERAE Sibley, Ahlquist and Monroe, 1988
DROMORNITHIDAE Fürbringer, 1888
GENYORNIS Stirling and Zietz, 1896
GENYORNIS NEWTONI Stirling and Zietz, 1896
Referred material
Lake Callabonna Fossil Assemblage. SAMA P59516, articulated
cranium, rostrum, quadrates, pterygoids, elements of the hyoid
apparatus, and mandible associated with a complete postcranial
skeleton of individual 1 of CB2018.75 field collection number.
SAMA P59517, flattened rostrum and part mandible associated
with several postcranial elements from individual 2 of CB2018.75.
SAMA P59520, fragmented, articulated partial left cranium, quad­
rate, and caudal mandible, disarticulated rostral mandibular frag­
ment, and other associated cranial fragments, associated with
postcranial elements and gizzard stones of individual 3 of
CB2019.14. SAMA P59521, complete rostrum associated with post­
cranial elements from either individual 1 or 2 of CB2019.14. NMV
P256893, partial articulated left side of cranium, jugal arch, cerato­
branchial, quadrate and caudal mandible, disarticulated right par­
tial quadrate, jugal arch, and cranial and mandibular fragments, all
associated with the near complete, articulated postcranial skeleton
of individual CB2018.23. SAMA P53830, right partial quadrate
with disarticulated caudal part of the jugal arch and condylus
occipitalis, associated with postcranial elements of individual
CB2014.Geny5. Skull fossil previously attributed to Genyornis new­
toni include SAMA P10838, a fragmented skull, SAMA P10788, a
partial mandible and unregistered quadrates (Stirling 1913).
Descriptions and comparisons
Fossils
Complete fusion of the constituent bones of each of the
cranium, rostrum and mandible is evident in all specimens
signifying that they belonged to mature individuals (Zusi
1993). This is supported for specimen SAMA P59516 by
histological interpretations by Chinsamy and Worthy (2021:
Table 2) of the leg bones. All fossils described herein are
variably affected by crushing, deformation, and fragmentation
(see Table 1), which limits morphological interpretation,
although together they allow for a reliable and detailed
description of nearly all features of the skull of G. newtoni.
Descriptions are facilitated by comparisons with the morphol­
ogy of other dromornithid fossils throughout.
Proportions
The articulated skull of specimen SAMA P59516 (Figure 1), pro­
vides a basis for proportional estimates of the skull in this species.
Specimen SAMA P10838 (see Appendix One: Figure A1) was,
according to the original descriptions, 290 mm long from the con­
dylus occipitalis to the symphysial apex of the mandible, and 150
mm high (Stirling 1913, p. 112). Comparatively, the most complete
skull described here, specimen SAMA P59516, is slightly larger,
approximately 296 mm in length from the rostral apex of the
rostrum to the condylus occipitalis. This is perhaps slightly fore­
shortened by the disarticulation of the craniorostral hinge (sensu
Mourer-Chauviré and Balouet 2005) and the caudal movement of
the rostrum to overlap the cranium by maximally 33 mm. The
maximum height of the cranium from the dorsal surface to the
ventral tip of the processus paroccipitalis is 135.5 mm. Our obser­
vations support Stirling’s contention that only a total length and
maximum depth of the skull could be ascertained from specimen
SAMA P10838 due to poor preservation limiting identification of
the margins of important structural features. However, Murray and
Megirian (1998) inferred goose-like proportions of the skull with
the rostrum accounting for 50% of complete rostrocaudal skull
length. Contrary to this, specimen SAMA P59516, instead shows
that the rostrum is closer to 1.8 times the length of the cranium.
The proportions of the cranium and rostrum within the skull
of G. newtoni differ from other dromornithids; the rostra of
Dromornis stirtoni and D. planei are near three times as long as
the cranium. This is most extreme in D. stirtoni, wherein the
entire skull would have been approximately just over 500 mm
long with 460 mm taken up in beak length (Murray and
Megirian 1998: fig. 28). There are currently no rostra known
for species of Ilbandornis or Barawertornis, precluding compar­
isons. Marked rostrocaudal compression of the cranium is char­
acteristic for dromornithids, with the dorsoventral depth of the
cranium considerably greater than the rostrocaudal length
(Worthy et al. 2016a; Handley and Worthy 2021). This feature
is increasingly extreme in Dromornis species; least in
D. murrayi, greater in D. planei, and especially so regarding
D. stirtoni, wherein the cranium length of the latter is only half
the dorsoventral depth, with the impact of this also evident in
the subtle signs of brain compression (Handley and Worthy
2021). Although, as in Ilbandornis, some rostrocaudal compres­
sion is evident, the cranium of G. newtoni lacks such exagger­
ated compression and, as no cranium is known for the early
Miocene Barawertornis tedfordi, a trend, if any, in non-
Dromornis dromornithids cannot be ascertained.
Craniorostral hinge
The skull of G. newtoni is prokinetic (the rostrum is inflexible
but bending occurs at a craniofacial hinge: see Hofer 1949;
Frazatta 1962; Bock 1964; Bühler 1980; Bühler et al. 1988;
Gussekloo et al. 2001; Pecsics et al. 2017). The zona flexoria
craniofacialis is one of the primary flexion zones associated
with cranial kinesis in birds (Fisher 1955; Bock 1964; Bühler
1980; Zusi 1984, 1993; Bühler et al. 1988; Baumel and Raikow
1993; Bailleul et al. 2017), generally composed of overlapping,
thinned premaxillary and frontal processes of the nasalia
(nasals) and praemaxillaria (premaxillaries), respectively,
from the rostrum, which are fused with the rostral ends of
the frontalia (frontals) and the dorsal lamina of the meseth­
moidale of the cranium, and sometimes stabilised by laterally
bounding lacrimalia (lacrimals; Fisher 1955; Bock 1964; Bühler
1980; Zusi 1984; Baumel and Raikow 1993). This region is
instead represented by a completely mobile synovial joint
(Bühler 1980, p. 449, 451–452; as defined by Baumel and
Raikow 1993: Arthr. Intro., annot. 1, 4, 46) in Genyornis new­
toni, other dromornithids, gastornithids, and sylviornithids
among galloanserans (Witmer and Rose 1991; Andors 1992;

Murray and Megirian 1998; Worthy 2000; Mourer-Chauviré
and Balouet 2005; Worthy et al. 2016a). As such, specific
referral to this articulation as a craniorostral hinge (Murray
and Vickers-Rich 2004; craniorostral joint, sensu Mourer-
Chauviré and Balouet 2005) throughout this document, is
considered more appropriate for these taxa. This morphology
is rare among bird families and is comparable to that seen in
some Psittaciformes (and others e.g. Podargidae and
Microcarbo melanoleucos; Andors 1992; Zusi 1993).
The only specimen which preserves the craniorostral hinge
in near articulation is SAMA P59516, although the joint is
overlapped by the dorsocaudally displaced rostrum, and thus
not visible in its entirety. The articular surface on the dorso­
caudal rostrum is preserved in specimens SAMA P59521 and
SAMA P59517 (Appendix One: Figure A2), although signifi­
cantly damaged in the latter. The caudal regions of the prae­
maxillaries and the nasals (and the lacrimals, see below) are
synostosed to form a surface for articulation with the cranium.
This is less robust and appears proportionally more dorsoven­
trally narrow and caudally flattened than that of Dromornis
planei (NTM P9973–2), in which, robust, bulbous and caudally
prominent lateral articular ‘condyles’ (internal processes of
nasolacrimals, sensu Murray and Vickers-Rich 2004: fig. 181,
184) lie either side of a distinct medial fossa. These condyles
were considered to be reciprocal structures to the deep lateral
fossae seen on several dromornithid crania (which also were
identified for Sylviornis neocaledoniae Poplin 1980 and
Megavitiornis altirostris Worthy 2000 by Mourer-Chauviré and
Balouet 2005; D. planei and D. stirtoni, by Murray and Vickers-
Rich 2004: figs. 52, 76, 77), and may be formed in part from the
frontal processes of the nasal bones. Unlike the completely
mobile, true diarthrosis in S. neocaledoniae, the dromornithid
form would have provided relative lateral stability for the cra­
niorostral hinge, while allowing significant free movement of
the rostrum in the occlusal plane (Murray and Vickers-Rich
2004, p. 235; as was similarly observed for M. altirostris by
Mourer-Chauviré and Balouet 2005). Genyornis newtoni appears
to have homologous condyles on the dorsocaudal rostrum (see
4.6), although they are much smaller evidencing less stability
was required for the relatively smaller rostrum.
The medial section of the dorsocaudal-most rostrum is
occluded in specimens SAMA P59516 and SAMA P10838, and
post-mortem damage is present in this region in specimen
SAMA P59521. Thus, the presence or absence of a sulcus to
receive a median prominence as observed in D. planei (see
median process/tuberosity of Murray and Vickers-Rich 2004:
figs. 52, 76, 77, 181–184), cannot be confidently assessed.
However, the median part of the dorsocaudal rostrum of
SAMA P59521 has an abraded surface approximately rostro­
caudally-level with the previously described articulatory con­
dyles and suggests that in a better-preserved state it would
have caudally exceeded these condyles and dorsally overlapped
any median prominence of the dorsal cranium. In SAMA
P10838 and P59516, there is no evidence of an obvious dorsal
depression on the rostral cranium that participates in the cra­
niorostral articulation, opposing the median dorsocaudal ros­
trum (see ‘frontal groove’ of Murray and Vickers-Rich 2004,
p. 235), however, this area is not fully exposed in either speci­
men. The craniorostral hinge divides the rostrocaudal skull
length into two dorsally convex sections (as viewed laterally),
HISTORICAL BIOLOGY 1099
the rostrum and the cranium; when the rostrum is occluded
with the mandible, the hinge would form a shallow dorsal notch
in lateral aspect, becoming deeper as the upper jaw is opened
(also see Murray and Vickers-Rich 2004). In contrast, recon­
structions of Gastornis giganteus (Cope 1876) suggest the tran­
sition in lateral aspect across the hinge is flat (see Witmer and
Rose 1991: fig. 1).
The caudal foreshortening of the anteorbital region of the
cranium and the concomitant caudal repositioning of the cra­
niorostral hinge in dromornithids (Murray and Megirian 1998;
Murray and Vickers-Rich 2004, p. 126; Worthy et al. 2016a) is
evident in G. newtoni. The result is a lack of preorbital zone in
the cranium, contrasting with all other galloanserans. The hinge
transects the orbit just rostral to the processus postorbitales in
G. newtoni but is located further caudal in species of Dromornis,
in rostrocaudal alignment with the process.
There have been no lacrimals previously unambiguously identi­
fied for G. newtoni; the lacrimal and its processes identified and
figured for the original skull (SAMA P10838) by Murray and
Vickers-Rich (2004: fig. 107, p. 60, 127) cannot be verified by direct
examination of the relevant material. Furthermore, we do not
recognise any separate and unfused lacrimals in the new skull
material presented herein. Murray and Vickers-Rich (2004,
p. 127–128) did not find evidence of orbital processes of the lacri­
mals in other dromornithids. However, they inferred that the
supraorbital processes may have fused to the dorsorostral cranium
as the ‘anterolateral processes’ and that the lacrimals contributed to
the lateral dorsocaudal rostrum as ‘nasolacrimal processes’ (Murray
and Vickers-Rich 2004, p. 235, fig. 180, 184). Instead, considering
the material at hand, we interpret the area caudolateral of apertura
nasi ossea as representative of the synostosis between the head of
the lacrimal and the caudal rostrum, whereby the caudoventral
projection on the rostrum dorsal of angulus tomialis and laterally
adjacent to processus jugalis of the maxillare, may be homologous
with the processus orbitalis of the lacrimal. This interpretation is
supported by the distinct, dorsoventrally elongate process on the
caudal margin of the Dromornis planei rostrum NTM P932–2. The
latter is linked with the more ventral arcus jugalis by a narrow
channel and appears ventrally separate from the more rostral
nasal bar, forming an opening (i.e. fenestra antorbitalis), and is
consistent with a ventrocaudal process of the lacrimal.
Thus, we do not support the hypothesis regarding fusion of
the lacrimal to the cranium in dromornithids. This lack of
synostosis is typical of basal galliforms, such as megapodiids,
and some basal anseriforms, i.e. Anhimidae,
Anachronornithidae, Anseranatidae, Presbyornithidae, and
even basal anatids, e.g. species of Biziura, Dendrocygna, and
Coscoroba (see Olson and Feduccia 1980; Zelenkov and
Stidham 2018; Tambussi et al. 2019; De Mendoza et al. 2020;
Field et al. 2020: SI p. 26–27; Houde et al. 2023). Consequently,
this state is considered plesiomorphic for members of
Galloanserae (Tambussi et al. 2019). However, the lacrimals of
S. neocaledoniae, gastornithids, Danielsavis nazensis Houde
et al. 2023, and some galliforms are fused to the frontals and
span the craniorostral hinge to articulate with the nasals (pers.
obvs. photos of specimen AMNH 6169 of Gastornis giganteus;
Matthew and Granger 1917; Andors 1992; Murray and Megirian
1998; Mourer-Chauviré and Balouet 2005; Houde et al. 2023;
Mayr et al. 2023). The fusion of the lacrimal to the frontals
which occurs in anatids is considered a feeding adaptation (see
Fisher 1955; Dzerzhinsky 1982; Zelenkov and Stidham 2018).
1100 P. L. MCINERNEY ET AL.

Figure 2. Genyornis newtoni caudal orbit, NMV P256893, rostral view showing the caudal orbit and quadrate articulation with the skull and mandible: A. Image; B.
Annotated outline. Annotations: ap.zyg.oss., aponeurosis zygomatica ossificans; cr.or., crista supraorbitalis; f.pseu., fossa pseudotemporalis; mand., mandible; m.lev., origin
for m. levator palpebrae dorsalis; n.for., foramen neurovasculare; quad., quadrate. Scale bar: 10 mm. Dark grey shading indicates regions where damage precludes
morphological assessment, and light grey indicates foramina and fossae. Dotted lines provide approximate region corresponding to labelled area, and do not indicate exact
boundaries.

In the megapodiid species with the largest rostrum, Cranium

Macrocephalon maleo, no synostosis of the lacrimal is present and
instead, there is a very small frontal articulation and a much larger
one with the rostrum. In G. newtoni specimens which retain the
craniorostral hinge, there are no apparent laterally facing articula­
tory facets, processes or osseous lobes on the cranium that could
articulate with a more lateral lacrimal head or supraorbital process;
nor have any been identified in the crania of other dromornithids
(Murray and Megirian 1998; Worthy et al. 2016a). The only articu­
latory surfaces on the cranium are those rostrally facing, as part of
the hinge joint (i.e. anterolateral processes of Murray and Vickers-
Rich 2004, p. 235). Considering the form of M. maleo, an alternative
hypothesis to that above is that the lateral edge of the caudal
rostrum in G. newtoni may have been a mediolaterally thin, dorso­
ventrally tall facet that adjoined an unfused lacrimal. However, an
implication of this separate or discrete lacrimal scenario would have
been a severely restricted orbital space in G. newtoni, and since no
separate lacrimals have been reliably identified for any dromor­
nithid, this hypothesis is considered unlikely.
Orbit and rostrodorsal region of cranium
The cranium of G. newtoni, as for other dromornithids, is marked
by large (relative to the cranium), widely separated orbits extending
caudolaterally from the craniorostral hinge, that occupy the rostral
45% of total cranial rostrocaudal length and 35% of cranial dorso­
ventral depth, in lateral view. Specimen SAMA P59516 preserves
both orbits with an estimated interorbital width across the dorsal
surface of the frontals of minimally 74.4 mm, although poor pre­
servation of the cranium likely underestimates that of the undis­
torted skull, and precludes standard interorbital width
measurements (measurements are restricted to the region just cau­
dal of the hinge). As for all dromornithid crania, there is no
depressio frontalis, the roof of the cranium is smoothly curved in
sagittal section, contrary to most galloanserans.
The crista supraorbitalis (Figure 2, 3; sensu Livezey and
Zusi 2006) of G. newtoni is a sharp crest along its entire length
that encloses about a third of the circumference of the orbit
 HISTORICAL BIOLOGY 1101

Figure 3. Lateral view of the left side of the cranium of Genyornis newtoni: A. SAMA P59516 with shaded indication of focus region; B. SAMA P59516 image and annotated
outline, digitally removed from the image of the entire skull; C. NMV P256893 image (includes part of the mandible and quadrate ventrally) and annotated outline; D. Left
rostrolateral view of Anhima cornuta specimen NMV B12574, box denotes region of focus in E., Quadrate disarticulated; E. Rostrolateral view of the left lateral cranium of
Anhima cornuta specimen NMV B12574. Annotations: an.tymp., annulus tympanicus; ap.dep.m., an aponeurotic site of origin of m. depressor mandibulae; ap.zyg.oss.,
aponeurosis zygomatica ossificans; cr.zyg., crista zygomatica; cr.nu.trans., crista nuchalis transversa; cr.or., crista supraorbitalis; cr.art., crista aponeurosis articularis; f.pseu.,
fossa pseudotemporalis; imp.dm., impressio m. depressor mandibulae; imp.sup., impressio m. AME superficialis; jug., jugal arch; mand., mandible; m.a.e., osseous meatus
acusticus externus; pr.par., processus paroccipitalis; pr.post., processus postorbitalis; pr.sup., processus suprameaticus; quad., quadrate; t.pseu., tubercle
for m. pseudotemporalis (specifically aponeurosis pseudotemporalis superficialis). Scale bars: A. 50 mm, B., C. 20 mm, D., E. 10 mm. Dark grey shading indicates regions
where damage precludes confident morphological assessment, and light grey indicates foramina and fossae. Dotted lines provide approximate regions corresponding to
labelled areas, and do not represent accurate boundaries.
1102 P. L. MCINERNEY ET AL.
from the hinge to the processus postorbitalis. In rostral view,
the crest flares laterally as a convex arc far more than in
species of Dromornis. When viewed laterally, the caudal part
of the crest appears straight as in Ilbandornis woodburnei, and
less rounded than in D. planei. This lack of curvature of the
orbit in lateral aspect is enhanced by the medially convergent
rostral half of the supraorbital crests that are also relatively
dorsoventrally flattened. Its dorsal and ventral surfaces are
penetrated by numerous neurovascular foramina, in addition
to the unique vesicular surface texture (most apparent on
specimen NMV P256893, see Figure 3C, see also Stirling
1913, p. 113). The crista supraorbitalis of Gastornis giganteus
is a robust, laterally-prominent crest of bone (Matthew and
Granger 1917; also pers. observ. from photographs), more so
than that of the dromornithids. The wide mediolateral width
between supraorbital crests contrasts with the narrower form
typical of Anseres, as well as the late Paleocene anseriform
Anachronornis anhimops Houde et al. 2023 (see Houde et al.
2023). Contrary to Murray and Vickers-Rich (2004, p. 263), we
find no evidence of an impression or fossa for ‘nasal salt
glands’ (fossa glandulae nasalis, also see King 1993) on the
dorsal frontal area in any G. newtoni specimens. However, we
recognise the presence of a vascularised depression in species
of Dromornis, where it is largest and most obviously depressed
in D. planei (see Murray and Vickers-Rich 2004, p. 263),
which may or may not be associated with a nasal gland. The
anhimid Chauna torquata has a morphology comparable with
the latter, especially.
Caudal orbit
Just ventromedial of the caudal part of the crista supraorbitalis, on
the dorsocaudal surface of the orbit, is a dorsoventrally tall, rugose
surface, pitted by numerous foramina, that likely corresponds to the
origin of m. levator palpebrae dorsalis (m.lev., Figure 2 see also
Shufeldt 1890, p. 55–56; Fisher and Goodman 1955: fig. 2;
Elzanowski 1987). This is consistently present among dromornithid
crania that preserve this region.
The caudoventral part of the orbit, which includes the area
muscularis aspera, is marked laterally by a distinct fold of bone
formed by the processus postorbitalis and aponeurosis zygomatica
ossificans (see ‘Temporal region’ for discussion of the osteology and
myological correlates of this region). The area muscularis aspera is
not completely visible in any specimen of Genyornis newtoni, but is
well-preserved in specimens of Dromornis murrayi, D. planei and
Ilbandornis woodburnei. In these taxa, a rostrally projected tubercle
sits, dorsal to the foramen n. maxillomandibularis (identified in
Worthy et al. 2016a), and medial of a distinct fossa; this fossa is
identifiable in G. newtoni specimen NTM P256893 but not the parts
more mesad. The tuberculum likely hosts the aponeurosis pseudo­
temporalis superficialis (see also Davids 1952a, p. 89–90, fig. 10 a, b,
‘aponévrose 10, tuber 10’; sensu Dzerzhinsky and Potapova 1974;
Weber 1996), in association with the m. pseudotemporalis super­
ficialis (Vanden Berge and Zweers 1993: annot. 19), which is
a significant component of the adductor mandibulae internus
group (e.g. Dzerzhinsky 1982; Vanden Berge and Zweers 1993;
Murray and Vickers-Rich 2004; Holliday and Witmer 2007), in
recognition of a similarly located crest or attachment region in
other galloanserans (e.g. Anhima cornuta, Anseranas semipalmata,
Alectura lathami). Prominent cristae associated with the apo­
neuroses of m. pseudotemporalis, have been identified in other
birds that also have proportionally heavy mandibles. These include
heavy-billed finches (e.g. Geospiza fortis [Thraupidae], see
Genbrugge et al. 2011: fig. 4, Cr7, Cr8), species of Phoenicopterus
[Phoenicopteridae], and Porphyrio [Rallidae] (pers. observ.; Baumel
and Witmer 1993, annot. 89). Discussion on the variation in the
morphology of this region between species of Dromornis and
Ilbandornis is covered by Worthy et al. (2016a), although we suggest
a different musculature arrangement following a reassessment of
the relevant osseous correlates.
We also associate the aforementioned fossa
with m. pseudotemporalis superficialis (e.g. Dzerzhinsky 1982,
also see above references), and the feature termed the ‘fossa pseu­
dotemporalis’ by Murray and Vickers-Rich (2004, p. 240, figs. 52,
76, 77, 92, 188). Murray and Megirian (1998) attributed this fossa to
‘mm. protractor quadratus and pterygoideus’ and Worthy et al.
(2016a), to ‘mm. AME medialis et superficialis’. This particularly
well-developed cavity in dromornithids has been recognised as
characteristic of this family (Murray and Vickers-Rich 2004,
p. 76), but is also typical of anseriforms, in support of systematic
affinities with that group (Murray and Megirian 1998, p. 80, 96;
Murray and Vickers-Rich 2004, p. 240). An especially similarly deep
fossa is present in anhimids (pers. observ., also see Dzerzhinsky
1982). Other anseriforms, such as Anseranas semipalmata and
Anser caerulescens, have a distinct depression for the muscle origin
in the same position. That in the latter species is notably more like
that of Dromornis murrayi, in which the fossa is shallower than in
other dromornithids.
In association with the fossa pseudotemporalis and the medi­
ally adjacent tubercle, the impression for the origin
of m. pseudotemporalis superficialis appears to maximally extend
to the laterally adjacent aponeurosis zygomatica ossificans and
processus postorbitalis on the caudal orbit in all dromornithids.
The area of origin extends to a rostroventral portion of the
lateral cranium (including the temporal fossa, see
‘Nomenclature’) in several avian clades, e.g. palaeognaths
(Elzanowski 1987, p. 83; Holliday and Witmer 2007: and refer­
ences therein), although, is typically restricted or confined behind
the eye to the area muscularis aspera in anseriforms and galli­
forms (e.g. Lakjer 1926; Hofer 1950; Davids 1952a; Starck and
Barnikol 1954; Goodman and Fisher 1962; Fujioka 1963;
Dzerzhinsky and Belokurova 1972; Zweers 1974; Dzerzhinsky
1982; Weber 1996; Matsuoka et al. 2008). The latter, in effect,
forms the fossa pseudotemporalis, similar to the apparent dro­
mornithid condition. The incidence and variable development of
cristae on the area muscularis aspera, and expansion of the
pseudotemporal fossa presumably acts to increase the attachment
surface area for this muscle within its restricted region of origin.
The relative expression of the muscle origin area appears to
correlate with size of the mandible, being deeper in species
with larger mandibles.
Temporal region
As in all galliforms and anhimids, the dromornithids have relatively
small processus postorbitales that project ventrally and slightly
rostrally; in contrast, in non-anhimid anseriforms they are more
elongate and rostrally directed and contribute to the ventral margin
of the orbit (Baumel and Witmer 1993, annot. 14; Zusi and Livezey
2000; Field et al. 2020). The basal anseriform Anachronornis anhi­
mops also has postorbital processes that project only slightly ros­
trally and are not as rostrocaudally extensive as those in anatoids
(however, see below, Houde et al. 2023).
The aponeurosis zygomatica (sensu Elzanowski 1987; Weber
1996; Zusi and Livezey 2000) is one of the major aponeuroses of
the external adductor musculature complex (specifically m. AME

profundus, pars zygomaticus) in Neornithes, although its ossifica­
tion to form aponeurosis zygomatica ossificans (AZO; sensu Zusi
and Livezey 2000, p. 166) only occurs in some neornithine species
(Zusi and Livezey 2000). Additionally, the interaction and develop­
ment of the ossified and unossified aponeurosis, processus zygo­
maticus and processus postorbitalis, and the consequent impacts on
the arrangement of the associated musculature, varies across
Galloanserae, and has systematic importance (see Zusi and
Livezey 2000; Murray and Vickers-Rich 2004: fig. 136). To ade­
quately illustrate the morphology presented in G. newtoni and other
dromornithids, contextual description of the state in other galloan­
serans is required. The myology of the adductor chamber in galli­
forms has been extensively researched (e.g. Burggraaf 1954a;
Fujioka 1963; Dzerzhinsky and Belokurova 1972; Dzerzhinsky
1974, 1980; Weber 1996; Zusi and Livezey 2000). In most galli­
forms, ossification of aponeurosis zygomatica occurs and the resul­
tant AZO extends rostrally from the processus zygomaticus; in
many, the aponeurosis connects rostrally with the ventral tip of
the processus postorbitalis (see Kirikov 1944; Olson and Feduccia
1980; Dzerzhinsky 1982, 1995; Zusi and Livezey 2000). Complete
connection forms the orbitozygomatic junction (sensu Elzanowski
and Mayr 2017) and encloses a secondary temporal fenestra (see
Elzanowski and Mayr 2017). This morphology is notably extreme in
gastornithids, in which the AZO forms a large, robust bridge,
traversing more than half the side of the cranium (Matthew and
Granger 1917; Troxell 1931; Andors 1992; Murray and Vickers-
Rich 2004; Elzanowski and Mayr 2017).
In the basal anseriform family Anhimidae, the aponeurosis
zygomatica also ossifies to form the AZO, however, unlike galli­
forms, there is no zygomatic process (Zusi and Livezey 2000). The
aponeurosis zygomatica has a linear attachment along the rostro­
caudal length of the lamina lateralis cranii (sensu Zusi and Livezey
2000), and passes medial to, and rostrally beyond, the processus
postorbitalis (see Dzerzhinsky 1982; Zusi and Livezey 2000: fig. 6, 7;
Murray and Vickers-Rich 2004, p. 166–167, fig. 136). The apo­
neurosis is ossified along much of this length in adult anhimids,
except for in its rostral-most part (Zusi and Livezey 2000, p. 173–
175). Through myological, ontogenetic and homological studies
relating to the osseous structures of the adductor chamber in
these birds (Dzerzhinsky 1982, 1995; Zusi and Livezey 2000), it is
currently understood that the wedge-shaped temporal region cor­
responding to the AZO and postorbital process is related to the
evolutionary medial migration of the origin of m. AME profundus,
pars coronoideus (sensu Zusi and Livezey 2000; terminology of
Holliday and Witmer 2007), relative to a typical galliform state.
This medial retreat of fibres that would normally arise from the
dorsotemporal fossa effectively truncates the lateral squamosum
rostrally from the caudoventral orbit, while impressio m. AME
superficialis (caudotemporal fossa) remains on the caudolateral
squamosum. Consequently, the AZO is closely associated with the
processus postorbitalis in anhimids and contributes to the laterally
adjacent crista m. AME superficialis (sensu Zusi and Livezey 2000;
using terminology of; Holliday and Witmer 2007), and laterally
delimits the impressio m. AME profundus, pars coronoideus
(Dzerzhinsky 1982; crista zygomatica sensu Zusi and Livezey 2000).
Dzerzhinsky (1982, p. 1031) originally interpreted this complex
as a sesamoidal ossification and superimposition upon the proces­
sus zygomaticus (i.e. AZO), fused to a caudoventrally expanded
base of the processus postorbitalis, effectively closing the dorsotem­
poral fossa, and termed the resultant composite osseous structure
the processus sphenotemporalis (see also Dzerzhinsky 1995). Using
ontogenetic evidence, Zusi and Livezey (2000, p. 175–177, 180–181)
further honed interpretations on the homology of this structure,
regarding the contribution of the AZO. They noted that the
HISTORICAL BIOLOGY 1103
processus zygomaticus does not contribute to this morphology in
anseriforms and that the morphology was better explained as an
evolutionary product of the rostral extension of the origin for
aponeurosis zygomatica to the processus postorbitalis and the
aforementioned ventromedial migration of the origin of m. AME
profundus, pars coronoideus (see above). Recently, Houde et al.
(2023) dismissed Zusi and Livezey’s interpretations and advocated
for a sphenotemporal process, and the presence of a processus
zygomaticus that abuts the laterosphenoidale ventrally at sutura
laterospheno-squamosa in anseriforms. We instead find the
hypotheses stemming from the comprehensive homological and
ontogenetic study of this region by Zusi and Livezey (2000) to be
more compelling, especially in consideration of the following
important observations: (1) identification of a potential incipient
processus zygomaticus at the caudal-most area corresponding to
the origin of the aponeurosis zygomatica in some immature anseri­
forms; (2) the recognition that in anatids, the rostroventral exten­
sion of squamosum and its shape in its interaction with the
laterosphenoidale (including processus postorbitalis) at sutura
laterospheno-squamosa can falsely resemble a processus zygomati­
cus; and (3) the region of the squamosum which interacts with the
laterosphenoidale is non-homologous with the area from which the
processus zygomaticus arises in many other Neornithes (see Zusi
and Livezey 2000, p. 170, fig. 4, 5; Mayr and Manegold 2021). We
thus opt to follow the homological interpretations of Zusi and
Livezey (2000), herein, owing to lack of evidence that rejects the
findings of their focused study, nor robustly supports alternative
hypotheses. Zusi and Livezey (2000) further noted that while the
impressio m. AME profundus, pars coronoideus, was also similarly
medially located in other anseriforms, its relationship to osseous
structures was different in non-anhimid anseriforms. There is little
to no ossification present for the zygomatic aponeurosis (crista
zygomatica sensu Zusi and Livezey 2000, see below) in addition to
a distinct processus zygomaticus being absent (the homologous
locus may be represented by a tubercle, see Zusi and Livezey
2000), and the processus postorbitalis (of the laterosphenoidale
bone) is rostroventrally developed and orientated. The
impressio m. AME profundus, pars coronoideus, effectively has
greater association with the processus postorbitalis in Anseres
(that is anseriforms exclusive of anhimids, see Livezey 1997;
Worthy et al. 2017b), compared to arising from the ventromedial
AZO as in species of Chauna and Anhima.
In dromornithids, it is clear that considerable ossification of the
aponeurosis zygomatica occurs and produces a morphology,
including a distinct, conspicuously rostrally projected AZO, which
we interpret to be a near identical osteological arrangement to that
of anhimids (also previously mentioned by Murray and Vickers-
Rich 2004; Worthy et al. 2016a). This morphology is observed in all
dromornithid crania with adequate preservation. As a result, in
G. newtoni, the m. AME profundus, pars coronoideus, is inter­
preted to originate ventromedially on the AZO (also inferred for
species of Dromornis and Ilbandornis by Murray and Vickers-Rich
2004: fig. 188) and the impressio m. AME superficialis is represen­
tative of the only part of the m. AME complex to remain on the
lateral squamosal (and the ‘temporal fossa’). The various features of
this region are best preserved in specimen NTM P256893 (Figure 2)
of G. newtoni, as well as Ilbandornis (OMV2000:GFV:20 pers.
observ.) and D. planei (NTM P9464–106, see Murray and Vickers-
Rich 2004).
In G. newtoni, the impressio m. AME profundus, pars coronoi­
deus, is a wide and deep, mediodorsally expanded depression,
which is more similar in form to Chauna torquata than Anhima
cornuta, and although slightly differs in osteological homologies
(see below), is similar in shape to the same impression in Anseranas
1104 P. L. MCINERNEY ET AL.
semipalmata and Anachronornis anhimops. It is more dorsoven­
trally expansive than in Ilbandornis woodburnei, and more compar­
able to Dromornis planei. The impression is bordered
dorsomedially by a rostrocaudally elongate crest, and laterally by
the crista zygomatica which may be nearly confluent and contin­
uous with the crista m. AME superficialis in the rostral-most area,
but not caudally (see below, also Zusi and Livezey 2000: fig. 6, 7,
p. 175). The crista zygomatica is formed from the crest-like ossifi­
cation of the aponeurosis zygomatica where it meets the cranium,
ventral or ventromedial of the dorsolateral bounds of the origin
of m. AME superficialis, and is not mutually exclusive with respect
to the more extensive aponeurotic ossification that characterises
AZO in many galloanserans (see Zusi and Livezey 2000: fig. 6,
p. 175, Table 2). Scarring, present both laterally and medially on
the rostroventral AZO in specimen NMV P256893 of G. newtoni,
suggests that it predominantly or exclusively supported the unossi­
fied aponeurosis zygomatica. Comparably, in D. planei, the unossi­
fied aponeurosis zygomatica appears to have additionally emanated
from the postorbital process (see Murray and Vickers-Rich 2004:
fig. 188, ‘crista AME superficialis’). In anatids (e.g. Anser caerules­
cens, and Tadorna tadornoides), the minor ossification of the apo­
neurosis zygomatica (if present) typically produces only a low,
ridge-like crista zygomatica which is ventrally separated from, and
medial to, the crista m. AME superficialis (Zusi and Livezey 2000,
p. 175, fig. 6).
It is likely that aponeurosis zygomatica emanated from the
rostrocaudal length of lamina lateralis cranii in the late Paleocene
anseriform Anachronornis anhimops, as it does in anhimids, and
a small caudal impression, rostrally adjacent to processus supra­
meaticus would have accommodated the origin for m. AME super­
ficialis. The latter state is also comparable to that of anhimids, while
the relatively expansive impressio m. AME profundus, pars coro­
noideus, best resembles the anseranatid and dromornithid condi­
tion. Like anseranatids, but unlike anhimids and dromornithids,
there is no conspicuous rostral projection with apical ossified fibres
or scarring, and related mediolateral thickening of the ventral
lamina lateralis cranii, that would be typical of an extensively
ossified aponeurosis zygomatica. This is further supported by the
observable sutura laterospheno-squamosa ventrally on
Anachronornis anhimops, which comparatively appears obscured
by the development of AZO in anhimids and dromornithids, but
importantly denotes that the more rostral area corresponds to the
laterosphenoid and processus postorbitalis (see Houde et al. 2023:
fig. 1 A). Ontogenetic evidence suggests that rostral development of
the laterosphenoid bone, not associated with the AZO, is character­
istic of Anseres, where rostral projection of this region is addition­
ally accomplished through the development of the AZO in
anhimids and dromornithids (Zusi and Livezey 2000).
Additionally, as evidenced by some incomplete fusion of cranial
elements in the cranium of A. anhimops, its cranium shape and
pattern of suture arrangement closely resembles the skull of juvenile
Chauna torquata, before the AZO has entirely developed (compare
Zusi and Livezey 2000: fig. 5 A; Houde et al. 2023: figs. 1D, S3).
While the rostral projection of processus postorbitalis in
A. anhimops is superficially similar to the state of anhimids and
dromornithids, it is thus likely analogous, in support of this taxon
possessing an intermediate condition between extant anhimids and
Anseres (Houde et al. 2023, p. 16).
A foramen temporale venosum (sensu Mayr et al. 2021) has been
recognised within the region of origin for m. AME profundus, pars
coronoideus, in Galliformes, leading this feature to be considered
a cranial autapomorphy of the group (see Mayr et al. 2021: fig. 4).
Despite variation in the location of the non-ossified aponeurosis
zygomatica, a seemingly homologous foramen is also present in the
region of attachment for this muscle in all anatids compared in this
study, and possibly Presbyornis pervetus Wetmore 1926 (USNM
299846). This foramen was not identified in dromornithids or
Anhima cornuta. Considering this foramen is aligned with the
sutura laterospheno-squamosa, most visible in younger individuals
(e.g. Cygnus olor, see Houde et al. 2023: fig. S2), it may be associated
with the fusion of the two bones in adulthood. Presence or absence
in dromornithids and anhimids cannot be confirmed potentially
due to the AZO deforming this region.
The partes superficialis et zygomatica of m. AME profundus
(Zusi and Livezey 2000; Holliday and Witmer 2007; Appendix
Three) originate on the lateral and medial sides of the unossified
aponeurosis zygomatica, respectively (Zusi and Livezey 2000,
p. 169), and are located well rostrad in galloanserans compared to
the condition in other avian orders due to the relatively large size
of m. AME superficialis (see Zusi and Livezey 2000, p. 177, fig. 8). In
anseriforms, these two distinct bellies, partes superficialis et zygo­
maticus, uniquely originate from the unossified aponeurosis zygo­
matica rostral of its attachment to the processus postorbitalis (Zusi
and Livezey 2000, p. 177, figs. 6 and 8). The morphology in dro­
mornithids appears similar to anhimids (see Murray and Vickers-
Rich 2004: fig. 136), so the bellies are deemed likely to arise from the
unossified aponeurosis zygomatica in G. newtoni, in a similarly
rostral position. Any attachment surface(s) corresponding
with m. AME medialis cannot be identified and may be interrelated
with those of m. AME profundus, considering that this muscle is
not easily distinguishable from the latter muscle part in birds
(Holliday and Witmer 2007, p. 481; Appendix Three).
As aforementioned, the impressio m. AME superficialis (caudo­
temporal fossa) is the only part of the temporal fossa retained on the
lateral squamosal in dromornithids (see ‘Nomenclature’, and also
Murray and Megirian 1998; Murray and Vickers-Rich 2004;
Worthy et al. 2016a). It is characterised by a depression that extends
caudally from processus postorbitalis and the rostral end of the
AZO to a variable position dorsal of the dorsocaudal margin of the
osseous meatus acusticus externus (external auditory canal) and is
relatively medially positioned like in Anhimidae (see Zusi and
Livezey 2000). The dorsal bounds of this muscle origin are rounded
and poorly defined on the lateral squamosal in all dromornithids,
and clearly dorsolateral of crista zygomatica (see Figure 4). In
contrast, crista m. AME superficialis is pronounced ventrally, crest-
like in its association with aponeurosis mediosuperficialis (sensu
Dzerzhinsky 1982; Dzerzhinsky and Grintsevichene 2002), and
nearly confluent with the crista zygomatica across most of its
rostrocaudal length in both anhimids and anseranatids, and also
apparently in anachronornithids (see above and Zusi and Livezey
2000, p. 175; Houde et al. 2023). Presumably, the lack of a distinct
crista m. AME superficialis caudal of the AZO in dromornithids
may have myological implications with regards to superficial apo­
neurotic attachment and proportional forces acting in this region
(Bernhard 1924; Bryant and Seymour 1990 and references therein).
While the degree of ossification of aponeurosis zygomatica in
anatids clearly differs from that of dromornithids and anhimids
(and anseranatids and anachronornithids to a lesser extent, see
above), the dorsoventral separation between the dorsal bounds of
impressio m. AME superficialis and crista zygomatica in anatids is
more comparable to the dromornithid condition. The caudal extent
of the impressio m. AME superficialis in dromornithid taxa is
variously defined by a pronounced tubercle or dorsoventrally
aligned ridge. This tubercle (in part) relates to an aponeurotic
attachment of the caudal m. AME superficialis (crista aponeurosis
articularis, sensu Murray and Vickers-Rich 2004: fig. 188; fig. 4).
Compared to the large and rugose form of D. planei, the crest is less
pronounced in Ilbandornis woodburnei and even less distinct, and
 HISTORICAL BIOLOGY 1105

Figure 4. Soft-tissue attachment sites on the cranium of a dromornithid, illustrated using a digitally modified representation of Ilbandornis woodburnei (QMV 2000:gfv:20)
due to the relative lack of deformation in this skull and the conservative nature of dromornithid skull morphology: A. Rostral aspect; B. Caudal aspect; C. Ventral aspect; D.
Right lateral aspect; E. Oblique (rostrolateral) aspect. Abbreviations: ap.art., origin site of aponeurosis articularis; ap.dep.m., an aponeurotic site of origin of m. depressor
mandibulae; ap.med.sup., origin site of aponeurosis mediosuperficialis; ap.sup., origin site of aponeurosis superficialis; ap.zyg., origin site of aponeurosis zygomatica (muscle
fibres of m. AME profundus, pars zygomaticus, and m. AME profundus, pars superficialis, originate from the medial and lateral surfaces of this aponeurosis, respectively); lig.
oc.mand., attachment area corresponding to the origin of ligamentum occipitomandibulare, continuous with that of membrana postmeatica; lig.post.orb., attachment area
of origin of ligamentum postorbitale; m.AME.prof.cor., origin area of m. AME profundus, pars coronoideus; m.AME.sup., origin area of m. AME superficialis; m.biv.cerv.,
insertion area of m. biventer cervicis; m.c.c., origin area of m. cucullaris capitis; m.comp., insertion area of m. complexus; m.dep.mand., origin area of m. depressor
mandibulae; m.pro.pter.quad., origin area of m. protractor pterygoidei et quadrati; m.ps.tem.s., origin area of m. pseudotemporalis superficialis; m.rec.cap.dors., insertion
areas for slips of m. rectus capitis dorsalis; m.rec.cap.lat., insertion area of m. rectus capitis lateralis; m.rec.cap.vent.lat., area for aponeurosis of insertion of m. rectus capitis
ventralis, pars lateralis; m.rec.cap.vent.med., insertion area of m. rectus capitis ventralis, pars medialis; m.spl.cap., insertion area of m. splenius capitis; mem.at.oc.dors.,
attachment area of membrana atlantooccipitalis dorsalis; mem.at.oc.vent., attachment area of membrana atlantooccipitalis ventralis; mem.pm., attachment site of
membrana postmeatica, which incorporates and is not separatable with respect to the more ventromedial ligamentum occipitomandibulare. Scale bar: 50 mm.
Illustrated attachment areas on the cranium are non-extensive estimates; only the sites corresponding to selected muscles, aponeuroses, membranes and ligaments
are indicated.
1106 P. L. MCINERNEY ET AL.
more of a low ridge in G. newtoni. Several galliform taxa have an
angular, rostrally protruding ossified aponeurotic fibre mass asso­
ciated with this region caudal of the processus suprameaticus (e.g.
Lagopus lagopus), whereas in anseriforms, including anhimids, the
caudal-most attachment of the m. AME superficialis is represented
by an incurvate and variably pronounced ridge, dorsal of the supra­
meatic process.
The inferred myological topology associated with the dromor­
nithid adductor chamber (Figure 4 C–E), as evidenced by patterns
of fossae and cristae upon the skull (discussed above), are similar to
the anhimid condition, as explored and illustrated by several
authors (e.g. Dzerzhinsky 1982; Zusi and Livezey 2000;
Dzerzhinsky and Grintsevichene 2002). The primary variations
between these two lineages, aside from greater rostrocaudal fore­
shortening of the dromornithid cranium, is present in the propor­
tional sizes of the processus postorbitalis and AZO, which are
enlarged and more robust, the dorsoventral separation between
crista zygomatica and the dorsal bounds of m. AME superficialis,
and the development of crista aponeurosis articularis. The greater
size and depth of the impression m. AME superficialis (caudotem­
poral fossa), the proportionally enlarged rostral end of the AZO,
and evidence for further attachment for the non-ossified portion of
the aponeurosis zygomatica on the postorbital process in species of
Dromornis, may be also linked to the larger, heavier mandible
relative to skull size, compared to G. newtoni and species of
Ilbandornis. In D. planei, we associate the relative reduction of the
processus postorbitalis and the process tip not ventrally overhan­
ging the ossified zygomatic aponeurosis with the requirement for
greater adductor musculature throughout this region; this may
imply that the ligamentum postorbitale was absent, as observed
for the large-billed finch Geospiza fortis (see Genbrugge et al.
2011, p. 692). This ligament is also absent in the erismaturine anatid
Oxyura jamaicensis (see Goodman and Fisher 1962). Specimen
SAMA P59516 of G newtoni, shows the ventral extent of the pro­
cessus postorbitalis was variable in this lineage; we recognise three
potential drivers: (a) intraspecific variation associated with an
absence of selection pressures on the shape of the processus post­
orbitalis and presence of the ligamentum postorbitale, (b) sexual
dimorphism in the mass of the muscles in this region, as SAMA
P59516 is considered a likely female (Chinsamy and Worthy 2021)
and NMV P256893, sex unknown, and (c) ontogenetic variation.
Although the ligamentum postorbitale serves to coordinate motion
of both upper and lower jaws as the mandible is depressed, it is not
required as it is only one of several means of doing this (Zusi 1967;
Bout and Zweers 2001).
Lateral aspect of the caudal cranium
In all three partially preserved crania of G. newtoni, specimens
SAMA P59516, NMV P256893, and SAMA P59520, the otic
head of the quadrate remains articulated in the quadratic cotyla
of the cranium, ultimately obscuring it. For other dromor­
nithids where it is visible, e.g. I. woodburnei specimen
QMV:2000:GFV:20, I. ?lawsoni NTM P907–27, and D. planei
specimen QM F57947, we use the term, cotyla quadratica squa­
moso-otica (named for the close approximation of the two
quadratic cotylae which, together form a single articular facet,
see also Baumel and Raikow 1993, annot. 25; also termed
recessus quadratica, sensu Worthy et al. 2016a). In species of
Ilbandornis and Barawertornis, the residual homolog of recessus
tympanicus dorsalis extends dorsolaterally as a shallow fossa,
partly separating the constituent cotylae, at their ventromedial
margins, although it is thinnest and deepest in B. tedfordi
(specimen QM F58013). An exceedingly small foramen is
present centrally on the cotyla quadratica squamoso-otica in
D. murrayi, D. planei, and I. woodburnei, likely a remnant of
the associated foramen pneumaticum dorsale (Baumel and
Witmer 1993: annot. 25; Mayr 2020) along with a second
small foramen at the medioventral margin of the aforemen­
tioned dorsal tympanic depression. A large foramen just medial
of this and between the articulatory surfaces is associated with
the foramen for ramus occipitalis of arteria ophthalmica
externa. Due to similarities in quadrate morphology (see
‘Quadrate’), and relatively consistent morphology for this region
across all dromornithid specimens where it can be observed, we
assume that G. newtoni likely had a similar cotyla quadratica
squamoso-otica and associated features. This morphology is
uncommon within Galloanserae and differs dramatically from
the condition in non-anhimid anseriforms, and megapodes
which typically possess two distinct quadratic cotylae, cotyla
quadratica squamosi and cotyla quadratica otici, separated by
a deep recessus tympanicus dorsalis, and an associated, gener­
ally large foramen pneumaticum dorsale (sensu Mayr 2020; e.g.
see Anseranas semipalmata, Tadorna tadornoides, and Alectura
lathami). In many non-megapodiid galliforms and
S. neocaledoniae (see Mourer-Chauviré and Balouet 2005), the
cotylae are only separated by a thin depression continuous with
a dorsally located recessus tympanicus dorsalis consistent with
the more closely applied condyles on the quadrate head (e.g.
Acryllium vulturinum and Lagopus lagopus).
In Genyornis newtoni, SAMA P59516, the quadratic capsule is
bounded laterally by a thin osseous wall, perforated by a small,
ovular (9.4 mm high, 7.1 mm wide) fenestra, through which the
pars otica of the quadrate, still in articulation, can be viewed. This
lamina extends caudoventrally from the AZO, to join the thin, bar-
like annulus tympanicus (processus postglenoidalis, sensu
Stellbogen 1930; see also Baumel and Witmer 1993: annot. 77;
Mayr 2020). There is little evidence for this lamina being an artefact
of preservation or damage in specimen SAMA P59516, and damage
to both SAMA P59520 and NMV P256893 precludes a non-
ambiguous assessment of this feature in these G. newtoni speci­
mens. This feature may then reflect unusual intraspecific variation,
arising from an extension of the squamosum, or ossification of
aponeurosis zygomatica, m. AME superficialis, a membrane, liga­
ment, or cartilage. Alternatively, the absence of this character in
other dromornithid taxa, and all other assessed galloanserans, may
imply that this character is autapomorphic to G. newtoni.
The processus suprameaticus is mediolaterally flattened in
G. newtoni, as in D. planei, and less prominent than the more
rounded, ventrally protruding process in I. woodburnei, which is
more distinct from the annulus tympanicus, a bony ridge connect­
ing the process ventrally with the ala parasphenoidalis (Stellbogen
1930; Mayr 2020). In all dromornithids, the lateral overhang of the
small processus suprameaticus relative to the caudolateral margin
of the cotyla quadratica squamoso-otica is minimal (D. murrayi) to
lacking (D. planei, I. woodburnei, G. newtoni). This is uncommon
within Galloanserae, even those with an annulus tympanicus (see
below), due to variation in the rostrocaudal location of the process
relative to the cotyla. The processus suprameaticus of Gastornis
giganteus and S. neocaledoniae, considerably laterally overhangs
the caudolateral margin of the cotyla quadratica squamosi.
In species of Ilbandornis and Dromornis, the annulus tympani­
cus is especially robust compared to G. newtoni. Ossification of this
bar also occurs in some anatids (Anser caerulescens, Cereopsis
novaehollandiae, and Cnemiornis calcitrans; see also Worthy et al.
1997, 2017b: app. 1, char. 16), odontophorids (Callipepla califor­
nica), and phasianids (Perdix perdix and Syrmaticus soemmerringii,
for the latter see Mayr 2020: fig. 8), suggesting variable development

among galloanserans. In dromornithids, the ventromedial exten­
sion of the annulus tympanicus from processus suprameaticus, to
fuse to the enlarged and laterally projecting ala parasphenoidalis,
encloses the osseous meatus acusticus externus ventrorostrally. The
enclosed nature of the osseous meatus acusticus externus, differs
considerably from most galloanserans, including the gastornithids
and sylviornithids, which appear to lack an ossified annulus tym­
panicus associated with the processus suprameaticus. In G. newtoni,
the lateral opening of the ear (the osseous meatus acusticus exter­
nus) is relatively small and circular, approximately 10.8 mm high
and 10.05 mm wide in SAMA P59516, although it is more oval
shaped in NMV P256893. Species of Dromornis and Ilbandornis
have a proportionally larger and less enclosed (especially ventrally)
osseous meatus acusticus externus relative to the rostrocaudal
length of the cranium, and the laterally flaring and broadly curved
margins give a more funnel-shaped appearance in these taxa
(Murray and Megirian 1998; Worthy et al. 2016a). Much of the
otic region, and the more medial cavum tympanicum proper (see
Witmer 1990; Baumel and Witmer 1993: annot. 21), is obscured by
sediment and compression in all specimens of G. newtoni and,
therefore, cannot be described in further detail (see Mayr 2020 for
an assessment of the otic region in Neornithes), although as is
typical for Galloanserae, the pila otica is observed to be non-
trabeculated (see Mayr 2020).
The processus paroccipitalis projects far ventrally and slightly
laterally, terminating 58.5 mm past the ventral margin of the oss­
eous meatus acusticus externus in specimen SAMA P59516 of
G. newtoni. The process is mediolaterally compressed and flattened
with subparallel rounded rostral and caudal edges. In lateral view,
the process is proportionally slenderer and more dorsoventrally
elongate than in other dromornithids. In these species, e.g.
I. woodburnei, more extensive fusion of the process with the ala
parasphenoidalis creates a rostrocaudally wide and robust triangu­
lar structure, when viewed laterally, like that of non-anhimid
anseriforms; anhimids and galliforms have far smaller processus
paroccipitalis, which extend maximally to a point level with the
ventral floor of the otic region. The elongated length of the proces­
sus paroccipitalis in G. newtoni, resembles that of the processes in
Ga. giganteus and S. neocaledoniae. In G. newtoni, a marked attach­
ment region for the ligamentum occipitomandibulare on the ros­
trolateral surface of the ventral part of the processus paroccipitalis is
largely obscured or damaged in all specimens that preserve this
area, but it is visible on the left processus paroccipitalis of SAMA
P59516. This attachment is an ovoid, well defined mark in
G. newtoni but is comparatively better developed in both
D. planei and D. murrayi as a circular, flattened surface. In
I. woodburnei, the shallow depression indicating the ligamentous
attachment area is relatively smaller and less clearly delimited than
in all these species.
On the dorsolateral surface of the processus paroccipitalis, the
fossa subtemporalis, or more specifically, the
impressio m. depressor mandibulae (van Gennip 1986; Weber
1996; terminology of Zusi and Livezey 2000; see ‘Nomenclature’),
is a dorsoventrally elongate, impressed surface, variably obscured
by taphonomic damage in all G. newtoni specimens. In most dro­
mornithids, including G. newtoni, the impression is well-defined
rostrally in partial association with crista aponeurosis articularis
(of m. AME superficialis, see above); however, in D. planei the
impression is especially obvious and well delimited on all sides,
such as in specimen NTM P9464–106, where the origin
of m. depressor mandibulae expands dorsoventrally and is exten­
sively scarred. It is positioned rostrally adjacent to the crista nucha­
lis transversa as in most birds (Baumel and Witmer 1993, annot.
104). In galliforms, the attachment surface is small, more dorsal
HISTORICAL BIOLOGY 1107
relative to the bony external auditory canal, and lacks defined
margins, making it seem confluent with the temporal fossa in
some taxa. Comparatively, anseriforms have a much larger
impressio m. depressor mandibulae. In the most extreme forms
(e.g. Anhima cornuta, Anseranas semipalmata, Melanitta perspicil­
lata, Aythya australis), the impression extends dorsally along the
crista nuchalis transversa to near its dorsal terminus with raised
ridges delimiting all sides. Although predominantly hosting
the m. depressor mandibulae (van Gennip 1986; Vanden Berge
and Zweers 1993), the impression m. depressio mandibulae may
also have hosted fibres relating to the caudal-most origin of
the m. AME superficialis, as it does in most neognaths, including
Galloanserae (see Holliday and Witmer 2007). Concomitantly, in
G. newtoni as in other dromornithids, the rostral-most extent of the
origin of m. depressor mandibulae is indicated in part by crista
aponeurosis articularis. Comparatively, in Anhima cornuta, the
crista associated with the rostral-most origin of m. depressor man­
dibulae is distinct, while the crest indicating the caudal extent of
the m. AME superficialis located further rostrally. The rostrocaudal
foreshortening of the caudal part of the cranium in dromornithids
has seemingly narrowed the distance between the two muscles. This
suggests that the greater size and number of associated cristae is
likely to accommodate both the caudal-most origin of the m. AME
superficialis and the dorsorostral-most origin of the m. depressor
mandibulae (see 4.4.4 for further descriptions of
impressio m. depressor mandibulae). Unlike in anhimids, the
caudal m. AME superficialis and rostral m. depressor mandibulae
are closely spaced in anseranatids, and in anatids they abut each
other at a single common crest similar to the dromornithid state
(see above).
In G. newtoni, the rostral bounds of impressio m. depressor
mandibulae are also continued ventrorostrally by a pronounced
crest that ventrocaudally borders the osseous meatus acusticus
externus. The neck muscle m. cucullaris capitis attaches to the
cranium just rostrad and superficial of the rostral-most origin
of m. depressor mandibulae in anseriforms, in close association
with the rim of the bony external auditory canal (m. dermotempor­
alis sensu Goodman and Fisher 1962; Dzerzhinsky 1982; Vanden
Berge and Zweers 1993: annot. 7; Dzerzhinsky and Grintsevichene
2002) and is perhaps related to this crest in G. newtoni.
The m. cucullaris capitis originates more dorsorostrally in many
galliforms (e.g. Dzerzhinsky 1980; Weber 1996). Presumably, the
relatively pronounced crest that separates impressio m. depressor
mandibulae from impressio m. AME superficialis (see above) may
have hosted this attachment in other dromornithids.
Occipital region, caudal view
Delimiting the caudal extent of the impressio m. depressor mandi­
bulae, in dorsoventral alignment with the condylus occipitalis, at
the lateroventral junction of the crista nuchalis transversa and crista
ventralis (crista occipitalis, sensu Dullemeijer, 1951; sensu Ghetie
et al. 1976; crista occipitalis, sensu Landolt and Zweers 1985; linea
nuchalis transversa, sensu Weber 1996), are bilaterally paired pro­
minentiae exoccipitales (Owen 1873, p. 512; Mivart 1896: fig. 9;
Murray and Vickers-Rich 2004; Worthy et al. 2016a: fig. 1; SI T.4).
The two crests form a V-shaped notch ventrally upon each promi­
nentia, likely associated with the insertion of the m. rectus capitis
lateralis, which attaches here in many bird groups including anseri­
forms and galliforms, superficial to the insertion of the lateral most
part of m. splenius capitis on the caudal cranium (see Boas 1929;
Davids 1952b, 1952c; Goodman and Fisher 1962, p. 118; Zusi and
Storer 1969; Ghetie et al. 1976; Landolt and Zweers 1985; Vanden
Berge and Zweers 1993: annot. 43, 44; Lautenschlager et al. 2014;
1108 P. L. MCINERNEY ET AL.
Jones et al. 2019; Böhmer et al. 2020). The prominences are well
developed in Anhima cornuta and many other anseriforms (e.g.
Anseranas semipalmata, Aythya australis), but are to a greater
extent in the dromornithids, especially species of Dromornis,
where they contribute to the laterocaudal flare of the cranium
margins.
As in all dromornithids, anhimids, and most galliforms, the
crista nuchalis transversa is identifiable, although it is not par­
ticularly distinct in G. newtoni. Comparatively, the crest is more
prominent in anseriforms including Anachronornis anhimops
(see Houde et al. 2023: fig. 1E), enhanced by deeper fossae
dorsal and ventral to it (for an extreme, see Oxyura australis),
and the developmental effects of the forces associated with
muscles which insert upon or near it, i.e., the m. complexus
(m. cucullaris caput part, sensu Goodman and Fisher 1962,
p. 116) along its edge, the deeper m. splenius capitis, and the
dorsally situated m. biventer cervicis (Ostrom 1961, p. 88, and
references therein; Goodman and Fisher 1962, p. 116–117;
Landolt and Zweers 1985; Vanden Berge and Zweers 1993:
annot. 41–43; Lautenschlager et al. 2014; Böhmer et al. 2020).
The shape of the crest, although distorted, suggests that the
caudal dorsal profile of the cranium of G. newtoni is rounded,
like all other dromornithids. With the exception of some speci­
mens such as NTM P9464–106, the mid-dorsal parietal region
of the caudal cranium in dromornithids is not well-known as
most specimens either do not preserve this region (e.g. NTM
P907–27) or have damage precluding visibility (e.g. QMV:2000:
GFV:20, QM F57974, SAMA P59516). Based on all available
evidence, this region in all dromornithids appears to be dis­
tinctly flattened, and lacks (1) a prominentia cerebellaris, (2)
a crista nuchalis sagittalis (develops to differing degrees primar­
ily in goose-type non-anhimid anseriforms, and generally absent
in other galloanserans, Landolt and Zweers 1985; Tambussi
et al. 2019), and (3) fonticuli occipitales (considered
a synapomorphic feature for most anseriforms; lateral occipital
fontanelles sensu Beddard 1898; Landolt and Zweers 1985;
Baumel and Witmer 1993: annot. 87; Ellrott and Schmitz
2010; Tambussi et al. 2019).
The crista ventralis is well developed in dromornithids, as in
most anseriforms, although to a lesser extent than in the gastor­
nithid Ga. giganteus. The crista ventralis distinctly separates relative
dorsal and ventral regions of the caudal face, as it extends laterally,
from a short distance above the dorsal margin of the foramen
magnum, to converge with the crista nuchalis transversa at the
prominentia exoccipitalis (described above). The crista ventralis
overhangs the ventral region of the caudal cranium which is steeply
angled rostroventrally and encloses a deep depression on either of
its ventrolateral sides by the more lateral prominentia exoccipitalis
and processus paroccipitalis. This depression is more common in
anseriforms than in galliforms in which the crista ventralis is less
distinct and more rounded, although some, e.g. Sylviornis neocale­
doniae, and to a slight extent, Alectura lathami, also have this region
depressed and the crista, rostroventrally angled. Differing from
dromornithids however, this region in anseriforms appears almost
dorsoventrally compressed, and in many (e.g. Melanitta perspicil­
lata), the condylus occipitalis is ventrally displaced nearer to dor­
soventral alignment with the lamina parasphenoidalis.
The condylus occipitalis is rounded laterally and ventrally and
has a shallow incisura mediana condyli flattening the dorsal surface
and creating a smooth transition into the ventral floor of the fora­
men magnum (see also in Stirling 1913: plate XXXVI, Figure 3).
The incisura is intermediate in depth in G. newtoni compared to
other dromornithids; in I. woodburnei, this dorsal concavity is
extremely shallow, yet in D. planei and D. murrayi, it is distinct,
more like the deep, curved incisura in many extant and extinct
galloanserans, including S. neocaledoniae and Ga. giganteus
(Andors 1988, p. 138). The condylus occipitalis is pedestalled on
a slightly constricted column in G. newtoni, to approximately the
same extent as in I. woodburnei, although slightly less so than in
species of Dromornis. Compared to the large, rounded foramen
magnum found in most galloanserans, that of all dromornithids is
a dorsally tall, rectangular opening with sub-parallel sides and a size
that only slightly exceeds that of the condylus occipitalis. Ga.
giganteus also has a foramen magnum proportionally small com­
pared to the condylus occipitalis (Matthew and Granger 1917,
p. 312; Andors 1988, p. 183; pers. obvs). As in I. woodburnei (see
Worthy et al. 2016a) and D. planei, the condylus occipitalis and
foramen magnum are orientated directly rostrocaudally on the
caudal cranium of G. newtoni. The placement of the condylus
occipitalis and foramen magnum in D. murrayi is unique among
dromornithids in that it is more ventrorostrally positioned and
directed, similar to the plane observed in anhimids. In all dromor­
nithids, a fossa subcondylaris is lacking or very shallow; there is no
identifiable depression in the region ventral of the condylus occi­
pitalis. The depth of the fossa in other galloanserans is variable with
some species developing a distinct cavity (e.g. Tadorna
tadornoides).
The paired fossae parabasalis (Figure 5C) are shallow in
G. newtoni and most other dromornithids; in D. planei they are
deeper. The fossae are bordered medially by a distinct crest, likely
the crista fossae parabasalis, which is considered present in many
anseriforms (Baumel and Witmer 1993: annot. 86), as it separates
the foramina within the fossa parabasalis from those more medial.
It is less distinct in D. planei and species of Ilbandornis and indis­
tinct in D. murrayi. The two foramina for the n. hypoglossi (XII) are
represented by a larger foramen just medial to the fossa parabasalis,
near the condylus occipitalis, and a second smaller foramen, more
ventrally, on the medial margin of the crista fossae parabasalis.
Within the fossa, are the foramen n. vagi (X), the foramen
n. glossopharyngealis (IX), the ostium canalis carotici (for the
carotid and the branch of cranial nerve VII), and the ostium canalis
ophthalmici externi (for venae and arteria ophthalmica externa;
Baumel and Witmer 1993; Weber 1996: fig. 6; Worthy et al.
2016a; Handley and Worthy 2021). The foramen n. vagi and ostium
canalis ophthalmici externi sit dorsally within the fossa parabasalis,
with the former being the largest of the two, and both medial-most
and dorsal-most of three foramina that are nearly aligned dorso­
ventrally, including the foramen n. glossopharyngealis, and ostium
canalis carotici most ventrally. Damage and sediment infilling limit
further interpretation on the morphology of this region in
G. newtoni. The placement and arrangement of the fossa parabasalis
and the associated foramina does not appear to vary much from
that of other dromornithids. Variation is present between dromor­
nithids and Ga. giganteus, as well as extant galliforms and anseri­
forms, where the fossa is located further lateroventrally from the
condylus occipitalis; in some anseriforms, the location transitions
to be nearly on the ventral surface of the cranium (e.g. Oxyura
australis, Cygnus atratus, Tadorna tadornoides, and Melanitta
perspicillata).
Notably like the condition observed in Anhima cornuta, ventral
of the fossa parabasalis in all dromornithids, the crista basilaris
transversa and crista basilaris lateralis (see Landolt and Zweers
1985, p. 625) are relatively indistinct, although where they intersect
is a prominent (10.4 mm wide in G. newtoni) and rounded tuber­
culum basilare. The bilaterally paired structures here identified as
tubercula basilaria (see Baumel and Witmer 1993: annot. 83) are
synonymous with the mamillar tuberosities as described by Worthy
et al. (2016a). Each represents the area for attachment of the heavy
 HISTORICAL BIOLOGY 1109

Figure 5. Caudal and ventral views of the cranium of Genyornis newtoni, digitally cropped from the images of the entire skull: A. SAMA P59516 image and annotated outline
in caudal view; B. SAMA P59516 fossa parabasalis and associated foramina for nerves, focus region outlined in A.; C. Complete ventral view of SAMA P59516 with box
indicating region featured in D.; D. Ventral cranium, SAMA P59516 image and annotated outline, rotated anti-clockwise from C. Annotations: can., canal tentatively
identified as canalis orbitalis (see Baumel and Witmer 1993: annot. 95); cfp., crista fossae parabasalis; coe., ostium canalis ophthalmici externi; cond.oc., condylus occipitalis;
cr.bas.lat., crista basilaris lateralis; cr.nu.trans., crista nuchalis transversus; cr.ven., crista ventralis; f.para., fossa parabasalis; fng., foramen n. glossopharyngealis (IX); fnh.,
foramen n. hypoglossi (XII); fnv., foramen n. vagi (X); inc.mc., incisura mediana condyli; for.mag., foramen magnum; i.para., lamina parasphenoidalis; m.rec., insertion for
musculus rectus capitis dorsalis; occ., ostium canalis carotici and branch of nerve VII; pr.basi., processus basipterygoideus; pr.par., processus paroccipitalis; prom.ex.,
prominentia exoccipitalis; r.para., rostrum parasphenoidalis (basis rostri parasphenoidalis); s.int., septum interorbitale; tu.au., tuba auditiva communis; tubc.bas.,
tuberculum basilare. Scale bars: A., D. 20 mm, B. 10 mm, C. 40 mm. Dark grey shading indicates regions where damage precludes morphological assessment, and light
grey indicates foramen and fossae. Dotted lines in D. indicate the alignment and distortion of the sagittal plane of each region.

aponeuroses for insertion of m. rectus capitis ventralis, pars later­
alis, and for slips of the m. rectus capitis dorsalis on the caudal side
(see Landolt and Zweers 1985, p. 646–649; see Figure 4). The
tuberculum basilare forms a ventral protuberance of the basiocci­
pitale at its rostrolateral junction with the exoccipital, prooticum,
and basisphenoidale (Worthy and Scofield 2012), which are over­
laid during ontogenetic development by the lamina parasphenoi­
dalis (Jollie 1957; Baumel and Witmer 1993). Differing from our
use, Landolt and Zweers (1985, p. 625, 1987) instead used the term
tuberculum basilare to refer to the bulla basitemporalis (sensu
Davids 1952b). This emanates from the lamina parasphenoidalis
in many anseriforms, e.g. Anas platyrhynchos, and galliforms, e.g.
Alectura lathami, and is here considered absent in dromornithids.
We choose to follow Baumel and Witmer (1993) for nomenclatural
consistency, and as disparity exists over whether the term ‘mam­
millary processes’ (sensu Pycraft 1902) refers to the tubercles, or to
the processus medialis parasphenoidalis (Bock 1960b; sensu Baumel
and Witmer 1993: annot. 97; Livezey and Zusi 2006: char. 0123,
p. 44, fig. 8); confusion is likely due to the two coinciding in some
birds (see Baumel and Witmer 1993: annot. 83). Bock (1960b)
considered the processus medialis parasphenoidalis to be absent
in all Galloanserae, and thus we also assume its absence in
dromornithids.
The attachment surfaces of the cervical muscles on the basis
cranii externa (basicranium) are not well-pronounced in
G. newtoni although they are in I. woodburnei, specimen
QMV:2000:GFV:20 (see Figure 4), and D. planei, specimen
NTM P9464–106, so inferences of muscle attachments are facili­
tated by these two specimens (also see Boas 1929; Davids 1952b;
Goodman and Fisher 1962: fig. 6–7, p. 118–122; Fujioka 1963:
pl. VI; Zweers et al. 1987; Böhmer et al. 2020; Worthy et al.
2016a). The ventral surface of the tuberculum basilare acts as
the insertion point of m. rectus capitis ventralis, pars lateralis.
The m. rectus capitis ventralis, pars medialis, attaches further
1110 P. L. MCINERNEY ET AL.
rostrally on a large, round area of the flattened surface of the
lamina parasphenoidalis. A distinct impression for the insertion
of one of three paired branches of m. rectus capitis dorsalis is,
in D. planei, on the caudodorsal surface of the tuberculum
basilare, and appears as a slight depression in G. newtoni. The
remaining two paired heads of the m. rectus capitis dorsalis
each have distinct, small, and ovular correspondingly paired
attachment depressions (also see Lautenschlager et al. 2014;
Worthy et al. 2016a). The most rostroventral of the two sits
medially to the tuberculum basilare, on the caudal side of crista
basilaris transversa, whereas the more caudomedial insertions
are just ventral of the condylus occipitalis, in an area that would
relate to just rostral of fossa subcondylaris if it was marked,
near the mediolateral midline of the skull. In non-anhimid
anseriforms, aponeurotic attachment marks associated with the
insertions of the m. rectus capitis dorsalis branches are formed
on the caudal side of, and medial to the tubercula basilaria, and
in some taxa, e.g. Cygnus atratus, these structures become con­
tinuous along the crista basilaris transversus. Additionally, each
tuberculum basilare is ventrolaterally extended along each
respective crista basilaris lateralis, relating to a more developed
region of insertion for the m. rectus capitis ventralis, pars
lateralis. Comparatively, in galliforms, the two cristae, the pair
of tubercula basilaria, and the aponeurotic attachment surfaces
associated with insertion of the rectus capitis ventralis muscles,
are relatively flattened or less defined/pronounced. The crista
basilaris transversa extends mediolaterally, forming a low or
indistinctive, near-straight band across the caudal basicranium,
as defined by the rostral boundary of fossa subcondylaris (see
Landolt and Zweers 1985, p. 625).
Ventral aspect
The lamina parasphenoidalis (Figure 5 C-D) presents a smooth
uninterrupted surface, curving laterally onto each parasphenoid
process of the ventral cranium. In several other dromornithids,
such as D. planei and I. woodburnei, the lamina parasphenoidalis
merges rostrolaterally with ala parasphenoidalis, however, this tran­
sition is relatively abrupt in G. newtoni, specimen SAMA P59516,
and is most similar to D. murrayi in this regard. Damage and
obfuscation in all other available specimens of G. newtoni prevents
more thorough assessment of this condition. In I. woodburnei, the
lamina parasphenoidalis tapers rostrally into a point ventrorostral
of the tuba auditiva communis (see Worthy et al. 2016a). This is
inferred for G. newtoni and species of Dromornis, as although the
region is damaged, the remaining bone suggests a similar morphol­
ogy. Aside from specimen SAMA P59516 of G. newtoni, the ros­
trum parasphenoidalis is only preserved in D. planei (NTM P9464–
106), in which it is mediolaterally wider, dorsoventrally flatter, and
proportionally more robust. The rostrum parasphenoidalis is trun­
cated just rostral of both processus basipterygoidei in G. newtoni,
whereas it ends caudal to the rostral terminus of the processes in
D. planei. Additionally, unlike in D. planei, in ventral view, the
rostrum parasphenoidalis in G. newtoni does not extend rostrally
past the craniorostral hinge, which is likely a result of the lesser
rostrocaudal compression in the cranium relative to D. planei. The
septum interorbitale extends rostrally and dorsally from the rostral-
most rostrum parasphenoidalis as a thin osseous wall between the
orbits and is well developed in SAMA P59516. The processus
basipterygoidei in G. newtoni are smaller relative to cranium size
than in D. planei although, in both, they are widely separated by the
rostrum parasphenoidalis, with flat, elliptical articular surfaces,
which protrude laterally from the rostrum parasphenoidalis on
very short, stout pedestals. The location, shape, and sessile nature
of the processus basipterygoidei in dromornithids is a conclusive
feature supporting their placement within Galloanserae
(Elzanowski 1977; Baumel and Witmer 1993: annot. 93; Weber
1993; Dzerzhinsky 1995; Livezey 1997; Murray and Vickers-Rich
2004, p. 163; Mayr 2017, p. 107) and has been used several times to
assist in the placement of other fossil birds within this radiation
(e.g. S. neocaledoniae, see Mourer-Chauviré and Balouet 2005).
Quadrate
The quadrates (Figure 6) of G. newtoni that were described by
Stirling (1913: pl. XXXVI, fig. 4–8), and again by Murray and
Vickers-Rich (2004, p. 60–62, fig. 38), were near complete.
However, they could not be located for this study so, consequently,
we redescribe the quadrates using new material (specimens SAMA
P59516, SAMA P59520, NMV P256893: fig. 6, and SAMA P53830)
in conjunction with the description and plates published by Stirling
(1913). The specimens appear to be of comparable size, although
differential preservation restricts quantitative comparisons.
Previously, Murray and Megirian (1998) suggested that the quad­
rates of G. newtoni correspond most closely with those of D. planei
although we recognise clear proportional differences between the
two species; the quadrate of G. newtoni is less robust, more tapered
dorsally, and has a more elongate pars otica and processus orbitalis.
The proportions and morphology of the quadrates of G. newtoni
are, instead, more like those attributed to species of Ilbandornis.
Regardless, all dromornithid quadrates are medially concave, form­
ing a deep c-shape from the head of the pars otica down the medial
side of the quadrate. This curvature is extreme in D. stirtoni (NTM
P3202). Only the quadrates of Anhima cornuta appear to show
a similar curvature although this is coupled with mediolateral
flattening that is absent in dromornithids.
As per the quadrate morphology of Ilbandornis sp. (NTM
P3237), D. stirtoni (NTM P5401, NTM P3202) and D. planei
(NTM P9464–100, NTM P9464–118; for all see Murray and
Megirian 1998; Murray and Vickers-Rich 2004; Worthy et al.
2016a), the head of the pars otica in G. newtoni also has a single,
sub-oval and dorsally convex compound structure comprising two
adjoined capitula, capitulum squamoso-oticum (new term, refer­
ring to the fused capitulum squamosum and capitulum oticum).
Paired with the receiving singular joint cavity of the cranium (see
cotyla quadratica squamoso-otica and recessus quadratica, above)
this is characteristic of articulatio quadrato-squamoso-otica, as in
species of Gallus, for example (see Baumel and Raikow 1993: annot.
25). Due to the lack of a distinct vallecula intercapitularis, only
a dorsally raised angularity, which represents the rostrolaterally
orientated lateral rim of the capitulum oticum articular surface,
indicates the location of the junction between the two capitula in
G. newtoni. Similarly, only the variable pronunciation of the edges
of the articular surfaces of the capitula in rostral view reveal the
extent of each capitulum in other dromornithids. In extant galloan­
serans, the capitula are distinct with a variably deep and wide
vallecula intercapitularis. In Sylviornis neocaledoniae and species
of Gastornis, the vallecula intercapitularis is shallow and the capi­
tula are closely abutting (Matthew and Granger 1917; Andors 1988;
Mourer-Chauviré and Balouet 2005; Bourdon et al. 2016). In many
anatids, the pars otica is functionally single-headed (Zelenkov and
Stidham 2018).
The shape of the conjoined capitulum in G. newtoni is similar to
that of the smaller quadrates attributed to D. planei (NTM P9464–
118) and D. stirtoni (NTM P5401); more rounded mediolaterally
and dorsally than the quadrates of Ilbandornis sp. (NTM P3237,
NTM P3235). In G. newtoni, Ilbandornis sp., and D. stirtoni (NTM
P5401), a ridge runs ventrally from the capitulum squamosum to
 HISTORICAL BIOLOGY 1111

Figure 6. Genyornis newtoni right quadrate, NMV P256893: A. Lateral view, image and annotated outline; B. Medial view, image and annotated outline; C. Dorsal view; D.
Ventral view; E. Rostral view; F. Caudal view. Annotations: cap.s-o., capitulum squamoso-oticum; cond.l., condylus mandibularis lateralis; cond.m., condylus mandibularis
medialis; con.pt., condylus pterygoideus including facies articularis pterygoidei; cr.med., crista medialis; cr.tymp., crista tympanica; d.caud., depressio caudomedialis; d.rost.,
depressio rostromedialis; d.sup., depressio supracondylaris; for.rost., foramen pneumaticum rostromediale; f.basi., fossa basiorbitalis; fov.q., fovea quadratojugalis; musc.,
tubercles for insertion of m. protractor pterygoidei et quadrati; o.cap., otic part of capitulum squamoso-oticum; p.oti., pars otica; p.sub., prominentia submeatica; s.cap.,
squamosal part of capitulum squamoso-oticum; t.sub., tuberculum subcapitulare; v.inco., vallecular intercotylaris. Scale bars: 10 mm. Dark grey shading indicates regions
where damage precludes morphological assessment, and light grey indicates the foramen pneumaticum rostromediale.

become confluent with the tuberculum subcapitulare (sensu
Elzanowski and Stidham 2010: Table 2; see also SI 2). However,
the quadrates of D. planei and D. stirtoni contrast with those in
species of Ilbandornis and G. newtoni, in that the capitulum squa­
mosum variably overhangs the shaft of pars otica; this is to the
greatest extent in D. stirtoni, where it markedly overhangs laterally
the tuberculum subcapitulare.
The tuberculum subcapitulare (eminentia articularis, sensu
Lowe 1926; Weber 1996; adductor crest, sensu Murray and Vickers-
Rich 2004) is considered a typical galloanseran character, despite its
variability in shape and size, and absence in some species of Crax
(as discussed by Elzanowski and Stidham 2010; Elzanowski and
Boles 2012; Field et al. 2020: SI, p. 28). In G. newtoni, the tubercu­
lum forms a long crest, starting from near to the lateral capitulum
squamoso-oticum and spanning up to one third of the dorsoventral
height of the quadrate. This may be associated with a well-
developed aponeurosis articularis (sensu Weber 1996), as described
for several extant galloanserans (e.g. Burggraaf 1954a; Dzerzhinsky
and Belokurova 1972; Dzerzhinsky and Potapova 1974;
Dzerzhinsky 1982; Weber 1996; Zusi and Livezey 2000; see SI 2)
and aponeurosis quadrata (sensu Dzerzhinsky and Potapova 1974;
Dzerzhinsky 1982; aponeurosis Q2, sensu Weber 1996), related to
a large point of origin for a slip of the m. AME superficialis (sensu
Holliday and Witmer 2007), that also originates on the dorsally
adjacent cranium (see ‘Cranium’; e.g. Hofer 1950; Zusi and Storer
1969; Dzerzhinsky 1982; Weber 1996; Zusi and Livezey 2000;
Holliday and Witmer 2007 and references therein; Elzanowski
and Boles 2012). This muscle origin area is particularly large in
galliforms and anhimids (Weber 1996). Comparatively, in Anseres,
the m. AME superficialis does not originate upon the quadrate.
Instead, a similarly located, but non-homologous, tubercle
predominantly corresponds to the origin of m. adductor mandibu­
lae posterior lateralis (Goodman and Fisher 1962; Vanden Berge
and Zweers 1993; Zusi and Livezey 2000; Holliday and Witmer
2007; Appendix Three). In galliforms and anhimids,
the m. adductor mandibulae posterior muscle bellies instead arise
ventrally and rostrally near the base of the orbital process (e.g.
compare Davids 1952a; Fuchs 1954a; Fujioka 1963: figs. 3–8, 10;
Dzerzhinsky 1982; Weber 1996; Zusi and Livezey 2000: fig. 8;
Dzerzhinsky and Grintsevichene 2002: fig. 7). Considering the
many similarities observed between anhimids and dromornithids,
particularly pertaining to the external adductor musculature (see
‘Cranium’), we expect homological association of the tuberculum
subcapitulare with m. AME superficialis to be more likely. In sup­
port of this assessment, the G. newtoni quadrates with complete
orbital processes figured by Stirling (1913: pl. XXXVI) possess
a well-developed crest near the basal orbital process which is con­
sistent with the more typical origin of m. adductor mandibulae
posterior (see below; also e.g. Davids 1952a; Weber 1996; Holliday
and Witmer 2007).
An especially extensive tuberculum subcapitulare is typical for
dromornithids and is proportionally shorter but still elongate in
species of Presbyornis and some megapodiids, compared to the
smaller form of other galloanserans (Elzanowski and Stidham
2010). The dromornithid form is also generally more robust
(Murray and Vickers-Rich 2004; Worthy et al. 2016a), and although
proportionally variable in size, it is uniquely unpronounced on the
quadrate of D. murrayi (see Worthy et al. 2016a: fig. 4 A, B). The
bipartite tubercle of anhimids is potentially associated with
a fragmented ancestral crest (Elzanowski and Stidham 2010;
Elzanowski and Boles 2012), which may be derived from
a structure not dissimilar to that of G. newtoni. In species of
1112 P. L. MCINERNEY ET AL.
Gastornis, the tuberculum (‘process for m. AME profundus and
associated aponeuroses’, sensu Andors 1992; tuberculum musculi
adductor mandibulae, sensu Bourdon et al. 2016) is dorsoventrally
shorter than that of dromornithids, developing more as
a prominent, conical tubercle which is hooked ventrally in
G. parisiensis (see Bourdon et al. 2016; Mourer-Chauviré and
Bourdon 2020). This tuberculum is also likely related to the origin
of m. AME superficialis in gastornithids and many galloanserans,
compared to the relatively atypical association with m. adductor
mandibulae posterior that is characteristic of Anseres (see above).
In specimen SAMA P53830 (see Appendix One: Figure A3 C),
on the medial side of pars otica, a small foramen is present, which is
absent in other specimens. The medial face in this specimen appears
flat relative to both SAMA P59520 (see Appendix One: Figure A3 F)
and NMV P256893 (see Appendix One: Figure A4) which both
have a distinct depressio rostromedialis (sensu Elzanowski et al.
2001), covering most of the medial side of the pars otica. In speci­
men NMV P256893, yet absent in SAMA P59520, a large (7.3 mm
high, 3.6 mm wide), rostrally situated and oval foramen is present
within this depression. A similarly located and obvious foramen is
also found in D. planei, D. murrayi and Ilbandornis sp. and absent
from the imperforate quadrates of D. stirtoni (see Worthy et al.,
2016a: fig. 4F). In several of the dromornithid quadrates, there
appears to be a ridge which traverses dorsally from the region of
articulation with the pterygoid and curves rostrally around the
caudal edge of fossa basiorbitalis prior to continuing along the
foramen’s ventrocaudal margin towards the caudal-most part of
the capitulum squamoso-oticum on the head of the pars otica. This
strut-like ridge is distinct in some specimens (e.g. D. planei NTM
P5401; Ilbandornis sp. NTM P3237), including in specimens of
G. newtoni (SAMA P59520; NMV P256893, SAMA P53830, as
well as the complete quadrates described by Stirling 1913: pl.
XXXVI), but is rounded and inconspicuous in others (e.g.
D. planei NTM P9464–118, only visible by manipulating specimens
under a light source). Should this ridge be homologous with the
crista medialis, then the foramen can be matched with that of the
galliform foramen type (following descriptions by Elzanowski et al.
2001; Elzanowski and Stidham 2010) and thus, following Worthy
et al. (2016a), we tentatively assign the term foramen pneumaticum
rostromediale.
Comparably, most extant anseriforms have a foramen pneuma­
ticum caudomediale; in anhimids, the foramen is considered vesti­
gial and is often absent (Elzanowski and Stidham 2010, p. 315).
While typically in the caudomedial position, in some anseriforms it
appears rostrally displaced (e.g. species within Anserinae and
Mergini, Elzanowski and Stidham 2010). In fossil taxa,
Anachronornis anhimops and Presbyornis species have a variably
developed foramen pneumaticum caudomediale or deep associated
depression (Elzanowski and Stidham 2010: fig. 5; Houde et al. 2023,
p. 17, fig. 1 L – M); Conflicto antarcticus Tambussi et al. 2019 has
both a rostromedial as well as a caudomedial pneumatic foramen
(Tambussi et al. 2019: fig. 6); early Eocene Danielsavis nazensis has
neither foramen but forms a deep depressio caudomedialis (Houde
et al. 2023: fig. 5 H – I, 7 D, p. 29; Mayr et al. 2023: fig. 3); the Upper
Cretaceous Asteriornis maastrichtensis Field et al. 2020, of near-
galloanseran affinities (Field et al. 2020) has only the rostromedial
pneumatic foramen.
The proposed homology of the foramen is further supported by
the confident identification of the crista tympanica on the caudal
pars otica in some dromornithid specimens, which is especially
pronounced in G. newtoni SAMA P53830 (see also Elzanowski
and Stidham 2010). On specimen NTM P256893, a short (6.4
mm), component of the crista tympanica is present approximately
4.8 mm caudolateral of the crista medialis and laterally delimits
depressio caudomedialis (sensu Elzanowski et al. 2001), ventral to
the compound capitular heads. The small, shallow, and triangular
depressio caudomedialis is comparatively more dorsoventrally
extensive in specimen SAMA P53830. In other dromornithids, the
caudomedial depression is ventrally confluent with a broad depres­
sion on the caudal pars mandibularis, that is especially bowl-like in
species of Dromornis. In quadrates of G. newtoni, this ventral
depression is variably deeper.
The foramen pneumaticum basiorbitale is common among
crownward birds, including those in the genera Ichthyornis,
Conflicto, Pelagornis, Presbyornis (see Field et al. 2020: SI p. 28),
Danielsavis and Anachronornis (see Houde et al. 2023; Mayr et al.
2023), as well as galliforms (Elzanowski and Stidham 2010). The
foramen is occasionally present in D. planei (e.g. NTM P9464–100)
and absent in D. stirtoni yet remains unknown for G. newtoni and
Ilbandornis sp. due to taphonomic damage. Variably located and
often vestigial foramina have been identified in a similar region of
the derived quadrates of anhimids, although the foramen pneuma­
ticum basiorbitale is absent in anatoids (Elzanowski and Stidham
2010). The foramen is also absent in gastornithids (Mourer-
Chauviré and Bourdon 2020) and likely absent in S. neocaledoniae
(see Mourer-Chauviré and Balouet 2005: fig. 6).
Small tubercles in G. newtoni just ventral to the foramen pneu­
maticum rostromediale are equivalent to what was interpreted as
the attachment for the ‘posterior ligament of the quadrate’ by
Murray and Megirian (1998) for dromornithids. This medial area,
and the more rostral fossa basiorbitalis, corresponds to the insertion
of m. protractor pterygoidei et quadrati and interrelated apo­
neuroses in many studied avian taxa (e.g. Fuchs 1954b; Starck and
Barnikol 1954), including the megapodiid Aepypodius arfakianus
by Weber (1996: Q3). Regardless of the exact homology, this rugose
attachment surface is observed in all dromornithid specimens,
although the shape of the attachment region slightly varies. For
example, in D. planei, the attachment region is small and triangular.
Despite consistent damage to the caudal side of all G. newtoni
quadrates, the presence of a well-developed prominence projecting
dorsocaudally from the pars quadratojugalis of the processus later­
alis (sensu Elzanowski et al. 2001) is evident, most especially in
specimen SAMA P59520. This correlates with the massive quad­
ratojugal eminence described by Murray and Vickers-Rich (2004,
p. 62), is visible in the quadrate images figured by Stirling (1913: pl.
XXXVI), and is also indicated by the preserved structure of the
quadrates of species of Ilbandornis and Dromornis despite also
having consistent damage to the region. Articulated quadrates of
G. newtoni (SAMA P59516, NMV P256893, SAMA P59520), show
this projection to be directed towards the dorsal processus parocci­
pitalis and aligned with the ventrocaudal bounds of the osseous
meatus acusticus externus. The position and orientation of this
structure suggests it is homologous with the prominentia submea­
tica (sensu Elzanowski 1987) of anatoids (also noted by Elzanowski
and Boles 2012, p. 908), however, it is a comparatively inflated
structure in these taxa, instead of the strong projection that appears
characteristic of dromornithids (pers. observ.; see also Elzanowski
and Stidham 2010). The prominence likely provided attachment for
the membrana postmeatica (Elzanowski 1987; sensu Baumel and
Raikow 1993: annot. 37) and was closely associated with
the m. depressor mandibulae and incorporated aponeuroses
(Elzanowski 1987). Together, these soft-tissue structures would
have effectively participated in the caudal wall of the external
auditory canal, as it does in anatoids and other taxa which possess
a homologous prominentia, e.g. tinamous (see Davids 1952a;
Elzanowski 1987 and references therein; Elzanowski and Stidham
2010). A more medial processus submeaticus (sensu Elzanowski
and Stidham 2010), a feature common among anseriforms

including anhimids, which hosts tissues that connect with the
processus suprameaticus of the cranium (and more closely asso­
ciated with the rostral bounds of meatus acusticus externus), is
absent in G. newtoni and all dromornithids (see ligamentum post­
quadratum, sensu Dzerzhinsky 1982; Elzanowski 1987). In galli­
forms, the prominentia submeatica occurs only in cracids, and
a processus submeaticus is absent (Elzanowski and Stidham
2010). Additionally, quadrates of Presbyornis species (see
Elzanowski and Stidham 2010), Ga. parisiensis (see Bourdon et al.
2016: fig. 2), Ga. giganteus (see Matthew and Granger 1917: Pl.
XXII, fig. 2a) and S. neocaledoniae (see Mourer-Chauviré and
Balouet 2005: fig. 6 E-F) all lack both the prominentia submeatica
and processus submeaticus.
The only complete orbital processes of the quadrate known
for G. newtoni are those presented by Stirling (1913), from
which this updated description is based. The processus orbitalis
of D. stirtoni (specimen NTM P3202) and the quadrates
described by Stirling (1913) for G. newtoni are similar in length
to that of the pars otica from which it extends rostrally at
a wide angle. The rostral tip of the orbital process in
G. newtoni is slightly curved ventrally and medially in associa­
tion with the origin of m. pseudotemporalis profundus along its
edge (e.g. see Davids 1952a; Fuchs 1954a). The straight dorsal
margin intercepts the pars otica just ventral of the capitulum
squamoso-oticum, the latter is only comparable to that of
Anseranas semipalmata; in anatids (but not Dendrocygna), this
dorsal margin is markedly concave. Comparably, the right angle
between the processus orbitalis and pars otica in gastornithids
and S. neocaledoniae, is like many other anseriforms, and smal­
ler than most galliforms (see also Elzanowski and Stidham
2010).
As in many galloanserans, including species of Anhima,
Anseranas, Oxyura and Cygnus, and G. parisiensis (see Bourdon
et al. 2016), the fossa basiorbitalis on the medial side of the proces­
sus orbitalis is large and deeply excavated for insertion
of m. protractor pterygoidei et quadrati (e.g. Davids 1952c; Fuchs
1954b; see Stirling 1913). Rostrally, a second depression, likely the
depressio protractoris (sensu Elzanowski et al. 2001) is evident and
separated from the fossa basiorbitalis by a prominent, rostrocaud­
ally narrow ridge. On the lateral surface of the orbital process,
a sharply-ridged crista orbitalis (sensu Elzanowski et al. 2001)
traverses longitudinally along the long axis from the rostral apex
before turning sharply obliquely to extend close to the dorsal
margin of the process. While probably associated
with m. pseudotemporalis profundus rostrally, the latter structure
and flattened surface in between evidences an especially well-
developed attachment site predominately corresponding to the
origin of m. adductor mandibulae posterior (see above, also e.g.
Davids 1952a; Fuchs 1954a; Goodman and Fisher 1962; Weber
1996).
There is no evidence for a distinct facies articularis pter­
ygoidea ventrally on the processus orbitalis. The morphology
of this region is instead more like the anseriform and cracid
conditions: the raised condylus pterygoideus is dorsally con­
fluent with, or adjacent to, the facies articularis pterygoidea
(see Dzerzhinsky 1982; Elzanowski and Stidham 2010;
Zelenkov and Stidham 2018). In G. newtoni, the condylus
pterygoideus is a small oval tuberosity (Murray and
Megirian 1998; Murray and Vickers-Rich 2004), less distinct
than in other dromornithid quadrates (see Worthy et al.
2016a). In all, the ventral margin of the condylus pterygoideus
meets the rostral edge of the condylus mandibularis medialis
(see also Stirling 1913, p. 119; Elzanowski and Stidham 2010;
Worthy et al. 2016a).
HISTORICAL BIOLOGY 1113
All dromornithids have the galloanseran bicondylar form of the
mandibular process (Murray and Vickers-Rich 2004; Mayr 2017;
Worthy et al. 2017b), although the pars mandibularis is consider­
ably larger proportionally in species of Dromornis than it is in
G. newtoni and Ilbandornis species. The complete articular surface
is narrow, the condyles are near equal in size, and both are rostro­
caudally elongate ovate shapes, tapering towards the midline of the
articular surface with overlap of their long axes just greater than half
the length of the condyles. Unlike other galloanserans, the articular
surface is relatively flat, the two condyles are not well defined
individually, and the vallecular intercotylaris is shallow and dorso­
ventrally wide. The condylus mandibularis lateralis is markedly less
ventrally convex and the condylus mandibularis medialis, slightly
more so (see also Worthy et al. 2016a). Comparatively, in gastor­
nithids and S. neocaledoniae, both condyles are ventrally convex,
separated by a deep vallecula intercondylaris (Bourdon et al. 2016;
Worthy et al. 2016a; Mourer-Chauviré and Bourdon 2020).
For articulation with the jugal arch, galloanseran quadrates,
including in Ga. parisiensis and S. neocaledoniae (see Bourdon
et al. 2016; Mourer-Chauviré and Bourdon 2020) typically have
a deep bowl-like fovea quadratojugalis linking the distinct facies
quadratojugalis ventralis and facies quadratojugalis dorsalis.
Comparatively, in G. newtoni and other dromornithids, the
fovea quadratojugalis is a large, shallow, plate-like structure at
the base of the prominentia submeatica with no distinct facies
quadratojugalis dorsalis. Only in Ilbandornis sp. specimen NTM
P3235 is there a small, raised shelf which indicates the potential
presence of the facies articularis ventralis. In G. newtoni, an
angular ridge defines the ventral margin of the fovea quadrato­
jugalis and overhangs the lateral mandibular condyle. Whether
the rim extends dorsally cannot be determined due to tapho­
nomic damage in all quadrates we observed for G. newtoni. An
embayment, separated from the depressio subcondylaris by
a ridge, is present between the fovea quadratojugalis and the
lateral mandibular condyle. This is relatively shallow in
G. newtoni, as in species of Ilbandornis, whereas in D. stirtoni,
it is especially distinct. A depressio supracondylaris (sensu
Elzanowski and Stidham 2010) appears to be present just ros­
tromedial to the fovea quadratojugalis in the Stirling (1913: pl.
XXXVI, fig. 4–8) quadrates of G. newtoni; however, equivalents
cannot be verified in any available specimens.
Pterygoid
There are two poorly preserved, disarticulated pterygoids in the
caudal area of specimen SAMA P 59516, which are visible from
ventral perspective, on left and right sides of the skull
(Figure 1). The right pterygoid appears to present a cup-like
facies articularis quadratica, similar to the pterygoids of other
dromornithids. The fragmentary nature of these elements pre­
vents more comprehensive description.
Rostrum
The rostrum (Figure 7) of Genyornis newtoni is composed of
completely synostosed elements (praemaxillaria, maxillaria,
nasalia, palatina, and vomer), indicating that all studied speci­
mens represent adult individuals. The lacrimalia also appear
to be completely fused to this structure. There is no complete
arcus jugalis preserved for any specimen although fragments
of the jugal arches and caudal maxillare (i.e. mainly contrib­
uted to by the processus jugalis of the maxillare) are used to
interpret morphology (see Table 1). In dorsal view, the ros­
trum is nearly three times longer than its maximum width
1114 P. L. MCINERNEY ET AL.

Figure 7. Genyornis newtoni rostrum, lateral views: A. SAMA P59521 left lateral view image and annotated outline; B. SAMA P59521 right lateral view; C. SAMA P59517 left
and right lateral views. Annotations: ang., angulus tomialis; ca., casque; cr.tom., crista tomialis; fov., foveae corpusculorum nervosorum; lac., lacrimal ; margo.lat., margo
lateralis palatini; nas., apertura nasi ossea; pala., palatinum; p.o.l., processus orbitalis of lacrimale; pr.jug., processus jugalis of the maxillare; pr.max., processus
maxillopalatinus of the maxillare; pr.n.m., processus nasalis of the maxillare; sul.para., sulcus paratomialis. Scale bars: 20 mm. Dark grey shading indicates regions
where damage precludes morphological assessment, and light grey indicates fenestrae, foveae, and aperturae.

(SAMA P59521, see Appendix Two) and has a broad and
rounded tip. The lateral margins narrowly diverge and are
notably convex, the culmen is sharply angled over the mid-
length zone, the sides are flat or slightly shallowly concave,
and in lateral aspect, the tomial surface is flat. The rostrum of
specimen SAMA P59521 is larger than SAMA P59517, and
greater in dorsoventral height although the height of both
specimens is proportionate to length. SAMA P59517 is likely
from a male (Chinsamy and Worthy 2021), with the SAMA
P59521, thus presumed to be of the smaller female morph.
Dorsal and lateral view: os praemaxillare et os nasale
On the dorsocaudal surface of the rostrum, a well-defined,
lowly-raised, triangular structure, interpreted as a ‘casque’,
abuts the entire width of the craniorostral hinge and rostrally
terminates in a rounded point at the dorsal culmen. Complete
synostosis of the processus praemaxillaris nasalis (sensu Livezey
and Zusi 2006) and the processus frontalis praemaxillaris (sensu
Livezey and Zusi 2006) contribute to the casque. The casque
may be associated with the conchal and or maxillary diverticula
of the antorbital sinus (Witmer 1990; Mayr 2018b) and the
presence of paired depressions covered with dimples on the
dorsal surface, bilateral to the midline, may suggest this area,
at least, was highly vascularised. Although the casque is absent
in other dromornithid species with identified rostra (Dromornis
planei and D. stirtoni), two specimens of D. planei (NTM
P9973–2 and NTM P9464–107) have distinct grooves which
follow the same triangular shape of the casque from the dorsal
borders of the aperturae nasi osseae (external nares; sensu Zusi
1993) to meet rostrally along the culmen. The triangular space
within is also heavily vascularised in D. planei specimens, most
conspicuously in NTM P.9973–2 and NTM P932–2. The casque
of S. neocaledoniae is similarly located on the rostrum (rather
than the cranium), although it is a much larger, dorsally pro­
minent, ornament (Mourer-Chauviré and Balouet 2005). Few
other galloanserans have such bony ornamentation on the skull
and rostrum with exceptions being species of Melanitta, and
some species of Numididae and Cracidae (e.g. Mitu mitu; see
Mayr 2018b for a review).

Figure 8. Dromornis planei rostrum, specimen NTM P932–2, shows distortion and cavitation on the surface of the bone: A. Right side; B. Left side; C. Close-up image of the
distortion on the rostrodorsal-most part of the left side of the rostrum. Annotations: ar.jug., arcus jugalis; cr.tom., crista tomialis; fen., fenestra; lac., lacrimal; p.o.l., processus
orbitalis of lacrimal. Scale bars: A., B. 50 mm and C. 20 mm.
The rostrum, as per other dromornithids, is dominated by large,
flattened plates of the paired praemaxillaria and maxillaria, which
fuse into a single robust structure, a feature shared with other giant
galloanserans such as the gastornithids and sylviornithids (see
Andors 1988, 1992; Mourer-Chauviré and Balouet 2005), as well
as anseriforms, and some cracids. Comparatively, most galliforms
have weakly fused upper jaw elements (see Field et al. 2020). Aside
from taphonomic fragmentation, especially prominent in SAMA
P59517, the surface texture of the plates is smooth and likely were
shallowly depressed (accommodating for slight taphonomic defor­
mation). There is no evidence of the extensive cavitation and fold­
ing that is prominent on specimen NTM P932–2 of Dromornis
planei (Figure 8) and NTM P9245 of D. stirtoni (Murray and
Megirian 1998: fig. 15; Murray and Vickers-Rich 2004: fig. 195).
Due to differential preservation on both sides of these rostra, the
cause of this surface texture remains unknown, although it could be
associated with a soft-tissue structure on the dorsal rostrum, or
alternatively be pathological in nature. On the Genyornis rostrum,
there is no indication of the extent of the rhamphotheca – unlike the
HISTORICAL BIOLOGY 1115
horizontal groove present in Dromornis specimen NTM P.9464–
107. This may suggest that the rhamphothecal sheath covered the
complete rostrum aside from the casque.
The rostrum is widely triangular in transverse section, contra
Murray and Megirian (1998, p. 59, 67) who described the rostrum
of G. newtoni as laterally compressed, reminiscent of the extremely
narrow rostrum attributed to species of Dromornis. However, reas­
sessment of the latter, suggests taphonomic mediolateral compres­
sion may have been underestimated. In specimen NTM P.9973–2
(D. planei), the only seemingly uncompressed rostrum of
Dromornis, the angle at which the lateral sides diverge from the
culmen increases ventrally, suggesting the maximum mediolateral
width of the ventral rostrum (not preserved), would be far greater
than that of the preserved part, and so at least equal in width to the
cranium. Should this be the case, the relative width of the rostrum
may not vary so drastically from that of G. newtoni.
The culmen in lateral aspect, gently curves ventrally towards its
tip, lacking the hemispherical curvature of the dorsal profile char­
acteristic of species of Dromornis (see Murray and Megirian 1998:
1116 P. L. MCINERNEY ET AL.

Figure 9. Genyornis newtoni rostrum, specimen SAMA P59521, and comparison of palatine and jugal arch morphology: A. Ventral/Palatal view image and annotated
outline; B. Dorsal view image and annotated outline; C. Caudal view image and annotated outline; D. Hypothetical reconstruction of the palatines of Genyornis newtoni
(pale orange), pars lateralis palatini (dark orange), and the jugal arch (brown) from specimen SAMA P59521; E. the galliform condition, Ortalis canicollis (see Zusi and
Livezey 2006: fig. 7A); F. the anseriform condition, Anser albifrons (see Zelenkov and Stidham 2018: fig. 2A). Annotations: ang., angulus tomialis; bulb., bulbous regions
largely associated with each palatine including complete synostosis with the processus maxillopalatinus of the maxillare, the processus rostralis of the palatines, and the
processus palatinus of the praemaxillary; ca., casque; cho.na., choana nasalis ossea; cr., craniorostral hinge; cr.tom., crista tomialis; cul., culmen; fen.pal., fenestrae palatina;
fos.cho., fossa choanalis palatini; fov., foveae corpusculorum nervosorum; lac. – lacrimal; iam.d. – lamella dorsalis, pars choanalis palatini; margo.lat., margo lateralis palatini;
os.pal., ossa palatina; pal., palatum osseum; pons.mj. – pons maxillaro-jugalis; pr.cho. – processus choanalis palatini; pr.jug., processus jugalis of the maxillare; pr.max.,
processus maxillopalatinus; sep.na., septum nasale osseum; sn.pl., supranarial plate (of processus maxillopalatinus); sul.para., sulcus paratomialis and associated depression;
sut.Vm., sutura vomeromaxillaris; vom., vomer. Scale bars: 20 mm, D.-F. not to scale. Dark grey shading indicates regions where damage precludes morphological
assessment, light grey indicates sulci, fenestrae, foveae, and aperturae, dotted lines follow the rostral continuation of the sulcus paratomialis, and the white arrows in B and
C indicate the lateral articular ‘condyles’ (internal processes of the nasolacrimals, sensu Murray and Vickers-Rich 2004).

fig. 6, 7, 15, 28, 30) and the relatively planar deep gastornithid rostra towards the rostral apex is absent in Dromornis. The rostral apex
(Andors 1988: pl. 1; Andors 1992: fig. 1, 2; Bourdon et al. 2016: is mediolaterally wide and rounded with numerous foveae corpus­
fig. 1). Conversely, the dorsoventral shallowing of the culmen culorum nervosorum perforating it, as common among

anseriforms (Baumel and Witmer 1993: annot. 41), which Murray
and Megirian (1998) used to argue the presence of a thinly cornified
and well-developed nail on the bill tip. Although absent or very
subtle in G. newtoni, there is a slight rostral hook on the end of the
rostrum of D. stirtoni (see Murray and Megirian 1998). Both
Sylviornis neocaledoniae (see Mourer-Chauviré and Balouet 2005:
fig. 5), the juvenile gastornithid Omorhampus storchii (previously
synonymised with Ga. giganteus by Andors 1988; Louchart et al.
2021), and specimens of Ga. parisiensis show a ventral curve of
crista tomialis suggesting a small hook-like structure. A rostrally
hooked bill is common among the galliforms and some anseri­
forms, including Anhima cornuta, Anseranas semipalmata,
Presbyornis species (see Zelenkov and Stidham 2018),
Dendrocygna species and Oxyura australis, although in non-
anhimid anseriforms the hook is not a sharp point but instead
generally forms a rounded lip.
The external nares (aperturae nasi osseae) are holorhinal (the
caudal border of the nostrils end rostral to the craniofacial hinge
and are generally oval in shape, e.g. Bock 1964), as is common
among galloanserans (Sibley and Ahlquist 1990; Dzerzhinsky 1995).
Relative to the size of the rostrum, they are exceedingly small,
circular, and located dorsocaudally, differing in location from the
further ventral but similar shaped nares of gastornithids (Matthew
and Granger 1917; Andors 1988, 1992). The external nares are
slightly more rostrocaudally elongate in SAMA P59516, compared
to that of SAMA P59517 and P59521. Three shallow grooves radiate
rostrally from a neurovascular foramen, located on the lateral sur­
face, just rostrad of the nasal aperture. These correspond well with
passages for nerves and blood vessels (see Murray and Vickers-Rich
2004, p. 237, fig. 185), potentially terminal ramifications of the
ophthalmic (V1, rami rostri maxillaris from praemaxillary rami of
ramus medialis and rami nasales interni of ramus lateralis) and
maxillary nerves (V2, specifically rami nasales externi of nervi
nasopalatinus), which can be extensive and well developed in this
area of the rostrum in anseriforms (see Bubień-Waluszewska 1980).
A second foramen and deep, dorsoventrally aligned vascular groove
are located caudolaterally from the nares, on each post-narial bar
composed of the fused processus nasalis of the maxillare, processus
maxillaris of the nasal, and the lacrimal. The region surrounding
and rostral of the nares is more heavily vascularised in species of
Dromornis, with a ‘narial groove’ Murray and Vickers-Rich (2004)
attribute to functional ‘nasal salt glands’ (i.e. fossa glandulae nasa­
lis). These grooves may be better compared with the sulcus nasi that
is typical of gastornithids (see Bourdon et al. 2016; Mourer-
Chauviré and Bourdon 2020). We do not interpret any furrow as
related to nasal salt glands in G. newtoni (see also ‘Orbit and
rostrodorsal region of the cranium’).
Viewed laterally, the fused processus nasalis of the maxillare,
processus maxillaris of the nasal, and lacrimal (post-narial bar) is
aligned obliquely with respect to the dorsoventral plane. Fused
laterally to the casque, the dorsal-most lacrimal appears to lack
a distinct processus supraorbitalis, instead terminating caudally in
line with the craniorostral hinge (see ‘Craniorostral hinge’). The
processus orbitalis (also termed the descending or ventrocaudal
process of the lacrimal; Cracraft 1968; De Mendoza et al. 2020)
follows the alignment of processus maxillaris, mediolaterally thin­
ning ventrally and projecting caudally as a distinct angular flange of
bone at its ventral terminus, immediately dorsal and lateral to the
co-ossified rostral-most processus jugalis of the maxillare (see
below). In D. planei NTM P932–2, there is contact between the
lacrimal component and the jugal arch, and a sulcus formed
between them on the lateral side. Rostral of their intersection, an
obvious fenestra is evident (Figure 8), which has not formed in
G. newtoni. The homological nature of the structural link between
HISTORICAL BIOLOGY 1117
the descending orbital process of the lacrimal and the jugal arch in
dromornithids is unknown, due to the completely fused morphol­
ogy of the former part; whether it was derived from ossification of
ligamentum jugolacrimale (e.g. os lacrimale communicans of
Cariamiformes, see Degrange et al. 2015; Degrange 2021) or con­
tributed to by other bones (Mayr 2022b). In anatids, the jugo-
lacrimal ligament provides a strong connection between two well
developed processes on both the lacrimal and jugal arch (Davids
1952c; Zweers 1974).
Palatum osseum
Following Murray and Vickers-Rich (2004), we here, consider the
dromornithid palate, desmognathous (as described by Huxley
1867). Desmognathy is typical for anseriforms, and some basal
galliforms, including some cracids and megapodiids (Dzerzhinsky
1995; Zusi and Livezey 2006; Mayr 2018a), and is considered to
have derived several times from a relatively plesiomorphic schizog­
nathous palate (Hofer 1945), which is observed in some galliforms
and Asteriornis maastrichtensis (see Field et al. 2020: SI, p. 22).
The ventral rostrum of Genyornis newtoni consists of
a continuous, plate-like palatum osseum, comprising synostosed
structures relating to the praemaxillaria, maxillaria, palatina, and
vomer (see Huxley 1867; Zusi and Livezey 2006), as in species of
Dromornis, gastornithids (Matthew and Granger 1917; Andors
1988), and sylviornithids (Worthy 2000; Mourer-Chauviré and
Balouet 2005). Similarly, all anseriforms have extensive palatal
development due to the central fusion of the paired processus
maxillopalatinus of the maxillaria, although their palate retains
a central fenestra. The development of a bony palate is variable in
galliforms though restricted to the rostral-most region of the upper
bill. In G. newtoni, the shallow nature of the palate suggests a large
internal cavity within at least the caudal ¾ of the rostrum.
The palatal region of Dromornis planei was described from
specimen NTM P.9464–107 (see Murray and Megirian 1998),
although direct observation suggests the compression and subse­
quent damage to the specimen had obscured most features, and the
palatines are not preserved. The preserved osseous plate present on
the ventral aspect of the D. planei rostrum NTM P. 9973–2, and
partially retained in NTM P.932–2, is not considered the palatum
osseum; instead, we identify this as the ventral surface of the pila
supranasalis (composed of the praemaxillary processes of the nasale
and the frontal processes of the praemaxillare) which forms the roof
of the internal nasal cavity. This is for several reasons: a) in the
D. planei rostrum NTM P.932–2, the dorsolateral face of the prae­
maxillaria extend considerably further ventral than the roof; b) the
external nares open internally within the rostrum, ventral to this
osseous roof; and c) neither specimen retains any traces of the
palatines or associated structures which would be expected should
this be the palatum osseum.
Two sutures on the palatum osseum, at margo medialis palatini,
are each representative of the intersection between the ventral
processus maxillopalatinus of the maxillare (sensu Baumel and
Witmer 1993: annot. 58; Zusi and Livezey 2006; Mayr 2018a,
p. 3), and the more lateral, raised and synostosed structure formed
of the processus praemaxillaris, processus rostralis of the palatines,
and processus palatinus of the praemaxillary bone. These sutures
traverse rostrad in parallel towards the apex rostri, and possibly
supported a median palatal ridge (ruga palatina mediana) between
them (see McLelland 1993). The medial fusion of the paired pro­
cessus maxillopalatinus in G. newtoni, an anseriform feature
(Baumel and Witmer 1993: annot. 58), creates a single suture rostral
of the vomer (see below), contributing to the rostrocaudally con­
tinuous, shallowly concave, palatum osseum. Three small, oval
1118 P. L. MCINERNEY ET AL.
fenestrae palatina (sensu Baumel and Witmer 1993: annot. 13) are
mediolaterally aligned with this central suture and spaced caudor­
ostrally upon the palatum osseum. The median depression is likely
analogous to the ‘median suture’, that is described as dividing the
palatines and maxillopalatine process in gastornithids (see Matthew
and Granger 1917). The pits on the palatum osseum in the juvenile
gastornithid specimens, identified as neurovascular canals by
Louchart et al. (2021), are absent in dromornithid specimens,
although these authors associated the presence and size with the
age of the individuals, which suggests that, even if they were present
through the ontogenetic cycle of G. newtoni, they would be absent
in the adult specimens available.
Most galloanserans and the gastornithids (see Andors 1988;
excluding Omorhampus storchii, see Louchart et al. 2021: fig. 1
and G. parisiensis) have a variably thin and sharp crista tomialis.
In G. newtoni, the crista tomialis is instead goose-like (e.g. Anser
anser) with an angular junction between the dorsally ascending
lateral plates of the rostrum (praemaxillaria). It is continued by
a distinctly flattened tomial surface, which widens towards the
rostral apex, and is characterised by rugose, osseus material. In
some anatids, similar osseous material is observed on macerated
skulls, which support the ventrally overlying rhamphothecal lamel­
lae rostri on the tomium maxillare (e.g. Zweers et al. 1977).
However, Murray and Vickers-Rich (2004, p. 168) considered it
unlikely that dromornithids ‘would have developed any but the most
rudimentary lamellae’ on their tomia. The sulcus paratomialis
(sensu Livezey and Zusi 2006) in G. newtoni shallowly depresses
the rostrocaudal length of the tomial shelf, is confluent with con­
cavitas palati, and becomes deepest caudally towards the processus
jugalis of the maxillare, just lateral of the palatines. Unlike a typical
anatid rostrum, the crista tomialis (and angulus tomialis) migrates
dorsally in the caudal area, corresponding with this change in sulcus
depth. This is notably similar to the morphology of Anhima cornuta
in which a deep depression is present just rostral of the processus
jugalis, lateral of the processus maxillaris of the palatines.
Compared to G. newtoni, the sulcus upon the tomial surface in
D. planei (see Murray and Megirian 1998) is more mediolaterally
narrow in the rostral half, widening and becoming shallower later­
ocaudally in effect of the dorsal retreat of crista tomialis. Regardless,
they are overall similar in the sense of relating to the overlapping
caudal tomia of the articulated jaws, described by Murray and
Vickers-Rich (2004), which was functionally attributed to
a sheering facet and differentiated from the more rostral tomia in
species of Dromornis.
Maxillare, septum nasale osseum and arcus jugalis
The processus maxillopalatinus of the maxillare is not visibly tra­
beculated as in Anseranas semipalmata, however, the process is
enlarged and shows extensive co-ossification, a feature common
among anseriforms but not galliforms (Mayr 2018a). This region of
the rostrum is best preserved on specimen SAMA P59521, and
therefore, forms the basis of the following descriptions. As in
anseriforms, the paired maxillopalatine processes of the maxillaria
are fused to one another along the mediolateral midline of the
rostrum, however, this is more extensive in G. newtoni and appears
to continue dorsocaudally towards the craniorostral hinge to form
an apparent supra-narial plate (sn.pl., Figure 9), notably similar to
parrots. The plate appears to join the dorsal-most nasal bones
caudodorsally although crushing is evident in this region, and so
this may be a taphonomic feature. By comparing the morphology of
the synostosed ventral nasal and praemaxillary bones (pila supra­
nasalis) in specimens of Dromornis, it is likely that these plates were
not entirely fused to the ventral casque-region, but instead formed
a shelf which air could pass over and into a cavity within the
rostrum. Caudally along the mediolateral midline, a thin ridge of
bone traverses from the craniorostral hinge to the vomer; we inter­
pret this ridge to be an incorporated septum nasale osseum, which
also co-ossifies to the processus maxillopalatinus of the maxillare in
parrots (Mayr 2018a). While damaged, the lack of any prominent
caudal extension of the septum nasale osseum suggests that it was
not connected to the more caudal mesethmoidale and the inter­
orbital septum, thereby creating a hiatus craniofacialis septi ventrad
of the craniorostral hinge (Jollie 1957; Baumel and Witmer 1993).
This is entirely consistent with the high mobility of the rostrum
relative to the cranium as predicted by the mobile craniorostral
hinge (Craniorostral hinge Murray and Vickers-Rich 2004, p. 235).
Laterally, this supra-narial plate of the processus maxillopalati­
nus contacts the caudomedial side of the combined processus
maxillaris and processus nasalis for their complete dorsoventral
length (see Figure 9). This forms a distinct crest which medially
borders a dorsoventrally elongate and flat, caudally facing depres­
sion that is laterally delimited by the caudal edge of the orbital
process of the lacrimal. The depression resembles that on the caudal
lacrimal of Balaeniceps rex (Balaenicipitidae) which similarly has
lacrimals that are solely fused to the caudal rostrum (Bühler 1980;
see ‘Craniorostral hinge’). Notably, this region in anhimids is also
flattened rostrocaudally (see Cracraft 1968). This depression is
visible in rostrum specimens of D. planei, although it is shallower.
The ventral-most part of the orbital process of the lacrimal (and
associated structures, see above) meets the processus jugalis of the
maxillare laterally and forms a caudal angular flange which demarks
the ventrocaudal most extent of the depression. Immediately med­
ial, the depression contacts a prominent and deep sulcus (see light
grey filled outline, Figure 9C) which travels mediorostrally beneath
a bridge of the maxillare formed between the processus jugalis and
the medially adjacent post-narial bar (possibly processus nasalis of
the maxillare), onto the caudal surface of the supra-narial plate, just
caudodorsal of the palatines. This sulcus on the supra-narial plate is
delimited medially by a sharp crest and laterally by a sheet of
processus maxillopalatinus which is co-ossified with the dorsome­
dial surface of lamella dorsalis of pars choanalis of the palatines,
enclosing the choana nasalis ossea laterally (dashed outline,
Figure 7A). The caudolateral margin of this sheet is marked by
a robust bar, the pons maxillaro-jugalis (sensu Livezey and Zusi
2006), which buttresses the palatines dorsally and is likely part of
the maxillare. A deep cavity is present on the lateral surface of
lamella dorsalis, confluent with the sulcus paratomialis (see
above) and extends dorsocaudally onto a small triangular angulus
tomialis (sensu Livezey and Zusi 2006). Although this flange could
derive from the praemaxillare as in galliforms (see Livezey and Zusi
2006; Mayr 2018a), due to the similarities with anseriforms in this
region, it appears more parsimonious to identify it as an extension
of the maxillare.
The processus jugalis of the maxillare in SAMA P59521 is
broken caudally, at the point where arcus jugalis becomes dis­
tinct with respect to the rostrum and lacks the more caudal
region relating to fusion with the jugal bone. In anatids, and to
a lesser degree anseranatids, the processus jugalis uniquely
projects from the maxillary bone from a point ventral of, and
coincident with, the rostrolateral margin of the mediolaterally
expanded processus rostralis of the palatinum (see Zusi and
Livezey 2006, p. 155; Zelenkov and Stidham 2018).
Comparatively, in galliforms, gastornithids, and sylviornithids,
the processus jugalis is positioned far lateral and more dorsal of
the processus rostralis, so there is no interaction present
between the two processes. Neither of these states are directly
comparable to the rostrum of G. newtoni, which appears

intermediate, i.e. the processus jugalis is more mediorostral of
the galliform-position but not to the extent of the typical ana­
toid condition. Resultantly, processus jugalis extends from the
maxillary bone just dorsolateral of the processus rostralis of the
palatinum and contributes to the aforementioned depression
associated with the sulcus paratomialis (see ‘The palate: palatum
osseum’). This is very similar to the intermediate state also
observed in Anhima cornuta.
Fragmented parts of the arcus jugalis of G. newtoni can be
described from specimens SAMA P59516, SAMA P53830
(Appendix One: fig. A3a), and SAMA P10838, which corresponds
to the jugale and quadratojugale bones. The caudal part of the arcus
jugalis, which would have largely corresponded to the quadratoju­
gale bone, has a slender, slightly mediolaterally compressed shaft
which bows ventrally and then laterally towards the caudal region
to participate in the articulatio quadrato-quadratojugalis. Caudal of
the poorly developed condylus quadraticus, the terminus of the
jugal arch is rounded caudally and flattened mediolaterally as per
the description and figure of this region by Stirling (1913).
Ventral of all other processus, including the processus jugalis, on
palatum osseum, the maxillaria and praemaxillaria join the pala­
tines through robust and complete synostosis of the processus
praemaxillaris maxillaris, the processus rostralis of the palatinum,
and the processus palatinus praemaxillaris. This region is defined
rostrally by paired, rostrocaudally elongate, bulbous surfaces (see
Figure 9A: bulb.; described as ‘bulges of spongey bone’ for D. planei
by Murray and Megirian 1998) which lack a flexible zona flexoria
palatina. We assume the absence of a processus palatinus of the
maxillare (Zusi and Livezey 2006; sensu Mayr 2018a, SI 2) as this
process is not common among neognaths (nor within Galloanserae,
see Mayr 2018a: p. 7–8).
Palatinum et vomer
The pars choanalis et lateralis palatini of the palatine together form
mediolaterally flattened, and largely dorsoventrally aligned, osseous
wings. The pars lateralis palatini extends ventrolaterally from the
ventral margin of lamella dorsalis, pars choanalis palatini (espe­
cially visible in G. newtoni SAMA P59517, P59516), ventral to the
level of margo medialis palatini, a character shared with D. stirtoni
(see Murray and Megirian 1998, p. 67 and 80, fig. 15). In anseri­
forms, the pars lateralis palatini is mediolaterally narrow along the
ventral margin of lamella dorsalis, whereas in galliforms, it is absent
or represented only by a thickened crista lateralis on the lateral
margin of pars choanalis palatini (Zusi and Livezey 2000, p. 158,
fig. 7). This variation has been utilised previously to support close
association of fossil taxa to either lineage (Sylviornis neocaledoniae,
see Mourer-Chauviré and Balouet 2005). The relatively mediolat­
erally widened structure of the ventrolateral palatines in G. newtoni
is considered more anseriform-like although pars lateralis palatini
appears to be proportionally greater in its ventrolateral extension
(Figure 7A; e.g. compared with species of Anser), which may be
functional (see Discussion) and/or related to an enlarged surface
area associated with the origin of musculus pterygoideus (e.g.
Lakjer 1926; Goodman and Fisher 1962; Zusi and Livezey 2006;
Holliday and Witmer 2007). Unlike those of G. newtoni, the pala­
tines of most extant taxa do not project far below the plane of the
crista tomialis, if at all; only anhimids appear to have a palatine that
extends noticeably ventrad of the crista tomialis (Figure 3D).
Additionally, the angulus caudolateralis on pars lateralis of the
palatine of anseriforms is also absent in galliforms (Zusi and
Livezey 2006, p. 147, fig. 7). The palatines in G. newtoni specimens
SAMA P59521, P59516, and P59517 are eroded at their ventrocau­
dal-most extremities, although appear to terminate in angulus
HISTORICAL BIOLOGY 1119
caudolateralis palatini which clearly did not project much more
caudally than what is preserved, based on the positions of other
bones and articulatory surfaces (e.g. processus basipterygoideus, see
Figure 6). The processus pterygoideus palatini, and the region of
articulation with the pterygoids (facies articularis pterygopalatina),
has been damaged or lost in all specimens and cannot be commen­
ted upon. The relatively short rostrocaudal length of the palatines in
dromornithids, is entirely expected given the very short crania in
these birds. That observed in specimens of G. newtoni, is more like
grazing anseriforms than the filter feeding taxa (Marugán-Lobón
and Buscalioni 2006; Pecsics et al. 2017). Regardless of the simila­
rities in the dromornithid palatal structure to that of anseriforms,
the general shape of the palatines appears to be unique among
galloanserans, even when compared to gastornithids and sylvior­
nithids (for comparison see Mourer-Chauviré and Balouet 2005:
fig. 2).
The rostrocaudally short ossa vomeris are fused along their
entire length, forming a single rod-like central element, here
referred to as the vomer, that defines the medial margins of the
bilaterally paired choana nasalis ossea. Caudally, the vomer is fused
laterally to pars choanalis palatini, specifically processus choanalis
palatini. A sutura vomeropalatina is not visible between the area
corresponding to the vomer and the palatines, suggestive of com­
plete synostosis between these two elements. This is reminiscent of
an anseriform state, where the suture is indiscernible in adults. By
contrast, no fusion occurs between these elements during matura­
tion in galliforms and the vomer is comparatively easily disarticu­
lated (Baumel and Witmer 1993: annot. 78; Zusi and Livezey 2006,
p. 162). The margin of choana nasalis ossea is continued caudolat­
erally, and then rostrally by the margo choanalis palatini of the
lamella dorsalis, pars choanalis of each palatinum, and defines the
rostral limits of fossa choanalis palatini on each palatine (see Zusi
and Livezey 2006). The vomer is connected to the processus max­
illopalatinus of the maxillare dorsorostrally, where a narrow ros­
trally directed triangle identifies the sutura vomeromaxillaris. This
marking is perhaps contributed to by the linea that demarks the
synostosis of the left and right maxillopalatine processes, typical of
anseriform birds (see above; Baumel and Witmer 1993: annot. 58).
Unlike some galliforms, anhimids and anseranatids, there is no
contact between processus choanalis palatini or the vomer with
rostrum parasphenoidale (Zusi and Livezey 2000, p. 161–162); the
terminus of the latter is far caudad in dromornithids. As in adult
anatids (Zusi and Livezey 2000, p. 162), there is complete separation
between palatum osseum and rostrum parasphenoidale in
G. newtoni. The vomer of gastornithids (see Matthew and
Granger 1917) and Sylviornis neocaledoniae (see Mourer-Chauviré
and Balouet 2005) is small and not a major contributor to the
palatal roof (Baumel and Witmer 1993: annot. 78).
Mandible
The partial mandible originally described by Stirling (1913: pl.
XXXVII, fig. 1–3; SAMA P.10788, Figure 10), differs a little from
the mandible of the near-complete skull specimen, SAMA P59516.
The former is longer from the rostral apex to the caudal-most point
on the fossa articularis quadratica, and the symphysial region is
proportionally shorter, 7% of length compared to 16%. This may be
indicative of intraspecific variation and potentially sexual dimorph­
ism. The mandible fragments of SAMA P59517 (Appendix One:
Figure 4) represent the partes intermedia et caudalis, and have
suffered much damage, rendering most measurements uninforma­
tive, although the pars caudalis and the fossa articularis quadratica
are relatively intact. There are two additional isolated specimens
that represent the rostral part of the mandibles, NMV P256893 and
1120 P. L. MCINERNEY ET AL.
SAMA P59520, which preserve in good condition the left and right
pars symphysialis, respectively.
The specimens are typically larger than that of the incomplete
mandible attributed to Ilbandornis sp (NTM P2774–2; Worthy et al.
2016a), and dorsoventrally taller and more robust than that attrib­
uted to Barawertornis tedfordi (QM F57895; Worthy et al. 2016a);
they are proportionally rostrocaudally longer with respect to max­
imum dorsoventral height than mandibles attributed to all species
of Dromornis which are comparatively far taller. Mandibles of
species of Dromornis are considered more massive and deeper
than those of species of Gastornis (see Angst and Buffetaut 2013),
although the opposite is true for all other dromornithid genera.
Mandibular rami of anseriforms are generally dorsoventrally taller
than those of galliform birds (Mayr et al. 2023), however, this
character appears to be homoplasious when considering giant gal­
loanserans (e.g. Sylviornis neocaledoniae).
Mandibles of G. newtoni are overall robust, with complete fusion
of elements eliminating indications of the respective boundaries
and areas of interactions between elements, even along the man­
dibular symphysis. The rostrocaudal length of the symphysis, 44.2
mm in specimen SAMA P59516, varies slightly between dromor­
nithid species, although, all are far shorter than the extreme length
of the gastornithids, that have a unique rostrocaudal elongation of
the symphysial region accounting for 44% to 48.1% of total mand­
ible length (Matthew and Granger 1917; Witmer and Rose 1991;
Angst and Buffetaut 2013). A similarly great length was also noted
for the Sylviornithidae (Worthy 2000, p. 357; Mourer-Chauviré and
Balouet 2005: fig. 8; Worthy et al. 2016b). The apex rostri of the
rostrum mandibulae in G. newtoni is curved with a notably blunt tip
which approaches the shape of some anseriforms, e.g. Anseranas
semipalmata, more so than in other dromornithids, although it
lacks the distinct dorsoventral flattening of this region, notably
extreme in Cygnus atratus and Melanitta perspicillata. The density
of foveae corpusculorum nervosorum both dorsally and ventrally is
low, relatively comparable to Anhima cornuta and galliforms.
The rostral mandible of G. newtoni, as evidenced by SAMA
P10788, NMV P256893, and SAMA P59520, has two symmetrical
depressions or broad sulci on the caudoventral part of the dorsal
symphysis that extend caudally from this region along the medial
side of their respective mandibular ramus (Figure 10 D). They are
divided at the midline by a low, wide, rugose elevation. Dromornis
planei (specimen QVM:2000:GFV:440 and NTM P9464–112) has
similar sulci. This morphology is likely associated with the gular
and extrinsic lingual apparatus musculature, and adjoining fasciae
and glands that contribute to the sublingual floor of the mouth
cavity (see Homberger and Meyers 1989); the rostrolateral ridge
that dorsally bounds each depression across their rostrocaudal
length (dotted line, Figure 10 D) may indicate the origin of
the m. mylohyoideus (m. mylohyoideus anterior, sensu Burggraaf
1954b; m. intermandibularis, sensu Fujioka 1963; Homberger and
Meyers 1989). The m. genioglossus originates in the symphysial
area in several avian taxa, as has been inferred for gastornithids
(Homberger 1986, and references therein; Andors 1988; Angst and
Buffetaut 2013: fig. 2; Jones et al. 2019), however, this muscle is
absent in adult galloanserans, including Gallus, Aepypodius (see
Fujioka 1963; Homberger and Meyers 1989; Weber 1996) and
Anas (see Zweers et al. 1977). Instead, in addition
to m. mylohyoideus, these sulci in G. newtoni/dromornithids are
likely associated with the fascia intermandibularis and fascia sub­
lingualis, and also potentially m. branchiomandibularis rostralis
which arises from the rostral region of the medial mandible
(Gadow and Selenka 1891; Goodman and Fisher 1962; Fujioka
1963; Zweers 1974; Homberger and Meyers 1989: p. 235–239,
table 21, regarding clarification, synonymy and homology; Weber
1996: fig. 2). The sulci in G. newtoni and D. planei are, as in Anseres,
rostrocaudally elongate and relatively flat compared to galliforms
and anhimids. They are especially similar in form and position to
that of Cereopsis novaehollandiae and to a lesser extent, Anseranas
semipalmata, as other anseriforms generally possess less well-
defined and mediolaterally narrower depressions and retain
a dorsally overhanging medial plate (sensu Burggraaf 1954b), the
latter of which is comparatively absent in dromornithids. In galli­
forms (Burggraaf 1954b: fig. 31) and anhimids, these sulci are
restricted to the caudal margin of the symphysial region and meet
along the midline, just ventral of the medial plate.
Paired foramen neurovasculare (Stirling 1913: p. 114; Baumel
and Witmer 1993; sensu Livezey 1997: char. 16; lingual median
symphysial foramina, sensu Stidham 1998) are located on the dorsal
surface of the rostral mandible, either side of the symphysis, and sit
at the rostral-most point of an associated shallow, predominately
rostrocaudally aligned sulcus (potentially homologous with the
canalis primordialis, see Burggraaf 1954b; and the longitudinal
vascular groove, see Currie et al. 1993), as best seen in G. newtoni
specimens NMV P256893 and SAMA P10788. Like in galliforms
and anhimids, the foramina are each positioned within an elongate
depression which extends caudally, whereas in the mandibles of
D. planei and non-anhimid anseriforms, the foramen is located
relatively further rostromedially on the rostrum mandibulae than
these depressions and the associated sulci extend further caudally.
In this region of the rostral mandible, the crista tomialis of
G. newtoni is thin. Caudally a rostrocaudally long, wide, flat surface
of bone, extends medially from the near indistinct crista tomialis, as
described for Ilbandornis sp (see Worthy et al. 2016a). The medial
margin of the shelf is truncated sharply as it drops ventrally towards
the lateral margins of the aforementioned paired depressions which
house the foramen neurovasculare. The shelf decreased in medio­
lateral width caudally (see also Stirling 1913). Such a flat tomial
surface is also identified in the mandible of gastornithids, although
the crista tomialis is more distinct, sharper and more blade-like
across its entire length (Troxell 1931; Angst and Buffetaut 2013:
fig. 2). Extant galloanserans appear to lack the shelf, and instead
have a typically thin and sharp (as in many galliforms) or wide and
rounded (i.e. Anseres) crista tomialis. A similar medial crest and
associated tomial shelf is only present in the anhimids although it is
far less developed, restricted in its rostral extension, and may not be
homologous in origin (see Previatto 2012: fig. 21, 23).
The mandibular rami in G. newtoni and other dromornithids
diverge caudally from the symphysis only slightly (fig. 1 g;
Murray and Megirian 1998: figs. 11, 18, 20), at a much smaller
angle than in mandibulae attributed to most galliform and
gastornithid taxa (although see Stirling 1913, p. 115). The dor­
sally elevated angulus dorsalis mandibulae (ADM, sensu Livezey
and Zusi 2006: characters 0662, 0679, 0680) is just rostral to the
pars caudalis of the mandible and is coincident with the same
ridge that hosts the processus coronoideus as in all dromor­
nithids – the former marking the caudal-most limit of the
rhamphothecal sheath (Baumel and Witmer 1993: annot. 43),
and broadly correspondent with aponeurosis superficialis (sensu
Dzerzhinsky and Potapova 1974; Dzerzhinsky 1982; see
Figure 10; see also Weber 1996). In all dromornithids this
appears dorsally raised, although it is confluent with, rather
than distinctly dorsally produced, the dorsal edge of the ros­
trum mandibulae. Among extant galloanserans, the ADM is
only clearly dorsally prominent with respect to the level of the
more rostral tomium in Anseres, in most galliforms it is subtle.
As in crown group Anseres, the dorsal elevation of angulus
dorsalis mandibulae can be clearly distinguished from the
more rostral crista tomialis in late Paleocene Anachronornis
 HISTORICAL BIOLOGY 1121

Figure 10. Genyornis newtoni mandible morphology: A. Caudal view of skull part NMV P256893, partially preserving ramus mandibulae, pars caudalis ventrally, which
retains the medial process of the mandible; B. Image and annotated outline of SAMA P59516 in right lateral view, digitally removed from the image of the entire skull; C.
Ventral view of the symphysial region of mandible specimen SAMA P59516; D. Caudal view of symphysial part of mandible specimen NMV P256893; E. Dorsal view of
symphysial part of mandible specimen NMV P256893; F. Proposed arrangement of insertions of the adductor muscle complex on the lateral side of the mandible for
dromornithids (drawn from specimen NTM P2774–2 of Ilbandornis sp. due to the limited distortion and conservative nature of dromornithid skull morphology) compared
with re-drawn examples of extant galloanserans, Anseres – Anas platyrhynchos (Davids 1952a: fig. 3b), Anhimidae – Anhima cornuta (NMV B12574), Megapodiidae –
Aepypodius arfakianus (Weber 1996: fig. 2a) and Phasianidae – Gallus gallus (Fujioka 1963: fig. 7). Annotations: c.lat., cotyla lateralis.; cr.par., crista paracoronoidea rostralis
and crista paracoronoidea caudalis.; ADM, angulus dorsalis mandibulae; AVM, angulus ventralis mandibulae; exc., excavation associated with the region of recessus conicalis
in anatids; f.neur., foramen neurovascularis.; m.AMEPc, m. AME profundus, pars coronoideus; m.AMEs, m. AME superficialis; m.AMEPs, m. AME profundus, pars
superficialis; m.AMEPz, m. AME profundus, pars zygomaticus; m.AMP, m. adductor mandibulae posterior; m.ps., abraded prominence and approximate area for insertion
of m. pseudotemporalis profundus; pr.cor., processus coronoideus (approximate position indicated, not visible as it is obscured by associated cranium); pr.lat.m., processus
lateralis mandibulae; pr.med.m., processus medialis mandibulae; pr.ret., processus retroarticularis; sym., pars symphysialis, rostrum mandibulae; tom., crista tomialis; Scale
bars: A., C.-E. 10 mm, B. 20 mm, F. not to scale. Dark grey shading indicates regions where damage precludes morphological assessment, and light grey indicates fenestra.

anhimops, whereas in early Eocene Danielsavis nazensis, there is
comparatively no clear transitional change in dorsoventral
height (as in galliforms, see Houde et al. 2023: figs. 1, 7; Mayr
et al. 2023). The adjacency of the processus coronoideus and
ADM in Anseres confines the insertions of the m. AME pro­
fundus partes zygomaticus et superficialis and m. AME super­
ficialis to within a relatively caudal area (e.g. Davids 1952a,
1952b; Goodman and Fisher 1962; Zusi and Livezey 2000).
Their similarly close proximity in dromornithids, likely results
in the same musculature arrangement, as also interpreted for
D. planei (see Murray and Vickers-Rich 2004: fig. 189). This
contrasts with a typical galliform state, whereby the ADM is
relatively far rostral, and the attachment surfaces for these
external adductor muscles extend more rostrally on each man­
dibular ramus (relative to its total length, e.g. Kirikov 1944;
Dzerzhinsky and Belokurova 1972; Dzerzhinsky 1982; Weber
1996; Zusi and Livezey 2000). Anhima cornuta appears inter­
mediate between Anseres and galliforms with respect to proxi­
mity between processus coronoideus and ADM, and interrelated
myology (see Figure 10; Dzerzhinsky 1982).
The gentle rostral transition from the ADM to the tomial margin
of G. newtoni differs markedly from Anseres, in which the steep
rostral incline of the ADM is comparatively near perpendicular,
such as in Anser caerulescens, and even the fossil anseriform,
Anachronornis anhimops which also has a relatively steep incline.
The rounded rostral profile of the ADM in G. newtoni is largely
reminiscent of that in D. murrayi (see Worthy et al. 2016a), and is
less consistent with the more sloped form of Ilbandornis sp. NTM
P2774–2. Both rami of Barawertornis tedfordi (QM F57895) are
damaged in this area, although the left ramus mandibulae is better
1122 P. L. MCINERNEY ET AL.
preserved in this region and does not appear to differ substantially
from other dromornithids, although more detailed comparisons are
not possible. The form of the ADM in transition to the tomial
margin in G. newtoni is less hyperdeveloped than in species of
Dromornis (see Murray and Megirian 1998).
A lack of symmetry in the rostrocaudal curvature of the ridge
that hosts the ADM is evidenced in the G. newtoni mandible
described by Stirling (1913), whereby the more dorsocaudal angle
on the caudal part of this ridge represents the processus coronoi­
deus. This region is obscured (SAMA P59516) or damaged (SAMA
P59517) in other specimens although the visible shape is consistent
with this morphology. This is also consistent with other dromor­
nithids of which G. newtoni is largely similar to that of D. murrayi
(see Worthy et al. 2016a: fig. 3). In Ilbandornis sp., Barawertornis
tedfordi (see Worthy et al. 2016a: fig. 3), and especially D. stirtoni
and D. planei (larger in the latter taxon proportionally, see Murray
and Vickers-Rich 2004: fig. 109), the processus coronoideus pro­
jects more sharply and prominently dorsocaudally. This may evi­
dence a less well-developed aponeurosis coronoidea (sensu
Iordansky 1970; aponeurosis rostralis, sensu Dzerzhinsky 1982;
also following Weber 1996) in association with m. AME profundus
coronoideus (Weber 1996; Zusi and Livezey 2000; Holliday and
Witmer 2007; Matsuoka et al. 2008) in both G. newtoni and
D. murrayi, compared to other dromornithids. The processus cor­
onoideus in dromornithids (except for species of Dromornis) is
comparable to the small caudally protruding angle in Anhima
cornuta or Oxyura australis, lacking the more extensive lateral
flare of most Anseres (e.g. Cereopsis novaehollandiae), and differs
from the prominent tubercle-like protrusion of most galliforms.
In other dromornithids, especially species of Dromornis, the ridge,
just caudal of the processus coronoideus, shows greater attenuation
in height towards the fossa articularis quadratica than in G. newtoni.
Such variation may be associated with the attachments of corre­
sponding musculature; m. adductor mandibulae posterior inserts
on the lateral surface between the processus coronoideus and the
fossa articularis quadratica in neognaths, a pattern galloanserans –
and most likely dromornithids – also adhere to (e.g. Dzerzhinsky
1982; Zusi and Livezey 2000; Holliday and Witmer 2007). Variable
development of areas related to insertions of mm. pseudotemporalis
superficialis et profundus and m. pterygoideus on the medial surface,
and the origin of m. branchiomandibularis caudalis (m. geniohyoi­
deus, lateral slip, sensu Goodman and Fisher 1962, only reported for
Mergini among anatids) on the ventral edge and ventral half of the
lateral caudal ramus (e.g. Goodman and Fisher 1962; Dzerzhinsky
and Belokurova 1972; Homberger and Meyers 1989; Weber 1996),
also likely contribute to this morphological disparity. Species of
Dromornis additionally have a deeply depressed medial mandibular
surface correlated with the large attachment site of
the m. pterygoideus (Murray and Vickers-Rich 2004, p. 241, fig.
189), in association with their relatively dorsoventrally deep caudal
mandible. A low, rounded, and subtle bulge on the dorsal part of the
ramus mandibulae between the processus coronoideus and fossa
articularis quadratica – likely homologous with the area for insertion
of m. pseudotemporalis profundus in other galloanserans (Davids
1952a; Fujioka 1963; Zweers 1974; Weber 1996; Matsuoka et al.
2008) – can be observed in Ilbandornis sp., Dromornis murrayi, and
is especially pronounced for B. tedfordi, however, its form is not
accurately assessable in any specimens of G. newtoni. While similarly
located in some birds, this bulge is not to be considered equivalent to
the tuberculum praearticulare (sensu Weber 1996), for aponeurosis
praearticularis caudalis (sensu Weber 1996) of m. pterygoideus dor­
salis, which is instead positioned just rostral of the fossa articularis
quadratica on the medial mandible in dromornithids, as is especially
evident on the right caudal ramus of G. newtoni (SAMA P59517, see
Appendix One: Figure A4) and Ilbandornis sp. NTM P2774–2 (see
Weber 1996: fig. 26; Worthy et al. 2016a: fig. 3A).
The damaged lateral surfaces of all mandibles known for
G. newtoni, restrict a confident examination of some structures
pertinent to galloanseran systematics, such as the exact nature of
the crista(e) paracoronoidea (lateral coronoid process, sensu
Goodman and Fisher 1962; see also Weber 1996; terminology of
Weber and Hesse 1995, p. 295; Matsuoka et al. 2008; Mourer-
Chauviré and Bourdon 2020; SI 2), which is a laterally shifted homo­
logue of the tuberculum paracoronoideum of many birds (see Weber
1996, p. 27). In galliforms and anhimids, the crista paracoronoidea is
separated into two crests, distinguished as the crista paracoronoidea
rostralis and caudalis, which are perpendicular to near parallel to one
another and closely approximated, or touching, at their dorsocaudal-
most points (sensu Weber and Hesse 1995; Weber 1996). In the
Megapodiidae (including Alectura lathami) and Acryllium vulturi­
num (Numididae), these two cristae are more widely separated, and
the rostral part is well-defined and rostrally extended into a process
or flange (Weber and Hesse 1995, p. 295; Weber 1996, p. 27, 28).
A similarly located, but undivided crista paracoronoidea forms
a prominent processus paracoronoideus (sometimes mistakenly
referred to as the processus coronoideus, see Weber and Hesse
1995; Weber 1996) in non-anhimid extant anseriforms
(Figure 10F), and is analogous to the well-developed crista paracor­
onoidea rostralis in Megapodiidae (Weber 1996).
As far as can be discerned for G. newtoni, a distinct rostrocaud­
ally elongated crista paracoronoidea rostralis is present, caudoven­
tral of the processus coronoideus, corresponding to the aponeurosis
paracoronoidea externa (sensu Weber 1996) and the associated
insertion of m. AME superficialis (see Figure 10F; also
Dzerzhinsky and Potapova 1974; Dzerzhinsky 1982; Zusi and
Livezey 2000). The crista paracoronoidea caudalis is less well-
marked, yet is likely represented by a relatively swollen, dorsocaud­
ally-ventrorostrally elongated ridge just caudoventral of its rostral
counterpart. This is likely associated with the aponeurosis paracor­
onoidea interna (aponeurosis caudalis interna, sensu Dzerzhinsky
and Potapova 1974; Dzerzhinsky 1982; sensu Weber 1996), which
receives some fibres from, and caudally bounds, the insertion
of m. AME superficialis (Weber 1996, p. 46, 53). The form and
position of this structure may also be contributed to by the devel­
opment of the caudally adjacent insertion of m. adductor mandi­
bulae posterior (Weber and Hesse 1995, p. 58; Weber 1996;
‘adductor fossa’ Zusi and Livezey 2000; Murray and Vickers-Rich
2004: fig. 56; see above). A similar morphology is observed in
Barawertornis tedfordi, with regards to both the rostral and caudal
cristae. Comparatively, Ilbandornis sp. (NTM P2774–2) has dis­
tinct, lowly raised, narrowly separated, and parallel rostral and
caudal paracoronoid crests (Murray and Megirian 1998: fig. 12;
Worthy et al. 2016a). This also appears to be the case in well-
preserved D. planei and D. stirtoni specimens (e.g. NTM P9464–
112, P98107, see Murray and Megirian 1998; Murray and Vickers-
Rich 2004; Worthy et al. 2016a). The nearly parallel orientation of
the cristae in dromornithids appears to resemble that of the
Anhimidae and Phasianidae, more than that of the Megapodiidae.
In contrast, the late Paleocene ‘screamer-like’ Anachronornis anhi­
mops, Wyoming, preserves a laterally projected processus paracoro­
noideus, as is typical of Anseres. However, this is comparatively less
laterally developed than crown group representatives (see Houde et al.
2023). The early Eocene Danielsavis nazensis has a well-developed,
flange-like crista paracoronoidea rostralis that compares well with the
aforementioned megapodiid condition (Houde et al. 2023; Mayr et al.
2023). In gastornithids, the crista paracoronoidea rostralis is potentially
homologous with a long, predominately dorsoventrally orientated
ridge which extends caudodorsally to meet the processus coronoideus.

This ridge is further rostral of the processus coronoideus compared to
the relatively caudal placement and rostrocaudal extension of this crest
in dromornithids; a second faint ridge is evident in some gastornithid
specimens, notably Ga. parisiensis MHNT.PAL.2012.1.1, which is
caudally displaced from, but in near parallel dorsoventral orientation
with the aforementioned crest, and may be homologous to the crista
paracoronoidea caudalis in this taxon (see Matthew and Granger 1917,
p. 312, pl. XXI; Angst and Buffetaut 2013: fig. 3C; Angst et al. 2014: fig.
4; Mourer-Chauviré and Bourdon 2020: figs. 1, 2).
The rounded angulus ventralis mandibulae (AVM; sensu Livezey
1998; Livezey and Zusi 2006) of G. newtoni is positioned at a point
noticeably caudal to the processus coracoideus, as in B. tedfordi and
D. stirtoni. In contrast, in Ilbandornis sp. NTM P2774–2, and speci­
mens of D. murrayi, and especially D. planei, the AVM is relatively
more rostrad. In D. planei and D. stirtoni (see Murray and Megirian
1998: fig. 12; Worthy et al. 2016a: fig. 3), the AVM is more pro­
nounced ventrally, and more robust than in other species. This would
support greater attachment regions for relevant musculature on
medial and lateral sides (e.g. m. AME externus partes superficialis
et profundus, m. adductor mandibulae posterior and m. pterygoideus
ventralis, and m. branchiomandibularis caudalis, see Figure 10F, also
Goodman and Fisher 1962; Vanden Berge and Zweers 1993; Murray
and Vickers-Rich 2004; Holliday and Witmer 2007; Matsuoka et al.
2008; Holliday 2009). In contrast to mandibles attributed to
D. stirtoni, D. planei and Ilbandornis sp., there is less of
a dorsoventral shift from the AVM to the ventral edge of the rostral-
most processus retroarticularis in both G. newtoni and B. tedfordi,
although not as level as in D. murrayi. Although less accentuated than
in the dromornithids, the AVM is only distinct as a rounded structure
in goose-like anserines including Cereopsis novaehollandiae, Cygnus
atratus, Branta canadensis, and Anser caerulescens. The AVM is
indistinct in species of Gastornis.
The ventrocaudal edge of the AVM of G. newtoni continues as
a low ridge dorsocaudally onto the lateral mandibular face and
forms the caudal and ventral margins of a shallow depression just
rostroventral to the processus lateralis mandibulae (lateral mandib­
ular process), which likely corresponds to the bounds of the inser­
tion of m. adductor mandibulae posterior (Figure 10 F; Dzerzhinsky
1982; Weber 1996; Zusi and Livezey 2000; Holliday and Witmer
2007). This fossa is deeper and more distinct in mandibles attrib­
uted to species of Dromornis (e.g. see ‘fossa for external adductors’,
Murray and Megirian 1998: fig. 16; ‘adductor fossa’, Murray and
Vickers-Rich 2004: fig. 56). Murray and Vickers-Rich (2004: figs.
54, 56, 189) labelled an area near the ventral margin of the mandible
of D. stirtoni, close to this fossa, as a ‘groove for m. mylohyoideus’
for ‘m. mylohyoideus posterior’ (similarly labelled by Davids 1952a,
1952c). In this sense, this more appropriately relates to the passage
for m. serpihyoideus and m. stylohyoideus, passing across the
ventral mandible towards the lingual apparatus, from the caudal
processus retroarticularis, and is involved in retraction of the ton­
gue and the raising of the tongue and trachea (see Davids 1952a,
1952c; Goodman and Fisher 1962; Fujioka 1963; Zweers 1974; sensu
Homberger and Meyers 1989; Weber 1996; Matsuoka et al. 2008).
This form of the ventral mandible may also be influenced by the
origin of m. branchiomandibularis caudalis on this area of the
caudal ramus (see above; Fujioka 1963; Homberger and Meyers
1989; Weber 1996; SI 2). The path of the former muscles on the
ventral edge of the caudolateral mandible, especially, may contri­
bute to the development of the low ridge that continues from the
ventral margin onto the lateral face in G. newtoni and D. stirtoni.
As in B. tedfordi, D. stirtoni, and gastornithids (Angst and
Buffetaut 2013; Worthy et al. 2016a), the mandible of G. newtoni
is imperforate, lacking both fenestra caudalis mandibulae and
fenestra rostralis mandibulae. Contrastingly, Ilbandornis sp. NTM
HISTORICAL BIOLOGY 1123
P2774–2, D. planei, and D. murrayi all have a caudal fenestra (see
Worthy et al. 2016a: fig. 3). Although considerable variation in the
size and shape of the fenestra caudalis mandibulae is present within
galloanserans, the presence and absence of the fenestra among some
dromornithids is seemingly unusual. Only in anhimids, is this
fenestra exclusively absent. Intercepting the caudal fenestra on the
medial side of the mandible of Ilbandornis, an obvious ridge
stretches rostrocaudally along pars caudalis towards the processus
medialis mandibulae, representing the ventral edge of the fossa
aditus canalis neurovascularis (see Murray and Megirian 1998: fig.
12; Worthy et al. 2016a: fig. 3). This is well-developed in mandibles
attributed to Dromornis planei (e.g. NTM P9464–112), also present
in B. tedfordi and to a far lesser extent, in G. newtoni.
A medial mandibular process of G. newtoni is well-preserved only
in the fragmentary caudal ramus of SAMA P59517 and NMV
P256893.4 and is partially visible on the skull of SAMA P59516. The
medial process is larger than its lateral counterpart although propor­
tionally to a far lesser extent than that of all extant galloanserans. In
G. newtoni, like that of B. tedfordi, the process is short, truncated
medially, and invaginated caudally as it curves to form a deep and
mediolaterally wide depression on the caudal surface. The medioven­
tral edge of this depression likely accommodated the attachment of
ligamentum occipitomandibulare (Davids 1952a, 1952c; Goodman
and Fisher 1962, p. 133; Bock 1964; Zweers 1974: fig. 14;
Dzerzhinsky 1982: fig. 3; ligamentum neurocranio-mandibulare,
sensu Baumel and Raikow 1993). Although there are no deep,
depressed recessus conicalis as in anatids, nor an excavation as deep
as that in Anseranas semipalmata, Anachronornis anhimops, Conflicto
antarcticus or Presbyornis pervetus, the depression (Figure 10A: ‘exc’.)
on the caudal surface of the process appears like the shallow depression
in the region on mandibles of Anhima cornuta. This is perhaps also
comparable to the mandible of galliform-like stem anseriform, or
anseriform-like stem galliform, Danielsavis nazensis, which also lacks
a distinct fossa caudalis and recessus conicalis (see Houde et al. 2023:
fig. 5C; Mayr et al. 2023). Comparatively, this side of the process is
dorsoventrally thinner and caudally rounded in most galliforms and
has a horizontal mediolaterally oriented crest in some megapodes (e.g.
Alectura lathami).
The rostromedial surface of the processus medialis mandibulae is
gently sloped dorsorostrally-caudoventrally in G. newtoni, and would
have primarily related to insertions of parts of m. pterygoideus ven­
tralis (see following for references). Rostrally, a large, flattened, or
shallow attachment surface likely primarily supported the insertion
of m. pterygoideus dorsalis (Davids 1952a; Fuchs 1954a; Fujioka
1963; Zweers 1974; Dzerzhinsky 1982; Weber 1996; Holliday and
Witmer 2007; Matsuoka et al. 2008; Bianki et al. 2013; see Appendix
Three). The medial process is also well-preserved in Barawertornis
tedfordi QM F57985 (Worthy et al. 2016a: fig. 3) and on the left
ramus of Dromornis planei NTM P9464–112 (Murray and Megirian
1998: fig. 11–12). In these taxa, the process largely conforms with that
of G. newtoni with regards to shape. Immediately dorsal of
the m. pterygoideus dorsalis insertion area, an apparent rostral con­
tinuation of the processus medialis mandibulae as an obvious ridge or
tubercle corresponds to tuberculum praearticulare for the attachment
of aponeurosis praearticularis caudalis (see above; Weber 1996). The
pattern of pterygoideus muscle attachments also appears similar
across all dromornithids, although the surface for insertion
of m. pterygoideus dorsalis, rostral of processus medialis mandibulae,
is slightly more excavated in Ilbandornis sp. NTM P2774–2, and well-
impressed in D. planei and D. stirtoni (see Murray and Vickers-Rich
2004, p. 241). Medially adjacent to cotyla medialis mandibulae,
a foramen pneumaticum articulare is obvious dorsally on the medial
process of D. planei (NTM P9464–112), Barawertornis tedfordi and
Ilbandornis sp., although its presence is unknown for G. newtoni.
1124 P. L. MCINERNEY ET AL.
Notably, the articulatory region differs considerably from those on
gastornithid mandibles, the latter of which instead have greater
morphological similarity to with S. neocaledoniae: the crista inter­
cotylaris is sharper and more distinct, processus medialis mandibulae
is proportionally larger and more medially elongate, and the proces­
sus lateralis mandibularis is also proportionally larger, as
a prominent, convex tubercle (see Mourer-Chauviré and Balouet
2005: fig. 8; Angst and Buffetaut 2013: fig. 2).
The noticeable lateral overhang of the cotyla lateralis of processus
lateralis mandibulae in G. newtoni appears more like that of some
galliforms and anhimids than the relative lack of lateral projection in
species of Dromornis, B. tedfordi, and Ilbandornis sp (Worthy et al.
2016a: fig. 3). In most non-anhimid anseriforms, the edge of the
cotyla is more prominent, and in some derived forms, develops
a lateral or rostral angularity (e.g. Melanitta perspicillata and Anas
superciliosa). A caudally cambered ridge caudally borders the sloped
rostral surface of the process; the morphology of this surface likely
developed in relation to the insertion of the m. adductor mandibulae
posterior (see above). In some galloanseran taxa where this muscle
can be divided into distinct bellies, the lateral part consistently inserts
on this area of the caudodorsal mandible (Holliday and Witmer
2007), and is associated with the attachment of a strong aponeurosis
(e.g. aponévrose 15, Davids 1952a; m. quadratomandibularis ‘10A’
Fujioka 1963; ‘a. 11’ Dzerzhinsky and Belokurova 1972). A distinct
lateral part of this muscle, the corresponding attachment to this area
is absent in anhimids (Dzerzhinsky 1982; Zusi and Livezey 2000), but
present in anseranatids and is typical of anatids (e.g. Goodman and
Fisher 1962; Dzerzhinsky and Grintsevichene 2002; Bianki et al.
2013), consequently preventing a confident assessment of its presence
in G. newtoni and other dromornithids (contra Murray and Vickers-
Rich 2004: e.g. fig. 189). The potential lack of the ligamentum post­
orbitale in G. newtoni and the rest of the dromornithids, may be
a factor in the variable development of the lateral mandibular process.
Additionally, a small tubercle is variably developed just rostral of the
lateral mandibular process where the ligamentum postorbitale
attaches (Baumel and Raikow 1993: annot. 42) in galliforms and
anseriforms but not in dromornithids. The ligamentum lacrimoman­
dibulare also attaches in this region in most anseriforms and is
associated with filter feeding in anatids due to its absence in non-
filter feeders, e.g. Anseranas semipalmata (see Davids 1952c;
Goodman and Fisher 1962, p. 131, 132; Baumel and Raikow 1993:
annot. 41; Bout and Zweers 2001; Zelenkov and Stidham 2018). The
lack of evidence supporting such specialised feeding in dromor­
nithids suggests this particular ligament was absent in this family.
A feature common to all Galloanserae and present in
dromornithids, is the bicondylar articulation with the quad­
rate (Dzerzhinsky 1995; Ericson 1996; Mayr et al. 2018; Field
et al. 2020). This has assisted in supporting the galloanseran
affinities of other fossil taxa including Conflicto antarcticus
(see Tambussi et al. 2019). Of the articulatory region of
G. newtoni, the caudal part of the right mandibular ramus is
visible in SAMA P59517 and allows a dorsal view of the fossa
articularis quadratica, while the cotyla lateralis is visible on
the left side of SAMA P59516, and cotyla medialis is retained
in specimen MNV P256893. These specimens show a similar
morphology regarding the separation of the cotylae in other
dromornithids and galloanserans (see Worthy et al. 2016a).
Both cotylae are oval, angled obliquely to the main axis of the
mandibular ramus, separated by a low crista intercotylaris
although seeming to overlap for much of their long axis,
and the cotyla medialis is deeper than cotyla lateralis (see
also Worthy et al. 2016a). The cotylae in G. newtoni,
Ilbandornis sp., and B. tedfordi are more rostrocaudally elon­
gate compared to those in species of Dromornis (see Worthy
et al. 2016a), although in all, the proportional sizes of the two
cotylae to the surrounding pars caudalis mandibulae are more
like those of Anhima cornuta and megapodes than to other
galloanserans (in Anseres the medial cotyla is greatly
expanded rostrocaudally and near equal in mediolateral
width with the lateral cotyla, and in non-megapodiid galli­
forms, the lateral cotyla is larger mediolaterally).
The presence of a caudally prominent and laterally compressed
processus retroarticularis is characteristic of Galloanserae (Baumel
and Witmer 1993; Ericson 1996; Murray and Vickers-Rich 2004;
Mayr 2017; Mayr et al. 2018; Tambussi et al. 2019; Field et al.
2020). While all caudal mandibles attributed to G. newtoni are
incomplete, the processus retroarticularis is relatively visibly rostro­
caudally short, lateromedially compressed, and robust with a caudal
extension almost equal to its dorsoventral depth, similar to the
morphology of other dromornithids (Worthy et al. 2016a) and
Gastornis giganteus (Matthew and Granger 1917: pl. XXI; AMNH
6169; Angst and Buffetaut 2013: fig. 4). The process is directed
caudally and has a slight dorsal upturn at its end creating a clear
angle on the dorsal margin, immediately caudal to the fossa articu­
laris quadratica. All dromornithid mandibles are somewhat damaged
in this region, and so the extent of the dorsal upturn of the retro­
articular process, and its caudal extremity, cannot be precisely ascer­
tained. However, remnants of the rostral origin of the retroarticular
process suggest some variation is present among dromornithids:
D. planei has a steeply curved dorsal margin; D. stirtoni, a steeply
curved ventral margin, yet a shallower dorsal curve; and D. murrayi
and B. tedfordi are considered to lack dorsal deflection, as their
retroarticular processes only project caudally. No mandible of species
of Dromornis preserves a complete processus retroarticularis, yet
Murray and Vickers-Rich (2004, p. 81 and 105, fig. 57) speculated
that D. stirtoni had a high, angular, caudodorsal tip. With new
evidence from G. newtoni, if we are to assume D. stirtoni had similar
morphology, such recurvature and caudodorsal apex would not be
approximated. The processes on the mandibles of G. newtoni are
dorsocaudally rounded and only have a small dorsal projection
(SAMA P59516, right side). The process is caudally orientated as in
most anseriforms (e.g. Anseranas semipalmata), not laterally
deflected as in many galliforms (e.g. Pavo muticus and Gallus gallus).
Hyobranchial apparatus
Specimen SAMA P59516 retains, in association, two disarticulated,
near complete ceratobranchials (ceratobranchiale), the rostral-most
part of the basihyal (basihyale), broken just rostral of the articulatio
ceratobasihyalis, and a partial, crushed paraglossum. Part of the left
ceratobranchial is also preserved on the ventral-most caudal mand­
ible of NMV P256893. These are the first osseous elements of the
hyoid skeleton identified for any dromornithid.
The basihyal articular surface for the articulatio paraglosso-
basihyalis in dorsal and lateral view, is similar to that of Anser
caerulescens, where the basihyal thins bilaterally and expands dor­
soventrally to form a curved saddle-type joint (see Baumel and
Raikow 1993: annot. 51). The dorsal surface appears flat and ven­
trally the articular surface extends rostrally, to a blunt, taphonomi­
cally worn tip; this shape is present in Alectura lathami although
proportionally much smaller than those of anseriforms that possess
this structure, e.g. Anseranas semipalmata and Anser caerulescens.
The body of the basihyal is rostrocaudally long and bilaterally
narrow, proportionally more so than other anatid and anseranatid
taxa (e.g. Anser caerulescens, pers obvs.), as is typical of Anhimidae
and Galliformes, and lacks the wide form and wing-like projections
of Danielsavis nazensis (see Mayr et al. 2023: fig. 3). The lateral
expansion of the basihyal expected for the articulatio ceratohyalis

(articulation with the ceratobranchials), including any indication of
each process lateralis of the basihyal (sensu Zweers 1974), cannot be
identified on the specimen. Additionally, no urohyal (urohyale) is
preserved, fused (as in Anser caerulescens) or separate (Gallus
gallus, see Homberger and Meyers 1989: fig. 1) to the basihyal.
The paraglossum (Figure 11B) is present in specimen SAMA
P59516, near the rostral end of the mandible, crushed between
the medial aspect of the right mandibular rami, and the ventral
surface of the attached rostrum. This bone is perforated by
a large, obvious central foramen (for., Figure 11B), and is overall
a mediolaterally symmetrical, triangular, arrowhead shape. The
paired cornua, or processus paraglossus caudalis (sensu
Homberger and Meyers 1989), extend caudally away from the
body of the bone but have lost much of their caudal length due
to taphonomic damage. The general morphology of the paraglos­
sum is similar to the form of most birds (see Baumel and
Witmer 1993: annot. 80) including galliforms (see McLelland
1968; Homberger and Meyers 1989). The paraglossum of non-
anhimids is comparatively rostrocaudally longer and more oval
or rectangular (e.g. Zweers et al. 1977: fig. 6). The overall shape
of this bone in Anhimidae is intermediate in form, being slightly
rostrally tapered, but also bilaterally wide.
The ceratobranchials have a shaft minimally 115.7 mm in
rostrocaudal length. They are displaced from the mandible
and rostrum and disarticulated. Additionally, the articular
ends are both damaged, preventing identification of either
end. The shaft does not gently curve in one direction as in
many galloanserans but instead curves twice to be slightly
sigmoidal, potentially a result of taphonomic distortion.
Phylogenetic analyses
Parsimony analysis
The heuristic search resulted in 2 MPTs (Most Parsimonious
Trees) with a length of 786 steps, where a total of 122,871,532
rearrangements were tested. Both MPTs (phylograms displayed
in SI 5) have a Consistency Index of 0.2799, Homoplasy Index
of 0.7201, Retention Index of 0.6939 and a Rescaled Consistency
Index of 0.1942. The MPTs vary only in the exact topological
relationship of Anas superciliosa (Anatini), Aythya australis
(Aythini) and Melanitta perspicillata (Mergini) to one another,
as evidenced by the associated formation of the polytomy in the
strict consensus tree (Figure 12A). All other relationships,
including those described below, were recovered in both MPTs.
Since modern taxa were topologically constrained based on
genetic data (see Materials and methods: Phylogenetic analyses),
only the phylogenetic relationships of fossil taxa are described
herein. A branch to the monophyletic clade containing all
crown group Anseriformes, Asteriornis maastrichtensis,
Conflicto antarcticus, Presbyornis pervetus, Nettapterornis
oxfordi (Olson 1999) and Dromornithidae, was recovered in
44.69% of bootstrap replicates and is supported by two unam­
biguous (see SI 5 for all state changes discussed here) and 18
ambiguous character state changes. This clade to the exclusion
of Asteriornis maastrichtensis was resolved in 76.57% of repli­
cates and supported by three unambiguous and five ambiguous
apomorphic state changes. A branch to the Anhimidae +
Dromornithidae clade was found consistently across MPTs
and received moderate bootstrap support (76.4%). The
Anhimidae + Dromornithidae clade is supported by 13 unam­
biguous and 10 additional ambiguous character state changes.
There are 24 character state changes that distinguish
a monophyletic dromornithid clade with respect to the
HISTORICAL BIOLOGY 1125
Anhimidae (20 unambiguous and four ambiguous), the branch
to which received high bootstrap support (96.73%). The
Genyornis newtoni + Ilbandornis species clade was supported
by just one unambiguous and three ambiguous character state
changes (bootstrap support for this branch is 45.61%), and the
Dromornis clade distinguished by four unambiguous changes
(bootstrap support is 64.59%).
A branch to the clade including all non-anhimid crown group anseri­
forms¸ Conflicto antarcticus, Presbyornis pervetus and Nettapterornis
oxfordi was recovered sister to the Anhimidae + Dromornithidae clade
in all MPTs, with high bootstrap support (98.75%), in association with 16
character state changes (11 unambiguous and five ambiguous). A sister-
taxon relationship between Conflicto antarcticus and a clade including
Nettapterornis oxfordi + Anseres was recovered in 46.41% of bootstrap
replicates, related to nine character state transitions (three unambiguous
and six ambiguous). The branch including Presbyornis pervetus and
crown group Anseres, sister to Nettapterornis oxfordi, has low-
moderate bootstrap support (42.95%), associated with two unambiguous
and three ambiguous character state changes. The branch to crown
group Anseres, sister to Presbyornis pervetus, is related to eight unambig­
uous and five ambiguous character changes and received moderate
bootstrap support (79.32%).
A branch comprising a clade of crown group Galliformes,
Gastornis giganteus, and Sylviornis neocaledoniae was recovered
as sister to the Anseriformes clade in 54.03% of bootstrap repli­
cates, supported by eight unambiguous and six ambiguous char­
acter transitions. A poorly supported (48.01%) branch that
includes a clade of Gastornis giganteus, Sylviornis neocaledoniae,
and all non-megapodiid galliforms is associated with three unam­
biguous character state changes and four ambiguous ones.
A branch to a clade including the Gastornithidae and
Sylviornithidae was resolved in 72.76% of bootstrap replicates;
13 unambiguous and two ambiguous character state changes
support the monophyly of this clade as sister to a one comprising
crown group Phasianoidea. The branch to the clade containing
crown group Phasianoidea as sister to Gastornis giganteus and
Sylviornis neocaledoniae received moderate bootstrap support of
71.92% and was distinguished by three unambiguous character
state changes and five ambiguous ones.
Bayesian inference
MrBayes output was analysed in part using Tracer v. 1.7.2 (Rambaut
et al. 2018). All four independent runs converged and achieved
stationarity. All runs had a Potential Scale Reduction Factor (PSRF,
Gelman and Rubin 1992) of 1.000. The Standard Deviation of Split
Frequencies (SDSF) approached 0.0 as MCMC runs converged, with
an average value across runs of 0.004799. After burn-in (20%), the
mean log likelihood statistics (LnL) and Effective Sample Size (ESS)
values for each run from one to four were −2505.3998 and 7601.4,
−2505.2851 and 7750.2, −2505.33 and 8001, and −2505.4065 and
7817.2, respectively. Post-burn-in trees of the four independent
runs were combined to produce a consensus tree, which is displayed
with posterior probability (PP) support values (PP in Figure 12B). As
above, only the topological relationships of fossil taxa and the asso­
ciated support for their resolved positions will be focused on here.
The topological relationships are similar to those recovered in
parsimony analysis with respect to the clade that is associated with
modern and fossil anseriforms. The most inclusive anseriform
clade, formed of Anhimidae, Dromornithidae, Conflicto antarcti­
cus, Nettapterornis oxfordi and Anseres, is robustly recovered with
a posterior probability (PP) of 1. The posterior distribution sup­
ports the monophyly of a Anhimidae + Dromornithidae clade, as
1126 P. L. MCINERNEY ET AL.

Figure 11. The hyoid elements of Genyornis newtoni specimen SAMA P59516: A. Complete specimen SAMA P59516 in ventral view; B. paraglossum, arrows for orientation
(dorsal, ventral); C. Right lateral view of the basihyal and both left and right ceratobranchials in situ, image rotated 90 degrees clockwise from A.; D. Annotated outline of
hyoid elements, arrow for orientation of basihyal only (clockwise: dorsal, rostral, ventral, caudal). Annotations: a.cb., damaged region just rostral of the articulatio
ceratobasihyalis; a.pb., surface corresponding to articulatio paraglosso-basihyalis; bh., basihyal; cb., ceratobranchials; cor., cornua (processus paraglossus caudalis); for.,
central foramen. Scale bars: A. 40 mm, B. 10 mm, C., D. 20 mm.

a distinct group sister to other anseriforms, but is poorly supported

(PP of 0.46, resolved in 49.25% of post-burn-in trees). A clade
including all dromornithid species is supported with PP values of
0.77 and recovered in 80.82% of post burn-in trees. This is sister to
a poorly supported clade comprising of Anhimidae (PP of 0.67),
resolved in 65.24% of post-burn-in trees; the low support and
frequency for the Anhimidae clade may be linked to a clade com­
prising Dromornithidae + Chauna chavaria as sister to Anhima
cornuta which occurred in 26.58% of post burn-in trees, but was
not produced in the majority-rule consensus tree (Figure 12B). The
exact interrelationships of dromornithid taxa within this clade
receive low PP values for their resolved positions. A clade compris­
ing Genyornis newtoni and species of Ilbandornis is resolved but is
weakly supported (PP of 0.54). Other fossil and modern anseri­
forms are resolved in a strongly-supported clade (PP of 1). In
contrast to parsimony analysis, the fossil taxon Nettapterornis
oxfordi is resolved basal to Conflicto antarcticus, the latter of
which is resolved in a monophyletic clade including the Anseres
with weak support (PP of 0.69). The clade including Presbyornis
pervetus and crown group Anseres is supported by a PP of 0.85,
while the posterior probability of the clade comprising all non-
anhimid crown Anseriformes receives strongly support (PP
of 0.99).
All fossil and modern Galliformes, as well as Gastornis giganteus
and Asteriornis maastrichtensis, are united in a poorly supported
clade (PP of 0.47). The base of this clade is characterised by a clade
including fossil taxa Gastornis giganteus and Sylviornis neocaledo­
niae, which is also weakly supported (PP of 0.65), and one compris­
ing all crown group galliforms and Asteriornis maastrichtensis (in
contrast to the position of A. maastrichtensis in the analysis parsi­
mony optimality criterion) that is resolved with low support (PP of
0.53). The posterior distribution recovers fossil Asteriornis maas­
trichtensis as sister to the megapodiid taxa in a monophyletic clade
(PP of 0.41). A monophyletic clade, sister to the aforementioned,
comprising all non-megapodiid crown galliforms received weak
support (PP of 0.65).
Remarks
Dromornithidae are phylogenetically resolved within Anseriformes,
with close affinities to Anhimidae across both phylogenetic meth­
ods, as evidenced in both consensus trees (see Figure 12).
Phylograms illustrate relatively little variation in branch lengths
across dromornithid species, in the context of the wider
Galloanserae, indicating only minor morphological change with
regards to the characters assessed (Figure 12 and SI Figure 1). It
should be noted that while a clade of Anseriformes including
dromornithids are strongly supported in Bayesian analyses (PP
of 1), bootstrap support for this clade (excluding Asteriornis maas­
trichtensis) is only moderately supported (76.57%). This can likely
be attributed to the nature of bootstrapping, where alternative
 HISTORICAL BIOLOGY 1127

Figure 12. Galloanseran consensus trees derived from skull characters to assess the relationships of dromornithids: A. Parsimony strict consensus tree cladogram of 2 MPTs
(length = 786, corresponding phylograms are displayed in SI 5), bootstrap support values are displayed below each corresponding branch; B. the consensus tree phylogram
based on Bayesian inference (majority-rule, undated). The scale bar in B. relates to degree of morphological change across branch lengths. Posterior Probability values are
specified next to respective nodes. The suborders Anhimae and Anseres are indicated, as is the superfamily Phasianoidea. Branches are differentially coloured
corresponding to Galliformes (green) and Anseriformes (blue). The additional colour gradient across branches is indicative of support values in both consensus trees
displayed (bootstrap values and posterior probabilities, respectively). Fossil taxa are shown in bold.
1128 P. L. MCINERNEY ET AL.
topologies can be preferred in the random resampling of characters
with replacement in each pseudoreplicate. This is due to the poten­
tial occasional exclusion of characters that are strong drivers of the
resolved topology in the extensive heuristic search and may also
represent characters multiple times in the ‘bootstrap matrix’
(Felsenstein 1985). All dromornithids are united in
a monophyletic clade to the exclusion of non-dromornithids in all
analyses, associated with high bootstrap support and low posterior
probability values. The latter may be reflective of shared character­
istics among dromornithids and anhimids, and resultant alternative
branching arrangements within the wider clade comprising all of
these taxa (such as a Dromornithidae + Chauna chavaria clade,
sister to Anhima cornuta, see above). Similarly, the low posterior
probability values estimated for the monophyly of a clade including
dromornithids and anhimids can be associated with alternative
topological placements of Dromornithidae and Anhimidae, variably
as sister to a strongly supported clade (PP of 1) that includes
Nettapterornis oxfordi, Conflicto antarcticus and Anseres (including
Presbyornis pervetus). Specifically, the clade comprising Anhimidae
+ Nettapterornis oxfordi, Conflicto antarcticus and Anseres occurred
in 32.87% of post burn-in trees estimated by the posterior distribu­
tion, compared to a frequency of 17.85% for a Dromornithidae +
Nettapterornis oxfordi, Conflicto antarcticus and Anseres clade.
These topological arrangements were comparatively associated
with low bootstrap support values of 13.77% and 6.10% respectively,
under parsimony optimality criterion.
Consensus trees derived from maximum parsimony and
Bayesian phylogenetic approaches differ topologically in the posi­
tions of Nettapterornis oxfordi and Conflicto antarcticus, where the
latter taxon is more basal in Anseriformes in all MPTs, whereas
Bayesian phylogenetic inference resolved them in a relatively more
derived position as sister to Anseres. Their close relationship, as
supported in both analyses here, was inferred by the MPT presented
by Tambussi et al. (2019), although these authors specifically
resolved these taxa as sister to one another within a monophyletic
clade. These topological differences are likely highly contributed to
by our focus on skull characters exclusively, where the additional
incorporation of postcranial ones will likely better resolve their
phylogenetic relationships.
Inconsistencies across parsimony and Bayesian inference meth­
ods are also evident in the phylogenetic positions of Asteriornis
maastrichtensis and the clade containing Gastornis giganteus and
Sylviornis neocaledoniae (see Figure 12). A clade comprising the
latter two taxa is consistently resolved in a basal position among
Galliformes, on a basal branch sister to a clade that includes all
galliforms in Bayesian analysis, while under parsimony criterion it
is resolved in a relatively more derived position, sister to a branch
comprising Phasianoidea, as part of an inclusive clade that is itself,
sister to Megapodiidae. The associated low branch support under
both methods is illustrative of this topological inconsistency, likely
contributed to by the presence of only three unambiguous character
state changes supporting the resolution of these taxa in a clade sister
to non-megapodiid crown galliforms in parsimony analysis. As for
dromornithids, these results contrast with those presented by
Worthy et al. (2017b) and Tambussi et al. (2019), whereas a sister
taxon relationship between Gastornithidae and Dromornithidae is
not supported by parsimony analysis or Bayesian phylogenetic
inference using skull characters. Our results thus recover
Gastornithiformes (sensu Worthy et al. 2017b) as a paraphyletic
clade.
The poorly supported basal galliform affinities for Asteriornis
maastrichtensis resulting from Bayesian inference, is comparable to
relationships reported for an unconstrained parsimony analysis by
Field et al. (2020) which was associated with low bootstrap support
values, as well as positions moderately and strongly supported by
the posterior distribution in both tip-dated and undated Bayesian
analyses (PP of 0.93 and 1.0, respectively), and favoured by their
stepping-stone analyses. In contrast, Asteriornis maastrichtensis is
resolved sister to the most inclusive crown group anseriform clade
in parsimony analyses herein, yet the branch to the clade including
this taxon and anseriforms only received low support values
(42.98% bootstrap support). Field et al. (2020) cited the lack of
a postcranial skeleton in this taxon and the inability to appraise
other fossil taxa for certain characteristics as contributing factors in
it being resolved among stem galliforms in undated Bayesian ana­
lysis. The inability to consistently and strongly resolve the topolo­
gical relationships of Asteriornis maastrichtensis in this study using
exclusively skull characters, is similarly likely contributed to by
limitations regarding the number of codable characters and the
effects of missing data (e.g. Weins 2003, 2006), and notably a lack
of information pertaining to the lateral and caudal cranium in this
taxon. More extensive testing using a wider sample of modern and
fossil taxa, and postcranial characters, is needed to better under­
stand how Asteriornis maastrichtensis relates to modern and fossil
Galloanserae.
In general, the differing topologies and inability to strongly
resolve or support the placement of several taxa, especially near
the base of Galliformes and Anseriformes, is likely reflected in the
limited sample of characters (and to a lesser degree, taxa) used in
these precursive analyses (skull features only; see Materials and
methods: Phylogenetic analyses), missing data associated with fossil
taxa, related taxon instability among trees, and conflicting signal of
relationships between some taxa. The phylogenies presented herein
are primarily presented as a means of phylogenetically testing the
influence of morphological characters described and compared in
this study on higher-level taxon associations and are not intended
to represent an exhaustive test of phylogenetic relationships. We
acknowledge the preliminary nature of these analyses, and the need
to further test the phylogenetic placement of Dromornithidae
among Galloanserae in a more focused phylogenetic study invol­
ving a more complete morphological character set (across the
postcranial and cranial skeleton). We perceive benefits of further
elaboration and comparison of Bayesian-inferred and parsimony-
based phylogenetic methods, more consideration of their appropri­
ateness in resolving the interrelationships of fossil taxa with regards
to crown group Galloanserae, and a more complete evaluation of
topological arrangements, however, these are outside of the scope of
this study.
Discussion
The fossils of Genyornis newtoni described above, are nearly incom­
parable to those used to describe the skull of this species in 1913
(Stirling 1913; see Appendix One: Figure A1), 1998 (Murray and
Megirian 1998), and again in 2004 (Murray and Vickers-Rich
2004). The new material is better preserved, less fragmentary, and
provides a more complete basis for interpreting the skull morphol­
ogy of G. newtoni and creating an updated reconstruction
(Figure 13). Some original interpretations of the skull are sup­
ported, e.g. the lack of the large dorsally convex shape of the
rostrum described for species of Dromornis. However, the new
specimens reveal different proportions of the cranium and rostrum
and that the apparent narrow width of the rostrum is unsupported.
We additionally illuminate further details of the morphology not
previously recognised, reported on, and/or preserved.

Figure 13. An artistic reconstruction of the skull of Genyornis newtoni, based on all available fossil material, left lateral view. Illustration by Jacob C. Blokland. Scale bar is
equal to 50 mm.
The Dromornithidae
Following the findings of Murray and Vickers-Rich (2004, p. 328)
and Worthy et al. (2016a), the overall structure of the dromornithid
skull is highly conserved and varies little amongst the species,
G. newtoni included. The results of the phylogenetic analyses herein
also support this (see above, Figure 12 and SI Fig, 1). Although no
feature is necessarily unique to the family, together all features form
a characteristic morphology. This is inclusive of, but not limited to:
the structure of the synovial craniorostral hinge and its transection of
the rostrodorsal portion of the orbit; a hemispherical dorsal cranial
surface in caudal view; the reduction in cranium rostrocaudal length;
the rostral extension of aponeurosis zygomatica ossificans along the
ventrolateral postorbital process and the medial retreat of
impressio m. AME profundus, pars coronoideus; the presence of an
annulus tympanicus and a rostroventrally enclosed osseous meatus
acusticus externus; the enlarged and ventrally elongate processus
paroccipitalis; the fossa pseudotemporalis on the caudal wall of the
orbit; the lacrimal fused to the caudal rostrum; the desmognathous
palate; the fused elements of the mandible which eliminate flexion
therein; the small medial mandibular processes; the oblique, sheering
tomial margin of the caudal rostrum and corresponding part of the
mandible; the broad shape of the retroarticular process; and the
laterally arched and single-headed quadrate.
The skull morphology of G. newtoni appears to be most similar to
that of Ilbandornis species, as supported by results of the phylogenetic
analyses, and suggests a closer relationship to one another (and
possibly to Barawertornis tedfordi), than to those within the mono­
phyletic Dromornis clade (see Figure 12). Many features of the skulls
of G. newtoni and species of Ilbandornis are more slender than those
attributed to species of Dromornis. This is especially noticeable in the
quadrates which seemingly show two forms: a robust Dromornis-
type, and the slender G. newtoni/Ilbandornis-type with a more elon­
gate pars otica, less lateral overhang of the pars otica by the capitulum
squamoso-oticum, and a proportionally smaller pars mandibularis
(see Figure 6; Worthy et al. 2016a: fig. 4). Additionally, although the
foreshortening of the frontal and parietal bones of the cranium is
HISTORICAL BIOLOGY 1129
shared by all dromornithids, the most extreme rostrocaudal compres­
sion of the cranium occurred only in species of Dromornis, with
successive shortening from D. murrayi to D. planei and D. stirtoni,
with the cranium of the latter twice as bilaterally wide and three times
dorsoventrally taller than its rostrocaudal length (see Worthy et al.
2016a; Handley and Worthy 2021).
Regardless of such general similarities in morphology among
dromornithids and further similarities to species of Ilbandornis, the
skull of G. newtoni does show specific features which characterise
the taxon and give support to its generic distinction. This includes
the smaller, more enclosed osseous meatus acusticus externus and
the especially long and slender processus paroccipitalis, both of
which, may have relevance to functional adaptations for increased
musculature attachment on these areas. Another feature, so far
unique to G. newtoni, is the presence of the triangular casque on
the rostrum, and the hypothesised extent of the rhamphotheca
which covers most of the upper beak – apparently more than in
species of Dromornis. This may suggest a potentially significant
difference in communication and sexual displays, and there are
several possibilities as to what the function of such a structure
may be, similar features are known in other galloanserans such as
Sylviornis neocaledoniae, and in curassow cracids (Mayr 2018b).
Brightly coloured casques on the bill have a clear sexual display
function. Furthermore, the rostrum of G. newtoni also differs from
those of species of Dromornis in its wide, and rounded, more
spatulate shape, with far shallower depth (see ‘Rostrum’).
Although the upper bill morphology is unknown for species of
Ilbandornis, it has a dorsoventrally shallow mandible compared to
species of Dromornis, and given the similarities in the cranium,
quadrate, and mandible with G. newtoni, we may expect the ros­
trum to also be similar in morphology.
Gastornithiformes
Previous phylogenetic analyses resulted in the weak association of
dromornithids with the gastornithids in the Gastornithiformes
1130 P. L. MCINERNEY ET AL.
Stejneger 1885 (Worthy et al. 2017b). The morphological descrip­
tions and phylogenetic analyses presented herein, although preli­
minary, are not congruent with this hypothesis, warranting further
discussion on the evolutionary relationships of these groups with
larger datasets as more complete specimens become available
Skull morphology
Dromornithids and gastornithids share various aspects of cranial
morphology but differ markedly in others. Some identified shared
characters in both families are reflective of their galloanseran affi­
nities, for example, the sessile processus basipterygoideus of the
crania, the extensive processus retroarticularis of the mandible (see
Baumel and Witmer 1993; Weber 1993; Dzerzhinsky 1995; Ericson
1996; Livezey 1997; Murray and Vickers-Rich 2004; Mayr 2017;
Mayr et al. 2018; Tambussi et al. 2019; Field et al. 2020), the
prokinetic skull, and holorhinal nares (Sibley and Ahlquist 1990;
Dzerzhinsky 1995). Other morphological features conform to typi­
cal anseriform traits, such as the fused elements of the rostrum, the
flat, enclosed palatal roof, caudally restricted external nares, shape
and angle of the mandibular condyles on the quadrate, and the long
processus paroccipitalis (see also Andors 1988, 1992). Notably, in
the phylogenetic analyses, few, if any morphological character states
shared by both gastornithids and dromornithids, are only found in
galliforms (character 72, ambiguous for Anhimidae).
Other characters, such as the synovial craniorostral hinge, and
the reduction of the anteorbital/supra-orbital region of the cra­
nium, are less common in Galloanserae and yet are present in
dromornithids, gastornithids and sylviornithids. These features
likely inform more on a homoplasious functional adaptation to
the robusticity and large size of both upper and lower jaws, rather
than being representative of close relatedness phylogenetically. For
sufficient mobility of the prokinetic upper bill and flexion capabil­
ities of the zona flexoria craniofacialis to be maintained, the region
must be non-pneumatic, very flat, and in larger species, composed
of sheets of overlapping thin bone (Bühler 1980, p. 450, 451). This
limits the strength of the region against feeding and movement
stress, making it the weakest point of the prokinetic upper bill
(Bock 1966). The transformation of a flexion zone to a synovial
joint that is not spanned by any thin osseous tissues, allows greater
flexion between the cranium and rostrum; as seen in parrots
(Psittaciformes), for example, which have syndesmotic or synovial
articulations with cartilaginous enclosures (Bühler 1980, p. 450,
451). Witmer and Rose (1991) suggested the connective tissue
within a synovial joint would provide the stability required to
manage the stress associated with a large, heavy upper jaw, in
reference to the form in gastornithids. As described above
(‘Craniorostral hinge’), the dromornithids, especially the larger spe­
cies, and S. neocaledoniae, have evolved a ball and socket arrange­
ment to the hinge-like joint that confers much stability while
allowing rotation on the occlusal plane (Murray and Megirian
1998; Murray and Vickers-Rich 2004; Mourer-Chauviré and
Balouet 2005). The synovial hinge is then likely a size-scaling
phenomenon, allowing, or deriving from, the development of larger
skulls without the limitations associated with flexion zones. Should
this be the case, gastornithids, dromornithids, and sylviornithids
would have convergently evolved this hinge type as an adaptation to
gigantism; a hypothesis supported by the restriction of this hinge
type primarily to giant galloanserans. Reduction of the anteorbital
region of the cranium could provide additional hinge support, as
the increase in the robusticity and surface area available on the
cranium for the rostrum to articulate with, would assist in mana­
ging and distributing a stress load. Alternatively, or in association,
the synovial craniorostral hinge could have evolved as an essential
functional adaptation to a specific feeding strategy (see 6.4). The
various configurations between the lacrimals and the rostrum and
cranium in these taxa, being fused to the rostrum in dromornithids,
while they are totally fused to the frontals but articulate with the
rostrum in gastornithids and sylviornithids (pers. observ.; Andors
1992; Mourer-Chauviré and Balouet 2005), possibly conferred an
additional degree of lateral stabilisation in the latter families, as in
large anatids (Bühler 1980, p. 452), compared to dromornithids (cf.
Murray and Vickers-Rich 2004: fig. 184). The dromornithid state
instead better resembles the less laterally confined hinge of parrots
(except that the lacrimals exclusively articulate with the cranium
rather than the rostrum in these birds, see Tokita 2003)
In Anatidae, and especially in species of Dendrocygna, the pro­
cessus orbitalis of the lacrimal is ventrocaudally developed, coupled
with a similarly orientated ligamentum lacrimomandibulare that
likely prevents caudal displacement of the mandible during water
expulsion in feeding (Zelenkov and Stidham 2018, and references
therein). The fusion of the lacrimals to the frontal bones in many of
these birds provides strength against caudal pulling, in the action of
this ligament (Zelenkov and Stidham 2018). However, this ligament
is absent in galliforms, anhimids, presbyornithids and anseranatids
(although represented by extensive fascia in the latter, see
Dzerzhinsky and Grintsevichene 2002; Zelenkov and Stidham
2018), most of which do not have lacrimals that are fused to the
cranium or rostrum either (see ‘Craniorostral hinge’, character 32).
This is paired with the absence of a strongly ventrocaudally directed
orbital process in all the aforementioned taxa, which is also not
observed in dromornithids, sylviornithids or gastornithids. Thus,
the fusion of the lacrimals to the cranium in sylviornithids and
gastornithids cannot be an adaptation to the feeding specialisations
typical of Anatidae, in this regard. In anatids, however, the frontal-
fused lacrimals articulate with a distinct notch on the caudal ros­
trum, which acts as a stop, supporting the mobility of the upper jaw,
and reinforcing this joint against breaking and excessive upper bill
retraction (Fisher 1955; Zelenkov and Stidham 2018). In sylvior­
nithids and gastornithids, the strong and complete fusion of the
lacrimals with the frontals, and their corresponding clear articula­
tory structure on the caudal rostrum (pers. observ., see Matthew
and Granger 1917; Mourer-Chauviré and Balouet 2005) may have
provided strength in a similar manner.
The ventrally short processus orbitalis of the lacrimal in sylvior­
nithids and gastornithids also contrasts with the dromornithid
condition (character 33). The ventral end of this process supports
the ligamentum jugolacrimale which runs to the tuberculum lacri­
male on the rostral arcus jugalis, which is well developed in anatids
(Davids 1952c; Goodman and Fisher 1962; Bock 1964; Baumel and
Raikow 1993), and may limit protraction of the rostrum (Fisher
1955). The processus orbitalis in anhimids is ventrally descending
but not as robust as in anatids but is similarly coupled with
a ligamentum jugolacrimale that is stronger than what is typically
observed in galliforms (see Ghetie et al. 1976; Dzerzhinsky 1982,
p. 1031). The ventrally long and narrow orbital process in presby­
ornithids is also hypothesised to have supported this ligamentous
attachment to the jugal arch (Zelenkov and Stidham 2018). The
poorly developed and ventrally short processus orbitalis on the
lacrimals of sylviornithids and gastornithids probably did not host
a well-formed ligamentum jugolacrimale if it was present. In con­
trast, the strut-like complete ossification between the processus
orbitalis and the jugal arch in dromornithids presumably provided
additional strength across the rostrum and arcus jugalis, and
reduced bending at the rostral end of the latter (zona flexoria
arcus jugalis); comparatively, this region is a more mobile synovial

socket and joint articulation in sylviornithids and gastornithids
(character 96, Matthew and Granger 1917; Mourer-Chauviré and
Balouet 2005).
As discussed in the section ‘Temporal region’, the formation and
structure of the lateral cranium (including the AZO) and the loca­
tion of associated musculature (characters 7–12), is a major point of
difference between the gastornithid and dromornithid skulls. This
region is nearly identical between dromornithids and anhimids,
with respect to homologous structures. Comparatively, that of
gastornithids is more like the morphology of many cracids and
phasianids, especially regarding the robust orbitozygomatic junc­
tion and resultant secondary temporal fenestra (see Figure 14).
Elzanowski and Mayr (2017) recognised cracids and phasianids to
have independently evolved this morphology, because the closure
of the orbitozygomatic junction is not so complete in the stem
galliform S. neocaledoniae, and extant megapodiids. Therefore,
independent evolution in the gastornithids would be possible.
The extreme size of the ossified aponeurosis zygomatica and
secondary temporal fenestra in gastornithids is linked to a second
adaptation in this group for an increase in the surface area available
for the external adductor muscles, primarily related to the origin
of m. AME profundus, pars superficialis, on the lateral orbitozygo­
matic junction, but also m. AME profundus, pars zygomaticus,
medially, and m. AME profundus, pars coronoideus, within the
more medial, enclosed and well-excavated dorsotemporal fossa on
the squamosum, and possibly some fibres related to m. AME super­
ficialis dorsal of the quadrate articulation (see ‘Cranium’, also
Witmer and Rose 1991, p. 102; Weber 1996; Zusi and Livezey
2000; Holliday and Witmer 2007). The rostrocaudally compressed
cranium of species of Dromornis and the different morphology of
the lateral cranium in dromornithids allows for a comparatively far
smaller surface area for origin of m. AME profundus, pars coro­
noideus and would not have supported such a hyperdeveloped
musculature. However, numerous well-developed osseous struc­
tures of the jaw apparatus in dromornithids show some emphasis
on the m. AME superficialis (with some associated cristae on the
dorsal or dorsocaudal margin of the osseous meatus acusticus
externus), m. adductor mandibulae posterior, m. pterygoideus,
and the m. pseudotemporalis superficialis (associated fossa and
tubercle within the area muscularis aspera), indicative of consider­
able jaw power. The especially heavy and robust quadrates in
species of Dromornis compared to that of G. newtoni and species
of Ilbandornis, and the lack of a pneumatic foramen in this bone in
D. stirtoni, may be further adaptions for supporting and maintain­
ing large muscle mass in order to adduct the massive mandible.
Functional implications of the dromornithid jaw apparatus are
explored further in ‘Ecomorphology and niche adaptations’.
The contrasting conditions in both families appear to be driven
by the need for well-developed musculature, in part to support large
upper and lower jaws, but is limited by the variation in underlying
cranium morphology (especially influenced by the development of
the AZO). Further indications of large muscle size from analysis of
the crania of Gastornis species are evident in the crista nuchalis
sagittalis, prominentia exoccipitalis, and processus paroccipitalis.
Such adaptations for enlarged muscle attachments have led to the
suggestion that the adductors were unusually powerful (Andors
1988, p. 137) and were a driving factor in the carnivory hypothesis
(see also Witmer and Rose 1991; Mayr 2022a, p. 52), although it also
may support a folivorous diet including hard food items (Andors
1992; Mayr 2022b). It was not considered that the extensive mus­
culature may be required to manipulate such a large rostrum and
mandible without great force, a factor which further analysis of the
skulls of these taxa may shed light on. Witmer and Rose (1991)
HISTORICAL BIOLOGY 1131
additionally linked the development of massive palatines in gastor­
nithids to a requirement for large muscle attachments, specifically
the m. pterygoideus, which they propose to be one of three impor­
tant ways of increasing bite force and may explain the similarly
massive size of the palatines of dromornithids. Unfortunately, con­
firmation of the area involved, and development and use of these
muscles can only be made by assessing the formation of the separate
cranial bones in chicks (e.g. Mayr and Manegold 2021) in associa­
tion with myological dissections as per Zusi and Livezey (2000),
both of which are not possible for these fossil taxa (see also palaeo­
myological considerations as per Ostrom 1961, p. 88–89).
Aside from the dorsoventrally broad processus retroarticularis,
and the large size—intermediate between that of species of
Dromornis and G. newtoni—the mandible of all gastornithids varies
considerably from that of dromornithids. Instead, several features
of the mandible of Gastornis species bear more resemblance to that
of S. neocaledoniae, including the great proportional rostrocaudal
length of the symphysial region (character 59), and the morphology
of pars caudalis of the mandibular ramus, specifically the articula­
tory region. Similarly, the rostra of both Gastornis species and
S. neocaledoniae appear superficially like that of dromornithids,
with large palatines, flat, fused, plate-like lateral surfaces and
a casque structure (the latter only between S. neocaledoniae and
G. newtoni). However, both differ from dromornithids in their
maxillary articulation with the jugal arch, the small vomer, not
likely contributing to the palatal roof, and the sharp tomial margin
(Matthew and Granger 1917; Mourer-Chauviré and Balouet 2005).
In the cranium, neither gastornithids, nor sylviornithids, have
a rostrally enclosed osseous meatus acusticus externus (and
cavum tympanicum), and they both retain distinct capitula on the
head of the quadrate despite it being functionally single headed.
Phylogenetic relationships of the giant flightless
galloanserans as informed by skull morphology
Among the Galloanserae, three extinct, basal lineages diversified to
include giant terrestrial forms in the families Dromornithidae,
Gastornithidae, and Sylviornithidae (Worthy et al. 2016b; 2017b).
We exclude the Brontornithidae Moreno and Mercerat 1891, from
the comparative and phylogenetic analyses following the findings
on the phylogenetic affinities of the group by Worthy et al. (2017b).
The cranial material that can be associated, then only tentatively,
with this taxon is restricted to a fragmented mandibular symphysis
and a quadrate, which has a condylus mandibularis caudalis on pars
mandibularis (for most recent analyses see Worthy et al. 2017b: fig.
5; Agnolín 2021: fig. 6). This feature is absent in all quadrates of all
crown-group, and definitively placed fossil galloanserans which all
have a bicondylar articulation (e.g. see Dzerzhinsky 1974, 1982,
1995; Mayr 2017, 2022a; Field et al. 2020).
Recent phylogenetic analyses have aligned the dromornithids
with the gastornithids to include them in the order
Gastornithiformes, in effect, excluding them from the orders
Galliformes and Anseriformes, or being considered distinct at
ordinal level. However, this relationship was weakly resolved in
the phylogenetic analyses by Worthy et al. (2017b), and several
authors have since questioned it. Mayr (2022a), one of such
authors, listed several cranial and postcranial characteristics of
dromornithids which distinguish them from gastornithids, and
suggesting the relationship was supported only by convergent char­
acters. Our morphological comparisons and phylogenetic assess­
ments also find little support for a dromornithid-gastornithid clade
(see above); several major differences in the morphology of the
skulls are present, and many similarities can be explained by the
1132 P. L. MCINERNEY ET AL.

Figure 14. Two select morphological features which show variation across Galloanserae, and, in isolation as single character studies, indicate different phylogenetic
hypotheses for the evolutionary placement of the Dromornithidae: A. the presence of either the rostromedial or caudomedial foramen (dark orange) in relation to the crista
medialis (light orange) on the quadrate, in medial view; B. the relationship between the aponeurosis zygomatica and processus postorbitalis, in addition to the presence of
an ossified portion (dark orange) of the aponeurosis zygomatica (light orange). Dotted and dashed branches in B. illustrate alternative hypotheses discussed in text; the
unresolved position illustrated by the dotted line denotes the uncertainty of the order of divergence between the anhimids and dromornithids. Positions of Conflicto
antarcticus and the Presbyornithidae are based on Tambussi et al. (2019: fig. 14) and Houde et al. (2023: fig. 9B). Nodes do not correlate with time. Not all families within
Galloanserae are displayed. Morphologies drawn from photos and figures in literature: A.: Anseres – Anseranas semipalmata (see Elzanowski and Stidham 2010: fig. 5C),
Anhimidae – Anhima cornuta (see Elzanowski and Stidham 2010: fig. 8B), Presbyornithidae – (see Elzanowski and Stidham 2010: fig. 5B), Conflicto antarcticus – (Tambussi
et al. 2019: fig. 6A), Dromornithidae – Genyornis newtoni (NMV P256893), Megapodiidae – Megapodius freycinet (see Elzanowski and Stidham 2010: fig. 5A), Phasianidae –
Gallus gallus (FUR 119); B.: Anseres – Sarkidiornis melanotos (see Zusi and Livezey 2000: figs. 6D, 7D), Dromornithidae – Genyornis newtoni (SAMA P59516), Anhimidae –
Chauna torquata (see Zusi and Livezey 2000: figs. 6C, 7B), Megapodiidae – Alectura lathami (SAMA B2439), Phasianidae – Meleagris gallopavo (see Zusi and Livezey 2000: fig.
4 G). Silhouettes (designed by PLM) are illustrative representations of species in the same family or appropriate clade, as indicated. Images are not to scale.

galloanseran affinities of the lineages and convergent functional
adaptation to large body and skull size. Thus, we consider enough
evidence is present to warrant reconsideration of this relationship.
As an example, one of the most enticing similarities between the
dromornithids and the gastornithids is the synovial craniorostral
hinge. As aforementioned, there is a high likelihood that the shared
hinge type convergently evolved as a functional adaptation to feed­
ing or supporting large skull size. This suggests that the morphology
of this region may not be as phylogenetically important in inform­
ing upon the higher-level evolutionary relationships of these taxa.
The separation of dromornithids and sylviornithids in phylogenetic
analyses, and the sister-group relationship between gastornithids
and sylviornithids, further supports minimally two independent
evolutions of the craniorostral hinge. Additionally, this supports
our identification of morphological similarities between gastor­
nithids and the convincingly galliform, S. neocaledoniae, showing
the importance of recognising these similarities when reassessing
gastornithid relationships.
Andors (1988, 1992) recognised the mosaic structure of several
elements in gastornithids as evidence of ‘intermediacy between the
orders Galliformes and Anseriformes’ and considered them to be
the ordinal-level sister-group to anseriforms, and more broadly
related to galliforms. Similarities with galliforms, including cracids
and phasianids were considered symplesiomorphic, and some
shared features with anhimids were considered evidence of con­
sanguinity with basal anseriform stock. Comparatively, Murray and
Vickers-Rich (2004, p. 167) concluded that any relationship
between gastornithids and anseriforms is ‘undoubtedly remote’,
noting that they appear to have more in common with the
Megapodiidae. Our study of the skull morphology and phylogenetic
analyses support a closer relationship of gastornithids with basal
galliforms than anseriforms.
Dromornithids within Galloanserae
Bayesian and parsimony-based phylogenetic analyses by Worthy
et al. (2017b) variably placed Gastornithiformes at the ordinal level,
as sister to galliforms and anseriforms respectively with no signifi­
cant results favouring either placement. Ultimately, however, the
slightly higher support and 13 identified unambiguous characters in
their study, suggested the placement as sister to galliforms to be
most likely. Handley and Worthy (2021) additionally considered
aspects of the morphology of the brain, several cranial nerves, and
trigeminal ganglia to support this placement (see below). The
alternative pull of dromornithids to galliform or anseriform
lineages may be a function of the early diverging nature of the of
this group, resulting in plesiomorphic characters confounding the
results of analyses.
Galliformes
This study found little evidence to support the sister-group relation­
ship of dromornithids and galliforms due to dromornithids having
relatively few exclusively galliform skull characteristics. Typically
crown group galliform characters which are present, include the
apparent absence of a crista nuchalis sagittalis and fonticuli occipitalis
(character 19), the short and ventrally directed processus postorbitalis
of the cranium (character 7), the presence of paracoronoid crests on
the lateral mandible (character 80), and the foramen pneumaticum
rostromediale on the pars otica of the quadrate (character 38). Such
characters could evidence galliform affinities for the dromornithids.
However, many of these are also observed in anhimids (previously
discussed by Dzerzhinsky 1982), concordant with their early diver­
gence as basal anseriforms supported by all molecular phylogenetic
HISTORICAL BIOLOGY 1133
analyses and most/all morphological ones (see Figure 12; for addi­
tional examples, see Worthy et al. 2017b; Kuhl et al. 2021). These
traits may instead be plesiomorphic for Galloanserae and lost gradu­
ally associated with the evolution of Anseres and close relatives (see
SI 5). Indeed, many similarities between Galliformes and
Anseriformes are considered to be relatively primitive states (see
Murray and Vickers-Rich 2004, p. 168–169). Therefore, we infer
a priori polarisation for several characters.
For example, the paracoronoid crests on the lateral mandible
that are present in dromornithids, and typical of galliform birds, are
also present in anhimids (Weber and Hesse 1995; Weber 1996).
Anhimids also have a processus postorbitalis that lacks the distinct
rostral extension typical of other anseriforms (this is also true for
Presbyornis pervertus, specimen USNM.VP.299846, pers. observ.,
see also Olson and Feduccia 1980; Zelenkov and Stidham 2018:
fig. 3), and they lose the fonticuli occipitalis that are typical of
waterfowl in adulthood (Ericson 1997). Additionally,
a discernible, yet not prominent, crista nuchalis transversa is
observed in anhimids, galliforms and dromornithids, in contrast
with the hyper-developed form of many anseriforms. The location
of the condylus occipitalis at near mid-dorsoventral height on the
caudal surface of the cranium in most dromornithids (excluding
D. murrayi in which it is more ventrally positioned), is also a trait
shared with both galliforms and anhimids.
That several conditions considered typical of galliforms appear
to be symplesiomorphic for anseriforms is especially important to
consider with regards to the quadrate, as archaic fossil relatives such
as the near-galloanseran Asteriornis maastrichtensis (see Field et al.
2020), and the basal anseriforms Presbyornis pervetus and Conflicto
antarcticus, appear to share quadrate traits with basal galliforms
(see Elzanowski and Stidham 2010; Elzanowski 2014; Zelenkov and
Stidham 2018; Field et al. 2020: SI). The Upper Cretaceous
A. maastrichtensis possesses a foramen pneumaticum rostrome­
diale, typical of galliforms, although there is an apparent, deep
depression on the caudomedial side of the medial crest, which is
potentially a precursor to the state that characterises most anseri­
form taxa. Early Eocene Danielsavis nazensis appears to have
neither foramina (Elzanowski 2014; Houde et al. 2023; Mayr et al.
2023), although it also forms a deep depression on the caudomedial
surface (Houde et al. 2023: fig. 7; Mayr et al. 2023: fig. 3C). Despite
clear anseriform affinities, Conflicto antarcticus has both
a rostromedial and caudomedial pneumatic foramen (Tambussi
et al. 2019: fig. 6), while species of Presbyornis often have
a markedly deep depressio caudomedialis where a caudomedial
pneumatic foramen is occasionally present (Elzanowski 2014).
Similarly, late Paleocene Anachronornis anhimops has a deeply
grooved depressio caudomedialis that terminates in a diminutive
foramen pneumaticum caudomediale on one quadrate but may not
be present on the other (Houde et al. 2023). Additionally, morpho­
genetic instability in the development of the pneumatic diverticula
has been recognised for both anhimids (Elzanowski 2014) and
presbyornithids (Elzanowski and Stidham 2010), as the caudome­
dial foramen is vestigial and often absent. In anhimids, their parti­
cularly erratic pneumatic variability has been correlated with their
highly derived quadrate morphology and may be evidence of
a transitionary state with respect to the more derived evolution of
the caudomedial pneumatic foramen (Elzanowski and Stidham
2010). While some non-anhimid anseriforms possess the caudome­
dial foramen in the caudomedial position, in others it is rostrally
displaced (e.g. species within Anserinae and Mergini, Elzanowski
and Stidham 2010). This factor, in addition to the variable presence
of the rostromedial foramen in individuals of G. newtoni and other
dromornithids, demonstrates the diversity regarding the status of
the pneumatic foramina within Galloanserae and challenges the
1134 P. L. MCINERNEY ET AL.
weight of using the presence or absence of these foramina as
polarising traits, with respect to assigning fossil taxa to galliform
or anseriform clades. Regarding the dromornithids, this character
may preclude close affinities with the danielsavids, presbyornithids,
anachronornithids, anseranatids, and anatids, but has limited utility
for definitively supporting or excluding a close relationship with
any other galloanseran lineage (see Figure 14A).
A similar case can be made for the foramen pneumaticum
basiorbitale, which is typically present in crown group galliform
birds, as well as Asteriornis maastrichtensis, Danielsavis nazensis,
Conflicto antarcticus, Anachronornis anhimops, and Presbyornis
pervetus (Elzanowski and Stidham 2010; Tambussi et al. 2019;
Field et al. 2020; Houde et al. 2023; Mayr et al. 2023). However,
this region of the quadrate is associated with considerable pneu­
matic variability in both anhimids and cracids (Elzanowski and
Stidham 2010), and appears variably present in Dromornis planei,
yet absent or unverified in other dromornithids. Homoplasy
regarding this character in relatively basal galloanserans may addi­
tionally be affected by gigantism, considering the additional
absence of a basiorbital pneumatic foramen in both gastornithids
and sylviornithids.
Anseriformes
In recent decades, dromornithids have been considered to be basal
anseriforms based predominantly on cranial material (Olson 1985;
Vickers-Rich 1991; Murray and Megirian 1998). Murray and
Vickers-Rich (2004) provisionally refined this to a sister-group
relationship with the Anhimidae to the exclusion of other anseri­
forms, although they suggested that a full revision of the systematic
relationships of Anseriformes and anseriform-like birds, following
identification of additional early Palaeogene fossil representatives,
may result in dromornithids becoming an independent anseriform
Suborder. From our independent assessment of the skull morphol­
ogy of dromornithids, it is clear that the proposed anseriform
affinities of these birds are well-founded; we find support for the
skull synapomorphies identified by Murray and Vickers-Rich
(2004, p. 152–154) in addition to several more that further sub­
stantiate this relationship.
The goose-like rostrum of G. newtoni is a key factor supporting
placement among basal anseriforms (5 character state changes in
our phylogenetic analyses that support a clade containing crown
group anseriforms, including dromornithids, are associated with
the rostrum). The rostrum and mandible show morphological
similarities with several taxa which are representatives of other
independent occurrences of the polyphyletic ‘goose’ type (Li and
Clarke 2016; Olsen and Gremillet 2017; Pecsics et al. 2017), includ­
ing Cereopsis novaehollandiae. This is potentially linked to the
convergent evolution of a primarily herbivorous diet and grazing
behaviour (Olsen 2015), and the elevated rate of beak shape evolu­
tion compared to other bird clades (Cooney et al. 2017; Olsen and
Gremillet 2017). Both have ultimately contributed to the evolution
of diverse feeding ecologies among anseriforms (Li and Clarke
2016).
Of the cranium, the development of the paired exoccipital pro­
minences (for characters see Appendix Four) and the tubercula
basilaria, as well as the low, sessile basipterygoid processes are like
that of anseriforms, with the shared states in dromornithids and
anhimids unambiguously supporting their close relationship, as
found in both phylogenetic analyses. For the mandible, the low
angle of divergence of the rami, the presence and size of the dorsal
and ventral mandibular angles, the caudally directed processus
retroarticularis, and the inferred arrangement of the muscle attach­
ments are all important anseriform characteristics. The prokinetic
nature of the skull, the medial fusion of the paired maxillopalatine
process of the maxillare, and the fusion of elements within the
rostrum, with large, flat praemaxillary and maxillary plates broadly
fused with the nasals, are additional characters dromornithids share
with anseriforms. The desmognathous palate, which provides sup­
port (character 97) for the separation of these clades in the phylo­
genetic analyses, could further support anseriform affinities,
although this has been proposed to have evolved several times
independently from the schizognathous plesiomorphic condition
in anseriforms and galliforms (Hofer 1945; Dzerzhinsky 1982, 1995;
Zusi and Livezey 2006; Mayr 2018a; Field et al. 2020: SI).
Moreover, there are several similarities with the anseriforms in
the quadrates of dromornithids. These include the angle and size of
processus orbitalis compared to pars otica, the presence of the
prominentia submeatica (albeit less inflated, see character 53), the
deep fossa basiorbitalis, the angle of the medial condyle compared
to the lateral one on pars mandibularis, and the adjacent or con­
fluent condylus pterygoideus and facies articularis pterygoidea (see
‘Quadrate’). The latter differs from the condition in Presbyornis
pervetus, Asteriornis maastrichtensis, megapodiids and all phasia­
nids, which instead develop a separate facies articularis pterygoidei
on the caudoventral base of the processus orbitalis (Elzanowski and
Stidham 2010; Field et al. 2020). Dzerzhinsky (1982) and
Elzanowski and Stidham (2010) concluded that this latter state
was likely to be symplesiomorphic for galliforms and possibly all
Galloanserae.
As discussed in sections ‘Temporal region’ and ‘Skull morphol­
ogy’, the form of the AZO and the processus postorbitalis, and the
related structural arrangement and osseous correlates for the man­
dibular adductor muscles in dromornithids, show compelling simi­
larities with the anhimids. This morphology does not occur
anywhere in the galloanseran radiation aside from anhimids and
differs considerably from that of galliforms. However, dromor­
nithids also share some similarities in the shape of this region
with Anseranas semipalmata, as previously described. Concordant
with the basal relationship of anhimids relative to other crown
group anseriforms and several hypothesised character transforma­
tions (e.g. lamellae rostri, see Livezey 1997), the osteo-myological
condition in Anseres is potentially derived from a state more similar
to non-anserean Galloanserae (such as anhimids and dromor­
nithids), whereby more extensive ossification of the zygomatic
aponeurosis of the latter was secondarily lost in Anseres in associa­
tion with structural modification of processus postorbitalis to sup­
port the adductor musculature (see Zusi and Livezey 2000). Fossil
anseriforms such as Presbyornis pervetus and Conflicto antarcticus
also appear to lack such ossification, resembling the state typical of
Anseres, relative to the seemingly more plesiomorphic anhimid and
dromornithid conditions. In this sense, the adductor complex mor­
phology in birds like dromornithids and anhimids appears an
appropriate precursor to the structure of the true waterfowl,
Anseres. This may be a contributing factor to the low support values
in the phylogenetic analyses for the clade comprising dromor­
nithids and anhimids, as such morphology might instead suggest
the Dromornithidae to be an independent basal anseriform lineage
with morphologies somewhat intermediate between anhimids and
anseranatids (as per the hypothesis shown in Figure 14A: dotted
line; see ‘Remarks’). The homology of this region appears increas­
ingly important regarding the evolution of the anseriforms and
identification of character transitions across the Galloanserae
radiation.
An aponeurosis zygomatica ossificans is observed in many adult
crown group galliforms and may extend rostrad until near the level
of processus postorbitalis, which is comparable to anhimids in this
respect (Zusi and Livezey 2000). In some galliforms, such as

megapodiids and cracids, the ossified aponeurosis is non-extensive
and may not reach the postorbital process (Dzerzhinsky 1980; Zusi
and Livezey 2000). Zusi and Livezey (2000) documented similarities
in the location of the m. AME profundus, pars coronoideus,
between megapodes and anseriforms (i.e. the medial rotation and
reduction of impressio m. AME coronoidea). They used this as
evidence of the basal divergence of megapodes within the
Galliformes, as supported by all phylogenetic analyses using mole­
cular data (e.g. Ksepka 2009; Jetz et al. 2012; Worthy et al. 2017b;
Kuhl et al. 2021), and to hypothesise the nature of the transforma­
tion to the anseriform-form from a hypothetical, common ancestor
with galliforms. Considering the morphological similarities
between species of Megapodiidae and Anhimidae for this adductor
complex, the plesiomorphic state of this region may be intermediate
between that of megapodes and anhimids, potentially similar to the
morphology of the Eocene stem galliform Gallinuloides wyomin­
gensis Eastman, 1900. No ossification is present in this latter taxon
(see Mayr and Weidig 2004), which could be expected in stem
Galloanserae given the osteo-myological structure present in basal
galliforms is less hyper-specialised with regards to independent jaw
joint mobility and rostral positioning of the adductor muscles (e.g.
compared to phasianids, see Dzerzhinsky 1974, 1980, 1982; Zusi
and Livezey 2000).
An alternative scenario then, is that the apparent lack of both
a processus zygomaticus and ossification of the zygomatic apo­
neurosis in fossil anseriforms, such as Presbyornis pervetus and
Conflicto antarcticus, is the plesiomorphic state, which is present
in Anseres. This would resultantly infer the ossification of the
zygomatic aponeurosis in anhimids, and, considering their close
similarity, dromornithids, to be a derived state (see also Olson and
Feduccia 1980; Ericson 1997; Zelenkov 2011, p. 909; as per the most
parsimonious topology of Tambussi et al. 2019). The implication of
this hypothesis would imply that the rostral movement of the
adductor complex in Anseres was achieved through adaptation of
the structure of processus postorbitalis only, without any antece­
dent contribution by AZO. In this case, a sister relationship
between the Dromornithidae and Anhimidae would be best sup­
ported, as found in the phylogenetic analyses (Figure 12; and illu­
strated in Figure 14B: dashed line) with the morphology of the AZO
evolving early in their most recent common ancestor; similarities
with anseranatids would then be convergent.
Both hypotheses interpret the independent evolution of the
processus postorbitalis and AZO complex in galloanseran lineages
as a sufficient explanation for the variation in the structure.
Unfortunately, we do not yet have adequate cranial material for
some fossil galloanseran lineages to better test such hypotheses; this
adductor complex region is lacking in the skull of Asteriornis
maastrichtensis (see Field et al. 2020), although considering the
patterns of variation in this region within Galloanserae, and its
basal positioning, it appears unlikely that this taxon would have
developed an AZO (see above). Regardless of which hypothesis
ultimately takes precedence (dotted line or dashed line:
Figure 14B), the shared state of this unique character between
anhimids and dromornithids appears to be an important link evi­
dencing their close evolutionary relationship.
Prior to their formal description, several anseriform fossils were
historically reported on and described as anhimid-like, especially
with reference to their non-spatulate (‘fowl-like’) bill shape, and
contextually cited as pivotal in the understanding of anseriform
evolution (e.g. Houde 1996; Ericson 1997; Boles 1999; Olson 1999;
Mayr, 2022a). Of these, the recently described fossil anseriform
family, Anachronornithidae, is represented by Anachronornis anhi­
mops, of the latest Paleocene Willwood Formation, Wyoming,
which possesses several synapomorphies characteristic of Anseres
HISTORICAL BIOLOGY 1135
and anhimids, and was resolved ambiguously among Anseriformes
in phylogenetic analyses (Houde et al. 2023; also see Mayr et al.
2023). A taxonomically unassigned, fossil from the early-middle
Eocene Green River Formation, Wyoming, was considered more
closely related to Anseres than anhimids. Concomitantly described
fossils from the early Eocene London Clay Formation, England,
including Danielsavis nazensis, were collectively considered more
closely aligned with the Anhimae than Anseres (Houde et al. 2023;
also see Mayr 2022a). Mayr et al. (2023) recently refined attribution
of material to Danielsavis nazensis and demonstrated that many
shared characteristics with anseriforms were homoplasious, while
a relatively greater number of morphological aspects aligned with
galliforms, even considering that some may be plesiomorphic for
the group. Part of this included the recognition that the presence of
several characters which were used to exclude this taxon from
galliform affinities may in fact be plesiomorphic for Galloanserae
or Neognathae, rather than apomorphies exclusively relating to
anseriforms. It was therefore considered as either the most early
diverging anseriform, outside of the crown clade, or a stem group
galliform with some anseriform characteristics, warranting family-
level distinction (Danielsavidae, see Mayr et al. 2023). These fossils
significantly preserve an important combination of characters for
understanding the evolutionary trajectories of Anseriformes and
Galliformes, which additionally extends to interpreting the rela­
tionship between Anhimae and Anseres, and character-state polar­
ity. In particular, Ericson (1997, p. 477) specifically noted the
presence of a ‘laterally located coronoid process’ on the mandible
of the Wyoming fossils. This observation was confirmed for
Anachronornis anhimops and also reported for the unnamed
Green River Formation taxon by Houde et al. (2023) and appears
representative of the paracoronoid process (see ‘Mandible’), which
is characteristic of Anseres (processus paracoronoideus, see Weber
and Hesse 1995; Weber 1996). The relatively less laterally produced,
analogous structure on the lateral mandible of Danielsavis nazensis
is more similar to that in megapodiids (Mayr et al. 2023), particu­
larly, a well-developed crista paracoronoidea rostralis that is char­
acteristic of this clade (Weber 1996). The presence of the former
process in ‘screamer-like’ early birds has been used in support of the
hypothesis that the modern anhimid skull is derived with respect to
an ancestral condition that is typical of Anseres (including presby­
ornithids, Ericson 1997). However, current fossil evidence regard­
ing the existence of superficially similar, but non-homologous
structures in nearly contemporaneous fossil galloanserans, is in
conflict with this proposed evolutionary trajectory. Furthermore,
dromornithids, like the closely allied anhimids, and also galliform
birds, also possess paracoronoid crests (crista paracoronoidea ros­
tralis and crista paracoronoidea caudalis) rather than the singular
process of Anseres. To be consistent with the aforementioned
hypothesis, it is most parsimonious to assume that the dromor­
nithid-anhimid condition was present in their most recent common
ancestor, or alternatively, that both lineages independently evolved
this following their divergence. The large dromornithid foot cast of
(at least) Eocene age from Redbanks Plains Formation, Queensland,
temporally constrains both these scenarios (Vickers-Rich and
Molnar 1996).
More generally, the pattern of evolution whereby the anhimid
skull is derived, is also considered less parsimonious or improbable
and unsupported by our phylogenetic analyses (see also, Livezey
1997; Worthy et al. 2017b; Zelenkov and Stidham 2018; Houde et al.
2023). In consideration of the close relationship between presbyor­
nithids and anatids found here and in cladistic analyses (Ericson
1997; Livezey 1997), their shared similar complex jaw and lingual
morphology (see Olson and Feduccia 1980), and the ‘primitive’
presbyornithid postcranial osteology (see also De Pietri et al.
1136 P. L. MCINERNEY ET AL.
2016), it has been suggested that, rather than evidencing the basal
position of Presbyornithidae in Anseriformes, the unique suite of
morphological characters in this family may represent extreme
specialisation (Mayr 2022a) or the retention of plesiomorphic post­
cranial characters with respect to Neognathae. The latter is perhaps
evidenced by postcranial synapomorphies between presbyornithids
and some fossils closely associated with anatids (Romainvillidae,
e.g. Mayr 2008; De Pietri et al. 2016; Zelenkov 2018). In this case,
the anserean-type adductor complex would not need to be plesio­
morphic for all anseriforms and could have evolved from an anhi­
mid- and/or galliform-like condition, as supported by character
transitions across the phylogenetic trees (shown in Figure 12).
This is further evidenced by poor-specialisation for filter feeding
in Presbyornis pervetus, despite the indication of lamellae (Ericson
2000; Stidham 2001), which better supports Presbyornithidae as
a possessing a transitional state to filter-feeding Anseres, retaining
some plesiomorphic galliform-like characters, rather than being
itself plesiomorphic to Anseriformes (Zelenkov and Stidham
2018; Houde et al. 2023). In consideration of skeletal characters in
the early Wyoming anseriform fossils (above) that distinguished
them from presbyornithids, and clearly represented dissimilar ecol­
ogies, Houde et al. (2023) concluded that any shared character
states must be plesiomorphies of Anseriformes. Furthermore,
these authors draw comparisons in jaw apparatus morphology
between Anachronornis anhimops, Presbyornis pervetus, and
Conflicto antarcticus to suggest that the spatulate bill form was
gradually evolutionarily derived from ancestors with more galli­
form-like jaws, consistent with our hypothesis and phylogenetic
analyses. Better understanding of the phylogenetic polarities of
morphological characters important in the early evolution of
Anseriformes, and the exact relationship of these fossil clades
among extant counterparts, awaits a more focused investigation
(see also Field et al. 2020: SI).
In addition to those discussed in the section ‘Dromornithids
within Galloanserae’, other compelling links between the dromor­
nithids and anhimids are the absence of the fenestra caudalis
mandibulae, and the apparent absence of the foramen temporale
venosum. While pterygoids identified for G. newtoni are fragmen­
tary, we find it pertinent here to note that the facies articularis
pterygopalatina on the pterygoids of other dromornithids, appears
comparable to that of anhimids as described by Zusi and Livezey
(2006). They seemingly lack the more complex articulation
described for Anseres, e.g. the rostral development of the processus
palatinus du ptérygoïde (sensu Davids 1952b). Instead, the surface
is a concave, oblong facet, aligned nearly dorsoventrally along its
main axis, as it is in species of Anhima and Chauna (see Zusi and
Livezey 2006, p. 160). Unlike anhimids, however, the facies articu­
laris pterygoidea is positioned in the rostral half of the pterygoid,
and better compared to the condition in waterfowl (see Ericson
1997). Within anseriforms, the superficial shape of the mandible,
and the convex dorsal profile of the rostrum are additional simila­
rities that dromornithids share with anhimids and not Anseres
(originally noted by Murray and Vickers-Rich 2004, p. 149, 158).
Furthermore, in dromornithids and anhimids, the processus jugalis
of the maxillare is located dorsal and slightly lateral of the palatines,
which extend further rostral of the processus jugalis (Figure 9D–F).
This contrasts with the state in Anseres in which the palatines are
more dorsal of the processus jugalis and both converge rostrally.
Conversely, the presence of a prominentia submeatica is better
compared with several anatids, and Anseranas semipalmata
(although in form it is projected, rather than bulge-like), while
a lack of processus submeaticus contrasts with the anhimid condi­
tion (see ‘Quadrate’; Elzanowski and Stidham 2010; Elzanowski and
Boles 2012).
Despite concluding basal galliform similarities from the dro­
mornithid brain and associated nerves, the results of Handley and
Worthy (2021) could alternatively be interpreted as supporting
a basal- or sister-anseriform relationship, as is consistent with our
interpretation of the skull osteology and phylogenetic analyses. The
features assessed are variable in their likeness to those of the two
anseriforms and two galliforms included in the study, showing the
dromornithid brain to have a hybrid character set reflective of their
external skull morphology. Characters shared with the galliforms
and basal anseriforms, Anhima cornuta and Anseranas semipal­
mata, include the separation of the glossopharyngeal and vagus
nerves, the surface area ratios for the optic lobes, and the rostral
location of the wulst structures on the dorsal endocast. Several
characteristics were limited to dromornithids and Gallus gallus
(e.g. the transmission of the maxillomandibular branch of the
trigeminal ganglia through the cranium and the log length shape
ratio for the caudal telencephalon module), although these are likely
inconsequential phylogenetic indicators given Gallus gallus is
a derived phasianid (in the context of all Galloanserae). Only
morphological attributes which are shared between dromornithids
and Leipoa ocellata (Megapodiidae, a basal galliform, i.e. the surface
area ratio of the cerebellum module) could support galliform affi­
nities for the dromornithids, yet these are limited.
Considering both the numerous shared characteristics between
dromornithids and anhimids, and the discussed morphological
transformations towards the evolution of early Anseres, we inter­
pret the evolutionary placement of the Dromornithidae, within the
Anseriformes, as likely sister to the Anhimidae, or alternatively an
independent lineage branching more basal to anhimids and Anseres
or between anhimids and anseranatids. The former-most hypoth­
esis is resolved in both parsimony and Bayesian analyses and we
thus, advocate for the inclusion of Dromornithidae along with
Anhimidae, within the Suborder Anhimae (as per Nguyen et al.
2010). Both lineages then would together represent basal anseri­
forms in the gradual transition from galliform-like birds to those of
more anatoid character. They may also have relatively close affi­
nities to early anseriforms from Wyoming, particularly in consid­
eration of their mosaic, apparently somewhat relatively derived
morphology, seemingly intermediate between Anhimae and
Anseres (see Houde et al. 2023). Whether the postcranial morphol­
ogy of the dromornithids supports these inferences, is beyond the
scope of our study, although Murray and Vickers-Rich (2004)
found postcranial evidence supporting anseriform affinities.
Palaeogeographical considerations
The long fossil record of dromornithids show the endemic
Australian family obtained large size by the Eocene and essentially
all its characteristic morphological features by the late Oligocene
(Vickers-Rich and Molnar 1996; Worthy et al. 2016a).
Gastornithids are only known from the Paleocene to middle
Eocene of Europe, and the early Eocene of North America, and
Asia (for review, see Mayr 2022a). This discord in the localities of
the two groups has been proposed to be a limitation to the dro­
mornithid-gastornithid clade hypothesis. Mayr (2022a, p. 59) stated
no current palaeogeographical reconstructions show potential dis­
persal routes in the early Cenozoic for large flightless birds between
Eurasia and Australia. Regardless of our findings, this cannot be
used to rule out a close relationship between the two families as
there is currently no reason to assume gastornithids and dromor­
nithids did not share a volant common ancestor and evolve flight­
lessness, large body size, and concomitant morphological
adaptations independently. Given that the fossil record and several
molecular-based phylogenetic approaches support a pre-KPg

divergence estimate between Anseriformes and Galliformes (van
Tuinen and Dyke 2004; Clarke et al. 2005; van Tuinen et al. 2006;
Brown et al. 2008; Stein et al. 2015; Worthy et al. 2017b; Field et al.
2020; reviewed by Mayr 2022a), it is possible that the last common
ancestor between the dromornithids and other Galloanserae
diverged within the Upper Cretaceous. Additionally, fossil evidence
not yet discovered, may provide evidence linking or further separ­
ating the two lineages geographically.
Regarding anseriforms, an apparently overrepresentation of
basal lineages in Australasian faunas supports a Southern
Hemisphere origin for the radiation (Worthy et al., 2023). This
includes several extant crown group anseriform lineages such as
Anseranas semipalmata (Anseranatidae) in Australia, and species of
Chauna and Anhima (Anhimidae) in South America. Fossil anseri­
forms are comparatively widespread with an extensive record cov­
ering both the southern and northern Hemispheres (for a review,
see Mayr 2022a, p. 59–68; also Houde et al. 2023). Despite limited
terrestrial vertebrate fossil sites sampling the Eocene and Oligocene
of Australia (Worthy et al. 2017b), several fossil anseriforms have
been described, showing the lineage was present in Australia and
the Southern Hemisphere throughout this time. For example, pres­
byornithids Wilaru tedfordi Boles et al. 2013 (late Oligocene: De
Pietri et al. 2016) and Murgonornis archeri Worthy, De Pietri,
Scofield, and Hand, 2023 (early Eocene: Worthy et al., 2023), the
anseranatid Eoanseranas handae Worthy and Scanlon 2009 (late
Oligocene/early Miocene: Worthy and Scanlon 2009), and several
crown group anatids (Oligocene-Miocene: Worthy 2009).
Additionally, a putative fossil quadrate of an anhimid-like taxon
was described from the early Eocene Tingamarra Local Fauna by
Elzanowski and Boles (2012).
In a similar manner to the anseriforms, basal extant galliforms
(the Megapodiidae) are also restricted to the Southern Hemisphere.
However, conversely, the Southern Hemisphere fossil record for
galliforms is far less extensive, being restricted to a single fossil
megapodiid taxon Ngawupodius minya Boles and Ivison 1999,
described from the late Oligocene of Australia (Boles and Ivison
1999), and Namaortyx sperrgebietensis Mourer-Chauviré et al. 2011,
of unknown affinities from middle Eocene of Namibia, southern
Africa (Mourer-Chauviré et al. 2011). Most stem galliform lineages,
such as the Quercymegapodiidae, Paraortygidae, and
Gallinuloididae, and potentially the Danielsavidae (see Mayr et al.
2023), are known from the Northern Hemisphere (see Mayr 2022a,
2022, and references therein).
The occurrence of latest Cretaceous Asteriornis maastrichtensis
in Belgium challenges the hypothesis of a Gondwanan origin for
crown birds (Claramunt and Cracraft 2015), as phylogenetic ana­
lyses suggest a phylogenetic position as a stem galloanseran, prox­
imal to the divergence between Galliformes and Anseriformes, or
a basal galliform (Field et al. 2020; see Figure 12). However, the
early Paleocene anseriform Conflicto antarcticus, shows definitive
anseriforms had already diversified and dispersed into the Southern
Hemisphere shortly after this time (Tambussi et al. 2019). This
presence of anseriforms in the Southern Hemisphere by the early
Paleocene, and proposed austral origin of the anseriforms, contrast­
ing that of galliforms (Mayr 2022a) and the gastornithids, provide
appropriate support for our hypothesised anseriform affinities of
the dromornithids.
Ecomorphology and niche adaptations
The kinetic abilities of the dromornithid skull, specifically of
Dromornis species, have been explored previously (see Murray
and Vickers-Rich 2004). Here we further elaborate on the ecology,
cranial kinesis, and the functional application of dromornithid skull
HISTORICAL BIOLOGY 1137
morphology, although, given limitations, we recommend dedicated
kinetic and biomechanical analyses to appropriately test that which
is discussed below.
Bite force and jaw opening
Recent studies on dromornithids have concluded that they had
weak, or not especially powerful bite force capabilities due to the
inferred reduction in surface area on the lateral cranium for attach­
ment of the external adductor muscles (Murray and Vickers-Rich
2004; Worthy et al. 2016a). Furthermore, a negative relationship
between bite force and body mass identified for birds may implicate
proportionally low bite forces for dromornithids in consideration of
their large size (Dickinson et al. 2022). However, assessment of
biomechanical attributes associated with dromornithid jaws is mul­
tifaceted, and the interactions of all elements contributing to the
potential force exerted throughout jaw closure is complex, requir­
ing consideration of numerous factors. All galliforms and anseri­
forms share morphological adaptations which support powerful
bite force, i.e. the lack of the ligamentum jugomandibulare laterale,
and the variable rostral positioning of the processus postorbitalis
and/or AZO complex (Dzerzhinsky 1974, 1980, 1982; Dzerzhinsky
and Grintsevichene 2002), both explored in greater depth below.
Considering the relatively definitive galloanseran affinities for dro­
mornithids, specifically with anseriforms, and Dollo’s law, dromor­
nithids are phylogenetically constrained with respect to the
limitations imposed by inherited morphological adaptations, char­
acteristic of this group; some of which, such as the loss of the
aforementioned ligament, can be confidently assumed to have
existed in Dromornithidae but cannot be observed by direct exam­
ination of the fossils alone.
In most birds, the relative non-elasticity of the ligamentum
jugomandibulare laterale plays a role in locking the jaw joint and
is intrinsically connected to the condylus mandibularis caudalis on
pars mandibularis of the quadrate and the corresponding caudal
cotyla on the mandible, that acts as a pivot for abduction. The loss
of both the ligament and the condyle (and mandibular cotyla) in
Galloanserae is therefore equivalently linked, to allow freedom of
longitudinal sliding between the quadrate and mandible, with ros­
tral movement of the mandible limited by the caudally attaching
ligamentum occipitomandibulare (Dzerzhinsky 1974, 1980, 1982,
1995; Dzerzhinsky and Grintsevichene 2002). The loss of the lateral
jugomandibular ligament removes constraints on the functionality
of the pterygoid muscles, allowing them to effectively retract the
bony palate and quadrate with respect to the cranium, and adduct
the rostrum without causing automatic, responsive movement of
the mandible (Dzerzhinsky 1974, 1995, and references therein).
Concomitantly, adduction of the mandible is primarily and more
efficiently performed by the external adductor muscles and the
superficial pseudotemporal muscle (Dzerzhinsky 1974, 1995;
Dzerzhinsky and Grintsevichene 2002). This then has major func­
tional implications for the jaw closing mechanisms in Galloanserae,
and by extension, inferences of dromornithid jaw biomechanics,
due to the resultant relative independence of lower and upper jaw
movement, and consequential specialisation of osteo-myological
structures contributing to efficiency in their respective roles
(Dzerzhinsky 1980, p. 155–156).
In galloanserans, powerful jaw movement is especially afforded
by the rostral positioning of the processus postorbitalis and/or AZO
complex, and associated adductor musculature (especially m. AME
profundus, partes zygomatica et superficialis) from the point of jaw
rotation (i.e. the area of articulation with the quadrate), resulting in
a longer lever arm for efficient transmission of forces during lower
jaw adduction (Dzerzhinsky 1974). This increased space is also
1138 P. L. MCINERNEY ET AL.
associated with greater area for muscle attachment,
particularly m. AME superficialis and m. adductor mandibulae
posterior, interrelated with other shared biomechanical features
(as discussed by Dzerzhinsky 1974, 1980, 1982; see also Zusi and
Livezey 2000). Dromornithids have jaw adductor musculature ema­
nating in association with a rostrally developed processus postorbi­
talis and AZO complex, especially comparable to anhimids (see
above), and while their cranium is rostrocaudally compressed, this
is primarily with relation to the supraorbital area, and does not
appear to limit the available caudal attachment area for mandibular
external adductor muscles. Neither does it reduce the distance
between the quadrate and the orbit. This would likely somewhat
similarly achieve the functional benefit of increased bite force
produced from the aforementioned rostral positioning of the
adductor musculature that occurs in other galloanserans (see
Dzerzhinsky 1974). Accordingly, several structures corresponding
to the origin of the m. AME superficialis on the cranium and
quadrate of dromornithids (crista m. AME articularis, tuberculum
subcapitulare) are well-developed. The site of origin of m. adductor
mandibulae posterior on the quadrate is similarly well formed,
interrelated with a vast insertion site on the lateral caudal mandible,
and evidencing strong retraction power for adducting both the
mandible and the rostrum (Goodman and Fisher 1962;
Dzerzhinsky 1974). In close association with the origin
of m. AME profundus, pars coronoideus (Murray and Vickers-
Rich 2004, p. 240), the obviously excavated fossa, associated crests
and tuberculum in the area of origin for m. pseudotemporalis
superficialis in dromornithids marks an extensive and likely hyper­
trophied superficial pseudotemporalis muscle (Murray and
Vickers-Rich 2004) which also participates in the adduction of the
lower jaw (Dzerzhinsky 1974).
Furthermore, the large, dorsoventrally deep and wing-like pala­
tines in dromornithids provide an expansive attachment surface for
both dorsal and ventral bellies associated with m. pterygoideus
muscles (e.g. Dzerzhinsky 1982; Vanden Berge and Zweers 1993:
annot. 21; Carril et al. 2015). This has been considered one of
several mechanisms for increasing bite force (see Witmer and
Rose 1991, p. 100) as they provide a surface for powerful muscle
leverage (Murray and Vickers-Rich 2004, p. 244). Their insertion
area on the mandible also appears well developed in dromornithids
(see ‘Rostrum’; Murray and Vickers-Rich 2004, p. 241, fig. 189), and
so, in conjunction with the anchoring ligamentum occipitomandi­
bulare, contraction of these muscles from their fixed point on the
mandible provides a large force of retraction to caudoventrally pull
the quadrate and bony palate, and efficiently adduct the rostrum,
which is further facilitated by the mobility of the craniorostral hinge
in its relatively caudal position (Dzerzhinsky 1995; see also Murray
and Vickers-Rich 2004: fig. 190). The deep pseudotemporal muscle
(m. pseudotemporalis profundus) also acts to retract the rostrum,
with corresponding areas of origin on the rostrolateral processus
orbitalis (crista orbitalis) of the quadrate, and its insertion the
medial mandible, whereby the former, as a lever arm, is pulled
caudoventrally to lower the rostrum (Dzerzhinsky 1974, 1980;
Bühler 1980). This muscle also acts to adduct the mandible
(Goodman and Fisher 1962; Zweers 1974; Buhler, 1980). The sur­
face of origin is especially marked in dromornithid quadrates, when
preserved, and infers strong emanating musculature. Similarly, in
addition to the adduction of the lower jaw (see above),
the m. adductor mandibulae posterior, which originates on the
lateral quadrate and extends to the mandible, also participates in
active retraction of the rostrum in an arc-like path (rather than
purely ventrally, see Dzerzhinsky 1974: fig. 1A). This active low­
ering of the upper jaw thus assists in severing material between the
jaws, compared to only vertically clasping them (Dzerzhinsky 1974,
1980). Such functional adaptations, coupled with specialisations for
adducting the lower jaw (above), would have resulted in formidable
closing power of the jaws, although not as proportionally extreme as
those that may be inferred for gastornithids. Evidently, the lack of
the lateral jugomandibular ligament, and resultant muscle speciali­
sation with respect to function in the isolated movement of the
upper and lower jaws (Dzerzhinsky 1980), likely translated to
positively affecting potential bite force capabilities in
dromornithids.
The consumption of fixed, resistant foods which place greater
and differential stress loads on the upper or lower jaws, are related
to the significant evolution of independent jaw mobility in
Galloanserae (Dzerzhinsky and Grintsevichene 2002). In a bird
that possesses a ligamentum jugomandibulare laterale, such as non-
Galloanserae, upwards pressure exerted upon the upper bill would
automatically impact the jaw joint and depress the lower bill; this is
to be avoided during activities such as digging, as opening the beak
would result in an influx of sediment or water into the mouth
(Dzerzhinsky 1974). Isolation of the movement of each jaw relative
to the other can therefore be considered advantageous as it restricts
passive protraction, and is especially beneficial when feeding on
attached, coarse vegetation, or in association with mediums that
may exert resistance, such as digging through wet and dry sediment
(Dzerzhinsky 1974, 1980, 1982). A morphological feature of note
with regards to this, and jaw closing, are the mandibular cotylae in
dromornithids which are oblique with respect to the rostrocaudal
plane (as especially observed in Ilbandornis sp. NTM P2774–2 and
Dromornis planei NTM P9464–112, and perhaps in Genyornis new­
toni SAMA P59517), which can be compared with the orientation
of those in Penelope jacucaca (Cracidae). The angle of the cotylae in
this taxon, offset from the sagittal plane of the mandible, have been
interpreted as contributing to efficiently locking the quadrate-
mandible joint at the point of maximum retraction (Dzerzhinsky
1980, and references therein). The action of tearing off fixed vegeta­
tion that is clasped between these jaws creates additional forces of
protraction, and so this joint locking is hypothesised as
a mechanism of preventing passive opening of the jaws that is
effective even with a relatively weak jaw apparatus, especially
when additionally contributed to by dorsoventral compression
placed under the taut caudally attaching ligamentum occipitoman­
dibulare (Dzerzhinsky 1980). In conjunction with the oblique align­
ment of the cotylae, the transverse, lateromedial movement of the
quadrate during contraction of the m. pseudotemporalis profundus
and m. adductor mandibulae posterior, in adduction, provides
reinforcement to this lock (Dzerzhinsky 1974). While dromor­
nithids do not appear to have such a distinct mandibular crista
intercotylaris as P. jacucaca, the inclination of the cotylae may
indicate some similar adaptations for efficiently feeding on attached
vegetation while mechanically minimising the use of musculature
that would resist adduction. As P. jacucaca is a representative of
a basal galliform with relatively weak jaw musculature, the advan­
tage in foraging enabled by this structural specialisation has been
considered an archaic adaptation, important in the successful radia­
tion of Galloanserae (Dzerzhinsky 1980), and as such may be
symplesiomorphic with respect to Dromornithidae.
Analysis of the skull morphology of dromornithids also allows
some interpretations of other biomechanical features of the jaw
apparatus, particularly related to finer adjustment in upper and
lower jaw interaction. The aforementioned adaptations for
increased mobility of the quadrate are associated with several addi­
tional related adaptations in the dromornithids. The low crista
intercotylaris between the lateral and medial cotyla on the mandible
of dromornithids allows for additional rotational movement of the
quadrate upon this surface (Murray and Vickers-Rich 2004). In

conjunction with forces exerted by m. protractor pterygoidei et
quadrati, and the muscles involved in quadrate-mandible adduction
(m. pseudotemporalis profundus and m. adductor mandibulae
posterior), low intercotylar crests may have enabled the movement
of the quadrate that thus shifted the fulcrum across the mandible
surface during feeding, as is observed in species of Anas (see
Goodman and Fisher 1962; Zweers et al. 1994, p. 269; Murray and
Vickers-Rich 2004). The contraction of the m. protractor quadrati
et pterygoidei, via quadrate and pterygoid movement relative to the
cranium, is the primary driver of active protraction and elevation of
the rostrum (Bühler 1980), and is likely facilitated by the mobility of
the craniorostral hinge, however, the dorsal quadrate is further
freed for greater movement by the functional single-headedness of
the capitulum squamoso-oticum and supporting modifications to
the quadratic cotyla of the cranium in which it sits. With no
buttressing from a distinct otic capitulum, the quadrate can rotate
mediolaterally around its dorsoventral axis, rather than being
restricted to a primary rostrocaudal direction of movement (see
Dzerzhinsky 1974, 1982; Zweers 1974; Murray and Vickers-Rich
2004, p. 244; Zelenkov and Stidham 2018). Notably, movement of
the quadrate is further unhindered in dromornithids by the weak
condition of interlocking articulation points between the quadrate
and both the pterygoids and jugal arches (also see Samejima and
Otsuka 1987). In most galliforms, the pterygoid articulates at two
points on the quadrate, whereas in anseriforms (although to a lesser
extent in anhimids, see Elzanowski and Stidham 2010), cracids and
dromornithids, the simplicity of the single continuous quadrate-
pterygoid articulation and the lack of a fortifying secondary articu­
lation on the orbital process (see Dzerzhinsky 1982; Elzanowski and
Stidham 2010; Zelenkov and Stidham 2018), permits rotation of the
quadrate while maintaining contact with the pterygoid
(Dzerzhinsky 1974; Murray and Vickers-Rich 2004). Effectively,
this mobility in quadrate-pterygoid articulation would have limited
the capacity for transferral of forces to stress and distort the palatal
apparatus during adjustments of the quadrate (Zelenkov and
Stidham 2018). Less potential for deformation is also facilitated by
the very robust palatines of dromornithids, which, in conjunction
with this mobile, continuous articulation, would have been able to
effectively transmit protraction forces through rostral quadrate
movement (Zelenkov and Stidham 2018). Furthermore, dromor­
nithids lack a bowl-like fovea quadratojugalis for the articulation
with the quadratojugale and instead only retain a shallow, plate-like
articular facet. This weak articulation, in conjunction with the
robust jugal arch that would have been resistant to deformation
or lateral flexion, would have also been able to efficiently impart
rostrocaudal forces during upwards movement (protraction) of the
rostrum. This adaptation combined with the mediolateral rota­
tional mobility afforded by the ball and socket-like quadrate-
cranium joint morphology, may have thus been able to partly
compensate for the lack of flexibility in the jugal arch. These quad­
ratic adaptations are therefore vital for maintaining independent
movement of the upper bill of dromornithids, while also enabling
use of its robust and inflexible structure for other aspects of feeding.
This is in contrast to other galloanserans, especially Anseres,
wherein flexibility of the jugal arches particularly, is integral for
feeding (see Zelenkov and Stidham 2018).
Rotational mobility in jaw use is also complimented by other
aspects of the dromornithid skull. This importantly extends to the
craniorostral hinge, whereby the uniting ligamentous attachments
between the caudal rostrum and rostral cranium may have allowed
some small degree of rotation on transverse, frontal and sagittal
planes (Murray and Vickers-Rich 2004, p. 241). Given the obviously
impressed and extensive attachment of m. pterygoideus, Murray
and Vickers-Rich (2004) also suggested that independent
HISTORICAL BIOLOGY 1139
contraction of parts of this musculature may have been able to
slightly reposition the mandibles on the frontal plane, and hypothe­
sised that when supported by other musculature involved in adduc­
tion, this would allow dromornithids to control the edges of their
jaws transversely when forces were applied to resistant objects. The
deeply socketed origin of the superficial pseudotemporal muscle on
the caudal orbit of all dromornithids, near the level of, or just dorsal
to the tomial margins, indicates high levels of control over, and
resistance of, axial rotation and mediolateral displacement of the
mandible, but also maintenance of strong, stable adduction (see also
Murray and Vickers-Rich 2004, p. 244). This would be supported
and further enhanced by the aforementioned rostral positioning of
the external adductor muscles, and the active lowering of the
rostrum enabled by the m. adductor mandibulae posterior
(Dzerzhinsky 1974; Dzerzhinsky and Grintsevichene 2002). Based
on their considerable control over movement of the upper and
lower jaws independently, and proposed vectors involved in muscle
action, it has been suggested that dromornithids were well-adapted
for holding and manipulating food held in the tip of the beak, and
had relatively high static bite force in this region, as is achieved by
other goose-type anseriforms (Murray and Vickers-Rich 2004;
Olsen and Gremillet 2017; Pecsics et al. 2017). Considering these
adaptations, as well as those corresponding to a highly developed
lingual apparatus, Murray and Vickers-Rich (2004, p. 245) con­
cluded that the manipulative jaw abilities of dromornithids may
have been approaching, or comparable to, those observed in
parrots.
Dzerzhinsky (1974) noted that the extensive retroarticular pro­
cess of the caudal mandible in Galloanserae (specifically in refer­
ence to Tetrao urogallus) may be primarily related to the necessity
for a more caudal origin of lingual musculature on its lateral surface
(e.g. m. serpihyoideus and m. stylohyoideus). While a complete
retroarticular process is not known for any dromornithid, pre­
served areas indicate that it was large, dorsoventrally deep, and
extended well caudal of the fossa articularis quadratica, not unlike
other galloanserans, and certainly also accommodated lingual mus­
culature on its lateral surface. In Gallus gallus, the paraglossum is
small and relatively broad, as it is in Genyornis newtoni, which, in
the former taxon, supports a large, thick and somewhat flexible
tongue in the context of a feeding apparatus specialised for pecking
(Zweers et al. 1994). How this affected the function and manipul­
ability of the lingual apparatus in dromornithids is unknown,
however.
Because of the size and lever-like caudal extension of the retro­
articular processes, and the consequent oblique nature of insertion
of m. depressor mandibulae, contraction of this muscle produces
a strong protraction force, and efficient abduction of the lower jaw
(Dzerzhinsky 1974, 1980). In several galliforms, this morphology
has been associated with optimised forceful gaping, against soil
resistance or to overcome the friction of vegetation stuck between
the upper and lower jaws (Dzerzhinsky 1980; Zweers et al. 1994),
and has also been related to the rapid filter-feeding jaw movements
in anatids (Bühler 1980). The plausibility of the latter in dromor­
nithids is not well supported considering other aspects of their
anatomy. The development of the processus retroarticularis is gen­
erally considered an adaptation to depress the lower jaw against
resistance (Zweers et al. 1994, p. 246, 271), and the well-developed
retroarticular process in dromornithids would then imply that
forceful opening of the mandible was important in their feeding
or other aspects of their ecological niche.
Consequently, considering the aforementioned adaptations, we
do not support the hypothesis that dromornithid jaws had an
especially weak bite force, despite rostrocaudal compression of the
cranium. While this latter factor may have limited the potential
1140 P. L. MCINERNEY ET AL.
force generated in the closing of the jaws, dromornithids have
inherited adaptations such as independent rostrum-mandible
mobility that are characteristic of other galloanserans, which has
permitted associated subsequent osteological specialisation and
hyper-development of musculature involved in feeding. This likely
translated to powerful bite forces, jaw opening, and also notable
manipulative abilities in relation to feeding on plant matter.
However, despite the shared characteristics or comparisons that
can be drawn between dromornithids and other galloanserans, the
unique combination of traits observed in the dromornithid skull do
not present any close known analogue. The robust palatines and all
associated components form a single functional structure in dro­
mornithids, while another is formed by the adjacent articulations of
the quadrate and pterygoid, indicated by the elongate mandibular
condyles of the former that are longitudinally aligned with the axis
of the latter. Together with the jugal arch as a third functional unit,
dromornithids exhibit a structural condition of the palatal appara­
tus that is typical of anseriforms (Dzerzhinsky 1974, 1982; Zelenkov
2017; Zelenkov and Stidham 2018). In contrast, the dorsoventral
separation of the palatines and jugal arches where they interact with
the rest of the rostrum is unlike Anseres, nor especially comparable
to galliforms (for comparative description of the functionality of the
palate in Galloanserae, see Zelenkov and Stidham 2018: fig. 2). The
implication of this is that the functionality of the dromornithid
palatal apparatus is unique with respect to the condition of any one
galloanseran order (Dzerzhinsky 1974, 1982; Zelenkov and Stidham
2018: fig. 2). However, aside from the quadrate-quadratojugal
articulation, this distinctive character composition is in some
ways intermediate between that which is characteristic of each of
these galloanseran orders, as would be expected by a close evolu­
tionary affinity to anhimids, which also appear somewhat inter­
mediary in their morphology and functionality of the palatal
apparatus (see Dzerzhinsky 1982).
Feeding and aquatic adaptations
The typical galliform skull morphological arrangements support
feeding through digging, active tearing of attached and sometimes
coarse vegetation, or grasping of fruits and grains, compared to the
grazing, submerged dexterous manipulation of soft aquatic-based
plant matter, and filter-feeding that is typical of many anseriforms
(e.g. compare with Tetrao urogallus, see Kirikov 1944; Dzerzhinsky
1974, 1980, 1982, p. 1036–1039; Zweers 1974; Zusi and Livezey
2000). In anhimids, their structural arrangement combined with
a galliform-like bill, likely supports their consumption of coarser
foliage through grazing and aquatic macro-feeding, as well as fora­
ging in mud – although it is unknown if they filter-feed while
digging (Dzerzhinsky 1982; Naranjo 1986). Anseranas semipalmata
also feeds predominately on water-based macro-vegetation, and
filtration-feeds in mud (Davies 1963; Murray and Vickers-Rich
2004).
Dromornithids have long been known to be herbivorous (for
a review, see Murray and Vickers-Rich 2004). Genyornis newtoni is
no exception with the consistent presence of jasper, sandstone,
claystone, and quartz (Rich and van Tets 1984) gastroliths, found
associated with its skeletons, being a strong link to herbivory. These
gastroliths are small in size (most in range 4–8 mm in diameter,
with few over 12 mm) and overall mass ratio with respect to mean
body mass, suggesting only soft foods are being consumed and not
items such as twigs (Handley and Worthy 2021: SI, p. 36; see SI 6 for
additional data of G. newtoni gizzard stones). This is consistent with
the dromornithid diet as proposed by Murray and Vickers-Rich
(2004, p. 295–322), who extensively explored known dromornithid
habitat and available sources of food, and Handley and Worthy
(2021), who found the well-developed somatosensory and sensor­
imotor capabilities of dromornithids to be indicative of a selective
soft browse diet dominated by leaves, new growth, and fruit. The
morphology of the skull of Genyornis newtoni as described above,
further supports herbivorous niche occupation, in a manner not
unlike that of geese or the Anhimidae. The lack of several filter-
feeding adaptations that are contrastingly present in Anseres, nota­
bly the thin, flexible jugal arch, and the modified attachment of the
jugal arch and palatines to the rostrum (see Zelenkov and Stidham
2018), and the more similar nature of these regions to galliforms
and anhimids, supports an absence of specialised filter feeding in
the diet of dromornithids.
Relationships have been identified linking relatively rostrocaud­
ally short and dorsoventrally high skulls and rostrocaudally short
palatines with grazing and omnivorous diets, in contrast to the
more elongate, gracile skulls of filter feeding and piscivorous anseri­
forms (Goodman and Fisher 1962; Pecsics et al. 2017). In dromor­
nithids, the rostrocaudal compression and rostral foreshortening of
the cranium may then be associated with a capacity for grazing-type
behaviour and niche occupation. As in species of Dromornis,
Genyornis newtoni also possesses shearing planes at the caudal
region of the upper and lower bill, created by overlapping of the
lateral plate on each side of the rostrum and the corresponding
tomial shelf of the lower jaws in articulation. The nearly vertical
oriented planes of the interlocking jaws in this region may con­
tribute to the efficiency of clamping and tearing fixed vegetation (as
described for some galliforms, see above and Dzerzhinsky 1980),
and is additionally considered to be functionally important for
shearing of small and resistant objects (see Murray and Vickers-
Rich 2004). This caudal tomium is differentiated from the more
rostral crista tomialis in dromornithids, although the surface of the
latter is comparatively flat in G. newtoni and contrasts with the
more angular cristae in species of Dromornis, which probably
relates to different feeding ecologies (see Murray and Vickers-
Rich 2004; Worthy et al. 2016a). Dromornithids also appear to
have well-developed neck musculature, to support their large (and
heavy) heads. Their expansive, high attachment over the caudal
cranium that reaches the dorsal apex of the calvaria, more similar
to Anseranas semipalmata than that of anhimids, would have direc­
ted forces away from the occipito-atlas joint to be distributed
further down the neck, likely in association with pulling or tugging
at resistive vegetation, such as when browsing (see Murray and
Vickers-Rich 2004). The difference in dorsoventral location of the
foramen magnum between dromornithids (i.e. the dorsoventral
middle of the skull in Genyornis newtoni and Dromornis planei,
for example, compared to the more ventral position in D. murrayi)
suggests variability in postural holding of the head relative to the
neck, or in exact feeding behaviour between species.
The presence of the synovial craniorostral hinge in dromor­
nithids likely permitted greater upwards movement of the rostrum,
ultimately increasing the available gape width, and allowing for
food of larger size to be consumed (see Bock 1960a). According to
Murray and Vickers-Rich (2004, p. 246), the rostrum of species of
Dromornis was capable of much dorsoventral movement at the
craniorostral hinge, contributed to by the proportionally dorsoven­
trally thin articular surface, and possibly some slight axial rotation.
This is a key benefit of this hinge type for parrots in which it likely
evolved in association with feeding on hard and large items (Tokita
2003). The desmognathous, and inflexible upper jaw would have
structurally reinforced and strengthened the rostrum (Dzerzhinsky
1980). The enclosed bony palate, and interpretations on the tongue
mobility of dromornithids that may have mirrored parrot-like
manipulative abilities (see above, Murray and Vickers-Rich 2004,
p. 245), allowed some capacity to crush food with their tongue on

the palatal roof (as in some passerines, see Bock 1960a). It is
unknown whether the dromornithids may have been better able
to maintain the position of their eyes relative to their food, although
the rotational movement of the rostrum about the craniorostral
hinge would likely allow the cranium to maintain a constant food –
eye position, as is important for many birds (see Bock 1960a).
Many of the features which distinguish anseriforms from galli­
forms, are considered derived transformations intrinsically linked
to the transition to aquatic habitats (Dzerzhinsky 1982) early in the
evolution of anseriforms; given aquatic habitat preference for anhi­
mids (e.g. Naranjo 1986) and the aquatic-adapted morphology of
other fossil anseriforms (see Mayr 2022a). These characteristic
anseriform adaptations which are also present in dromornithids,
are more parsimoniously symplesiomorphic and evolved prior to
the divergence of the dromornithid lineage, rather than conver­
gently. The closure of the osseous meatus acusticus externus ven­
trorostrally on the dromornithid skull (in association with annulus
tympanicus) may additionally be associated with aquatic adapta­
tions. This is an inferred adaptation for isolating the ear from the
quadrate while the head is submerged in water, whereby movement
of this bone could periodically deform the ear cavity, making it
difficult to protect the ear from water ingress and create acoustic
interference (Dzerzhinsky 1982, p. 1039). In G. newtoni, the osseous
meatus acusticus externus is enclosed to an extreme degree and in
turn well separated from the quadrate, compared to other
dromornithids.
This then suggests two options for dromornithid habitat pre­
ference. First, that dromornithids were adapted to aquatic-
associated habitats (e.g. edges of wetlands, large lakes, and flood
plains, as discussed by Murray and Vickers-Rich 2004, p. 2, 157,
167; initially proposed by Stirling and Zietz 1900) and had a varied
grazing folivorous and frugivorous diet with relation to foraging in
the surrounding environment, not dissimilar to anhimids (see
above). This is additionally supported by the common discovery
of dromornithid fossils around former bodies of water, such as Lake
Callabonna and Alcoota (see Worthy and Yates 2017), and the
inferred tethering of G. newtoni to channel systems and floodplains
(Stirling and Zietz 1900; Smith 2009). The second hypothesis is that
dromornithids were constrained by or retained ancestral adapta­
tions to the sub-aquatic environments yet did not necessarily utilise
these in association with a sub-aquatic niche or environment.
Desmognathy in the rostrum, and the caudal position of the
choana nasalis ossea in Genyornis newtoni, may have additionally
offered protection against water ingress (achieved by fascia in
Chauna torquata, see Dzerzhinsky 1982). Furthermore, the flat
tomial surface of the rostrum in G. newtoni with associated rough
osseous material similar to that which supports rhamphothecal
lamellae rostri in some anseriforms, is consistent with the idea
that dromornithids may have possessed very rudimentary lamellae
on their tomia, most comparable to anhimids, to facilitate the
successful seizing and tearing of slippery attached aquatic vegeta­
tion (see Murray and Vickers-Rich 2004). Murray and Vickers-Rich
(2004) additionally suggested that the sulcus paratomialis of dro­
mornithids may have also allowed expelling of excess water in this
method of feeding. Many anseriforms also have adaptations for
straining water from the between their jaws in food acquisition
(Dzerzhinsky 1982). Alternatively, simple, non-extensive rham­
phothecal lamellae could be used in efficiently gripping and tearing
other types of fixed plant matter with the jaws and considering the
more blade-like rostral tomia in species of Dromornis, may have
supported diversification with regards to other dietary preferences,
including an inferred head-level browsing niche in this lineage (see
Murray and Vickers-Rich 2004, p. 192). It is also worth noting here
that we do not find convincing evidence for supraorbital ‘nasal salt
HISTORICAL BIOLOGY 1141
glands’ (fossa glandulae nasalis) in any dromornithid (see
‘Cranium’ and ‘Rostrum’), which Murray and Vickers-Rich (2004)
suggested were adaptations to more arid environmental conditions,
and against habitation of freshwater or rainforest regions.
The aquatically adapted skull may seem contradictory to pur­
ported terrestrial locomotion adaptations in the hindlimbs of
G. newtoni and other dromornithids. The hind limbs (including
the blunt, hoof-like, but mobile toes and inferred elastic recovery
system of the ankle joint), as described by Murray and Vickers-Rich
(2004, p. 221–225), were hypothesised to be adaptations for terres­
trial locomotion in dromornithids. In the context of the hypothesis
that dromornithids were at some point adapted to exploiting aqua­
tic-associated habitats, then such unusual morphology of the hind
limb may be a result of large birds which rely on aquatic systems
being forced to adapt to hard, dry ground. This would occur as they
become less confined to water and travelled increasingly long dis­
tances between appropriate habitats due to dry environmental
conditions, ephemeral lake systems, and increasing aridity in
Australia since the middle Miocene (for reviews of environmental
changes and their influence on the dromornithids, see Murray and
Vickers-Rich 2004; Angst and Buffetaut 2017). In this way, forced
range expansion may be related to diversification of their diet to
include soft browse not exclusively associated with near-freshwater
regions. Indeed, the variable hindlimb morphology of dromor­
nithids (especially late Miocene forms) demonstrate adaptations
to different modes along the cursorial-graviportal spectrum of
locomotion and have been paired with contrasting capacity to
expand their range, as well as potential for nomadic or migratory
lifestyles (Murray and Vickers-Rich 2004; Worthy and Yates 2015).
Variation of rostrum shape within the Dromornithidae
Across dromornithids, beak shape varies considerably when com­
paring G. newtoni to species of Dromornis and suggests divergent
evolutionary history and possibly niche adaptation. For example,
the tomial margin of G. newtoni rostral of the palatines is bordered
by a wide, flattened tomial surface more effective for crushing and
gripping than the thinner, grooved margin of Dromornis (see
Murray and Vickers-Rich 2004, p. 245–246). Additionally, the gen­
eral shape of the bill varies, with G. newtoni lacking the extreme
dorsoventral height and dorsal curvature of the rostrum noted for
species of Dromornis (see ‘Rostrum’ and Murray and Megirian
1998); such height may have increased resistance to reaction forces
from crushing seeds (as was identified in Darwin’s finches, see
Herrel et al. 2005). The shorter rostrocaudal rostrum length relative
to cranium length for G. newtoni, likely instead facilitated greater
dexterity and proportionally increased power of the jaw muscula­
ture (see above, Goodman and Fisher 1962, p. 169), and may be
associated with the wide shape and flattened tomial surface of the
beak of Genyornis newtoni, absent in species of Dromornis.
Goodman and Fisher (1962, p. 170) noted that force is applied
more effectively in birds with narrow bill tips (species of
Dromornis) compared to those with wide bill tips (G. newtoni).
There are currently no known rostra associated with species of
Ilbandornis or Barawertornis, although their relatively slender and
dorsoventrally narrow mandibles (see Worthy et al. 2016a), com­
pared to those of Dromornis species, may suggest that their upper
jaws were also not so dorsoventrally deep.
The synovial craniorostral hinge of dromornithids was sup­
ported by two lobes which would act to stabilise the joint and
restrict excessive movement at high force loads (see ‘Craniorostral
hinge’). As the lobes are proportionally larger in species of
Dromornis than in G. newtoni, less movement in the former
would have likely been possible, potentially driven by the large
1142 P. L. MCINERNEY ET AL.
size of the rostrum requiring increased stability. Although the
incorporation of the lacrimal into the rostrum would have
increased robusticity of the caudal rostrum itself, other than pro­
viding some lateral strengthening and stabilisation at the hinge, this
was likely limited since the dorsal lacrimal did not appear to have
caudally crossed the hinge in dromornithids.
Beak shape evolution in anseriforms has both facilitated and
been driven by expansion into new ecological niches (see Olsen
and Gremillet 2017) and this may have played a role in this varia­
tion. Olsen and Gremillet (2017) found that an increase in culmen
angle led to an increase in stress resistance, and in finches, bite force
has previously been positively correlated with depth of the beak (see
Herrel et al. 2005); the latter a factor likely contributing to beak
shape evolution in parrots. Such a driver may have been more
dominant in the Dromornis group resulting in such disparate ros­
trum shapes relative to the shape of G. newtoni. Alternatively, the
more goose-like shape of the rostrum of G. newtoni may be
a derived morphology resulting from a transition towards
a similar niche occupation as some geese. This could have been
driven by millions of years of adaptation to changing environmen­
tal conditions and food resources, linked to the expansion of scler­
ophyll and grassland habitats across Australia, which Murray and
Vickers-Rich (2004, p. 295–322) consider an important factor in
dromornithid evolution and extinction.
We thus propose two alternative hypotheses for such variation
in beak shape among dromornithids. Genyornis newtoni displays
a derived morphology driven by adaptations to Australia’s
Quaternary environment. Alternatively, G. newtoni may have
been descended from close relatives of Ilbandornis species and
retained an Ilbandornis-type bill. The difference between the
Genyornis- and Dromornis- type jaws then may have been driven
by divergent niche occupation, allowing geographic and temporal
overlap of species of Ilbandornis and Dromornis, of which two and
one species, respectively, were sympatric in Alcoota and perhaps
Bullock Creek local faunas. The two alternative hypotheses for
adaptation to aquatic environments (see ‘Feeding and aquatic adap­
tations’), may both be applicable but to different dromornithid
lineages, that is, one lineage retaining a water-based feeding habitat
and the other, only constrained by historical aquatic adaptations.
Having a large, heavy skull, as is especially characteristic of
Dromornis species, is a large energy investment (estimated to have
been between 5–8 kg in D. stirtoni, see Murray and Vickers-Rich
2004), and thus must have been integral to evolutionary fitness in
this group of birds, such as feeding on large and/or resistant items.
It is possible that the need for such large head size may have also
been a factor that necessitated the evolution of correspondingly
large body, since large body size alone is not a sufficient driver of
increased head size (as exemplified by the skull to body proportions
of flightless palaeognathous taxa). Irrespective of what drove large
body and head size in dromornithids, compensatory adaptations
for additional structural robusticity are required for supporting
such a large structure, and to also enable functional capabilities at
such extreme dimensions. As such, the large skull size of Dromornis
species may itself predominately explain their mediolaterally nar­
row and dorsoventrally deep rostrum that has capacity for reducing
vertical compression, torsion, and shear, to maintain rigidity in
such a large bony structure which is only supported by a single
suspension point (see Murray and Vickers-Rich 2004, p. 235).
Other functional roles which may have significantly contributed
to the variation includes sexual selection, and functions associated
with vocalisation, preening, and thermoregulation, which can lead
to trade-offs with feeding performance (e.g. Clayton et al. 2005;
Bright et al. 2016; Olsen and Gremillet 2017, and references
therein). As there is currently no rostrum attributed to any species
of Ilbandornis, it is difficult to deduce the exact relationships
between such factors and beak shape diversity among dromor­
nithids. The discovery of further cranial material especially, will
be vital in better elucidating the patterns of dromornithid evolution,
and how interspecific differences in morphology relate to environ­
mental changes across the Cenozoic of Australia.
Conclusion
The discovery of new skull fossils of the Pleistocene dromor­
nithid Genyornis newtoni, and their description and phyloge­
netic analysis in this study, have considerably furthered the
current understanding of this taxon, as well as the
Dromornithidae. Their morphology, based on their osteology
and inferred syndesmological and myological structures, likely
facilitated a wide gape, fine and independent control over
movement of the upper and lower bills, and adaptations for
water-associated habitats, potentially retained from the early
divergence of the anseriform lineage from stem Galloanserae.
We support a varied soft-browse, folivorous and frugivorous
diet, as previously proposed for dromornithids, although varia­
tion between Genyornis newtoni and species of Dromornis, sug­
gest differences in niche occupation and beak functionality.
This study additionally provides a more complete morphological
basis for addressing the relationships within this family, and with
other galloanserans. We find support for the generic distinction of
the genus Genyornis, and little evidence for close galliform or
gastornithiform affinities. There are few features of the skull
which could be inferred as confident indicators of such affinities
and the differing approaches to the evolution of large, deep mand­
ibles between dromornithids and gastornithids, appear to be driven
by functional adaptations to their respective ecological niches,
constrained by their contrasting galloanseran ancestry. Instead,
a more compelling hypothesis, and that which we advocate for, is
that the dromornithids have basal anseriform affinities, and are
likely sister to the Anhimidae, within the Anhimae. Even among
their diverse modern relatives within Galloanserae, and indeed all
of Aves, only the anhimids appear to be a close morphological
counterpart in skull anatomy, despite some notable divergences.
However, the complexity surrounding the hybrid concoction of
galliform and anseriform traits in the skull of dromornithids, pre­
sents an impediment in determining phylogenetic affinities using
morphology. Identifying ancestral or plesiomorphic conditions is
challenging for both observational and phylogenetic studies, espe­
cially considering the great age of the Galloanserae lineage, with
only a fragmentary Upper Cretaceous and early Palaeogene fossil
record. The early divergence of dromornithids likely contributes to
the combination of galliform and anseriform characters which has
produced conflicting findings, and unresolved placements in pre­
vious phylogenetic studies. Consequently, the dromornithid form
provides an unforeseen opportunity to consider character polarities
deep within the Galloanserae clade and indicates that future ana­
lyses of morphology in early diverging Anseriformes, especially,
must be considered within the greater context of the entire
Galloanserae radiation. Although we find phylogenetic support
for several hypothesised character polarities in our preliminary
phylogenetic study, there remains a need for a more extensive
analyses, to aid in better understanding and resolving the interre­
lationships of all early diverging fossil galloanseran lineages.
Acknowledgments
There are many people who contributed to this research, who we would like to
acknowledge. Firstly, Adam Yates provided access to many MAGNT specimens,

including NTM P932-2 (Figure 8), and patiently allowed us to keep the speci­
mens for longer than initially stated. Adam Yates and Sam Arman both assisted
with answering questions on the specimens available from MAGNT. Mary-
Anne Binnie (SAMA), Maya Penck (SAMA), Katie Date (NMV), Lisa Nink
(FU), and Kristen Spring (QM) also loaned us material and/or gave us access to
their collections. We thank Lawrence M. Witmer (Ohio University) for his
generosity in providing a micro-CT scanned skull of Presbyornis pervetus
(USNM 299846). We would additionally like to acknowledge Patricia Vickers-
Rich and Peter Murray for their extensive monograph on the Dromornithidae
(Murray and Vickers-Rich, 2004); Warren Handley (FU) contributed to data
analysis on gastroliths; Jacob van Zoelen (FU) who provided guidance in using
Blender v. 2.93.2 (Blender Online Community, 2023); Michael S. Y. Lee for
tutelage with regards to phylogenetic methodologies and photographs of dro­
mornithid fossil from MAGNT; Peter Trusler for interesting and useful discus­
sions; Mahala Fergusen for photographing specimen SAMA P10838; and Joseph
Bevitt who kindly took time to scan the specimen SAMA P59516 using the
neutron scanner at ANSTO which was a big contribution to understanding the
internal structure of the fossils and the scanning potential for the Lake
Callabonna Genyornis newtoni material. We also thank three anonymous
reviewers who contributed to improving the manuscript. We would like to
thank the other Chief Investigators for the Australian Research Council grant
DP180101913 (Extricating extinction histories at Lake Callabonna’s megafauna
necropolis) which funded the collection of material from Lake Callabonna,
namely Lee Arnold and Anusuya Chinsamy-Turan. Additionally, all the people
who conducted fieldwork at Lake Callabonna and contributed to preparing the
fossils, primarily Carey Burke who spent many months cleaning and preparing
the specimens use in this study, Aaron Camens, and Keith Cook (Maxim
Foundation) who made large contributions to the field work at Lake
Callabonna. Finally, thank you to Gerard and Karina Sheehan for the access to
Lake Callabonna that allowed this work to transpire.
Disclosure statement
No potential conflict of interest was reported by the author(s).
Funding
Funding was provided from the Australian Research Council grant
DP180101913 (Extricating extinction histories at Lake Callabonna’s megafauna
necropolis) and THW was sponsored by the Maxim Foundation Ltd in 2021.
ORCID
Phoebe L. McInerney http://orcid.org/0000-0001-7224-8636
Jacob C. Blokland http://orcid.org/0000-0002-3353-5533
Trevor H. Worthy http://orcid.org/0000-0001-7047-4680
Author contributions
PLM made the morphological observations, conducted the comparisons,
and created the initial drafts of the manuscript. THW provided physical
access to, and/or photographs of, many specimens, as well as the field work
and funding required to conduct this research (Australian Research Council
grant DP180101913, see below). THW and PLM conceived the project as
part of the larger project this work has contributed to, namely investigating
the biology of Genyornis newtoni, and forms a part of PLM’s PhD thesis.
PLM, JCB and THW contributed to research and interpretations regarding
relevant morphology and palaeobiology. JCB provided detailed and signifi­
cant contributions to the understanding of myological and syndesmological
correlates of osseous structures, and their homologues across the specimens,
taxa and publications studied (Appendix Three). The assembly of morpho­
logical character descriptions and states, and subsequent coding of all taxa
was undertaken by PLM, which was revised and edited by PLM and JCB.
JCB ran the phylogenetic analyses, and JCB and PLM interpreted results
and wrote associated sections. PLM created all figures aside from Figures 4
and 12 (and part of 14) which were produced by JCB. All authors partici­
pated in associated field work. All authors contributed to editing the manu­
script and approved the submitted version.
Data availability statement
All data generated or analysed during this study are included in this published
article, the appendices and supplementary information files.
HISTORICAL BIOLOGY 1143
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 HISTORICAL BIOLOGY 1149

Appendix 1: Additional figures of original and new fossils assigned to Genyornis newtoni

Figure A1. SAMA P 10838, Stirling (1913) original skull of Genyornis newtoni as it is currently: A. lateral view of the skull; B. dorsal view of the skull, showing the block in
which the skull has been set. Annotations: ap.zyg.oss., aponeurosis zygomatica ossificans; cond.oc, condylus occipitalis; cr.or., crista orbitalis; pr.post., processus postorbitalis.
Image a credit: Mahala Fergusen. Scale bar: 2 cm.
1150 P. L. MCINERNEY ET AL.

Figure A2. Laterally compressed rostrum, specimen SAMA P59517, transected by a fault in the clay matrix which has offset the caudal part from that more rostral: A. right
lateral; B. left lateral; C. dorsal; D. ventral. Annotations: ang.cl., angulus caudolateralis; ca., casque; cr.tom., crista tomialis; cul., culmen; margo.lat., margo lateralis; nas.,
apertura nasi ossea; pala., os palatinum. Scale bars: 20 mm.
 HISTORICAL BIOLOGY 1151

Figure A3. Additional skull specimens of Genyornis newtoni: A. NMV P256893, partial dorsal calvaria in dorsal and ventral views; B. SAMA P53830 jugal arch, right side,
medial, lateral, and dorsal views; C. SAMA P53830, right quadrate; D. SAMA P53830 partial condylus occipitalis and occipital region of the cranium; E. SAMA P59520, ramus
mandibulae, pars symphysialis, right side, dorsal and ventral views; F. SAMA P59520, partial left lateral cranium in medial (left) and lateral (right) views. Annotations: an.
tymp., annulus tympanicus; c. quad, condylus quadratica; cap.s-o., capitulum squamoso-otica; cond.oc., condylus occipitalis; cr.med., crista medialis; cra., cranium; d.rost.,
depressio rostromedialis; m.a.e., osseous meatus acusticus externus; mand., mandible; quad., quadrate. Scale bars: 10 mm.
1152 P. L. MCINERNEY ET AL.

Figure A4. Additional specimens of Genyornis newtoni: A. NMV P256893 partial left skull medial view; B. – G. SAMA P59517, fragmentary mandibular rami, B. right ramus
mandibulae, pars caudalis and pars intermedia in lateral view; C. and medial view; D. left ramus mandibulae, pars caudalis et intermedius, lateral view; E. and medial view;
F. fragmentary right ramus mandibulae, pars caudalis, dorsal view; G. and rostral view. Abbreviations: c.med., cotyla medialis mandibulae; cer., ceratobranchial; d.rost.,
depressio rostromedialis; imp.cor., impressio m. AME profundus, pars coronoideus; m.pt.dor., insertion area of m. pterygoideus dorsalis; p.lat., processus lateralis
mandibulae; p.med., processus medialis mandibulae; t.prae., tuberculum praearticulare. Scale bar: 10 mm.
 HISTORICAL BIOLOGY 1153

Appendix 2: Measurements of skull elements assigned to Genyornis newtoni

Table A1. Genyornis newtoni skull measurements. Rounded to the first decimal point.

Element and Measurements Specimens (mm)

Cranium SAMA P59516 NMV P256893/
SAMA P53830
Total rostrocaudal skull length, including both the cranium and rostrum. Note: This is a minimum estimate of 296 –/–
skull length considering the displacement of the cranium relative to the rostrum which has resulted in the
rostrum overlapping the most rostral part of the cranium.
Width of lateral side: from prominentia exoccipitalis to rostral margin of processus postorbitalis at ventral 61.6 59.3/–
base of crista orbitalis, in horizontal alignment.
Note: For specimen SAMA P59516, measured on left side of the cranium.
Height of caudal cranium: from ventral point of processes paroccipitalis to most dorsal point on crista nuchalis 126.8 –/–
transversus.
Dorsoventral height of caudal cranium along the mediolateral midline. 83.9 –/–
Fossa pseudotemporalis: mediolateral width and dorsoventral height (W/H). – 9.8/6.6
Processus paroccipitalis: maximum length from ventral base of osseous meatus acusticus externus. 55.1 52.5/–
Width of caudal side, occipital region: mediolateral width in alignment with the condylus occipitalis. 88.6 –/–
Osseous meatus acusticus externus: mediolateral width and dorsoventral height (W/H). 10.6/12.1 14.4/12.7
Condylus occipitalis: mediolateral width and dorsoventral height (W/H). 16.4/16.1 –/17.3
Foramen magnum: width and height (W/H). 16.9/24.4 –/–
Individual tuberculum basilare: mediolateral width and dorsoventral height (W/H). Note: For specimen SAMA 12.6/15.3 –/–
P59516, measurements from left tubercle.
Processus basipterygoideus: rostrocaudal length of the articular surface. 15.2 –/–
Quadrate NMV P256893 SAMA P53830
Maximum height from rostrocaudal midpoint of the ventral margin of pars mandibularis to dorsal-most point 47.2 –
on pars otica.
Pars otica: Height from dorsal margin of fovea quadratojugalis to dorsal-most point on pars otica. 28.1 31.0
Capitulum squamoso-otica: mediolateral width/rostrocaudal length. 12.8/12.8 –
Depressio rostromedialis: dorsoventral height. 12.5 –
Foramen pneumaticum rostromediale: dorsoventral height. 6.3 –
Pars mandibularis: maximum length of long axis including both articular surfaces. 26.2 –
Rostrum SAMA P59521 SAMA P59517
Rostrocaudal length of the rostrum from the centre of the craniorostral hinge to rostral apex. 212.4 237.1
Maximum dorsoventral height of the fused praemaxillare/maxillare plates, from the caudal end of the 72.9 (34.3% length) 81.2 (34.2% height)
culmen, just rostral of the casque, ventrally to the tomial surface.
Maximum mediolateral width of the rostrum between left and right margo tomialis. 76.05 NA
Rostrocaudal length of casque. 51.4 59.1
Mediolateral width of the casque. 76.3 N/A
Rostrocaudal length of the ventral rostrum, not including the vomer and palatines. 129.4 162.6
Dorsoventral height of the caudal post narial bar, not inclusive of the casque. 32.4 39
Os palatinum: maximum dorsoventral height from point of interaction with the os maxillare near the angulus 42.8 45.1
tomialis to the ventral-most extent of the palatinum.
Mandible SAMA P59516 NMV P256893
Note: all measurements from
right side of specimen.
Rostrocaudal length of mandible: caudal end of processus retroarticularis to rostral tip of ramus mandibulae, 293 N/A
pars symphysialis.
Depth of mandible: dorsoventral height from highest point on the angulus dorsalis mandibulae to the 61.5 N/A
immediately ventral point on ventral side. This is approximately the rostral-most point of the angulus
ventralis mandibulae.
Rostrocaudal length of processus retroarticularis from its caudal extent to the caudal margin of processus 39.7 N/A
lateralis mandibulae.
Ramus mandibulae, pars symphysialis, rostrocaudal length. 44.2 N/A
Maximum mediolateral width of mandible at processus medialis mandibulae. N/A 25.5
Rostrocaudal length of angulus ventralis mandibulae where it is entirely distinct ventrally with respect to the 81.8 N/A
rest of the ventral mandible margin.
Appendix 3: Homological correlations for the terminology associated with the adductor muscles of the mandible across relevant literature
1154

Table A2. (Part 1) Homological correlations for the terminology associated with musculus adductor mandibulae across examples of relevant literature.
Part 1

Dzerzhinsky and
Matsuoka et al. Grintsevichene Zusi and Livezey Vanden Berge and Dzerzhinsky and
+
Holliday and Witmer (2007) (2008) (2002) (2000) Weber (1996) + Zweers (1993) Potapova (1974) Zweers (1974)
P. L. MCINERNEY ET AL.

Musculus adductor mandibulae externus

Musculus adductor mandibulae Musculus Musculus Musculus adductor Musculus Musculus Musculus Musculus adductor mandibulae externus, pars
externus superficialis adductor adductor mandibulae adductor adductor adductor caudolateralis (ACL)
mandibulae mandibulae externus, pars mandibulae mandibulae mandibulae
externus externus articularis, caput externus, pars externus, pars externus
superficialis, profundus externa articularis profunda profundus, pars
A-portion caudalis (Aec) [caudalis]1 caudalis (Aec)
Musculus adductor mandibulae Musculus Musculus Musculus adductor Musculus Musculus Musculus Musculus adductor mandibulae externus, pars
externus profundus, pars adductor adductor mandibulae adductor adductor adductor rostromedialis (ARM)
superficialis mandibulae mandibulae externus, pars mandibulae mandibulae mandibulae
externus externus superficialis externus, pars externus, pars externus
superficialis, superficialis (Aes: zygomatica ventralis3 superficialis (Aes)
C-portion also see
Dzerzhinsky and
Potapova 1974;
Dzerzhinsky 1982)
^
Musculus adductor mandibulae Musculus Musculus Musculus adductor Musculus Musculus Musculus Associated with aponeurosis 2a (ap.2a)
externus profundus, pars adductor adductor mandibulae adductor adductor adductor
zygomaticus mandibulae mandibulae externus, pars mandibulae mandibulae mandibulae
externus externus medialis zygomatica + externus, pars externus, pars externus medialis
superficialis, (Aem) musculus adductor zygomatica ventralis (Aem)
B-portion mandibulae
externus, pars
articularis, caput
externa (in part)
Musculus adductor mandibulae Musculus Musculus Musculus adductor Musculus Musculus Musculus Associated with aponeurosis 4a (ap.4a)
externus profundus, pars adductor adductor mandibulae adductor adductor adductor
coronoideus mandibulae mandibulae externus, pars mandibulae mandibulae mandibulae
externus medialis externus coronoidea externus, pars externus, pars externus
profundus rostralis coronoidea rostralis profundus, pars
(Aer) [temporalis] * rostralis (Aer)

Musculus adductor mandibulae internus

Musculus pterygoideus dorsalis Musculus Musculus — Musculus Musculus Musculus Musculus pterygoideus dorsalis medialis
pterygoideus pterygoideus pterygoideus pterygoideus pterygoideus
dorsalis medialis dorsalis (various dorsalis medialis
subdivisions
reviewed)
(Continued)
Table A2. (Continued).

Part 1

Dzerzhinsky and
Matsuoka et al. Grintsevichene Zusi and Livezey Vanden Berge and Dzerzhinsky and
Holliday and Witmer (2007) + (2008) (2002) (2000) Weber (1996) + Zweers (1993) Potapova (1974) Zweers (1974)
Musculus pterygoideus Musculus Musculus — Musculus Musculus Musculus Musculus pterygoideus ventralis lateralis +
ventralis pterygoideus pterygoideus pterygoideus pterygoideus pterygoideus musculus pterygoideus dorsalis lateralis; musculus
ventralis lateralis ventralis lateralis; (various ventralis lateralis; pterygoideus ventralis medialis
+ musculus musculus subdivisions musculus
pterygoideus pterygoideus reviewed) pterygoideus
dorsalis lateralis; ventralis medialis; dorsalis lateralis +
musculus musculus musculus
pterygoideus pterygoideus pterygoideus
ventralis medialis ventralis caudalis ventralis medialis
+ musculus
pterygoideus
caudalis
Musculus pseudotemporalis Musculus Musculus — Musculus Musculus Musculus Musculus pseudotemporalis superficialis
superficialis pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis
superficialis superficialis (Pss: superficialis superficialis superficialis (Pss)
also see
Dzerzhinsky and
Potapova 1974;
Dzerzhinsky 1982)
^
Musculus pseudotemporalis Musculus — — Musculus Musculus Musculus Musculus pseudotemporalis profundus
profundus pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis
profundus profundus profundus/ profundus (Psp)
musculus
adductor
mandibulae
caudalis

Musculus adductor mandibulae posterior †

Musculus adductor mandibulae Musculus Fibres of musculus Musculus adductor Musculus Musculus Musculus adductor Musculus adductor mandibulae externus
posterior lateralis adductor adductor mandibulae adductor adductor mandibulae profundus (AEP)
mandibulae mandibulae externus, pars mandibulae mandibulae ossis posterior (Ap),
externus externus articularis, caput caudalis2 quadrati possibly in
profundus profundus interna [caudalis]† association with
caudalis, attached their mandibular
to musculus aponeurosis
adductor posterior
mandibulae insertionis
posterior (Aec*)
Musculus adductor mandibulae Musculus Musculus Musculus adductor Musculus Musculus Musculus adductor Musculus adductor mandibulae posterior (AMP)
posterior medialis adductor adductor mandibulae adductor adductor mandibulae
mandibulae mandibulae posterior mandibulae mandibulae ossis posterior (Ap),
posterior posterior (Ap) caudalis2 quadrati [caudalis] from aponeurosis
posterior
HISTORICAL BIOLOGY

quadratica (apq)
(Continued)
1155
Table A2. (Continued).

Part 2
1156

Holliday and Witmer Dzerzhinsky and Zusi and Storer Goodman and Davids (1952a,
(2007) Belokurova (1972) (1969) Fujioka (1963) Fisher (1962) b, c) Hofer (1950) Lakjer (1926)
Musculus adductor mandibulae externus
Musculus Musculus Musculus Musculus masseter medius (5B) Musculus Musculus Musculus Musculus adductor mandibulae externus
adductor adductor adductor adductor adductor adductor profundus (III kaud.)
mandibulae mandibulae mandibulae mandibulae mandibulae mandibulae
externus externus, pars externus, part 2 externus, externus (m. add. externus, pars
superficialis profunda caudalis superficialis, Ia ext) superficialis
(Aec) portion
Musculus Musculus Musculus Musculus masseter superficialis (5A) Musculus Musculus Musculus Musculus
P. L. MCINERNEY ET AL.

adductor adductor adductor adductor adductor adductor adductor mandibulae externus superficialis
mandibulae mandibulae mandibulae mandibulae mandibulae mandibulae (I)
externus externus, pars externus, part 1 externus, externus (m. add. externus, pars
profundus, pars superficialis (Aes) superficialis, Ic ext) occipitalis
superficialis portion
Musculus Musculus Musculus Musculus masseter superficialis (5A) Musculus Musculus Musculus Musculus adductor mandibulae externus
adductor adductor adductor adductor adductor adductor medialis (II)
mandibulae mandibulae mandibulae mandibulae mandibulae mandibulae
externus externus, pars externus, part 2 externus, externus (m. add. externus, pars
profundus, pars medialis (Aem) superficialis, Ib ext) medialis
zygomaticus portion
Musculus adductor Musculus adductor Musculus Musculus masseter profundus (6) Musculus Musculus Musculus Musculus adductor mandibulae externus
mandibulae mandibulae adductor adductor adductor adductor profundus (III rost.)
externus externus, pars mandibulae mandibulae mandibulae mandibulae
profundus, pars profunda rostralis externus, part 3 externus, medialis externus (m. add. externus
coronoideus (Aer) (M. II) ext) temporalis

Musculus adductor mandibulae internus

Musculus Musculus Musculus Musculus pterygoideus dorsalis medialis Musculus Musculus Musculus Musculus pterygoideus, pars dorsalis
pterygoideus pterygoideus, pars pterygoideus, pars (11Ab) pterygoideus pterygoideus pterygoideus, medialis
dorsalis dorsalis medialis dorsalis, lateral dorsalis, medial dorsalis medialis pars dorsalis
branch part medialis
Musculus Musculus Musculus Musculus pterygoideus dorsalis lateralis Musculus Musculus Musculus Musculus pterygoideus, pars ventralis
pterygoideus pterygoideus, pars pterygoideus, pars (11B) + musculus pterygoideus ventralis pterygoideus pterygoideus pterygoideus, lateralis + musculus pterygoideus, pars
ventralis ventralis lateralis; lateralis, medialis (9A); musculus pterygoideus ventralis, lateral ventralis lateralis pars ventralis dorsalis lateralis; musculus pterygoideus,
musculus dorsolateral ventralis lateralis (9B) + pterygoideus part + musculus + musculus lateralis + pars ventralis medialis; musculus
pterygoideus, pars branch + dorsalis medialis (11Aa) pterygoideus pterygoideus musculus pterygoideus, pars dorsalis 1
dorsalis lateralis + musculus dorsalis, lateral dorsalis lateralis; pterygoideus,
musculus pterygoideus, pars part; musculus musculus pars dorsalis
pterygoideus, pars lateralis, pterygoideus pterygoideus lateralis;
ventralis medialis ventromedial ventralis, medial ventralis medialis musculus
+ musculus branch + part pterygoideus,
pterygoideus, pars musculus pars ventralis
dorsalis caudalis pterygoideus, pars medialis;
dorsalis, medial musculus
branch pterygoideus,
pars caudalis
Musculus Musculus Musculus Musculus temporalis (7) Musculus Musculus — Musculus pseudotemporalis superficialis
pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis
superficialis superficialis (Pss) superficialis superficialis superficialis (m.
pseudo. sup.)
Musculus Musculus Musculus Musculus quadratomandibularis (10C) Musculus Musculus — Musculus pseudotemporalis profundus
pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis pseudotemporalis
profundus profundus (Psp) profundus profundus profundus (m.
pseudo. prof.)
(Continued)
Table A2. (Continued).

Part 2

Holliday and Witmer Dzerzhinsky and Zusi and Storer Goodman and Davids (1952a,
(2007) Belokurova (1972) (1969) Fujioka (1963) Fisher (1962) b, c) Hofer (1950) Lakjer (1926)
†
Musculus adductor mandibulae posterior
Musculus Musculus Musculus Musculus quadratomandibularis (10A) Musculus Musculus Musculus —
adductor adductor adductor adductor adductor adductor
mandibulae mandibulae mandibulae mandibulae mandibulae mandibulae
posterior lateralis posterior (Ap), to posterior externus, externus externus, pars
aponeurosis a.11 profundus (M. III) profundus (m. profunda
add. ext. prof.)
Musculus Musculus Musculus Musculus quadratomandibularis (10B) Musculus Musculus Superficial —
adductor adductor adductor adductor adductor musculus
mandibulae mandibulae mandibulae mandibulae mandibulae adductor
posterior medialis posterior (Ap), posterior posterior (‘a’ and posterior (m. add. mandibulae
from superficial ‘b’ parts in Mergus post.) externus, pars
aponeurosis a.12 merganser) profunda
— not treated in the particular study.

Notes.
+While we follow Holliday and Witmer (2007) for all muscles included here in general, the homologies of more specific parts of musculus pterygoideus, not addressed in-depth by these authors, are associated in this study herein
following the relationships inferred by Weber (1996), namely musculus pterygoideus ventralis lateralis, musculus pterygoideus ventralis medialis, musculus pterygoideus ventralis caudalis, and musculus pterygoideus dorsalis.
Also see Weber (1996), Tab. 11.
1
Because of the ambiguity in the displayed figure by Vanden Berge and Zweers (1993: fig. 6.5) corresponding to the insertions of musculus adductor mandibulae externus, pars profunda [caudalis], and musculus adductor
mandibulae externus, pars ventralis, the muscle synonymous with musculus adductor mandibulae externus superficialis (sensu Holliday and Witmer 2007) may be predominately equable to musculus adductor mandibulae
externus, pars profunda [caudalis], but may also relate to musculus adductor mandibulae externus, pars ventralis, in part.
2
While Weber (1996, p. 57) recognised that there is some differentiation in the exact insertion area of fibres relating to their musculus adductor mandibulae caudalis (to the aponeurosis paracoronoidea interna, and rostrad of the
articulatio quadrato-mandibularis), likely representative of parts recognised in other studies, beyond this mention there is no differentiation of these parts.
3
Vanden Berge and Zweers (1993: annot. 18) described part of the musculus adductor mandibulae externus, pars ventralis, as originating on the processus postorbitalis, but then erroneously or accidentally displayed this part
originating upon the caudal orbit on the diagram (fig. 6.5) of Podilymbus podiceps that was adapted from Zusi and Storer (1969: Figure 6). Instead, this area corresponds to the origin of musculus pseudotemporalis superficialis
(part 2) of the latter authors’ study.
*Zusi and Livezey (2000) suggested that their musculus adductor mandibulae externus, pars coronoidea, is synonymous with musculus adductor mandibulae externus, pars rostralis [temporalis], of Vanden Berge and Zweers
(1993). However, Holliday and Witmer (2007) suggested that their musculus adductor mandibulae profundus (and included parts) is equivalent to musculus adductor mandibulae externus pars ventralis of Vanden Berge and
Zweers (1993) and interpret musculus adductor mandibulae externus, pars coronoidea, of Zusi and Livezey (2000) as part of their musculus adductor mandibulae profundus. Musculus adductor mandibulae externus, pars
rostralis [temporalis], of Vanden Berge and Zweers (1993) was also considered synonymous to musculus adductor mandibulae externus superficialis (Holliday and Witmer 2007: Table 2). Based on the origins and insertions
illustrated and described in Zusi and Storer (1969), and those of the derivative adaptive diagram and annotations of Vanden Berge and Zweers (1993), it is revealed that musculus adductor mandibulae externus, pars rostralis
[temporalis], of Vanden Berge and Zweers (1993) is comparable to musculus adductor mandibulae externus profundus, pars coronoideus, of Holliday and Witmer (2007), and not equivalent to musculus adductor mandibulae
externus superficialis, contrary to the latter study. This is supported by reporting that the terminal insertion of musculus adductor mandibulae externus, pars rostralis [temporalis], of Vanden Berge and Zweers (1993) relates to
processus coronoideus (sensu stricto, see Baumel and Witmer 1993: annot. 44), which is comparable to musculus adductor mandibulae externus profundus, pars coronoideus, of Holliday and Witmer (2007), and can be
compared with muscles that insert on this feature in other studies (e.g. musculus adductor mandibulae externus profundus, pars rostralis, sensu Dzerzhinsky 1982, and others).
†Musculus adductor mandibulae externus, pars profunda [caudalis], of Vanden Berge and Zweers (1993) was interpreted as synonymous with musculus adductor mandibulae externus, pars articularis (likely caput externa) by Zusi
and Livezey (2000). The latter name was considered equivalent to musculus adductor mandibulae externus superficialis, while the former is considered synonymous with musculus adductor mandibulae posterior by Holliday
and Witmer (2007: Table 2), and thus there appears to be a conflict in this assignment of synonymy. An overarching issue is that musculus adductor mandibulae posterior may not exist in two separate parts in some birds but
may do (as medial and lateral parts) in some Galloanserae and Neoaves (see Zusi and Storer 1969; Holliday and Witmer 2007, p. 467). Musculus adductor mandibulae posterior was not divided into parts by Zusi and Storer (1969)
and was hence not labelled in separate parts on this same diagram that was adapted by Vanden Berge and Zweers (1993: fig. 6.5).
Vanden Berge and Zweers (1993: annot. 18) categorised their musculus adductor mandibulae, pars profunda [caudalis], as an external adductor muscle, and stated that it acts independently of the musculus
adductor mandibulae ossis quadrati [caudalis]. They did not explicitly say that their musculus adductor mandibulae, pars profunda [caudalis], originates on the neurocranium in part, while the associated
diagram of Vanden Berge and Zweers (1993: fig. 6.5), adapted from Zusi and Storer (1969: fig. 6), shows part of the origin in this region. The insertion displayed in this figure as labelled by Vanden Berge and
Zweers (1993: fig. 6.5) leaves ambiguity with regards to whether it is better associated with the musculus adductor mandibulae externus or the musculus adductor mandibulae posterior (sensu Holliday and
Witmer 2007). However, the original figure and related description of the muscle illustrated by Zusi and Storer (1969) would also imply this corresponds to part of the external adductor musculature.
HISTORICAL BIOLOGY

Furthermore, the annotation given by Vanden Berge and Zweers (1993: annot. 18) describes this muscle as deriving solely from the quadrate, which when their subsequent figure labels their musculus adductor
mandibulae, pars profunda [caudalis], as inserting on the caudal mandible in an area comparable to where part of musculus adductor mandibulae posterior may insert in some taxa, it can be interpreted
ambiguously, however, and may be considered as synonymous with part of musculus adductor mandibulae posterior, therefore (as per Holliday and Witmer 2007: table 2).
1157
 1158

Holliday and Witmer (2007: 467) equated musculus adductor mandibulae posterior with only musculus adductor mandibulae ossis quadrati [caudalis] of Vanden Berge and Zweers (1993) in text but associated
both this muscle and musculus adductor mandibulae externus, pars profunda [caudalis], with musculus adductor mandibulae posterior in their related table (table 2). Considering Holliday and Witmer’s (2007)
interpretation that musculus adductor mandibulae posterior lateralis is synonymous with musculus adductor mandibulae externus, pars articularis, caput interna (of Zusi and Livezey 2000), combined with the
suggestion by Zusi and Livezey (2000) that their pars articularis is equivalent to musculus adductor mandibulae externus, pars profunda [caudalis], while the latter authors’ musculus adductor mandibulae
posterior is the same as musculus adductor mandibulae ossis quadrati [caudalis] of Vanden Berge and Zweers (1993), it makes sense that musculus adductor mandibulae externus, pars articularis, caput externa
of Zusi and Livezey (2000) would then correspond well with musculus adductor mandibulae externus, pars profunda [caudalis], of Vanden Berge and Zweers (1993), and thus an external adductor muscle
(musculus adductor mandibulae externus superficialis, sensu Holliday and Witmer 2007). In contrast, Holliday and Witmer’s (2007) musculus adductor posterior medialis can be considered the same as musculus
adductor mandibulae posterior (sensu Zusi and Livezey 2000, when musculus adductor mandibulae externus, pars articularis, caput interna is present), and the same as musculus adductor mandibulae ossis
quadrati [caudalis] of Vanden Berge and Zweers (1993).
We thus consider musculus adductor mandibulae externus, pars profunda [caudalis], of Vanden Berge and Zweers (1993) as inequivalent to part of the musculus adductor mandibulae posterior, and instead
associate it with the musculus adductor mandibulae externus superficialis. It should be noted that this assigned correlation is dubious and the area labelled as associated with this muscle by Vanden Berge and
P. L. MCINERNEY ET AL.

Zweers (1993: fig. 6.5) may receive fibres from both musculus adductor mandibulae posterior and musculus adductor mandibulae externus superficialis.
^ Specific term not mentioned in the particular study; non-abbreviated terminology derived from closely related studies using the same labelled acronym (cited)

References
Baumel JJ, Witmer LM. 1993. Osteologia. In: Baumel JJ King AS, Breazile JE, Evans HE, Vanden Berge JC (Eds.), Handbook of Avian Anatomy: Nomina Anatomica Avium, 2nd Edn (pp. 45–132). Publications of the Nuttall
Ornithological Club, Cambridge, Massachusetts.
Davids JAG. 1952a. Etude sur les attaches au crâne des muscles de la tête et du cou chez Anas platyrhyncha platyrhyncha (L.), I. Proceedings of The Koninklijke Nederlandse Akademie Van Wetenschappen. Series C.
Biological and Medical Sciences. 55:81–94.
Davids JAG. 1952b. Etude sur les attaches au crâne des muscles de la tête et du cou chez Anas platyrhyncha platyrhyncha (L.). II. Proceedings Of The Koninklijke Nederlandse Akademie Van Wetenschappen. Series C.
Biological and Medical Sciences. 55:525–533.
Davids JAG. 1952c. Etude sur les attaches au crâne des muscles de la tête et du cou chez Anas platyrhyncha platyrhyncha (L.). III. Proceedings Of The Koninklijke Nederlandse Akademie Van Wetenschappen. Series C.
Biological and Medical Sciences. 55:533–540.
Dzerzhinsky FYa. 1982. Adaptivnʹіe čerty v stroenii čelyustnogo apparata nekotoryh guseobraznyh ptic i veroyatnye puti èvolyucii otrjada [Adaptive features in the construction of the jaw apparatus of some Anseriformes
and probable evolutionary paths of the order]. Zoologichesky Zhurnal. 56:1030–1041.
Dzerzhinsky FYa, Belokurova IN. 1972. Sravnitelnoy anatomii chelustnoy muskul tury ptits. Chelustnyye myshtsy glucharya (Tetrao urogallus) [Comparative anatomy of the jaw muscles of birds. Jaw muscles of capercaillie
(Tetrao urogallus)]. Zoologicheskiĭ Zhurnal. 51:555–564.
Dzerzhinsky FYa, Grintsevichene TI. 2002. O vozmozhnykh filogeneticheskikh svyazyakh guseobraznykh ptits [On possible phylogenetic relationships of Anseriformes]. Kazarka. 8:19–39.
Dzerzhinsky FYa, Potapova EG. 1974. Sistema sukhozhil’nykh obrazovaniy kak ob’yekt sravnitel’noy miologii chelyustnogo apparata ptits [The system of tendon formations as an object of comparative theology of the jaw
apparatus of birds]. Zoologicheskiĭ Zhurnal. 53:1341–1351.
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anatomy of the fowl. IV. Origins and insertions of muscles of the head and neck in the fowl. 1. Muscles of the head]. Nihon juigaku zasshi. The Japanese journal of veterinary science. 25(4):207–226.
Goodman DC, Fisher HI. 1962. Functional anatomy of the feeding apparatus in waterfowl (Aves: Anatidae). Southern Illinois University Press, Carbondale.
Hofer H. 1950. Zur Morphologie der Kiefermuskulatur der Vögel Zoologische Jahrbücher. Abteilung für Anatomie und Ontogenie der Tiere. 70:427–556.
Holliday CM, Witmer LM. 2007. Archosaur adductor chamber evolution: integration of musculoskeletal and topological criteria in jaw muscle homology. Journal of Morphology. 268(6):457–484.
Lakjer T. 1926. Studien über die Trigeminus-versorgte Kaumuskulatur der Sauropsiden. Reitzel, C.A. (Ed.), Buchhandlung, Kopenhagen. 154.
Matsuoka H, Kurosu H, Inglis MP, Kitagawa H, Kusuhashi N, Hasegawa Y. 2008. Myology and osteology of the Whooper Swan Cygnus cygnus (Aves: Anatidae) part 2. Muscles of the jaws, tongue and anteriormost neck.
Bulletin of the Gunma Museum of Natural History. 12:1–14.
Vanden Berge JC, Zweers GA. 1993. Myologia. In: Baumel, J.J. King, A.S., Breazile, J.E., Evans, H.E., and Vanden Berge, J.C. (Eds.), Handbook of Avian Anatomy: Nomina Anatomica Avium, 2nd Edn. Publications of the
Nuttall Ornithological Club, Cambridge, Massachusetts, pp. 189–247.
Weber E. 1996. Das skelet-muskel-system des kieferapparates von Aepypodius arfakianus (Salvidori, 1877) (Aves, Megapodiidae). Courier Forschungsinstitut Senckenberg. 189:1–132.
Zusi RL, Livezey BC. 2000. Homology and phylogenetic implications of some enigmatic cranial features in galliform and anseriform birds. Annals of Carnegie Museum. 69(3):157–193.
Zusi RL, Storer RW. 1969. Osteology and myology of the head and neck of the Pied-billed Grebes (Podilymbus). Miscellaneous Publications Museum of Zoology, University of Michigan. 39:1–49.
Zweers GA. 1974. Structure, movement, and myography of the feeding apparatus of the mallard (Anas platyrhynchos L.) A study in functional anatomy. Netherlands Journal of Zoology. 24(4):323–467.
 HISTORICAL BIOLOGY 1159

Appendix 4: Morphological character list

Characters for the phylogenetic analysis. Original character list derived from Worthy et al. (2017). Reassessment, modifications, and additions made to characters,
states and coding based off personal observations and assessment of other matrices from Andors (1988), Bourdon (2005; 2011), Clarke (2002), Cracraft and Clarke
(2001), Dyke et al. (2003), Elzanowski and Stidham (2010: char. 1), Ericson (1997), Field et al. (2020), Ksepka (2009), Livezey (1986; 1996; 1997), Livezey and Zusi
(2006), Mayr and Clarke (2003), Murray and Vickers-Rich (2004), Stidham (2001), Worthy and Lee (2008), Worthy and Scofield (2012), Worthy et al. (2017), Zusi
and Livezey (2000).
Note: Here we use the name Nettapterornis oxfordi for the taxon previously called Anatalavis oxfordi following recommendations in Houde et al. (2023).
Note: As we, and others (e.g., Worthy et al. 2016), are unable to assign the quadrates attributed to the genus Ilbandornis to species level, and since character state
coding across all Ilbandornis sp. quadrates observed does not vary, we have opted to code the quadrate for both species identically. We treat the mandible similarly.
Note: We do not code the retroarticular process for Asteriornis maastrichtensis, following changes to specimen interpretation as reported on at the 10th
International Meeting of the Society of Avian Palaeontology and Evolution by Crane (2023).
Ordered Characters:
1, 4, 7, 10, 12, 13, 14, 15, 16, 17, 18, 21, 24, 26, 30, 31, 33, 34, 35, 36, 37, 38, 41, 42, 44, 45, 48, 50, 52, 54, 55, 57, 58, 60, 61, 67, 69, 70, 73, 74, 76, 77, 79, 81, 85, 87, 88, 90,
92, 95, 98.
The above characters are ordered in recognition that the states in these characters represent a morphocline where the transition between relative extremes (i.e.,
states 0 to 2 or greater number) must pass through an intermediate state (or states). Assuming an unordered system of character state changes for such characters
would not be accurate to the morphology observed and would often be biologically implausible.

Complete skull
(1) Skull, lateral and dorsal perspectives, rostrocaudal proportions, rostrocaudal length of the rostrum (as measured between the rostral apex of the rostrum and the
craniofacial hinge, in line with the rostral-most extension of the os frontale) relative to the rostrocaudal length of the cranium (as measured between the
craniofacial hinge, in line with the rostral-most extension of the os frontale, to the most caudal point on the occipital region of the cranium): 0. Rostrum
rostrocaudally shorter than the cranium; 1. Rostrum near equal to or up to 1.5 times the rostrocaudal length of cranium; 2. Rostrum is between 1.5 to 2 times the
rostrocaudal length of cranium; 3. Rostrum is greater than 2 times rostrocaudal length of the cranium. Modified from Worthy et al. (2017: char. 1) and Field et al.
(2020: char. 1). Ordered.
(2) Skull, craniorostral hinge, formation, and hinge type: 0. Flexion zone absent or indistinct, no transverse sulcus, limited flexion; 1. Well-developed flexion zone
present and distinct; 2. A synovial joint, essentially a hinge. Modified from Worthy et al. (2017: char. 8), see also Mayr & Clarke (2003: char. 5).
(3) Skull, presence of an osseous casque or other elaborate osseous structure: 0. Absent; 1. Present on rostrum; 2. Present on cranium.

Cranium
(4) Cranium, lateral and dorsal perspectives, frontals, rostral truncation or rostrocaudal shortening, location of the craniorostral hinge (rostral termination of the
frontal bones) in relation to the orbital area: 0. Frontals are extended rostrally, craniorostral hinge is far rostral of the orbits; 1. Craniorostral hinge intersects the
most rostral point of the orbits; 2. Frontal bones are rostrally truncated, craniorostral hinge transects the dorsal-most point, at the supraorbital area of the orbits;
3. Frontal bones are substantially rostrally truncated, craniorostral hinge transects the orbits in near rostrocaudal alignment with, or caudal of, the processus
postorbitalis. Ordered.
(5) Cranium, lateral and rostral perspectives, caudal orbit (and/or lateral cranium), area muscularis aspera, origin of the m. pseudotemporalis superficialis and
presence of fossa pseudotemporalis, status and form: 0. Not distinct, area muscularis aspera is nearly flat; 1. Distinct, a shallow depression demarked dorsally by a
ridge; 2. Distinct, fossa pseudotemporalis present as a deep fossa within a larger depression; 3. Homologous origin of the m. pseudotemporalis superficialis is not
restricted to caudal orbit, present on lateral cranium. State 3 is typical of Palaeognathae (see Dzerzhinsky, 1983; Elzanowski, 1987).
(6) Cranium, lateral and rostral perspectives, caudal orbit (and/or lateral cranium), area muscularis aspera, tuberculum for origin of m. pseudotemporalis
(superficialis): 0. Absent; 1. Present and projected; 2. Low osseous crest; 3. Lowly-raised rugose region. Modified from Dyke et al. (2003: char. 4). Note:
Although Picasso et al. (2023) identified this crest as the origin of the m. pseudotemporalis superficialis in the Rhea, for tinamous, Elzanowski (1987) noted that
the tubercle is the origin of the ligamentum orbitoquadratum to which the m. pseudotemporalis is attached. As in either case, the crest is associated with the m.
pseudotemporalis, we opt to treat the crest as the same across Palaeognathae and code its presence in palaeognaths accordingly.
(7) Cranium, lateral perspective, postorbital temporal region, processus postorbitalis, orientation: 0. Projects ventrally; 1. Directed rostroventrally; 2. Projected
strongly rostrally. Modified from Worthy et al. (2017: char. 46). Also see Bourdon (2011, char. 57), Ksepka (2009, char. 11), Zusi & Livezey (2000: p. 175).
Ordered.
(8) Cranium, lateral perspective, temporal region, relationship between areas of origin of m. AME profundus and m. AME superficialis, associated aponeuroses and
their respective osteological structures (e.g., crista zygomatica, crista m. AME superficialis): 0. Origin area for m. AME profundus and associated structures are
wholly dorsal or rostrodorsal of those corresponding to m. AME superficialis; 1. Origin area for m. AME profundus and associated structures are rostral or
slightly rostroventral of those of m. AME superficialis; 2. Origin area for m. AME profundus and associated structures are clearly rostroventral of those of m.
AME superficialis, where the latter extends substantially dorsal of the former; 3. Not comparable, the deep m. AME is not well differentiated or clearly subdivided
with respect rostral (m. AME profundus, pars coronoideus) and caudal parts (m. AME superficialis). Note: In Palaeognathae, the combined regions of origin
associated with the m. AME occupies areas of processus postorbitalis, processus zygomaticus (including crista temporalis), processus orbitalis of the quadrate,
directly or via related membranes, ligaments or fascia (see Dzerzhinsky, 1983); Elzanowski, 1987). Interpretation of states for neoavians follows the array of
literature available for various taxa (e.g., Dzerzhinsky & Ladygin, 2004; Dzerzhinsky & Potapova, 1974; Dzerzhinsky & Yudin, 1974; Fisher & Goodman, 1955;
Korzun, 1998; Kuular, 2002; Sokolov, 1995).
(9) Cranium, lateral perspective, temporal region, processus zygomaticus, status: 0. Absent, osseous projection in area (if any) is homologous with aponeurosis
zygomatica ossificans; 1. Present. See Zusi & Livezey (2000) for details regarding this subject. Sylviornis neocaledoniae is coded as present following Mourer-
Chauviré & Balouet (2005). Despite the lack of juvenile skull material preserving this region for Gastornis giganteus, a processus zygomatica is also coded as
present considering the similarities in the morphology of the lateral cranium to Galliformes (e.g., secondary temporal fenestra formation) which is inconsistent
with the absence of this feature (see Zusi & Livezey, 2000).
(10) Cranium, lateral and ventral perspectives, temporal region, aponeurosis zygomatica ossificans, status and rostral development: 0. Absent or minor ossification;
1. Variably ossified and rostrally developed; 2. Extensively ossified, rostrally developed to intersect or exceed the processus postorbitalis. Modified from Worthy
et al. (2017: char. 18), see also Bourdon (2011, char. 88), Dyke et al. (2003: char. 19), Ksepka (2009, char. 12 and 13), Livezey (1997: char. 8), Murray & Vickers-
Rich (2004: char. 8, table 8), Zusi & Livezey (2000: p. 160, 175-177). Note: Numididae are coded as 0 as there is no ossification of the aponeurosis present, see
Hofer (1950) and Zusi & Livezey (2000: p. 160). Ordered.
(11) Cranium, lateral perspective, processus postorbitalis and aponeurosis zygomatica ossificans, formation of a secondary temporal fenestra, status: 0. Absent; 1.
Present. See Zusi & Livezey (2000) and Elzanowski & Mayr (2017).
(12) Cranium, lateral and ventral perspectives, temporal region, processus postorbitalis and/or aponeurosis zygomatica ossificans, location of the impression for the
origin of m. AME profundus, pars coronoideus: 0. M. AME profundus, pars coronoideus does not form a distinct belly of the m. AME, associated fibres and
interrelated parts of the m. AME originate on the lateral cranium; 1. Located laterally on the cranium, in a clear impression between the processus postorbitalis
and processus zygomatica and/or aponeurosis zygomatica ossificans, may additionally originate ventrolaterally on the medial aspect of the aponeurosis
zygomatica ossificans, if present; 2. Restricted to a medioventral location on the lateral cranium, lacking the significant dorsocaudal expansion onto the lateral
side of the cranium; 3. Originates ventrally or medioventrally on the aponeurosis zygomatica ossificans, mediolaterally narrow; 4. Located medioventrally,
medially expansive but confined laterally; 5. Located medioventrally, expansive both medially and laterally, often slightly visible from lateral perspective.
Associated with Bourdon (2005: char. 27, 44; 2011: char. 59), Ericson (1997: char. 4), and Worthy et al. (2017: char. 19). Note: see Hofer (1950), Fujioka (1963),
1160 P. L. MCINERNEY ET AL.

Dzerzhinsky & Belokurova (1972), Dzerzhinsky (1983), Elzanowski (1987), Weber (1996), and Zusi & Livezey (2000). Ordered – specific ordering of states is
imposed following observations of evolutionary transitions (i.e., medial movement of the m. AME profundus, pars coronoideus, among galliforms and
anseriforms) by Zusi and Livezey (2000) as also discussed in the main text. An unordered model is additionally not evolutionarily plausible (e.g., state 4 to state
0 in one step).
(13) Cranium, lateral and ventral perspectives, otic region, osseus meatus acusticus externus, annulus tympanicus bridges the ala parasphenoidalis and processus
suprameaticus to form a complete ring, enclosing the osseus meatus acusticus externus ventrorostrally, status: 0. Absent, no tympanic enclosure; 1. Partial or
complete tympanic enclosure, thin and fragile; 2. Complete closure, robust and obvious. Modified from Livezey (1997: char. 4) and Worthy et al. (2017: char.
16), see also Bourdon (2005: char. 28), Livezey and Zusi (2006: char. 0019), and Worthy and Lee (2008: char 17). Ordered.
(14) Cranium, lateral and ventral perspectives, otic region, processus suprameaticus, osseus meatus acusticus externus, cotyla quadratica otici (or otic part of the
cotyla quadratica squamoso-otica), location relative to the more laterally positioned processus suprameaticus: 0. Cotyla is located caudal or caudoventral of the
processus suprameaticus, the rostral bounds of osseus meatus acusticus externus are caudally located; 1. Cotyla is rostrocaudally aligned with the processus
suprameaticus; 2. Cotyla is rostral of processus suprameaticus, the rostral bounds of osseus meatus acusticus externus are rostrally located. See Mayr (2020: p.
80). Ordered.
(15) Cranium, ventral perspective, otic region, separation of the cotyla quadratica otici and cotyla quadratica squamosi, development: 0. Cotylae widely separated; 1.
Cotylae are close together medially, width of separation increased laterocaudally; 2. Cotylae close together for entire border, narrowly separated; 3. Two cotyla
adjacent, separated only by a thin ridge; 4. Cotylae form a single articular surface (cotylae quadratica-squamosi-otica); Modified from Bourdon (2005, char. 50;
2011, char. 72), Livezey and Zusi (2006: char. 0150). Ordered.
(16) Cranium, lateral perspective, os exoccipitale, processus paroccipitalis, length of ventral extension from the cranium and placement relative to the os
parasphenoidale, lamina basisphenoidalis: 0. Processus paroccipitalis is near absent or short, terminates dorsad of, or in dorsoventral alignment with, the
lamina basisphenoidalis; 1. Processus paroccipitalis is short, terminates just ventral of the lamina parasphenoidalis; 2. Processus paroccipitalis has minor
elongation, terminates noticeably ventral of lamina parasphenoidalis; 3. Processus paroccipitalis is conspicuously elongate, extends considerably further
ventrally past the lamina parasphenoidalis. Modified from Bourdon (2005: char. 43, 108; 2011, char. 56), Livezey (1997: char. 3), and Worthy et al. (2017: char.
45). Ordered.
(17) Cranium, lateral, caudal and ventral perspectives, os exoccipitale, processus paroccipitalis, position of the connection of the ala parasphenoidalis to the
processus paroccipitalis: 0. Connects to the ventral-most processus paroccipitalis; 1. Connects just dorsal to the ventral apex of processus paroccipitalis; 2. The
ventral apex of processus paroccipitalis extends considerably ventral of its connection with ala parasphenoidalis. Ordered.
(18) Cranium, lateral and caudal perspectives, caudal cranium, crista nuchalis transversa and areas associated with insertions of m. complexus, m. biventer cervicis,
and m. splenius capitis, form: 0. Crista nuchalis transversa is visible yet relatively flat and confluent with curvature of cranium; 1. Crista nuchalis transversa is a
partially distinct ridge, accentuated caudally by paired depressions of muscle insertions in dorsal area; 2. Crista nuchalis transversa is a distinct ridge or flange
for its complete dorsoventral length due to expanded and ventrally undercutting depressions of muscle insertions along caudal margins. Modified from Livezey
& Zusi (2006: char. 0033), see also, Bourdon (2005: char. 90). Ordered.
(19) Cranium, caudal perspective, os supraoccipitale, fonticuli occipitalis, presence in adult condition: 0. Absent, 1. Present. Livezey (1997: char. 5), see also
Bourdon (2005: char. 45), Ericson (1997: char. 1), Livezey (1986: char. 9), Mayr & Clarke (2003: char. 27), Murray & Vickers-Rich (2004: table 8: char. 9),
Worthy & Lee (2008: char. 8), Worthy et al. (2017: char. 30). Note: In Anhimidae, the fontanelles are lost in adults (Ericson 1997). We code Nettapterornis as 1
considering the relevant statement in Olson (1999) and Conflicto as 1 due to Tambussi et al. (2019) stating the presence of fonticuli despite it not being clear in
the figure.
(20) Cranium, lateral and caudal perspectives, occipital region, prominentia exoccipitalis, status and form: 0. Absent or indistinct from processus paroccipitalis; 1.
Present, lowly raised caudal prominences; 2. Present, well-developed caudal prominences with distinct bulbous appearance.
(21) Cranium, caudal and ventral perspectives, occipital region, area encompassing the occipital region, ventral of the crista ventralis, degree of convexity or
concavity: 0. Flat or convex; 1. Slightly depressed; 2. Noticeably concave; 3. Noticeably concave, substantially accentuated by the extreme caudolateral
development of the exoccipital prominences. Ordered.
(22) Cranium, ventral perspective, occipital region, lamina parasphenoidalis, caudal region, tuberculum basilare, form: 0. Low, ridge-like tubercles, undifferentiated
from the crista basilaris lateralis and/or crista basilaris transversa; 1. Prominent, ventrally raised tubercles joined by crests aligned with the crista basilaris
lateralis and transversa, often right-angled in shape or profile; 2. Prominent, ventrally raised, rounded tubercle in lateral area; 3. Developed in association with,
and coincident with, the processus medialis parasphenoidalis. Modified from Worthy et al. (2017: char. 25). Note: see also Livezey & Zusi (2006: chars. 0065 and
0123), Parker (1895), Pycraft (1900: 172), Mayr & Clarke (2003: char. 30), Worthy & Scofield (2012: char 18).
(23) Cranium, caudal and ventral perspectives, occipital region, foramen magnum, shape: 0. Rounded, oval or near circular; 1. Rectangular, both dorsal and ventral
margins near perpendicular to lateral sides; 2. Triangular, dorsal margin distinctly apexed dorsally. See Livezey & Zusi (2006: char. 0027, 0028).
(24) Cranium, caudal and ventral perspectives, os parasphenoidale, lamina parasphenoidalis, bulla basitemporalis (sensu Davids 1952a, b): 0. Absent or indistinct, 1.
Present, not well developed; 2. Present, well-developed. Ordered.
(25) Cranium, caudal and ventral perspectives, os parasphenoidale, lamina parasphenoidalis, degree of overall ventral convexity: 0. Flat; 1. Rounded or inflated.
From Livezey and Zusi (2006: char. 117).
(26) Cranium, lateral perspective, os parasphenoidale, ala parasphenoidalis, shape or profile in lateral view: 0. Dorsally arced or concave lateral margin, often
directed rostrodorsally immediately rostral of processus paroccipitalis, proceeds rostrally on the horizontal (dorsoventral) plane; 1. Linear lateral margin
directed rostrally or rostroventrally from processus paroccipitalis; 2. Ventrally curved or convex, rounded flange. See also Worthy & Lee (2008: char. 12), and
Worthy et al. (2017: char. 20). Ordered.
(27) Cranium, ventral perspective, rostrum parasphenoidale, processus basipterygoideus, presence as distinct and developed process: 0. Absent; 1. Present.
Modified from Worthy et al. (2017: char. 27). Note: The presence of a process is assessed regardless of the presence of an articulatory surface (see below).
(28) Cranium, ventral perspective, rostrum parasphenoidale, articulatio pterygobasipterygoidea and facies articularis pterygoidea, articulation and contact between
pterygoid and rostrum parasphenoidale (through facies articularis pterygoidea), status: 0. Absent; 1. Present. Note: Some birds (e.g., Accipiter fasciatus) possess
a relatively vestigial processus basipterygoideus that is clearly present yet does not articulate with the pterygoid (coded as 0 here).
(29) Cranium, ventral perspective, rostrum parasphenoidale, articulatio pterygobasipterygoidea, facies articularis pterygoidea, shape and development of articular
surface: 0. Facies articularis pterygoidea is absent or vestigial; 1. Present, large and clearly ovoid articulatory facet, may be slightly tapered. 2. Facies articularis
pterygoidea is irregularly shaped and not flattened. Modified from Mayr & Clarke (2003: char. 24).
(30) Cranium, ventral and lateral perspectives, rostrum parasphenoidale, facies articularis pterygoidea, degree of relief with respect to rostrum parasphenoidale: 0.
Facies articularis pterygoidea is absent or vestigial; 1. Facies articularis pterygoidea is sessile and confluent with the rostrum parasphenoidale; 2. Facies
articularis pterygoidea is sessile and slightly raised with respect to the rostrum parasphenoidale, well defined; 3. Facies articularis pterygoidea is present and
slightly pedestalled upon a short processus basipterygoideus; 4. Facies articularis pterygoidea is present upon a pedicellate (stalked) processus basipterygoideus,
elongate. Modified from Worthy et al. (2017: char. 29), see also Murray & Vickers-Rich (2004: char. 13, table 8) and Mayr (2022). Ordered.
(31) Cranium, ventral perspective, rostrum parasphenoidale, facies articularis pterygoidea and/or the processus basipterygoideus, position: 0. Facies articularis
pterygoidea is rostrolaterally adjacent to lamina parasphenoidalis, caudal to rostrum parasphenoidale; 1. Facies articularis pterygoidea is just rostral of, and
distinctly separated from the rostral lamina parasphenoidalis on caudal-most rostrum parasphenoidale; 2. Facies articularis pterygoidea is located far rostrally
from the lamina parasphenoidalis, and rostral on rostrum parasphenoidale. Modified from Worthy et al. (2017: char. 28). Note: based on embryological
evidence, Weber (1993) interpreted that the processus basipterygoideus of Galloanserae was non-homologous with that of other Neornithes. However, further
assessment using a larger range of taxa is deemed necessary to more adequately test this (see Dzerzhinsky, 1995; Ericson, 1997; Zusi & Livezey, 2006), and were
considered conversely homologous by Worthy et al (2017: char. 28). Ordered.
 HISTORICAL BIOLOGY 1161

Lacrimal
(32) Lacrimal, dorsal perspective, synostosis with frontale and/or nasale: 0. Lacking; 1. Synostosed to frontale, caudal of craniorostral hinge; 2. Synostosed to nasale,
rostral of craniorostral hinge; 3. Synostosed to both nasale and frontale, extends across the craniorostral hinge. Modified from Worthy et al. (2017: char. 9), see
also Ericson (1997: char. 3), Livezey (1986, char. 10; 1996, char. 6; 1997: char. 15), Murray & Vickers-Rich (2004: char. 5, table 8), Stidham (2001: char. 21),
Worthy & Lee (2008, char. 4).
(33) Lacrimal, lateral perspective, processus orbitalis and processus supraorbitalis, relative dorsoventral and rostrocaudal length, respectively: 0. Processus orbitalis
is dorsoventrally shorter than the rostrocaudal length of processus supraorbitalis; 1. Respective lengths are near equal or processus orbitalis is slightly longer; 2.
The dorsoventral length of processus orbitalis far exceeds the rostrocaudal length of the processus supraorbitalis. Ordered.
(34) Lacrimal, lateral perspective, processus orbitalis, dorsoventral length of the processus orbitalis relative to the dorsoventral height of the rostrum: 0. Short,
especially so, does not approach the dorsoventral midline of the rostrum height; 1. Intermediate, approaches the dorsoventral midline of the rostrum height; 2.
Long, length approaches the dorsoventral height of the rostrum, approaches or overlaps the jugal arch. Ordered.

Quadrate
(35) Quadrate, dorsal and caudal perspectives, pars otica, capitulum oticum and capitulum squamosum, separation of the capitulum by vallecula intercapitularis: 0.
Absent, the two capitulae are fused to form a single articular head, the capitulum squamoso-otica, vallecula intercapitularis is absent, vestigial or indistinct; 1.
Cotylae are close together and the vallecula intercapitularis is mediolaterally narrow; 2. A significant space between cotylae exists, the vallecula intercapitularis
is mediolaterally wide. Modified from Worthy et al. (2017: char. 49), see also Bourdon (2011: char. 30, 35), Elzanowski & Stidham (2010: char. 7), Mayr &
Clarke (2003: char. 34), Murray & Vickers-Rich (2004: char. 14, table 8). Ordered.
(36) Quadrate, lateral perspective, pars otica, capitulum squamosum, degree to which articulatory surface of capitulum squamosum extends ventrally onto the
lateral surface of the pars otica, status: 0. Absent, dorsally confined; 1. Present, slight curve of articular surface onto lateral side; 2. Present, significant ventral
extension of the articular surface. Modified from Worthy & Lee (2008: char. 21) and Worthy et al. (2017: char. 57). Ordered.
(37) Quadrate, medial perspective, pars otica, dorsorostral of crista medialis, sulcus pneumaticus, status: 0. Nondescript surface, absent, or indeterminate; 1. Sulcus
pneumaticus is present, but indistinct; 2. Sulcus pneumaticus is present, and distinctly depressed. May be associated with Stidham (2001: char. 14). Note: see
Elzanowski & Stidham (2010: figure 5) regarding the presence of the sulcus pneumaticus with regards to a rostromedially positioned foramen pneumaticum.
The sulcus pneumaticus appears indeterminant or absent in all non-anhimid Anseriformes because the pneumatic foramen is positioned caudal of the crista
medialis, which contrastingly corresponds to depressio caudomedialis. Ordered.
(38) Quadrate, medial and caudal perspectives, pars otica, foramen pneumaticum caudomediale and foramen pneumaticum rostromediale, presence and position:
0. Both foramina are absent; 1. Foramen pneumaticum rostromediale is present only; 2. Both foramina are present; 3. Foramen pneumaticum caudomediale is
present only; 4. Foramen pneumaticum caudomediale is present only, but is rostrally positioned. See Elzanowski & Stidham (2010), Worthy & Scofield (2012:
char. 50 and 52), and Worthy et al. (2017: char. 51 and 52). Note: we follow Worthy et al. (2017) for the coding of Anhima cornuta, although differ from their
coding for Cereopsis novaehollandiae following Elzanowski & Stidham (2010: p. 315) who recognised that some taxa within the Anserinae sub-family and
Mergini tribe show a rostral displacement of the caudomedial pneumatic foramen. Ordered.
(39) Quadrate, lateral perspective, pars otica, tuberculum subcapitulare associated with the m. AME superficialis (and aponeurosis articularis), development: 0. Not
applicable, no visible corresponding feature on the quadrate; 1. Crested, follows the dorsoventral long axis of the pars otica; 2. Well-developed conical-shaped
tubercle; 3. Large tubercle, extremely well developed, dorsoventrally expanded. See Bourdon (2005: char. 19; 2011: char. 53), Livezey (1997: char. 49), Mayr &
Clarke (2003: char. 35), Stidham (2001: char. 17), and Worthy et al. (2017: char. 50). Note: The m. AME superficialis does not originate on the tuberculum
subcapitulare in Anseres. Due to differences in positions and spans of muscle origins, the superficially similar feature on the pars otica in the latter taxa instead
corresponds to m. AM posterior, and thus should be treated separately (see char. 39, below).
(40) Quadrate, lateral perspective, pars otica and processus orbitalis, position of origin for m. AM posterior: 0. On, or close to, processus orbitalis; 1. Pars otica. Note:
The origin of m. AME superficialis is associated with the tuberculum subcapitulare on the pars otica of the quadrate in most Neornithes, however, this muscle
originates more dorsally on the lateral cranium in Anatidae and does not have this same attachment to the quadrate. Instead, a superficially similar tubercle is
related to the origin of m. AM posterior in anatids, which is comparatively more ventral on the quadrate in other birds (e.g., see Davids, 1952a; Goodman &
Fisher, 1962; Zusi & Livezey, 2000).
(41) Quadrate, lateral perspective, pars otica and corpus quadrati, tuberculum or prominence associated with the m. AM posterior on the pars otica, status and
development: 0. Non-descript surface; 1. Present, low ridge; 2. Present, clearly crested eminence; 3. Present, well-developed and expansive tubercle. See
Bourdon (2005: char. 19; 2011: char. 53), Livezey (1997: char. 49), Mayr & Clarke (2003: char. 35), Stidham (2001: char. 17), and Worthy et al. (2017: char. 50).
Note: The conspicuous eminence associated with m. AM posterior on the processus orbitalis corresponds to the medial part of m. AM posterior (m. AM
posterior medialis), whereas attachment on the quadrate body or pars otica is related to the lateral part (m. AM posterior lateralis), in taxa which have clear
differentiation of this muscle into medial and lateral parts (e.g., see Fujioka, 1963; Goodman & Fisher, 1962; Weber, 1996; Zusi & Livezey, 2000; Holliday &
Witmer, 2007). While overall large and well developed, this differentiation is not observed in anhimid species, which possess an m. AM posterior that is
characteristic of the medial part only (Dzerzhinsky, 1982: 1034). Comparisons here should strictly observe the influence of the origin of m. AM posterior upon
the corpus quadrati and pars otica (most often related to m. AM posterior lateralis), rather than any corresponding to the more rostral origin on processus
orbitalis (m. AM posterior medialis or m. AM posterior). Ordered.
(42) Quadrate, lateral and medial perspectives, pars otica and processus orbitalis, relative lengths: 0. Pars otica is longer than processus orbitalis; 1. Pars otica and
processus orbitalis are subequal in length; 2. Pars otica is shorter than processus orbitalis. Ordered.
(43) Quadrate, lateral perspective, pars otica and processus orbitalis, orbital angle (the angle between the long axes of pars otica and processus orbitalis, sensu
Elzanowski & Stidham 2010: p. 310): 0. Obtuse, greater than 100 degrees; 1. Right or acute angle, 90 degrees or less. From Elzanowski & Stidham (2010: char. 6).
Note: This does not refer to the curvature of the dorsal profile between pars otica and processus orbitalis, but rather the orientation of the long axes of pars otica
and processus orbitalis).
(44) Quadrate, lateral perspective, processus orbitalis and pars otica, dorsal profile, curvature of the dorsal margin of the processus orbitalis as it intersects with the
rostral side of pars otica: 0. Deep curvature of the dorsal profile; 1. Marked, but weak, curvature of dorsal profile; 2. Minor curvature of the dorsal profile, little
distinction between dorsal margin of processus orbitalis and rostral margin of pars otica. Related to Livezey (1986: character 15), Ericson (1997: char. 11),
Worthy & Lee (2008: char. 22), Worthy et al. (2017: char. 58). Ordered.
(45) Quadrate, lateral perspective, processus orbitalis, shape: 0. Processus orbitalis is dorsoventrally broad across its rostrocaudal length (superficially rectangular);
1. Processus is triangular, dorsoventrally wider caudally and tapering rostrally to the apex; 2. Processus orbitalis is dorsoventrally narrow across its rostrocaudal
length. Related to Bourdon (2005: char. 94, 112), Dyke et al. (2003: char. 22), Ericson (1997: char. 11). Ordered.
(46) Quadrate, medial perspective, corpus quadrati, foramen pneumaticum basiorbitale, presence: 0. Absent or vestigial; 1. Present. Modified from Stidham (2001:
char. 13), Elzanowski and Stidham (2010: char. 1), and Worthy et al. (2017: char. 54). Note: some Chauna torquata have a vestigial foramen, see Elzanowski and
Stidham (2010: table. 4).
(47) Quadrate, rostral and medial perspectives, processus orbitalis and pars mandibularis, condylus pterygoideus and facies articularis pterygoidea, association: 0.
Facies articularis pterygoidea is separated from the condylus pterygoideus, a narrow articular surface or ridge between is variably present; 1. Facies articularis
pterygoidea and condylus pterygoideus are adjacent. Modified from Livezey and Zusi (2006: char. 600), Elzanowski and Stidham (2010: char. 4) and Worthy et
al. (2017: char. 56). Note: Elzanowski and Stidham (2010) suggested that the condition where the articular surfaces are adjacent is the plesiomorphic condition
(see also Worthy et al. 2017: char. 56).
(48) Quadrate, medial perspective, processus orbitalis and pars mandibularis, facies articularis pterygoidea, position: 0. Facies articularis pterygoidea is conspicuous
on the medial face; 1. Facies articularis pterygoidea is on the ventromedial margin of the ventral processus orbitalis, visible in medial perspective; 2. Facies
articularis pterygoidea is ventrally or ventrolaterally located on rostral pars mandibularis and partially on the ventral margin of processus orbitalis, not visible in
medial perspective. Related to Livezey and Zusi (2006: char. 600). Ordered.
1162 P. L. MCINERNEY ET AL.

(49) Quadrate, rostral perspective, pars mandibularis, condylus pterygoideus, prominence: 0. Flat, not developed; 1. Bulbous, broadly rounded tubercle; 2. Sharply
defined, subangular or conical tubercle. Modified from Bourdon (2011: char. 34) and Worthy et al. (2017: char. 56).
(50) Quadrate, rostral perspective, pars mandibularis, processus medialis, condylus pterygoideus and condylus mandibularis medialis, distinction and dorsoventral
separation between the ventral margin of condylus pterygoideus and the rostral margin of condylus mandibularis medialis: 0. Ventral margin of the condylus
pterygoideus is confluent with the rostral margin of the condylus mandibularis medialis (inseparable); 1. The two condyles are dorsoventrally separated by a
narrow furrow on the rostral pars mandibularis; 2. Dorsoventrally wide separation of the two condyles, condylus pterygoideus is located markedly dorsally.
Ordered.
(51) Quadrate, ventral perspective, pars mandibularis, mandibular condyles, number and presence of condylus mandibularis caudalis: 0. Three mandibular
condyles, condylus caudalis is present; 1. Bicondylar mandibular articulation, only condylus mandibularis lateralis and condylus mandibularis medialis are
present. See Bourdon (2005: char. 25; 2011: char. 1), Livezey (1997: char. 51), Murray & Vickers-Rich (2004: char. 16, table 8).
(52) Quadrate, ventral perspective, pars mandibularis, condylus mandibularis lateralis and processus orbitalis, orientation of the long axis of condylus mandibularis
lateralis relative to that of the processus orbitalis (and/or the main rostrocaudal alignment of the quadrate): 0. The long axes of the lateral mandibular condyle
and processus orbitalis are approximately aligned and parallel; 1. The caudal part of the lateral mandibular condyle is slightly rotated laterally, with its rostral
terminus angled relatively rostromedially, compared to the orientation orbital process (less than 180 degree angle formed); 2. The lateral mandibular condyle
has its long axis near perpendicular to that of the orbital process, producing a near 90 degree angle with respect to the main rostrocaudal alignment of the
quadrate. See also Livezey (1997: char. 52) and Murray & Vickers-Rich (2004: char. 17, table 8). Ordered.
(53) Quadrate, lateral perspective, pars mandibularis, processus lateralis, processus submeaticus and prominentia submeatica, presence: 0. Both the process and the
prominence are absent; 1. Only the prominentia submeatica is present; 2. Both the prominentia submeatica and processus submeaticus are present; 3. Only the
processus submeaticus is present. From Elzanowski & Stidham (2010: char. 3), see also Ericson (1997: char. 12) and Worthy et al. (2017: char. 48). Presbyornis
pervetus is coded as 0. Note: Following Elzanowski & Stidham (2010: fig. 8), we code anhimids as 2, despite whether the process is homologous with the
prominentia submeatica, as noted by Worthy et al. (2017: char. 48).
(54) Quadrate, lateral perspective, pars mandibularis, processus lateralis, fovea quadratojugalis: 0. A flat articular surface; 1. A shallow depression; 2. Distinctly
depressed; 3. A very deep fovea. Ordered.
(55) Quadrate, lateral and caudal perspective, pars mandibularis, processus lateralis, cotyla quadratojugalis, development of caudal or caudoventral margin: 0.
Margin is absent, fovea quadratojugalis is completely open caudoventrally, conspicuously notched appearance as a result; 1. Fovea quadratojugalis is partially
closed caudoventrally at the base of the fovea (medially) by a notched caudoventral rim; 2. Fovea quadratojugalis is completely enclosed caudoventrally by a
thin osseous rim; 3. Fovea quadratojugalis is completely enclosed, rim is uniformly well developed around fovea quadratojugalis. Modified from Worthy et al.
(2017: char. 59). For a discussion on this character, see Elzanowski and Stidham (2010). Ordered.
(56) Quadrate, lateral perspective, pars mandibularis, processus lateralis, cotyla quadratojugalis, facies articularis quadratojugalis dorsalis and facies articularis
quadratojugalis ventralis, presence: 0. Absent or indistinct; 1. Present, distinct.

Mandible
(57) Mandible, dorsal and ventral perspectives, rostrum mandibulae, ramus mandibulae, pars symphysialis, apex rostri, shape: 0. Rounded and bulbous rostral apex,
bilaterally broad and flared with respect to the more caudal mandibular rami, producing a conspicuously spatulate shape; 1. Rounded and spatulate at the
rostral apex, but not bilaterally widened relative to more caudal mandibular rami; 2. Mandibular rami are tapered to a blunt rostral apex; 3. Mandibular rami
are tapered to a pointed rostral apex. See also Livezey (1997: char. 22). Ordered.
(58) Mandible, rostral perspective, rostrum mandibulae, ramus mandibulae, pars symphysialis, apex rostri, degree of ventral convexity or curvature of dorsal
surface: 0. Dorsal surface is deeply ventrally convex, lateral sides curve steeply dorsally; 1. Shallowly centrally convex, with weak lateral curvature; 2. The dorsal
surface of the symphysial region is almost flat. Modified from Clarke (2002: char. 44), Mayr & Clarke (2003: char. 43). Ordered.
(59) Mandible, dorsal perspective, rostrum mandibulae, ramus mandibulae, pars symphysialis, rostrocaudal length of the symphysis relative to complete mandible
rostrocaudal length: 0. Short, restricted to within the most rostral quarter of the mandible; 1. Extends significantly caudally, greater than one quarter the
rostrocaudal length of the entire mandible.
(60) Mandible, dorsal and ventral perspectives, ramus mandibulae, lateral divergence of the mandibular rami as continued caudally from pars symphysialis: 0. None
or very little, mandibular rami are near parallel, inter-ramal width is consistently narrow; 1. Minor, slight diverging, inter-ramal width is narrow but gradually
increases caudally; 2. Marked, inter-ramal width significantly increases caudally. Ordered.
(61) Mandible, lateral perspective, ramus mandibulae, curvature of the mandibular rami: 0. Mandibular rami are arched (convex) dorsally; 1. Mandibular rami are
not substantially curved (relatively straight); 2. Mandibular rami are arched (convex) ventrally. Modified from Livezey (1996: char. 4; 1997: char. 17), Worthy &
Lee (2008: char. 23), and Worthy et al. (2017: char. 61). Ordered.
(62) Mandible, dorsal perspective, rostrum mandibulae, ramus mandibulae, pars intermedia and pars symphysialis, mediolateral width: 0. Mediolaterally narrow,
thin; 1. Mediolaterally wide, robust mandibular rami.
(63) Mandible, dorsal, medial and lateral perspectives, rostrum mandibulae, pars intermedia, margo tomialis and crista tomialis, form: 0. Margo tomialis is not
expanded, often sharp and crest-like; 1. Margo tomialis is slightly mediolaterally expansive relative to the more ventral ramus, producing a robust, rounded
edge; 2. Margo tomialis is conspicuously lateroventrally expanded and deflected, often rounded; 3. Margo tomialis is medioventrally expanded and deflected,
producing a flat and obliquely sloped tomial surface; 4. Margo tomialis is laterally expanded as nearing pars symphysialis to create a flattened surface. Note: a
marked crest is observed caudally, medioventral of the tomial margin in both Anhima cornuta and Chauna chavaria specimens, absent in other taxa examined.
However, it does bear some resemblance to the medioventrally expanded tomial margin that is characteristic of dromornithids, and may represent an less well-
developed form of such a structure.
(64) Mandible, lateral and ventral perspectives, rostrum mandibulae, ramus mandibulae, pars intermedia, mediolateral slant across the dorsoventral height of the
rami, ventral of tomial margin: 0. Mandibular rami are slanted medially as continued ventrally; 1. Mandibular rami have no mediolateral slant as continued
ventrally.
(65) Mandible, lateral perspective, rostrum mandibulae, ramus mandibulae, pars intermedia, groove that extends rostrocaudally along each lateral mandibular
ramus, ventral of margo tomialis, status and form: 0. Absent or only a slight rostrocaudally elongate depression; 1. Present, groove is dorsoventrally thin, and
often not well-defined; 2. Present, groove is dorsoventrally thin and well-defined for the complete rostrocaudal length of ramus mandibulae, pars intermedia; 3.
Present, groove is dorsoventrally broad. Modified from Ericson (1997: char. 17).
(66) Mandible, medial and lateral perspectives, pars intermedia, fenestra rostralis mandibulae, presence and form: 0. Absent; 1. Fenestra rostralis mandibulae is
open medially although completely covered laterally; 2. Fenestra rostralis mandibulae is open and ovoid laterally although completely covered medially; 3.
Fenestra rostralis mandibulae is open and ovoid laterally and open as a thin slit medially; 4. Fenestra rostralis mandibulae is open as a thin slit both medially and
laterally; 5. Fenestra rostralis mandibulae is open and conspicuously ovoid both medially and laterally. Note: degree of medial and lateral opening is related to
the interaction between the os dentale, os spleniale, os praearticulare and os suprangulare.
(67) Mandible, lateral perspective, ramus mandibulae, pars caudalis and pars intermedia, angulus dorsalis mandibulae and rostrum mandibulae, relationship
between angulus dorsalis mandibulae and the caudal margo tomialis: 0. Confluent, a gradual slope connects angulus dorsalis mandibulae and margo tomialis; 1.
Abrupt, a distinct, dorsoventrally short and variously rounded step is observed at the transition between margo tomialis and angulus dorsalis mandibulae; 2.
Strongly abrupt, considerable dorsoventral displacement is observed at the transition between margo tomialis and angulus dorsalis mandibulae. Ordered.
(68) Mandible, lateral perspective, ramus mandibulae, pars caudalis, angulus dorsalis mandibulae and processus coronoideus, relative dorsal elevation: 0. The dorsal
apex of processus coronoideus is dorsal to angulus dorsalis mandibulae; 1. The dorsal apex of processus coronoideus and that of angulus dorsalis mandibulae
exist on approximately the same dorsoventral plane; 2. The dorsal apex of processus coronoideus is ventral to that of angulus dorsalis mandibulae.
(69) Mandible, lateral perspective, ramus mandibulae, pars caudalis and pars intermedia, processus coronoideus, angulus dorsalis mandibulae and angulus ventralis
mandibulae, rostrocaudal positioning of angulus ventralis mandibulae: 0. Positioned markedly caudal to processus coronoideus; 1. Positioned slightly caudal or
 HISTORICAL BIOLOGY 1163

rostrocaudally aligned with processus coronoideus; 2. Positioned rostral of processus coronoideus and caudal of angulus dorsalis mandibulae; 3. Positioned
noticeably rostral of angulus dorsalis mandibulae. Note: taxa that do not possess an unambiguous angulus ventralis mandibulae (see character below) should be
coded as ‘?”. Ordered.
(70) Mandible, lateral perspective, ramus mandibulae, pars caudalis and pars intermedia, angulus dorsalis ventralis, prominence: 0. Ambiguous and not defined; 1.
Variably rounded; 2. Well defined, prominent. Ordered.
(71) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, fossa articularis quadratica, cotylae fossae articularis, number of cotylae: 0. Three cotylae; 1.
Two cotylae. Modified from Ericson (1997, char. 13), and Worthy et al. (2017: char. 60).
(72) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, fossa articularis quadratica, cotylae fossae articularis, orientation of cotylae medialis
mandibulae and cotylae lateralis mandibulae: 0. Cotylae are parallel, both rostrocaudally aligned; 1. Cotylae are nearly perpendicular, cotylae medialis
mandibulae oriented rostrocaudally; 2. Cotylae parallel, cotylae medialis mandibulae oriented mediolaterally. Modified from Andors (1988: char. 20) and Mayr
& Clarke (2003: char. 38).
(73) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, fossa articularis quadratica, cotylae fossae articularis, cotyla medialis, shape: 0. Mediolaterally
wide, rounded oval; 1. Relatively equal mediolateral and rostrocaudal widths; 2. Rostrocaudally elongate. Ordered.
(74) Mandible, dorsal, rostral and caudal perspectives, ramus mandibulae, pars caudalis, processus medialis mandibulae and fossa articularis quadratica,
rostromedial projection of cotyla medialis with respect to the profile of the rostral processus medialis mandibulae and the more ventral ramus mandibulae:
0. Absent, not defined rostromedially; 1. Rostromedial margin of the cotyla medialis is marked in dorsal view, but indistinct with respect to the more ventral
ramus mandibulae; 2. Rostromedial margin of the cotyla medialis is conspicuous in dorsal view, and distinctly projects to overhang the more ventral ramus
mandibulae. Ordered.
(75) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, processus medialis mandibulae, shape of rostromedial-most cotyla medialis mandibulae: 0. Not
arced to hemispherical or broadly rounded; 1. Subangular, almost right-angled.
(76) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, processus medialis mandibulae, orientation of medial-most area: 0. Caudally oriented; 1.
Completely medially projected; 2. Rostrally oriented. Ordered.
(77) Mandible, rostral and caudal perspectives, ramus mandibulae, pars caudalis, processus medialis mandibulae, degree of dorsal orientation as continued
medially: 0. Level, projects medially, does not project dorsally; 1. Projects mediodorsally; 2. Projects strongly dorsally. Note: A long, narrow, and dorsally
oriented processus was listed as a synapomorphy of Galloanserae by Cracraft & Clarke (2001, char. 41). Mayr & Clarke (2003, char. 45) found this state to be
more widely distributed, to the exclusion of Palaeognathae (Worthy et al. 2017: char. 68). Also modified from Bourdon (2011: char. 51). Also see character
below. Ordered.
(78) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, processus medialis mandibulae, length of the process medial of cotyla medialis mandibulae: 0.
Short; 1. Long.
(79) Mandible, dorsal, rostral and caudal perspectives, ramus mandibulae, pars caudalis, processus lateralis mandibularis, form: 0. Absent, the cotyla lateralis does
not project laterally; 1. Distinctly projected laterally, but confluent with the more ventral and rostral mandibular ramus; 2. Well-developed laterally and distinct
from the more rostral mandibular ramus. Modified from Ericson (1997: char. 16), Livezey (1997: char. 24), Murray & Vickers-Rich (2004: table 8, character 20).
Ordered.
(80) Mandible, lateral and dorsal perspectives, ramus mandibulae, pars caudalis, insertion of m. AME superficialis, aponeurosis paracoronoidea and corresponding
ossified structures, status and form: 0. Aponeurosis paracoronoidea is absent and therefore, so is any associated paracoronoid tubercle or crest; 1. Tuberculum
paracoronoideum is present on the dorsal edge of the mandible, caudal of processus coronoideus, and may extend somewhat laterally onto the lateral mandible;
2. Located on the lateral mandible as separate rostral (crista paracoronoidea rostralis) and caudal (crista paracoronoidea caudalis) crests, crista paracoronoidea
is rostrally extended and process-like in its development; 3. Located on the lateral mandible, crista paracoronoidea rostralis and caudalis are close together
dorsally, or touching at their dorsocaudal-most points to form an acute angle, respective rostral and caudal crests are perpendicular to near parallel to one
another; 4. Crista paracoronoidea is undivided, forms a prominent processus paracoronoideus. Note: see Weber & Hesse (1995) and Weber (1996) for details
regarding the aponeurosis paracoronoidea and associated crests or tubercula. The prominent processus paracoronoideus in anatids is analogous to the
prominently developed crista paracoronoideus rostralis of megapodiids (Weber & Hesse, 1995; Weber, 1996). The aponeurosis paracoronoidea is primarily
related to the caudally located m. AME superficialis. This part of the external adductor musculature is not distinct in Palaeognathae to the same degree it is in
Neognathae, where it is instead interrelated with other portions of the musculature (specifically the fibres associated with the more rostral deep part, m. AME
profundus, pars coronoideus); the lack of this distinct subdivision equates to a lack of any osseous structure on the mandible associated with it (see
Dzerzhinsky, 1983; Elzanowski, 1987; Weber & Hesse, 1995; Weber, 1996).
(81) Mandible, lateral perspective, ramus mandibulae, pars caudalis, processus retroarticularis, caudal development and projection: 0. Absent or present only as a
rostrocaudally short, dorsoventrally aligned crest; 1. Present as a rostrocaudally short projection which extends caudally no more than its dorsoventral height at
its rostral origin; 2. Present as a caudally long osseous projection, where total caudal length is greater than the dorsoventral height at its rostral origin but less
than twice this height; 3. Present as a caudally long osseous projection, where total caudal length is greater than twice the dorsoventral height at its rostral
origin. Modified from Worthy et al. (2017: char. 64). Ordered. Note: Because of the wide taxonomic range being treated here, the lack of a processus
retroarticularis must be a considered state for the following characters related to this structure.
(82) Mandible, lateral perspective, ramus mandibulae, pars caudalis, processus retroarticularis, degree of dorsal margin curvature: 0. Not applicable, processus
retroarticularis is absent; 1. The dorsal margin is not curved dorsally as continued caudally; 2. The dorsal margin broadly or subtly curves dorsally as continued
caudally; 3. The dorsal margin is conspicuously curved dorsally as continued caudally, particularly towards the caudal tip. Modified from Bourdon (2011: char.
52), Livezey (1997: char. 20), Mayr & Clarke (2003: char. 44), Murray & Vickers-Rich (2004: table 8), Worthy & Lee (2008: char. 25). Note: observations should
be restricted to the dorsal margin of the retroarticular process for this character, and not the ventral margin.
(83) Mandible, lateral perspective, ramus mandibulae, pars caudalis, processus retroarticularis, shape: 0. Processus retroarticularis is absent; 1. Dorsoventrally
narrow and acicular (needle-like); 2. Dorsoventrally splayed and broad.
(84) Mandible, dorsal perspective, ramus mandibulae, pars caudalis, processus retroarticularis, direction/angle from which the processus retroarticularis extends
caudally: 0. Process retroarticularis is absent; 1. Directed laterocaudally; 2. Extends caudally in line with the angle of the mandibular rami. Modified from
Murray & Vickers-Rich (2004: table 8) and Worthy & Lee (2008: char. 25).
(85) Mandible, caudal perspective, ramus mandibulae, pars caudalis, processus medialis mandibularis, form of caudal surface, status of fossa caudalis and
development of recessus conicalis: 0. Fossa caudalis is absent, the caudal processus medialis mandibularis is flat, dorsoventrally narrow; 1. Fossa caudalis
absent, caudal processus medialis mandibularis is flat, dorsoventrally expansive; 2. Fossa caudalis is present and shallow, dorsoventrally expansive; 3. Fossa
caudalis is present and deep, dorsoventrally expansive; 4. Fossa caudalis is significantly depressed, a large and deep recessus conicalis is present, dorsoventrally
expansive. Modified from Ericson (1997: char. 14), Livezey (1997: char. 21), Stidham (2001: char. 22), Worthy & Lee (2008: char. 26), Worthy et al. (2017: char.
65). Note: we code Presbyornis pervetus and Nettapterornis oxfordi based on descriptions by Olson (1999). Ordered.

Rostrum
(86) Rostrum, co-ossification of rostral elements (particularly, maxillaria, nasalia and praemaxillaria): 0. Absent or partial; 1. Present.
(87) Rostrum, dorsal perspective, rostrum maxillae, apex rostri, shape: 0. Rostral end is broadly rounded, spatulate; 1. Rostral end is laterally spatulate or rounded,
with a pointed rostral apex; 2. Tapered to a point. Modified from Livezey (1997: char. 32), Bourdon (2005: char. 46), and Ericson (1997: char. 10). Ordered.
(88) Rostrum, lateral perspective, culmen, dorsal profile of caudal two-thirds of rostrum (caudal of rostrum maxillae): 0. Non-arced or linear to very subtly dorsally
convex; 1. Dorsally arced throughout the rostrocaudal length; 2. Strongly dorsally convex throughout the rostrocaudal length. Ordered.
(89) Rostrum, lateral perspective, rostrum maxillae, profile of the dorsal rostrum maxillae towards its rostral apex: 0. Dorsally convex, uniform with respect to the
more caudal rostrum; 1. Linear, not arced, uniform with respect to the more caudal rostrum; 2. Flattens and dorsoventrally levels with respect to more caudal
rostrum, may appear ventrally arced; 3. Conspicuously dorsally inflated at the rostral apex.
1164 P. L. MCINERNEY ET AL.

(90) Rostrum, lateral perspective, apertura nasi ossea, development and span: 0. Very large, comprises most of, or a large proportion of, the lateral sides of the rostrum;
1. Large, comprises nearly two-thirds of the lateral side of the rostrum, the rostrocaudal length of the rostrum rostral of the apertura nasi ossea is no more than half
the rostrocaudal length of the apertura nasi ossea itself; 2. Comprises approximately one-half of the lateral side of the rostrum, the rostrocaudal length of the
rostrum rostral of the apertura nasi ossea is equal to or slightly greater than the rostrocaudal length of the apertura nasi ossea itself; 3. Small, caudally restricted,
approximately one-third to one-quarter the rostrocaudal length of the rostrum; 4. Very small, well restricted to within the caudal half of the rostrum, less than one-
fifth of the rostrocaudal length of the rostrum. Modified from Murray & Vickers-Rich (2004: char. 2, table 8); Bourdon (2005: char. 101). Ordered.
(91) Rostrum, lateral perspective, apertura nasi ossea, positioned significantly dorsally on rostrum: 0. Absent, centrally positioned dorsoventrally; 1. Present.
(92) Rostrum, ventral perspective, palatal surface, palatum osseum, os palatinum, degree of palatal ossification and closure of the fenestra palatina: 0. Very little to
no ossification, the fenestra palatina is large; 1. Partial but marked ossification, fenestra palatina is large; 2. Near complete palatal ossification, the fenestra
palatina is slit-like or reduced to a small opening. Modified from Worthy et al. (2017: char. 7). Ordered.
(93) Rostrum, ventral perspective, palatal surface, palatum osseum, fenestra palatina, closure caudally: 0. Absent, fenestra palatina is open caudally, continuous with
choana nasalis ossea; 1. Present, fenestra palatina is caudally closed through fusion of the vomer, palatines, and/or praemaxillary bones; 2. Present, fenestra
palatina is caudally closed through medial fusion of the processus maxillopalatinus of the maxillary bones.
(94) Rostrum, lateral and ventral perspectives, ossa palatina and ossa vomeris, fusion along the mediolateral midline caudally (synostosis interpalatina, sensu Zusi &
Livezey, 2006), with or without the vomer: 0. Absent; 1. Present, medial palatines are directly fused to one another or fused to, and separated by, a mediolaterally
thin vomer; 2. Present, medial palatines are fused to the vomer, which widely mediolaterally separates the palatines. Note: medial fusion is characteristic of
anseriform taxa, although not universally, to the exclusion of galliforms (see Zusi & Livezey, 2006). Modified from Worthy et al. (2017: char. 36).
(95) Rostrum, ventral and lateral perspectives, os palatinum and os maxillare, position of the interaction between the processus rostralis of the palatinum and
processus palatinus of the maxillare relative to the processus jugalis of the maxillare: 0. The rostral processus rostralis palatini is medially separated and ventral
of the rostral processus jugalis of the maxillare; 1. The rostral processus rostralis palatini is medially separated and slightly ventral of the rostral processus jugalis
of the maxillare; 2. The rostral processus rostralis palatini converges with the rostral processus jugalis of the maxillare, and is slightly ventral to it; 3. The rostral
processus rostralis palatini converges with the rostral processus jugalis of the maxillare, and is dorsal to it. Modified from Ericson (1997: char. 9), Worthy et al.
(2017: char. 33). Note: This character is derived from observations of Zelenkov & Stidham (2018). State 2 can also be associated with a concavity that forms
lateral of the processus rostralis palatini. Ordered.
(96) Rostrum, lateral perspective, arcus jugalis, os quadratojugale and processus jugalis of the os maxillare, arcus jugalis forms a synovial articulation with the
rostrum: 0. Absent, the elements are fused; 1. Present.
(97) Rostrum, palatum osseum, palatal type: 0. Desmognathous; 1. Schizognathous; 2. Other. See Mayr & Clarke (2003: char. 11), Murray & Vickers-Rich (2004:
char. 10, table 8), and Zusi & Livezey (2006).
(98) Rostrum, ventral and lateral perspectives, os palatinum, pars lateralis, form and development: 0. Pars lateralis absent, a slightly enlarged lateral margin (crista
lateralis) may exist; 1. Pars lateralis is poorly differentiated; 2. Pars lateralis is present but not well developed, does not extend far laterally or ventrally; 3. Pars
lateralis is lateroventrally or ventrally flared, does not extend ventrally past the level of the tomial margin; 4. Pars lateralis extends considerably ventrally beyond
the level of the tomial margin. See Livezey & Zusi (2006: p. 158); also relevant is Bourdon (2005: char. 68), Livezey (1997: char. 41, 42), Mayr & Clarke (2003:
char. 16), and Worthy et al. (2017: char. 34). Ordered.
(99) Rostrum, ventral and lateral perspectives, os palatinum, pars lateralis, formation of angulus caudolateralis palatini: 0. Absent; 1. Present.
(100) Rostrum, lateral and ventral perspectives, os palatinum, fusion to os pterygoideum (in adult birds): 0. Absent, 1. Present. From Worthy et al. (2017: char. 41).
Also see Worthy & Scofield (2012), where the pterygoid is unfused or incompletely fused to the palatines in osteological immature dinornithiforms, and
Benito et al. (2022).

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