Warren D. Handley

Osteology Supports a Stem-Galliform Affinity for the Giant Extinct Flightless Bird Sylviornis neocaledoniae (Sylviornithidae, Galloanseres).

The giant flightless bird Sylviornis neocaledoniae (Aves: Sylviornithidae) existed on La Grande Terre and Ile des Pins, New Caledonia, until the late Holocene when it went extinct shortly after human arrival on these islands. The species was generally considered to be a megapode (Megapodiidae) until the family Sylviornithidae was erected for it in 2005 to reflect multiple cranial autapomorphies. However, despite thousands of bones having been reported for this unique and enigmatic taxon, the postcranial anatomy has remained largely unknown. We rectify this deficiency and describe the postcranial skeleton of S. neocaledoniae based on ~600 fossils and use data from this and its cranial anatomy to make a comprehensive assessment of its phylogenetic affinities. Sylviornis neocaledoniae is found to be a stem galliform, distant from megapodiids, and the sister taxon to the extinct flightless Megavitiornis altirostris from Fiji, which we transfer to the family Sylviornithidae. These two species form the sister group to extant crown-group galliforms. Several other fossil galloanseres also included in the phylogenetic analysis reveal novel hypotheses of their relationships as follows: Dromornis planei (Dromornithidae) is recovered as a stem galliform rather than a stem anseriform; Presbyornis pervetus (Presbyornithidae) is the sister group to Anseranatidae, not to Anatidae; Vegavis iaai is a crown anseriform but remains unresolved relative to Presbyornis pervetus, Anseranatidae and Anatidae. Sylviornis neocaledoniae was reconstructed herein to be 0.8 m tall in a resting stance and weigh 27–34 kg. The postcranial anatomy of S. neocaledoniae shows no indication of the specialised adaptation to digging seen in megapodiids, with for example, its ungual morphology differing little from that of chicken Gallus gallus. These observations and its phylogenetic placement as stem galliforms makes it improbable that this species employed ectothermic incubation or was a mound-builder. Sylviornis neocaledoniae can therefore be excluded as the constructor of tumuli in New Caledonia.

Sexual dimorphism in the late Miocene mihirung Dromornis stirtoni (Aves: Dromornithidae) from the Alcoota Local Fauna of central Australia

ABSTRACT The dromornithids were giant flightless birds endemic to Australia from the late Paleogene to the late Pleistocene. Dromornithids are generally considered to be divergent members of the Anseriformes, but they display many convergent features with extant ratites. In this study, we investigate Dromornis stirtoni from the Alcoota Local Fauna, a species for which little is known of its biology. We used traditional methods of comparative morphology, mass estimation, landmark-based morphometrics, and histological investigations to determine the presence of medullary bone, to assess the possible presence, form, and extent of sexual dimorphism in D. stirtoni. Two morphological groups were identified for each main leg element, differing primarily in relative robustness. Core samples from femora and tibiotarsi shafts revealed medullary bone in the less robust morph, indicating that these were females. Mass, as estimated by algorithms applied to our preferred measurement of least-shaft circumference of tibiotarsi, was significantly different between males (mean = 528 kg) and females (mean = 451 kg). Therefore, male D. stirtoni were more robust but not much taller than the females and challenge the elephant bird, Aepyornis maximus, for the title of the most massive bird to have existed. Sexual dimorphism in this largest of all dromornithids, therefore, was like that of extant Anseriformes. We infer long-term monogamy, mutual display, shared parental care, female incubation, and aggressive defense of nests in these birds. The techniques of geometric morphometrics applied in this study maximize the use of fragmentary material, helping to overcome the common paleontological challenge of limited sample sizes.

The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)

The extinct dromornithids, gastornithids and phorusrhacids are among the most spectacular birds to have ever lived, with some giants exceeding 500 kg. The affinities and evolution of these and other related extinct birds remain contentious, with previous phylogenetic analyses being affected by widespread convergence and limited taxon sampling. We address these problems using both parsimony and tip-dated Bayesian approaches on an expansive taxon set that includes all key extinct flightless and flighted (e.g. Vegavis and lithornithids) forms, an extensive array of extant fowl (Galloanseres), representative Neoaves and palaeognaths. The Paleogene volant Lithornithidae are recovered as stem palaeognaths in the Bayesian analyses. The Galloanseres comprise four clades inferred to have diverged in the Late Cretaceous on Gondwana. In addition to Anseriformes and Galliformes, we recognize a robust new clade (Gastornithiformes) for the giant flightless Dromornithidae (Australia) and Gastornithidae (Eurasia, North America). This clade exhibits parallels to ratite palaeognaths in that flight presumably was lost and giant size attained multiple times. A fourth clade is represented by the Cretaceous Vegavis (Antarctica), which was strongly excluded from Anseriformes; thus, a crucial molecular calibration point needs to be reconsidered. The presbyornithids Wilaru (Australia) and Presbyornis (Northern Hemisphere) are robustly found to be the sister group to Anatoidea (Anseranatidae + Anatidae), a relatively more basal position than hitherto recognized. South Americas largest bird, Brontornis, is not a galloansere, but a member of Neoaves related to Cariamiformes; therefore, giant Galloanseres remain unknown from this continent. Trait analyses showed that while gigantism and flightlessness evolved repeatedly in groups, diet is constrained by phylogeny: all giant Galloanseres and palaeognaths are herbivores or mainly herbivorous, and giant neoavians are zoophagous or omnivorous.

The extinct flightless mihirungs (Aves, Dromornithidae): cranial anatomy, a new species, and assessment of Oligo-Miocene lineage diversity

ABSTRACT Giant flightless fowl (Aves, Dromornithidae) similar to the Northern Hemisphere gastornithids and weighing up to 350–650 kg evolved on Gondwana and existed in what is now Australia from the Eocene to the late Quaternary. Understanding cranial morphology of dromornithids has until now been based almost wholly on species of Dromornis, with that of species in three other genera either previously unknown or very fragmentary. Here we rectify this deficiency and describe a well-preserved cranium from the middle Miocene Bullock Creek Local Fauna referred to Ilbandornis woodburnei, rich, fragmentary crania, quadrates, pterygoids, and mandibles for the Oligo-Miocene Barawertornis tedfordi Rich, and additional material of the species of Ilbandornis. The morphological similarity of this cranial material suggests that the emusized B. tedfordi is a smaller precursor to and differs little from species of Ilbandornis. Dromornis murrayi, n. sp., from late Oligocene—Early Miocene sites at Riversleigh, based on cranial and postcranial elements, is the oldest and smallest species in its genus. Placed in the context of other data, these observations suggest that the dromornithids comprised only two lineages throughout the Oligo-Miocene. The Barawertornis-Ilbandornis lineage attained maximum diversity in the middle Miocene Bullock Creek and late Miocene Alcoota local faunas (LF), with two species in each, but the Dromornis lineage seems to have been monotypic throughout its temporal range. The low diversity of these giant galloanseres in Australia mirrors that of the giant herbivorous ratites (ostriches and kin), which similarly have low diversity where they coevolved with diverse mammalian faunas.

Flightless rails (Aves: Rallidae) from the early Miocene St Bathans Fauna, Otago, New Zealand

While known for over a decade to exist, fossil rails of the early Miocene (19–16 Ma) St Bathans Fauna, from the South Island of New Zealand, have not previously been described taxonomically or studied in detail. Here we use qualitative osteological features and analyse measurements from wing and leg bones to determine the number of taxa represented, their flight ability, and the presence and nature of sexual dimorphism within the identified taxa. We describe two new rail species in monospecific genera from the St Bathans Fauna: Priscaweka parvales gen. et sp. nov., which is extremely common, and Litorallus livezeyi gen. et sp. nov., a distinctly larger, uncommon species. Priscaweka parvales exhibited a significant degree of sexual dimorphism and was tiny, being the size of the extinct Chatham Island Rail Cabalus modestus. Both newly described species exhibit skeletal proportions and osteological features that indicate they had reduced wings and were flightless. These observations reveal that flightless rallid species have been present in New Zealand for millions of years. The distinctiveness of the St Bathans rails from their closest geographical and chronological neighbours suggests some hidden diversity of volant rails in Australias fossil record. However, the combined data from Australasian and European records reveal no evidence for a diverse early Miocene crown rallid fauna as predicted by some molecular studies. A subsequent, middle Miocene radiation for crown rallids seems more likely, and appears to have produced the high taxonomic diversity seen in Holocene Australasian rail faunas. http://zoobank.org/urn:lsid:zoobank.org:pub:9F638A1E-C17D-4A85-9D6E-3D0F24D5AE1E

Correction to ‘The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)'

[This corrects the article DOI: 10.1098/rsos.170975.].

Details on the mass, diet and stratigraphic ages for the taxa analysed, Worthy et al Galloanseres from The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)

An excel file giving the mass, diet, and stratigraphic ages and supporting data sources for the taxa used in the analyses.

Apomorphy lists from the parsimony analyses for Figure 2B, Worthy et al. galloanseres from The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres)

A text document giving the apomorphy lists from parsimony analyses resulting in the tree shown in Figure 2B, where a backbone constraint was employed, no characters were weighted, and all 48 taxa were included.