![Saintandrea chenoides, gen. et sp. nov. from the late Oligocene of Saint-André in southern France (holotype [part], NMB Mar. 874a–j), coracoid (a–e) and carpometacarpus (h–k) in comparison . S. chenoides, omal half of left coracoid in dorsal (a), dorsolateral (b), medial (c), lateral (d), and ventral (e) view. Romainvillia stehlini, left coracoid (lectotype, NMB P.G. 38a) in dorsal (f) and medial (g) view. S. chenoides, proximal portion of right carpometacarpus in dorsal (h), caudal (i), cranial (j), and ventral (k) view. R. stehlini, left carpometacarpus (NMB P.G. 53) in cranial (l) and ventral](https://www.researchgate.net/profile/Vanesa-De-Pietri/publication/235105082/figure/fig1/AS:393367881043971@1470797619521/Saintandrea-chenoides-gen-et-sp-nov-from-the-late-Oligocene-of-Saint-Andre-in_Q320.jpg)
![Saintandrea chenoides, gen. et sp. nov. from the late Oligocene of Saint-André in southern France (holotype [part], NMB Mar. 874a–j), selected bones in comparison to large Oligocene anseriform and phoenicopteriform birds. S. chenoides (holotype [part], NMB Mar. 874a), omal half of left coracoid in dorsal (a) and medial (b) view. Cygnopterus affinis (holotype [part], IRSNB Av 2), extremitas omalis of left coracoid in dorsal (c) and medial (d) view. Adelalopus hoogbutseliensis (holotype [part], IRSNB Av 71), extremitas omalis of right coracoid in dorsal (e) and medial (f) view. g A. hoogbutseliensis (holotype [part], IRSNB Av 71), fragmentary distal end of left tarsometatarsus in dorsal view. h S.](https://www.researchgate.net/profile/Vanesa-De-Pietri/publication/235105082/figure/fig2/AS:393367881043972@1470797619595/Saintandrea-chenoides-gen-et-sp-nov-from-the-late-Oligocene-of-Saint-Andre-in_Q320.jpg)
![Tarsometatarsus of Saintandrea chenoides, gen. et sp. nov. from the late Oligocene of Saint-André in southern France (holotype [part], NMB Mar. 874a–j). Proximal end of right tarsometatarsus in cranial (a), caudal (b), lateral (c), medial (d), and proximal (e) view. f, g distal end of right tarsometatarsus in plantar (f) and dorsal (g) view. Distal end of left tarsometatarsus in dorsal (h), plantar (i), medial (j), lateral (k), and distal (l) view. m distal end of right tarsometatarsus of Romainvillia stehlini in distal view (NMB P.G.](https://www.researchgate.net/profile/Vanesa-De-Pietri/publication/235105082/figure/fig3/AS:393367881043973@1470797619631/Tarsometatarsus-of-Saintandrea-chenoides-gen-et-sp-nov-from-the-late-Oligocene-of_Q320.jpg)
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A goose-sized anseriform bird from the late Oligocene of France: The youngest record and largest species of Romainvilliinae
Abstract and Figures
We describe a new anseriform bird from the late Oligocene of Saint-André, Marseille, in southern France. Saintandrea
chenoides, gen. et sp. nov. is the first avian species reported from the locality, which is well known for its mammalian fossils. The new species belongs to the extinct Romainvilliinae and represents the latest occurrence of the taxon, which was before only known from the late Eocene and early Oligocene of Europe. S.
chenoides is also the largest species of Romainvilliinae and increases the known morphological diversity of the taxon. The identification of a goose-sized representative of the Romainvilliinae in the late Oligocene of Europe raises the possibility that some of the large late Paleogene or early Neogene Anseriformes with uncertain phylogenetic affinities also belong to this taxon.
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... Additionally, it lacks the prominent tubercle laterodistal to the pons supratendineus reported by Musser and Clarke (2022) for their new anseriform. "T." howardae also differs from the late Oligocene Romainvillidae Saintandrea chenoides (Mayr and De Pietri, 2013) and the crown Anatidae by possessing a proportionally broader, blunt, and less developed condylus medialis. Finally, "T." howardae differs from Cygnavus senckembergi (early Miocene) (Lambrecht, 1931) as its tibiotarsus is proportionally narrower, with condyles more medially located and with a broader pons supratendineus. ...
... As for the genus. (Cracraft, 1970) Mayr and De Pietri (2013). ...
... The mosaic of characters displayed by the tibiotarsus of Wunketru prevents an unquestionable familial assignment. Some characters, such as the distal opening of the canalis extensorius located in the middle of the incisura intercondylaris and the proportional width and length of the pons supratendineus, align it with observations in Anatidae and Romainvillidae (see Mayr and De Pietri, 2013). However, the condylus medialis is less developed than in these taxa. ...
Telmabates" howardae, identified from a tibiotarsus discovered in the early Eocene Las Flores Formation in Chubut Province, Argentina, was initially classified under the extinct family Presbyornithidae. However, this classification was later dismissed, and it was deemed incertae sedis. In this study, we suggest a new genus for "Telmabates" howardae and evaluate its relationships within Anseriformes. Its familial relationships remain unclear due to the location of the distal opening of the canalis extensorius, the shape of the sulcus m. fibularis (smooth and laterally bounded by a ridge, similar to that of Saintandrea or Dendrocygna), and the medial position of the condyles. However, unlike Anatidae and Romainvillidae, the condylus medialis is smaller and blunt, whereas in these taxa, it is large and narrow. This suggests that this species was more terrestrial and less adapted to swimming than Anatidae.
... Globally, the earliest appearance of undoubted anatids, albeit in low diversity, was in the late Eocene (MP20, ca 35 Ma)-early Oligocene of Europe and Asia, with species attributed to Romainvilliinae (Romainvillia stehlini Lebedinsky, 1927; Romainvillia kazakhstanensis Zelenkov, 2018; Saintandrea chenoides Mayr & De Pietri, 2013) (see Mayr 2008, 2022, Zelenkov 2018. These taxa are usually treated as the subfamily Romainvillinae and considered primitive anatids (e.g., Mayr & De Pietri 2013, Mayr 2022; however, Zelenkov (2018) suggested they should be recognized at the family level as they are more plesiomorphic than Dendrocygna and represent an intermediate form between presbyornithids and anatids. ...
... Globally, the earliest appearance of undoubted anatids, albeit in low diversity, was in the late Eocene (MP20, ca 35 Ma)-early Oligocene of Europe and Asia, with species attributed to Romainvilliinae (Romainvillia stehlini Lebedinsky, 1927; Romainvillia kazakhstanensis Zelenkov, 2018; Saintandrea chenoides Mayr & De Pietri, 2013) (see Mayr 2008, 2022, Zelenkov 2018. These taxa are usually treated as the subfamily Romainvillinae and considered primitive anatids (e.g., Mayr & De Pietri 2013, Mayr 2022; however, Zelenkov (2018) suggested they should be recognized at the family level as they are more plesiomorphic than Dendrocygna and represent an intermediate form between presbyornithids and anatids. It should be noted, however, that Mayr & De Pietri (2013) had already observed that romainvillines were too derived to be ancestral to anatids, and Mayr & Smith (2001) had reported that they lived alongside anatids in the early Oligocene of Belgium. ...
... These taxa are usually treated as the subfamily Romainvillinae and considered primitive anatids (e.g., Mayr & De Pietri 2013, Mayr 2022; however, Zelenkov (2018) suggested they should be recognized at the family level as they are more plesiomorphic than Dendrocygna and represent an intermediate form between presbyornithids and anatids. It should be noted, however, that Mayr & De Pietri (2013) had already observed that romainvillines were too derived to be ancestral to anatids, and Mayr & Smith (2001) had reported that they lived alongside anatids in the early Oligocene of Belgium. Nevertheless, these early anatids and other potential stemanatids (e.g., Cousteauvia kustovia Zelenkov, 2020b) could be evidence for their evolution from presbyornithids in Europe or Asia (Zelenkov 2018(Zelenkov , 2020a(Zelenkov , 2020b. ...
The taxa found in an Eocene deposit, near Murgon, Queensland, the only pre-Oligocene Paleogene site recording a terrestrial vertebrate fauna from Australia, are very significant for the insight they provide concerning the evolution of the Australian biota. Here we resolve the identity of fossils previously referred to the Graculavidae, waterbirds of then unresolved affinities. We taxonomically describe the first bird to be named from this fauna, Murgonornis archeri gen. et sp. nov., Presbyornithidae. Our findings reveal that presbyornithids were widespread globally in the earliest Eocene, and that this family had a history on Australia from at least 55 Ma until ca. 24 Ma, when they disappear from the fossil record.
Trevor H. Worthy [trevor.worthy@flinders.edu.au], College of Science and Engineering, Flinders University, GPO Box 2100, Adelaide 5001, SA, Australia; Vanesa L. De Pietri [vanesa.depietri@canterbury.ac.nz], University of Canterbury, School of Earth and Environment, Private Bag 4800, Christchurch 8140, New Zealand; R. Paul Scofield [pscofield@canterburymuseum.com], Canterbury Museum, 11 Rolleston Avenue, Christchurch 8013, New Zealand; Suzanne J. Hand [s.hand@unsw.edu.au], Earth and Sustainability Science Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW 2052, Australia.
... This structure of the hypotarsus is definitively more derived than in Anhimidae and fossil Presbyornithidae, which have only one medial sulcus (Mayr and Smith 2017). The presence of closed medial hypotarsal canal distinguishes the new taxon from fossil Paranyroca Miller andCompton, 1939 (Paranyrocidae, Mayr andSmith 2017), which otherwise have similar outlines of the hypotarsus, though a similarly-built hypotarsus with a canal might have been present in Saintandrea, a large-sized representative of Romainvilliidae, but the referred specimen is not sufficiently preserved to confirm it (Mayr and De Pietri 2013). Although affinities of the new taxon with Romainvilliidae seem probable, the type genus Romainvillia Lebedinsky, 1927 has a morphologically rather different tarsometatarsus, more similar to that of Anatidae (Mayr 2008). ...
... The shaft is notably narrowed relative to the proximal end, being roughly similar to the condition in Presbyornithidae and Anseranatidae but unlike Anhimidae, Romainvillia and Anatidae sensu lato. A similarly widened proximal end might have also been present in large romainvilliid Saintandrea De Pietri, 2013 (Mayr andDe Pietri 2013). The cross-sectional area of the mid-shaft is sub-trapezoid. ...
... The shaft is notably narrowed relative to the proximal end, being roughly similar to the condition in Presbyornithidae and Anseranatidae but unlike Anhimidae, Romainvillia and Anatidae sensu lato. A similarly widened proximal end might have also been present in large romainvilliid Saintandrea De Pietri, 2013 (Mayr andDe Pietri 2013). The cross-sectional area of the mid-shaft is sub-trapezoid. ...
A specialized diving lifestyle has repeatedly evolved in several lineages of modern and fossil waterfowl (Anseriformes), with the oldest previously known representative being the late Oligocene Australian oxyurine ducks Pinpanetta. However, diving specializations have never been previously documented for any of the primitive Paleogene anseriforms ( “stem-anatids”), and thus may be associated with the origin of modern anatid-like body plan. Here I describe a tarsometatarsus of a new duck-sized diving anseriform bird from the latest Eocene (late Priabonian) Kusto Svita in Eastern Kazakhstan, which predates the previously reported occurrence of diving specialization in Anseriformes by at least 6 MA. The new taxon Cousteauvia kustovia gen. et sp. nov. has an unusual and previously undocumented morphology, but partly resembles the stem-anatids Paranyrocidae and Romainvilliidae, thus representing the first known occurrence of diving adaptations in primitive non-anatid anseriforms. The evolutionary appearance of specialized waterfowl taxa in the late Eocene of Central Asia supports a view that this region might have played an important role in the evolution of morphologically derived Anseriformes. The structure of the intertarsal joint in basal and modern anseriforms is here further discussed in relation with adaptations for aquatic locomotion. The presence of elongate and evenly narrow condyles of the tibiotarsus in Anatidae and other swimming/diving birds allows a firm contact with the hyperprotracted tarsometatarsus at the initial phase of the propulsion. This morphology contrasts with the restricted condyles of Presbyornithidae, which indicate a different, strictly wading locomotory specialization. Cousteauvia obviously evolved diving specializations on the basis of a more primitive structure of the intertarsal joint.
... Today, the morphologically derived family Anatidae (ducks, geese, swans, and related forms, including the recently proposed Dendrocygnidae; Sun et al., 2017) is the most diversified and comprises at least 159 recognized living species (Dickinson and Remsen, 2013). Undoubted anatids are known since the early Oligocene (Harrison and Walker, 1979;Mayr and Smith, 2001;Mayr, 2009Mayr, , 2017, and some of the earliest (Oligocene to early Miocene) fossil forms differ morphologically from their modern counterparts and thus are often classified within the extinct family-level taxa Romainvilliinae and Paranyrocidae (Lambrecht, 1933;Miller and Crompton, 1939;Mayr, 2008;Mayr and De Pietri, 2013;Mayr and Smith, 2017;Zelenkov, 2018). Crown-group Anatidae have an especially rich fossil record that dates to at least the late Oligocene (Worthy, 2008;Zelenkov, 2012), and the oldest undoubted members of Anhimidae and Anseranatidae are also known since that time (Alvarenga, 1999;Worthy and Scanlon, 2009). ...
... In all Anseriformes and most extant birds, this process is notably more distally positioned, aligning with the proximodistal midpoint of the processus extensorius (e.g., Fig. 2F) or sometimes even closer to its distal margin. Another distinctive feature of the holotype carpometacarpus is that the trochlea carpalis is shifted somewhat caudally: in Anseriformes, including Presbyornithidae and Romainvilliinae (e.g., Kurochkin and Dyke, 2010;Mayr and De Pietri, 2013), the cranial margin of the trochlea carpalis is located level with the cranial margin of the major metacarpal or even protrudes somewhat more cranially. A caudally shifted trochlea carpalis is present (Olson, 1999) in Anatalavis oxfordi but not in extant Anseranas semipalmata. ...
... Naranbulagornis khun differs from Romainvilliinae (represented by Saintandrea chenoides and Romainvillia stehlini; Mayr and De Pietri, 2013) in the craniocaudally wider processus extensorius, in lacking the reduced dorsal rim of the trochlea carpalis, in the proximally shifted processus pisiformis, and in the caudally shifted trochlea carpalis. Thus, the overall organization of the proximal carpometacarpus in Romainvilliinae is like that in Anatidae, whereas the morphology in N. khun is more similar to that in Anhimidae and partly similar to that in Anseranas semipalmata. ...
The early evolutionary history of waterfowl (Anseriformes) is poorly understood. Aside from the morphologically aberrant Presbyornithidae, the only other known early Paleogene anseriform taxon is Anatalavis oxfordi from the early Eocene of England. Here, I describe two bones from the late Paleocene of southern Mongolia (localities Naran-Bulag and Tsagaan-Khushuu), which belong to a very large swan-sized swimming waterfowl (Anseriformes) and are described as Naranbulagornis khun, gen. et sp. nov. This is the first non-presbyornithid anseriform bird found in the early Paleogene (Paleocene through early Eocene) of Asia and one of the few currently known Paleocene birds from Central Asia. It is also the largest volant bird from the Paleogene of Asia. Naranbulagornis khun documents a mosaic of autapomorphic and plesiomorphic features that are otherwise present in modern Anhimidae, Anseranatidae, and Anatidae, indicating a greater morphological diversity of early Cenozoic anseriforms than previously thought. The new taxon is morphologically closer to Anatoidea than to Presbyornithidae and represents the earliest global evidence of a spatial coexistence between presbyornithids and other basal waterfowl during the early Paleogene.
... The oldest definitely "duck-like" anseriforms are Romainvillia stehlini from the Late Eocene of France (Lebedinsky, 1927) and Romainvillia kazakhstanensis from Kazakhstan (Zelenkov, 2018). Romainvilliinae are also known from the Early Oligocene of Belgium (Mayr and Smith, 2001), England (Mayr 2008) and France (Mayr and De Pietri, 2013). The oldest representatives of the crown group Anatidae date back to the late Oligocene. ...
... In any case, the unusual morphology of this bird suggests that, like Romainvilliinae, it does not belong to the crown group Anatidae. In the late Oligocene in Europe, stem ducks continue to exist and are represented, in particular, by large (goose-sized) representatives of Romainvilliinae (Mayr and De Pietri, 2013). However, in the second half of the Late Oligocene (~26-24 million years ago), representatives of the crown group of ducks also appear in the fossil record: in Australia, these are stem representatives of the Oxyurinae (Pinpanetta with the three described species) and the alleged oldest shelduck bird, Australotadorna (Worthy, 2009). ...
An analysis of the Eurasian Cenozoic (late Eocene-Neogene) fossil record of anatids (Anatidae s. l.; including Romainvilliinae and Dendrocygninae) is presented. The evolutionary origin of Anatidae s. l. may be associated with the appearance of large shallow waterbodies in Asia during the Late Eocene as a result of the fall in the global sea level and the resulting retreat of the epicontinental seas. Four major temporal stages can be recognized in the evolution of the Cenozoic Eurasian anatids communities, without any traceable continuity between particular stages (at the current stage of knowledge). Some recent anatid genera (e.g., Tadorna) first appear in the paleontological record at the level of the early and middle Miocene (17-15 Ma), but temperate faunas of essentially modern ecological composition became widespread only in the late Mio-cene (9-6 Ma). The details of transitions between various faunistic stages, as well as the origin of modern communities, remain largely unstudied.
... If the new Paranyroca specimen is of early Miocene rather than late Oligocene age (what we consider most likely, see introduction), it is one of the strati-graphically youngest European anseriform fossils, for which a position outside crown group Anatidae needs to be considered. Anseriformes outside crown group Anatidae are, however, dominant in the Paleogene of Europe (MAyr 2009(MAyr , 2017, and a phylogenetic placement in the anatid stem group was also assumed for the Romainvilliinae, which include Romainvillia stehlini from the late Eocene of the Paris Basin and Saintandrea chenoides from the late Oligocene of Marseille, France (MAyr 2008;MAyr & De Pietri 2013). Unfortunately, critical parts of the tarsometatarsi of R. stehlini and S. chenoides are damaged, but Paranyroca is distinguished from R. stehlini in that the trochlea metatarsi II forms a less well developed plantar projection, the trochlea metatarsi IV is mediolaterally narrower, and the distal end of the trochlea metatarsi IV runs more obliquely relative to the longitudinal axis of the bone. ...
A tarsometatarsus of a large anseriform bird from the late Oligocene/early Miocene of the Saint-Gérand-le-Puy area in France is assigned to the distinctive taxon Paranyroca, which was before only known from the early Miocene of North America. Paranyroca may be a stem group representative of Anatidae and its tarsometatarsus exhibits a peculiar character mosaic, with an Anhimidae-like hypotarsus and an Anatidae-like distal end. With regard to these features, the bone resembles the tarsometatarsus of another anseriform bird from Saint-Gérand-le-Puy, which was described as Cygnopterus alphonsi but has not yet been compared with Paranyroca in the original description. C. alphonsi was originally identified as a swan, but this classification has subsequently been contested and here it is proposed that the species is more closely related to Paranyroca.
We describe nearly complete skeletons of basal Anseriformes from the Latest Paleocene to the early Eocene of North America and Europe. Collectively, these birds appear to be representative of anseriforms near the divergence of Anhimae and Anseres, but their exact positions relative to these clades remains uncertain. A new family, Anachronornithidae nov. fam., is erected on the basis of one of these, Anachronornis anhimops nov. gen., nov. gen. et sp., to which the others cannot be confidently assigned. The new fossils augment a growing collection of early Pan-Anseriformes, which in their diversity do not paint an unambiguous picture of phylogeny or character state evolution on the path to or within crown-Anseriformes. Anachronornis nov. gen. is similar in some aspects of both cranial and postcranial anatomy to other well-represented early Paleogene Anseriformes and members of Anseres, such as Presbyornis Wetmore, 1926. However, it exhibits a more landfowl-like bill, like that of Anhimae and unlike the spatulate bill of Anseres. Additional specimens of similar basal Anseriformes of uncertain affinities from the early Eocene of North America and Europe further complicate interpretation of character state polarity due to the mosaicism of primitive and derived characters they exhibit.
Galloanseres, the clade including galliform and anseriform birds, is supported by virtually all analyses of different kinds of molecular data and also resulted from analyses of morphological data. A recent study obtained an early Eocene date for the split of galliform and anseriform birds, some 55 million years. However, this divergence estimate is in clear conflict with the fossil record, which includes morphologically disparate stem group representatives of both Galliformes and Anseriformes from deposits of that age (anseriform birds have an even earlier fossil record). Extant Galloanseres are mainly characterized by morphological apomorphies that concern skull features. The postcranial skeleton of extant Galliformes and Anseriformes is quite different, but the morphology of Paleogene stem group Galliformes bridges the morphological gap between the extant taxa. Various ecomorphologically disparate Paleogene taxa were assigned to the Galloanseres. The morphological and ecological diversity within Paleogene Galloanseres, therefore, appears to have been extraordinarily high, including giant flightless ground birds with greatly reduced wings, long-legged filter-feeders, and pelagic taxa with wingspans of 4–5 m.
A new species of the extinct duck Romainvilla from the Upper Eocene of Kazakhstan is described. This is the earliest duck from Asia, the first record of Romainvilla beyond France and the first reliable evidence of the presence of Romainvillinae (regarded here as family) in Asia. This occurrence shows a wide range of Romainvilla and suggests a greater similarity of Late Eocene faunas of Western Europe and Asia than it was previously thought. The origin of Romainvillidae and their presumable descendants Anatidae may be connected with adaptation to a new biotope, the shallowing Late Eocene epicontinental Asian basins (due to a global decrease in sea level).
We investigate the phylogenetic afnites of the New Zealand blue duck (Hymenolaimus malacorhynchos), a riverine specialist of uncertain relationships, using 2613 bp of DNA sequence data from 3 mitochondrial genes. Hymenolaimus has variously been considered an aberrant Anas species, or an ancient taxa in the tribe Anatini. Presently, it is placed in a highly-derived clade (Tribe Merganetini) with the shelducks. Our fndings show that Hymenolaimus forms a monophyletic clade, and does not ft within any of the other duck tribes around the world. Our study also confrms convergent evolution among duck species that inhabit fast fowing rivers.
Avian remains from the lowermost Oligocene of Hoogbutsel (Belgium) are described. The material includes a new species of loon (Gaviiformes, Gaviidae), Colymboides belgicus n. sp., which is the largest Lower Tertiary representative of the Gaviidae known so far. A new taxon of the Palaelodidae (Phoenicopteriformes) is described as Adelalopus hoogbutseliensis n. gen. n. sp., and is unusual in that the caudal surface of the extremitas sternalis of the furcula bears a deep cavity. A distal tarsometatarsus of a small falconiform bird is tentatively assigned to the Accipitridae. Another distal end of a tarsometatarsus belongs to a strigiform bird and most closely resembles Prosybris antiqua Milne-Edwards 1867-71. In addition, mousebirds (Coliiformes, Coliidae) are represented by a fragmentary distal tarsometatarsus. A new taxon of the Sylphornithidae (Aves incertae sedis) is described as Oligosylphe mourerchauvireae n. gen. n. sp.: It is the first Oligocene record of this family which before was only known from the Upper Eocene of France.