A new large representative of the Anseriformes, Garganornis ballmanni n. gen. et n. sp., from the Miocene of Gargano, Italy, is described from the distal end of a left tibiotarsus. G.ballmanni displays morphological features that are characteristic of the Anseriformes, but differs from all extant and fossil Anseriformes, as well as from taxa closely related to Anseriformes. G.ballmanni is characterized by a very wide and shallow fossa intercondylaris, a less pronounced difference in width of the lateral and medial condyle, reduced epicondyli, and a circular opening of the distal canal. G.ballmanni is larger than any living member of Anseriformes, and with an estimated body mass between 15.3 and 22.3 kg, it was most likely flightless. Although the observed similarities between G.ballmanni and basal taxa might indicate that G.ballmanni represents an insular relict of a stem lineage, it is considered more likely that G.ballmanni was a species of waterfowl highly adapted to a terrestrial lifestyle in an insular environment where mammalian carnivores are rare.
Un nouveau grand représentant des ansériformes, Garganornis ballmanni n. gen. et n. sp., est décrit sur la base de l’extrémité distale d’un tibiotarse gauche dans le Gargano (Miocène, Italie). G.ballmanni présente des traits morphologiques qui sont caractéristiques des ansériformes, mais diffèrent de tous les ansériformes actuels et fossiles, ainsi que des taxons étroitement liés aux ansériformes. G.ballmanni est caractérisé par une fosse antérieure intercondylienne large et peu profonde, une différence peu prononcée dans la largeur des condyles latéral et médial, des épicondyli réduits, et une ouverture circulaire du canal distal. G.ballmanni est plus grand que tous les ansériformes actuels, et, avec une masse estimée entre 15,3 et 22,3 kg, il était probablement incapable de voler. Bien que les similitudes observées entre G.ballmanni et les taxons de base puissent indiquer que G.ballmanni représente une relique insulaire d’une lignée primitive, il est considéré comme plus probable que G.ballmanni ait été une espèce d’oiseau aquatique très adaptée à un mode de vie terrestre, dans un environnement insulaire où les mammifères carnivores étaient rares.
Anseriformes, Gargano, Italy, Miocene, Insular evolutionAnsériformes, Gargano, Italie, Miocène, Évolution insulairepresentedPresented by Philippe TaquetIntroduction
Now firmly joined to the Italian mainland, the Gargano promontory (Fig. 1) once formed part of the Apulia platform and hosted a highly endemic insular fauna. Sediments from palaeokarst fissures have yielded extensive collections of Neogene and Pleistocene vertebrate remains (Abbazzi et al., 1996, Freudenthal, 1971, Freudenthal, 1976, Pavia et al., 2012 and Van der Geer et al., 2010). Although the timing and mode of colonization of the Apulia platform has been the subject of debate (see Freudenthal et al., 2013, Mazza and Rustioni, 2008 and Van den Hoek Ostende et al., 2009), the Neogene assemblage, also known as the ‘Mikrotia’ fauna after the abundant endemic murid (Freudenthal, 2006), is now considered to be Tortonian in age (Freudenthal et al., 2013 and Van den Hoek Ostende et al., 2009). It documents the arrival and evolution of an endemic fauna that, with the exception of the otter Paralutra garganensis, is devoid of mammalian carnivores and displays strong insular evolution. Examples of endemic lineages include the giant Gargano hedgehog Deinogalerix, the Gargano dormouse Stertomys and the deer-like ruminant Hoplitomeryx (Daams and Freudenthal, 1985, Freudenthal, 1972, Leinders, 1984, Masini and Fanfani, 2013, Van der Geer, 2008 and Villier et al., 2013).
The Neogene fossil bird assemblage of Gargano was first described by Ballmann, 1973 and Ballmann, 1976, who identified 16 avian taxa, most of them endemic, in the orders Galliformes, Anseriformes, Accipitriformes, Strigiformes, Columbiformes, Apodiformes, Piciformes and Passeriformes. Göhlich and Pavia (2008) described the small phasianid Palaeortyx volans, the most abundant species in the assemblage. Birds of prey were abundant and varied greatly in size, with the largest being the buteonine hawk Garganoaetus freudenthaliBallmann, 1973 (the size of a Golden Eagle) and the barn owl Tyto giganteaBallmann, 1973 (larger than the Eurasian eagle owl B.bubo (Ballmann, 1976)). With mammalian carnivores being rare (note that P.garganensis is thought to have fed on shellfish (Willemsen, 1983)), birds filled the empty niche of predator (Ballmann, 1973 and Ballmann, 1976). Recent studies by Pavia, 2011 and Pavia, 2013 added several new taxa to the Gargano avifauna, including a species of Calidris and two new anatids, and brought the avifauna of Gargano to at least 26 taxa. Here, I describe a hitherto undescribed species of Anseriformes from the Miocene of Gargano that was discovered during a reorganisation of the Gargano collection at the Naturalis Biodiversity Center in Leiden, the Netherlands.
Material and methods
The specimen, a distal left tibiotarsus (RGM 443307) was compared to a number of extinct and extant Anseriformes (see Appendix) in the collections of the Naturalis Biodiversity Center (RMNH) in Leiden (the Netherlands), the British Museum of Natural History (BMNH) in London (UK), and the Smithsonian Institution's National Museum of Natural History (NMNH) in Washington, D.C. (USA). Comparisons to fossil species were mostly based on the literature. Measurements were taken with sliding callipers. Terminology follows Baumel and Witmer (1993), and Howard (1929) for structures not named in Baumel and Witmer. To estimate a body mass for this specimen, the minimum circumference of the tibiotarsus was estimated by wrapping a thin strap of paper around the tibiotarsus at the most proximal point and measuring the minimum circumference with sliding calipers calibrated to the nearest 0.05 mm. The minimum circumference was then log10-transformed and body mass was estimated by using the least-squares linear regression for anseriform tibiotarsi as described by Iwaniuk et al. (2004).
Systematic Palaeontology
Class AVES Linnaeus, 1758
Order ANSERIFORMES Wagler, 1831
Family Insertae sedis
GARGANORNIS n. gen.
Type species. Garganornis ballmanni n. sp.
Derivation of name. Masculine, after the Italian region Gargano where the quarries are located, and ‘ornis’, Greek for bird.
Diagnosis. Monospecific genus, as for species diagnosis.
Garganornis ballmanni n. sp.
(Fig. 2)
Derivation of name. ‘ballmanni’ refers to Peter Ballmann who was the first to describe the fossil avifauna of Gargano.
Holotype. Distal left tibiotarsus (RGM 443307), Fig. 2.
Diagnosis. A very large representative of the order Anseriformes characterized by the following unique combination of characters; central placement of the canalis extensorius, medially displaced condylus medialis, fossa intercondylaris very wide and shallow, distal opening of canalis extensorius circular, a less pronounced difference in width of the condylus lateralis and condylus medialis, condylus medialis projecting further cranially than condylus lateralis, a reduced epicondylus lateralis and a modestly developed epicondylus medialis, a reduced sulcus m. fibularis, a deep sulcus extensorius and the lateral half of pons supratendineus sunken.
Type locality. Posticchia 5 fissure filling, Gargano pedemountain belt (41.8° N, 15.4° E), Province of Foggia, Italy.
Description
The holotype of Garganornis ballmanni (RGM 443307, Fig. 2) consists of the distal end and part of the shaft of the left tibiotarsus. The bone surface is cream coloured with reddish stains. Overall, the bone has a heavy and stout appearance. Measurements: maximal length of fragment: 64.4 mm; maximal distal width: 37.1 mm; (proximodistal) height of tendinal bridge: 6.8 mm; minimum width of shaft: 21.51 mm; minimum depth of shaft: 15.8 mm.
The cortical bone wall is thick, ranging from 3–3.5 mm on the facies caudalis to 7–8 mm on the facies cranialis. In cross-section, the facies cranialis is flat and the facies caudalis is convex caudally. The sulcus extensorius (Fig. 2) is deep and located on the central axis of the shaft. The sulcus extensorius is medially bordered by a pronounced tuberositas retinaculum extensoris medialis, which slightly overhangs the sulcus. The pons supratendineus is high and attains a slightly medial position on the bone. Although its orientation is horizontal, the lateral half of the pons supratendineus is sunken and forms a depression upon merging with the shaft. Disto-laterally of this depression lies the triangular tuberositas retinaculum extensoris lateralis. Proximal of the tuberositas retinaculum extensoris lateralis lays the tuberculum retinaculi m. fibularis medialis. The tuberculum retinaculi m. fibularis lateralis is hardly visible. Although there is very slight abrasion of the bone surface, it appears that the tuberculum retinaculi m. fibularis lateralis has not been very distinct. The canalis extensorius opens distally of the sulcus extensorius as a circular foramen centred immediately proximal to the fossa intercondylaris and at equal distances from either of the condyli. On the medial and lateral sides of the distal opening of the canalis extensorius lay the attachment sites for the medial and lateral crurae of the ligamentum meniscotibialis, with the medial point of attachment more pronounced than the lateral one. The fossa intercondylaris is very wide, approximately 2/3 of the maximal width across the condyli, and in distal view attains a relaxed U shape rather than a V shape. The condylus medialis is offset medially from the shaft. The caudal portions of both condyli are missing so that condylar shape and depth cannot be determined. Of the condylus lateralis also the cranial and distal portion are damaged. The epicondylus lateralis itself is eroded as well, but the depressio epicondylaris lateralis is still mostly intact. This suggests that only a limited part of the lateral side of the condylus lateralis is missing. The outline in distal view of the condylus lateralis is reconstructed in Fig. 2c. The condylus medialis is more intact, with only its caudal side missing. The epicondylus medialis is more pronounced, and the depressio epicondylus medialis is also deeper than the depressio epicondylus lateralis. The impressio ligamentum collateralis medialis is prominent and extends proximally well up the shaft. Despite the damage, the width of the condyles is interpreted as the condylus lateralis being somewhat wider than the condylus medialis. In distal view (Fig. 2c), the condylus medialis projects further anteriorly than the (reconstructed) condylus lateralis. The facies caudalis bears very few distinctive features as the condyli lack the caudal portion.
Discussion
The distal left tibiotarsus described here represents a new genus and species of Anseriformes, G. ballmanni, based on distinct morphological characteristics. G. ballmanni represents the third, and largest, anseriform from the Miocene of Gargano, in addition to Anas velox and Anatidae indet (Pavia, 2013). Little is known regarding the origin of the Miocene Gargano avifauna, but there are indications that the Gargano avifauna is related to that of the Middle Miocene avifauna of La Grive-Saint-Alban in France. Ballmann (1973) considered the buteonine eagle Garganoaetus murivorus, to be closely related to Aquila delphinensis of the La Grive avifauna. More recently, Göhlich and Pavia (2008) described a new species of phasianid from the Neogene of Gargano, Palaeortyx volans, which they considered to be closely related to P.grivensis from La Grive. Unfortunately, no species of anseriform is known from La Grive-Saint-Alban (Ballmann, 1969). G. ballmanni represents a large-bodied addition to the Miocene avifauna from Gargano. The tibiotarsus of G.ballmanni is 30% larger than that of the largest extant anseriform, Cygnus olor (Table 1). Based on the minimum circumference of the tibiotarsus fragment, the estimated body mass for G.ballmanni is 22.3 kg, which is much larger than that of C. olor (Dunning, 2008 and Iwaniuk et al., 2010). Note that the minimum circumference of the fragment is not the minimum circumference of the bone itself. Assuming a minimum (conservative) circumference for the bone of approximately 20% smaller than that of the fragment gives a body mass estimate of 15.3 kg, which is closer to, but still larger than, the range reported for Cygnus olor (9.2–14.3 kg, Dunning, 2008). No wing bones that could potentially belong to G. ballmanni have been found so far, but it is reasonable to assume that G. ballmanni led a mostly terrestrial lifestyle and may have had a limited ability to fly. The large size of G. ballmanni mirrors that of other Gargano birds, such as the hawk Garganoaetus murivorus and the barn owl Tyto gigantea (Ballmann, 1973), and is most likely an adaptation to an insular environment where mammalian carnivores are rare or completely absent. Note that while insular birds of prey seem to increase in size and fill the empty niche of mammalian predator, insular waterfowl often increase in size and become major terrestrial herbivores. The most remarkable examples of gigantism in insular waterfowl are the Hawaiian waterfowl, i.e. the moa-nalo and giant Hawaii geese, which developed gigantism, flightlessness and terrestrial herbivory in the absence of mammalian herbivores (Paxinos et al., 2002). G. ballmanni might have followed a similar scenario that led to large body size and a lifestyle characterized by terrestrial herbivory.
Phylogenetic relationships
The medially deflected condylus medialis, the cranial projection of the condylus medialis, and the central placement of the canalis extensorius over the fossa intercondylaris of G. ballmanni n. gen. et n. sp. are considered synapomorphies of Anseriformes (Ericson, 1997, Livezey, 1986), and place G. ballmanni firmly in this group. However, G. ballmanni also displays characters that are reminiscent of the large-bodied extinct basal lineages of Dromornithidae, Gastornithidae and Presbyornithidae. Whereas Gastornithidae (Late Palaeocene to Middle Eocene of Europe and Early Eocene of North America (Mayr, 2009)) are considered a sister group to Anseriformes (Andors, 1992), Dromornithidae (Palaeogene and Neogene of Australia (Murray and Vickers-Rich, 2004)) and Presbyornithidae (Palaeogene of the New World as well as Mongolia and the London Clay (Kurochkin and Dyke, 2010 and Mayr, 2009)) have been grouped within crown-group Anseriformes (Ericson, 1997, Livezey, 1997 and Murray and Vickers-Rich, 2004). However, it has been pointed out that Presbyornithidae share morphological features with more basal groups (Elzanowski and Stidham, 2010) and might be outside Anseriformes. Upper Paleocene and Early Eocene Gastornis tibiotarsi from France have been described by Buffetaut (1997; 2008). G. ballmanni shares several features with Gastornis, such as a wide fossa intercondylaris, the circular opening of the distal canal, a deep sulcus extensorius and a pons supratendineus with a depression on the lateral end (Buffetaut, 2008) (Fig. 3a), but it differs from Gastornis in lacking the strong peroneal ridge (the sulcus m. fibularis) and having the distal opening of the canalis extensorius placed at equal distances from both condyles (a more medial position in Gastornis). The similarities between Gastornis and G. ballmanni are interesting, but the presence of Gastornithidae in Miocene Gargano is considered unlikely, as there is no evidence for the survival of Palaeogene faunal elements in the Gargano fauna. Furthermore, this scenario would require Gargano to have been dry land since at least the Middle Eocene, which is unlikely in this tectonically active region. G. ballmanni shares with Presbyornithidae (based on the descriptions by Ericson (2000)) the medial thrust of the distal end, a pronounced tuberositas retinaculum extensoris medialis and a broad and steep pons supratendineus, a deep sulcus extensorius and a poorly developed epicondyli medialis and lateralis (although the epicondylus medialis is more distinct than the epicondylus lateralis in G. ballmanni), but G. ballmanni differs from Presbyornithidae in the absence of a well-marked sulcus m. fibularis with a foramen (present in Presbyornithidae) and a less pronounced difference in width of the condyli. Dromornithid tibiotarsi display the general anseriform bauplan with a medially displaced condylus medialis and a centrally placed distal opening of the canalis extensorius (Murray and Vickers-Rich, 2004). A more detailed study of the basal dromornithid Barawertornis tedfordi (Nguyen et al., 2010) shows a fossa intercondylaris that is deeper and narrower than in G. ballmanni. In addition, Nguyen et al. (2010) describe B. tedfordi as having condyli that are about equal in cranial extent, and the condylus medialis being wider than the condylus lateralis, which is in contrast to the character states observed in G. ballmanni.
Crown-group Anseriformes include the South American Anhimidae (screamers), the Anseranatidae (magpie geese) of Australia and New Guinea, and the globally distributed Anatidae (swans, geese and ducks) (Ericson, 1997, Hackett et al., 2008, Livezey, 1997 and Worthy et al., 1997). Despite their global distribution and range of sizes and shapes, G. ballmanni differs from all extant Anseriformes by a significantly wider and shallower fossa intercondylaris, a less marked difference in width between the condylus medialis and condylus lateralis, and a circular distal opening of the canalis extensorius. Both Anhimidae and Anseranatidae display distinct tuberculi retinacula m. fibularis medialis and lateralis, forming a clear sulcus m. fibularis, which is absent in G. ballmanni. Anhimidae also display characteristic skeletal pneumatisation (Alvarenga, 1999 and O’Connor, 2004), but there is no sign of pneumatisation in G. ballmanni. Interestingly, within extant Anseriformes, G. ballmanni mostly resembles the Anserinae; they display a relatively wide fossa intercondylaris, and a less distinct sulcus m. fibularis (Fig. 3b–d). In addition, in cranial view, the medial side of the shaft that flares towards the medial condyle starts more proximally in Anserinae than in other groups.
Several fossil Anserinae have been described from the Late Eocene to the Pleistocene of Europe, but only a few taxa have the tibiotarsus preserved. The earliest representatives of Anatidae are the Romainvilliinae from the Late Eocene to the early Late Oligocene (Lebedinsky, 1927, Mayr, 2008 and Mayr and De Pietri, 2013). A large, goose-sized representative of the Romainvilliinae from the Late Oligocene of France, Saintandrea chenoides, was described by Mayr and De Pietri (2013). The tibiotarsus of S. chenoides displays a distinct attachment for the distal fibula, absent in G. ballmanni, and a narrow fossa intercondylaris (wider in G. ballmanni). From the Lower Miocene of Germany comes Cygnavus senckenbergiLambrecht, 1931, but this species is much smaller than G. ballmanni (width at the height of the pons supratendineus is 16 mm (Lambrecht, 1931)), versus 27.4 mm in G. ballmanni. The Early Miocene Cygnopterus alphonsiCheneval, 1984, from Saint Gerand le Puy displays a very broad distal opening of the canalis extensorius, unlike G. ballmanni, and is smaller. Cygnavus formosus, from the Oligocene of Kazakhstan (Kurochkin, 1968), is smaller than G. ballmanni, has an elliptical distal opening of the canalis extensorius and a more narrow fossa intercondylaris. The Middle Pleistocene giant swan Cygnus falconeri, endemic to Malta and Sicily (Parker, 1867) is similar to G. ballmanni in size (Table 1), but differs from the Gargano specimen in a number of morphological features; the distal opening of the canalis extensorius is oval in C. falconeri and is set more distally with respect to the medial condyle than in G. ballmanni (Fig. 4). In addition, the intercondylar fossa in G. ballmanni is much wider than in C. falconeri, which becomes evident in distal view (Fig. 4e–f). Interestingly, G. ballmanni also shares similarities with the fossil geese from the Hawaiian archipelago. The Big Island Goose Branta rhuax (Olson, 2013) and the moa-nalo Thambetochen chauliodous are both large, flightless geese and display a relatively wide fossa intercondylaris (although less wide in T. chauliodous). Branta rhuax also displays a reduced difference in condylar width, similar to the condition in G. ballmanni. However, the tuberculum retinaculi m. fibularis lateralis is well developed in B. rhuax and T. chauliodous, and B. rhuax also displays a well-marked sulcus m. fibularis. A broad fossa intercondylaris and a reduced difference in condylar width were also observed in the extinct, flightless New Zealand goose Cnemiornis calcitrans.
Conclusion
G. ballmanni n. gen. et n. sp. displays morphological features that are characteristic for Anseriformes, but differs from all extant and fossil Anseriformes, as well as from taxa closely related to Anseriformes. Given the marked differences between G. ballmanni and related taxa, as well as the fact that important diagnostic features of the condyles are not preserved, I consider an “incertae sedis” position most appropriate. It should be noted however, that insular evolution can result in morphologies that deviate from the ancestral state, and make it difficult to trace a taxon's ancestry. The observed similarities between G. ballmanni and basal taxa might indicate that G. ballmanni represents an insular relict of one of these basal lineages. Although insular environments can serve as refugia and preserve stem lineages, an alternative hypothesis considers G. ballmanni a species of waterfowl highly adapted to an insular environment, with a morphology that is interpreted as adaptations to a terrestrial lifestyle in an environment where mammalian carnivores are rare. Given similar adaptations in waterfowl from other islands and the highly insular nature of the Gargano fauna, I consider the latter scenario most likely. G. ballmanni thus represents an example of convergent evolution of waterfowl in island ecosystems.
Acknowledgments
I am much indebted to the following people for their feedback and suggestions: E. Buffetaut, L.W. van den Hoek Ostende, A. Iwaniuk, H. James, G. Mayr, M. Pavia, J. Prieto, M. Spitzer, and J. de Vos. T.H. Worthy and an anonymous reviewer provided comments and suggestions that significantly improved the manuscript. This research received support from the SYNTHESYS Project (http://www.synthesys.info/) which is financed by the European Community Research Infrastructure Action under the FP6 “Structuring the European Research Area” Programme.
AnatidaeAix sponsa NMNH 500362; Alopochen aegyptiacus NMNH 431687; Amazonetta brasiliensis NMNH 630177; Anas p. platyrhynchos RMNH 106153, RMNH 106154; A. strepera RMNH 86424; Anser albifrons NMNH 610517; A. anser RMNH 33–2, NMNH 290499; A. caerulescens NMNH 319918; A. indicus RMNH 7480; Aythya fuligula RMNH 9390, NMNH 432303; A. ferina NMNH 431534, 560588; Branta b. bernicula RMNH 86422, RMNH 86421, NMNH 553108; B. canadensis NMNH 555497, NMNH 489759; B. leucopsis RMNH 34–1, RMNH 34–2, RMNH 13357; Bucephala clangula NMNH 488592; B.c. clangula RMNH 90380; B. islandica NMNH 488596; Cereopsis novaehollandia no number (RMNH); Clangula hyemalis NMNH 561033; Coscoroba coscoroba no number (RMNH); Cygnus atrata, RMNH 1215, NMNH 345212; C. bewickii RMNH 80788, RMNH 104875; C. buccinator RMNH 104876, NMNH 432534; C. cygnus RMNH 86426, RMNH 104867; C. olor RMNH 9408, RMNH 104864, NMNH 344839; Dendrocygna arcuata RMNH 1659; D. bicolor RMNH 2056, NMNH 488132; D. guttata NMNH 560773; Melanitta nigra RMNH 11316; Mergus merganser RMNH 86431; Neochen jubatus NMNH 346907, NMNH 620513; Netta rufina NMNH 621199; N. peposaca NMNH 227348; Nettapus coromandelianus NMNH 318553; N. auritus NMNH 347388; Oxyura dominica NMNH 430928; O. jamaicensis NMNH 492433, 492491; Somateria m. mollissima RMNH 80999; S. m. v-nigrum NMNM 431986; S. spectabilis NMNH 620821; Tachyeres brachypterus RMNH 34414; T. pteneres NMNH 490930; Tadorna tadorna RMNH 86425, NMNH 432291; T. variegata NMNH 621375.
PhoenicopteridaePhoenicopterus ruber RMNH 106142; P. chilensis RMNH 6517.
Fossil speciesCnemiornis calcitrans, Pleistocene of New Zealand, BMNH A 2078, BMNH A 2080, BMNH A1561, BMNH A 2079; Cygnus falconeri, Pleistocene of Malta, BMNH49323, BMNH49324; Branta rhuax, Holocene of Hawaii, NMNH 560329; Thambetochen chauliodous, Holocene of Hawaii, NMNH 560328.
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Map of Italy and the location of Gargano.
Carte de l’Italie et localisation du Gargano.
Garganornis ballmanni n. gen. et n. sp., holotype, distal fragment of left tibiotarsus, in cranial (a), caudal (b) and distal view (c). Abbreviations: ca = caudal side: cr = cranial side; ce = canalis extensorius; cl = condylus lateralis; cm = condylus medialis; dem = depressio epicondylus medialis; em = epicondylus medialis; fi = fossa intercondylaris; ps = pons supratendineus; se = sulcus extensorius; trel = tuberositas retinaculum extensoris lateralis; trem = tuberositas retinaculum extensoris medialis; trmfm = tuberculum retinaculi m. fibularis medialis.
Garganornis ballmann n. gen. et n. sp., holotype, fragment distal de tibiotarse gauche, en vues crâniale (a), caudale (b) et distale (c). Abréviations : ca = côté caudal ; cr = côté crânial ; ce = canalis extensorius ; cl = condylus lateralis ; cm = condylus medialis ; dem = depressio epicondylus medialis ; em = epicondylus medialis ; fi = fossa intercondylaris ; ps = pons supratendineus ; se = sulcus extensorius ; trel = tuberositas retinaculum extensoris lateralis ; trem : tuberositas retinaculum extensoris medialis ; tmfm = tuberculum retinaculi m. fibularis medialis.
Photos by E. Kruidenier.
Left tibiotarsi of selected anseriform taxa and Garganornis ballmanni n. gen. et n. sp. in cranial view. (a) Gastornis parisiensis, MDE-A18, Early Eocene of Saint-Papoul, Aude, France, reprinted with permission from Buffetaut, 2008; (b) Garganornis ballmanni n. gen. et n. sp. (c) Branta canadensis, NMNH 489759, (d) Cygnus buccinator, NMNH 432534. Scale bar for (a) 5 cm, scale bar (b–d) 2 cm.
Tibiotarses gauches de taxa ansériformes et de Garganornis ballmanni n. gen. et n. sp. en vue crâniale. (a) Gastornis parisiensis, MDE-A18, Éocène inférieur de Saint-Papou, Aude, France, reproduit avec l’autorisation de Buffetaut, 2008 ; (b) Garganornis ballmanni n. gen. et n. sp ; (c) Branta canadensis, NMNH 489759 ; (d) Cygnus buccinator, NMNH 432534. Barre d’échelle pour (a) 5 cm, pour (b–d) 2 cm.
Distal tibiotarsi of Cygnus falconeri from the Pleistocene of Malta and Sicily, and Garganornis ballmanni n. gen. et. n. sp. from the Miocene of Gargano. (a) Right distal tibiotarsus, holotype, of C. falconeri as illustrated in Parker, 1867 (reprinted here with permission); right distal tibiotarsi of C. falconeri, (b) BMNH49323 and (c) BMNH49324; (d) Garganornis ballmanni n. gen. et n. sp. RGM 443307; (e) C. falconeri, holotype, in distal view as illustrated in Parker, 1867; (f) Garganornis ballmanni n. gen. et n. sp. RGM 443307 in distal view. Images are scaled to match in size. Note that (d) and (f) are mirror images from the original.
Tibiotarses de Cygnus falconeri du Pléistocène de Malte et de Sicile et Garganornis ballmanni n. gen. et n. sp. du Miocène du Gargano. (a) Tibiotarse distal droit, holotype de C. falconeri tel qu’illustré in Parker, 1867 (reproduit ici avec autorisation) ; tibiotarses distaux droits de C. falconeri (b) BMNH49323 et (c) BMNH49324 ; (d) Garganornis ballmanni n. gen. et n. sp. RGM 443307 ; (e) C. falconeri, holotype en vue distale tel qu’illustré in Parker, 1867) ; (f) Garganornis ballmanni n. gen. et n. sp. RGM 44307 en vue distale. Les images sont ajustées pour être bien assorties en taille. À noter que (d) et (f) sont le reflet de l’original.
Measurements (in mm) of Garganornis ballmanni n. gen. et n. sp. and selected Anseriform taxa. Note that the distal articulation is damaged in G. ballmanni, and distal width is underestimated.
Mesures en millimètres de Garganornis ballmanni n. gen. et n. sp. et de taxons d’ansériformes sélectionnés. À noter que l’articulation distale est endommagée chez G.ballmanni et que la largeur distale est sous-estimée.