878
Journal of Paleontology 89(5):870–881
fragmentary fish dentary bone with a strong and conical tooth and a piece of a second tooth. The tooth is slightly bent back- wards and has its internal cavity exposed at the tip. The tooth surface is weathered, but a pedicle is developed at its base, and the tooth is inserted in a shallow longitudinal groove. Based on its gross morphology, the specimen may probably be assigned to the cod-icefish Mesetaichthys jerzmanskae Bieñkowska- Wasiluk, Bonde, Moller and Gazdzicki, 2013, a Notothenioidei (Perciformes) recently described (Bieñkowska-Wasiluk et al., 2013) for the Submeseta Formation (middle/late Eocene). The specimen in question (MLP 83-V-30-2) is not referable to Aves, much less Pelagornithidae.
Discussion and conclusions
Regarding their affinities, both morphotypes recognized for Antarctic pelagornithids share with the late Paleocene/early Eocene Dasornis the presence of several possible plesiomorphic characters (i.e., humerus: deep concavities for the musculus extensor carpi ulnaris surrounded by a thick and smooth ridge; rostral end of the beak: slightly down-curved, absence of transverse sulcus; narial region of the rostrum: well-defined dorsolateral constriction and dorsal ridge, palatal ridge slightly prominent, and with a marked medial sulcus; tarsometatarsus: corpus mediolaterally narrower with square cross section, nar- row and poorly excavated trochlea II, proximally positioned trochlea II with a processus medioplantaris, narrow dorsal opening of foramen vasculare distale). However, a number of characters also remind the more derived condition typical of the Neogene Pelagornis (i.e., humerus: flattened diaphysis, well- developed epicondylus dorsalis, vertically positioned and proxi- mally extended tuberculumsupracondylare ventrale; narial region of the rostrum: longitudinal sulcus more dorsally positioned; tar- sometatarsus: recessed and more distally located plantar opening of the foramen vasculare distale, slight plantar projection of tro- chlea II, wider and lower dorsal surface of trochlea III). A com- bination of plesiomorphic and derived character states, showing an ‘intermediate’ condition between Dasornis and Pelagornis was already reported in post-early Eocene and pre-late Oligocene specimens. Taxa showing such combination include the middle Eocene L. tethyensis from Ukraine (Mayr and Zvonok, 2011, 2012), Gigantornis eaglesomei Andrews, 1916 from Nigeria (Harrison andWalker, 1976;Mayr et al., 2008), Togo specimens referred to Gigantornis (Bourdon and Cappetta, 2012), and some other remains from Belgium tentatively assigned by Mayr and Smith (2010) to D. emuinus and Macrodontopteryx oweni Harrison and Walker 1976 (although they probably correspond to Gigantornis and Lutetodontopteryx, respectively; Mayr and Zvonok, 2012; see alsoBourdon et al., 2010).The late Eocene and ?early Oligocene materials from Oregon described by Goedert (1989) also seems to fit in this intermediate morphology (Mayr et al., 2013). A valuable element in order to recognize plesiomorphic-
derived conditions is probably represented by the tarsome- tatarsus. The progressive increase in the distal projection of the trochlea metatarsi II, and the reduction of its plantar extension are consistent with the rise of the more modern taxa studied (Fig. 5). The distal end of tarsometatarsus UCR 22176 from the middle Eocene of Seymour Island is morphologically more
similar to the middle Eocene L. tethyensis and the probably late Oligocene P. “Palaeochenoides” mioceanus than to any other pseudo-toothed birds. UCR 22176 belonged to a huge bird similar to P. “Palaeochenoides” mioceanus (Table 1) and markedly larger than L. tethyensis (similar in size to D. toliapica).
Pelagornithid size-types.—Otherwise, considering the world- wide pelagornithid record and according to the estimated wingspan, four approximate size-ranges were identified (Fig. 7). The small gannet-sized Dasornis abdoun Bourdon et al., 2010 (1.5–1.7m wingspan; Bourdon et al., 2010), the medium albatross-sized D. toliapica and L. tethyensis (2–3m wingspan; Bourdon et al., 2010), the large D. emuinus (3.5–4.5m wing- span; Mayr, 2009; Bourdon et al., 2010), and finally, the giant middle/late Eocene and Neogene Pelagornis-related taxa (5–6m wingspan; Olson, 1985; Mayr, 2009; Mayr and Rubilar Rogers, 2010; Boessenecker and Smith, 2011; Ksepka, 2014). The Antarctic material assigned to morphotype 1 (middle
Ypresian), including the humerus MLP 12-I-20-4 and the humeral shaft USNM 494035 (Jones et al., 2000) correspond to the large size-type (i.e., equivalent in size to D. emuinus, Table 1). The material assigned to morphotype 2 (Bartonian/? Priabonian, including the humerus recently described by Rubilar-Rogers et al., 2011) correspond to the giant size-type. As stated previously (see also Tonni, 1980; Cenizo, 2012), they may belong to birds larger than the huge Neogene taxa (i.e., P. chilensis; similar-sized to P. sandersi; Table 1), constituting one of the largest pelagornithid known so far. In this sense, two Antarctic morphotypes were recognized
by previous authors. However, it was thought that both morphs coexisted during the middle/late Eocene of Seymour Island (Tonni and Tambussi, 1985; Cenizo, 2012). This idea was conceived from the finding of the mandible MLP 83-V-30-1 assigned—at that time—to a pelagornithid. Its removal from pelagornithids implies that only giant pseudo-tooth birds (i.e., morphotype 2) are known for the Bartonian/?Priabonian strata of the Submeseta Formation. Across the world, a large diversity of body sizes of
pelagornithids has been recorded between the late Paleocene and middle Eocene (Mayr, 2009). Since the middle/late Eocene, only giant pelagornithids are known. These forms (more than 5m wingspan) show a trend toward acquiring huge sizes, reaching a maximum specialization during the late Neogene (Fig. 7). Even taking into account the incompleteness of the fossil record, it seems that large, medium, and small taxa (Fig. 7) become extinct after middle Eocene times. An ecological competition for food or breeding sites could be the main cause (Mayr, 2009). A large list of candidates for such a competition may include the oldest Procellariformes (Noriega and Tambussi, 1996; Tambussi and Acosta Hospitaleche, 2007; Tambussi and Degrange, 2013; Reguero et al., 2013) described for the Cucullaea II Allomember (Ypresian/Lutetian), and the extre- mely diversified and widely extended early penguin fauna (Acosta Hospitaleche and Reguero, 2010; and references therein). The onset and diversification of giant forms occurred at the same time both in penguins (Clarke et al., 2007; Ksepka and Clarke, 2010), and pseudo-toothed birds. Giant penguins are recorded until the latest Eocene/earliest Oligocene
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