The Cretaceous Neornithine record and new Vegaviidae specimens from the López de Bertodano Formation (Upper Maastrichthian) of Vega Island, Antarctic Peninsula

SOUZA, GEOVANE A. DE; BULAK, BRUNO A.; SOARES, MARINA B.; SAYÃO, JULIANA M.; WEINSCHÜTZ, LUIZ CARLOS; BATEZELLI, ALESSANDRO; KELLNER, ALEXANDER W.A.

doi:10.1590/0001-3765202320230802

person GEOVANE A. DE SOUZA

conceived and designed the study · performed fieldwork in the Antarctic Peninsula collecting several fossil vertebrates that are housed in the Museu Nacional/UFRJ · did the osteohistological analysis

schoolPrograma de Pós-Graduação em Zoologia, Universidade Federal do Rio de Janeiro, Museu Nacional, Parque da Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil

0000-0003-0979-1526

person BRUNO A. BULAK

conceived and designed the study · did the osteohistological analysis

schoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Laboratório de Sistemática e Tafonomia de Vertebrados Fósseis, Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil

0009-0003-5994-943X

person MARINA B. SOARES

conceived and designed the study · did the osteohistological analysis

schoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Laboratório de Sistemática e Tafonomia de Vertebrados Fósseis, Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, BrazilschoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Geologia e Paleontologia, Laboratório de Paleobiologia e Paleogeografia Antártica, Parque da Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil

0000-0002-8393-2406

person JULIANA M. SAYÃO

conceived and designed the study · did the osteohistological analysis

schoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Laboratório de Sistemática e Tafonomia de Vertebrados Fósseis, Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, BrazilschoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Geologia e Paleontologia, Laboratório de Paleobiologia e Paleogeografia Antártica, Parque da Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil

0000-0002-3619-0323

person LUIZ CARLOS WEINSCHÜTZ

conceived and designed the study · performed fieldwork in the Antarctic Peninsula collecting several fossil vertebrates that are housed in the Museu Nacional/UFRJ · conducted the geological study

schoolCentro de Pesquisa Paleontológica da UNC (CENPALEO), Universidade do Contestado, Av. Nereu Ramos, 89300-000 Mafra, SC, Brazil

0000-0002-8978-6120

person ALESSANDRO BATEZELLI

conceived and designed the study · performed fieldwork in the Antarctic Peninsula collecting several fossil vertebrates that are housed in the Museu Nacional/UFRJ · conducted the geological study

schoolUniversidade Estadual de Campinas, Instituto de Geociências, Departamento de Geologia e Recursos Naturais, Rua João Pandiá Calógeras, 51, Barão Geraldo, 13083-870 Campinas, SP, Brazil

0000-0001-5324-7638

person ALEXANDER W.A. KELLNER

conceived and designed the study

schoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Laboratório de Sistemática e Tafonomia de Vertebrados Fósseis, Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, BrazilschoolUniversidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Geologia e Paleontologia, Laboratório de Paleobiologia e Paleogeografia Antártica, Parque da Quinta da Boa Vista, s/n, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brazil

0000-0001-7174-9447

Correspondence to: Geovane Alves De Souza E-mail:
geoosoouza@gmail.com

Figures (5)
Tables (1)

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Figure 1 A generalized timecalibrated phylogeny of Mesozoic birds. The time of divergences of the non-neornithine nodes follows Wang & Lloyd (2016). The split of neornithines in the Early Cretaceous follows Hedges et al. (1996) and Ericson et al. (2006). The positioning of the neornithines is according to Clarke (2004) for Apatornis celer and Iaceornis marshi, McLachlan et al. (2017) and Field et al. (2020) for Vegaviidae, De Pietri et al. (2016) for Teviornis gobiensis. Neogaeornis wetzelli, Limenavis patagonica, Kookne yeutensis, Antarcticavis capelambensis, and Australornis lovei were not submitted to any cladistic analyses. The black horizontal thick lines represent approximated temporal ranges of each taxon. Dotted lines indicate alternative positions of Vegaviidae.

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Figure 2 Global paleogeographic maps showing the localities with records of Cretaceous Neornithes. (a) Early Campanian, (b) Late Campanian-Maastrichtian. 1 - Northumberland Formation, British Columbia, Canada (Maaqwi), 2 - Niobrara Formation, Kansas, US (Apatornis and Iaceornis) but see the text to alternative position for these taxa outside Neornithes, 3 - Allen Formation, Rio Negro, Argentina (Limenavis and Lamarqueavis), 4 - Maastricht Formation, Belgium (Asteriornis), 5 - Nemegt Formation, Mongolia (Teviornis, af. Phalacrocoracidae, af. Charadriiformes), 6 - Quiriquina Formation, Chile (Neogaeornis), 7 - Chorrillo Formation, Santa Cruz, Argentina (Kookne, stratigraphic unit ranges from late Campanian to Maastrichtian), 8 - Cape Lamb Member of the Snow Hill Island Formation, Vega Island, Antarctica (Vegavis, Neornithes indet., and Charadriiformes indet.) and López de Bertodano Formation, Seymour Island, Antarctica (Polarornis). Yellow dots mean the Northern Hemisphere neornithines and red dots indicate the Southern Hemisphere neornithines. Plate tectonic maps of Early Campanian (~80 Mya) and Maastrichtian (~70 Mya) by C. R. Scotese (2001), PALEOMAP Project (www.scotese.com).

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Figure 3 Map of the Antarctic Peninsula showing location of Vega Island (a). Copernicus Sentinel-2 image of ice-free portion of Cape Lamb, in Vega Island showing the locality (red dot) where the specimens studied here were collected (b). Stratigraphic interpretation of the Sandwich Bluff Member of the López de Bertodano Formation modified from Roberts et al. (2014) showing the levels where previous Vegavis specimens were found (marked by the red star) (c). Stratigraphic chart of the Marambio Group, James Ross Basin (d). Simplified schematic drawings showing all specimens unambiguously assigned to Vegavis iaai and the new avian materials studied here (white silhouettes indicate the preserved bones). See text for more details. Credit of figure b: European Union, Copernicus Sentinel-2 imagery. Abbreviations: SBM, Sandwich Bluff lithostratigraphic units recognized by Roberts et al. (2014), JRIVG, James Ross Island Volcanic Group.

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Figure 4 Synsacrum MN 7832-V (a-i), photographs of the specimen in (a) right lateral, (b) left lateral, (c) dorsal, and (d) ventral view. Micro CT-scan of MN 7832-V in (e) dorsal, (f) anterior, (g) left lateral, (h) posterior, and (i) ventral views. Scale bar for (a-d) = 10mm, (e-i) 5mm. Arrows indicate anterior side. Abbreviations: cs, canalis synsacri, css, crista spinosa synsacri, f, foramina intervertebralia, lsi, lumbosacral intumescence, lstc, lumbosacral transverse canals, pt, processus transversus.

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Figure 5 Microstructural pattern of MN 7833-V (a-h), proximal fragment of tarsometatarsus MN 7833-V in lateral view (a), whole cross section of MN 7833-V (b). Two cortical regions showing the large amount of intertrabecular spaces (c) and thick endosteal lamella (d) in different portions of the perimedullary cortex. High-magnification of the cortex showing the clusters of osteocyte lacunae, which characterize the woven-fibered bone matrix (e). Nutrient foramen in subperiosteal cortex (f). High magnification of subperiosteal cortex showing the longitudinal primary osteons (g). High magnification of trabeculae bounded by cement lines in perimedullary cortex (h). Red line in a corresponds to the level where the section was made. The outer cortex face upward in b-g. Scale bar in a is equal to 15 mm. Abbreviations: cl, cement line, el, endosteal lamellae, mc, medullary cavity, nf, nutrient foramen, nvc, neurovascular canals, po, primary osteons, oc, osteocyte clusters, rc, resorption cavities, so, secondary osteon, t, trabeculae, ts, intertrabecular spaces.

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Table I List of Neornithes and putative neornithines of the Mesozoic. Asterisks indicate holotype number.

Taxon	Collection number	Referred specimens	Provenance	Age	Taxonomy	References
Antarcticavis capelambensis	SDSM 78147*	-	Antarctica, Snow Hill Island Formation	Late Campanian-Early Maastrichtian	Pangalloanserae	Cordes 2001, 2002, Cordes-Person et al. 2020
Apatornis celer	YPM 1451*	-	USA, Niobrara Formation	Early Campanian	Basal Carinatae?	Marsh 1873, Clarke 2004
Asteriornis maastrichthensis	NHMM 2013 008*	-	Belgium, Maastricht Formation	Late Maastrichtian	Pangalloanserae	Field et al. 2020
Charadriiformes	MLP 98-I-10-25	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Neornithes, Charadriiformes indet.	Case & Tambussi 1999, Reguero et al. 2013
Galliform	PVPH 237	-	Argentina, Portezuelo Formation	Turonian-Coniacian	Neornithes cf. Galliformes	Agnolín et al. 2006
Iaceornis marshi	YPM 1734*	-	USA, Niobrara Formation	Early Campanian	Basal Carinatae?	Marsh 1880, Clarke 2004
Kookne yeutensis	MPM 21550*	-	Argentina, Chorrillo Formation	Maastrichtian	Anseriformes	Novas et al. 2019
Cimolopteryx rara	YMP 1805*	-	USA, Lance Formation	Maastrichtian	Neornithes Cimolopterygidae	Marsh 1892, Agnolín 2010
Ceramornis major	UCMP V-5620*	-	USA, Lance Formation	Maastrichtian	Neornithes Cimolopterygidae	Brodkorb 1963a, Agnolín 2010
Lamarqueavis australis	MML 207*	-	Argentina, Allen Formation	Campanian-Maastrichtian	Neornithes Cimolopterygidae	Agnolín 2010
Lamarqueavis minima	UCMP 53976*	-	USA, Lance Formation	Maastrichtian	Neornithes Cimolopterygidae	Brodkorb 1963a, Agnolín 2010
Lamarqueavis petra	AMNH 21911*	-	USA, Lance Formation	Maastrichtian	Neornithes Cimolopterygidae	Hope 2002, Agnolín 2010
Lamarqueavis sp.	UALVP 47943	UALVP 47944?	Canada, Dinosaur Park Formation	Late Campanian	Basal Carinatae?	Hope 2002, Longrich 2009
Limenavis patagonica	PVL4730*	MML 206	Argentina, Allen Formation	Campanian-Maastrichtian	Basal Carinatae?	Clarke & Chiappe 2001, Hope 2002, Clarke 2004, Agnolín & Novas 2012
Maaqwi cascadensis	RBCM.EH2008.011.01120.001*	-	Canada, Northumberland Formation	Late Campanian	Pangalloanserae, Vegaviidae	McLachlan et al. 2017
Neogaeornis wetzell i	GMPK 123*	Unnumbered	Chile, Quiriquina Formation	Maastrichtian	Pangalloanserae, Vegaviidae	Lambrecht 1929, Olson 1992, Mayr et al. 2018, Schneider 1940, Acosta Hospitaleche et al. 2023
Neornithes	AMNH 30920	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Neornithes indet.	Roberts et al. 2014
Neornithes	MACN PV RN 1115	-	Argentina, Los Alamitos Formation	Campanian-Maastrichtian	Neornithes	Agnolín & Martinelli 2009
Neornithes	MPEF-PV 7758	-	Argentina, La Colonia Formation	Late Campanian–lower Maastrichtian	Neornithes indet.	Acosta Hospitaleche et al. 2023
Neornithes	Unnumbered	-	Mongolia, Nemegt Formation	Early Maastrichtian	Neornithes af. Diomedeidae	Kurochkin 1995
Neornithes	IGM 100/1309	-	Mongolia, Nemegt Formation	Early Maastrichtian	Neornithes indet.	Clarke & Norell 2004
Neornithes	AMNH 30898	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Neornithes	Roberts et al. 2014
Phalacrocoracidae	Unnumbered	-	Mongolia, Nemegt Formation	Early Maastrichtian	Neornithes af. Phalacrocoracidae	Kurochkin 1995
cf. Polarornis	MN 7833-V	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Pangalloanserae, Vegaviidae	This study
cf. Polarornis	?	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Pangalloanserae, Vegaviidae	Roberts et al. 2014
Polarornis gregorii	TTU P 9265*	MLP 96-I-6-2	Antarctica, López de Bertodano Formation	Maastrichtian	Pangaviiformes	Chatterjee 1989, 1997, 2002, Feduccia 1999, Olson 1992, Hope 2002, Chatterjee 2002, Reguero et al. 2013, Acosta Hospitaleche & Gelfo 2015, Agnolín et al. 2017
Teviornis gobiensis	PIN 44991-1*	-	Mongolia, Nemegt Formation	Maastrichtian	Anseriformes, Plesbyornithidae	Kurochkin et al. 2002
cf. Vegavis	AMNH FARB 30913	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Pangalloanserae, Vegaviidae	Roberts et al. 2014
cf. Vegavis	MN 7832-V	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Pangalloanserae, Vegaviidae	This study
Vegavis iaai	MLP 93-I-3-1*	MACN-PV 19.748	Antarctica, López de Bertodano Formation	Late Maastrichtian	Pangalloanserae, Vegaviidae	Noriega & Tambussi 1995, Hope 2002, Clarke et al. 2005, Chatterjee et al. 2006, Clarke et al. 2016, Agnolín et al. 2017, Marsà et al. 2017, Acosta Hospitaleche & Worthy 202
Vegavis sp.	SDSM 78247	-	Antarctica, López de Bertodano Formation	Late Maastrichtian	Pangalloanserae, Vegaviidae	Case et al. 2006, West et al. 2019

The Cretaceous Neornithine record and new Vegaviidae specimens from the López de Bertodano Formation (Upper Maastrichthian) of Vega Island, Antarctic Peninsula

Abstract