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At the Root of the Early Penguin Neck: a Study of the Only Two Cervicodorsal Spines Recovered from the Eocene of Antarctica Piotr Jadwiszczak

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At the Root of the Early Penguin Neck: a Study of the Only Two Cervicodorsal Spines Recovered from the Eocene of Antarctica Piotr Jadwiszczak RESEARCH/REVIEW ARTICLE At the root of the early penguin neck: a study of the only two cervicodorsal spines recovered from the Eocene of Antarctica Piotr Jadwiszczak Institute of Biology, University of Bialystok, Swierkowa 20B, PL-15-950, Bialystok, Poland Keywords Abstract Antarctic Peninsula; La Meseta Formation; Palaeogene; early Sphenisciformes; The spinal column of early Antarctic penguins is poorly known, mainly due to cervicodorsal vertebrae. the scarcity of articulated vertebrae in the fossil record. One of the most interesting segments of this part of the skeleton is the transitional series located Correspondence at the root of the neck. Here, two such cervicodorsal series, comprising rein- Piotr Jadwiszczak, Institute of Biology, terpreted known material and a new specimen from the Eocene of Seymour University of Bialystok, Swierkowa 20B, Island (Antarctic Peninsula), were investigated and contrasted with those PL-15-950 Bialystok, Poland. of modern penguins and some fossil bones. The new specimen is smaller E-mail: [email protected] than the counterpart elements in recent king penguins, whereas the second series belonged to a large-bodied penguin from the genus Palaeeudyptes. It had been assigned by earlier researchers to P. gunnari (a species of ‘‘giant’’ penguins) and a Bayesian analysis*a Bayes factor approach based on size of an associated tarsometatarsus*strongly supported such an assignment. Morphological and functional studies revealed that mobility within the aforementioned segment probably did not differ substantially between extant and studied fossil penguins. There were, however, intriguing morphological differences between the smaller fossil specimen and the comparative material related to the condition of the lateral excavation in the first cervicodorsal vertebra and the extremely small size of the intervertebral foramen located just prior to the first ‘‘true’’ thoracic vertebra. The former feature could have resulted from discrepancy in severity of external pneumatization. Both fossils provided valuable insights into the morphology and functioning of the axial skeleton in early penguins. The spinal column, a hallmark feature of vertebrates, the dorsal (thoracic) series. However, two vertebrae at the encloses the spinal cord, stiffens the body and provides root of the neck (i.e., the 14th and 15th ones, counting attachment for girdles and muscles. In tetrapods, it is from the cranial end), that in most cases bear ribs not regionalized into presacral (cervicalthoracic), sacral reaching the sternum (a feature not limited to penguins), and caudal series of vertebrae, which results from dif- have sometimes been counted among the cervical seg- ferent Hox gene expressions along the anteroposterior ment (e.g., Shufeldt 1901; see also Bertelli & Giannini body axis (Mu¨ ller et al. 2010, references therein). In birds, 2005; Livezey & Zusi 2006). Importantly, these transi- the total number of vertebrae varies considerably, with tional spinal elements, termed ‘‘cervicodorsal vertebrae’’ most interspecific variation occurring in the cervical (Baumel & Witmer 1993, references therein; figure 5.10 spine (Baumel & Witmer 1993). in Baumel & Raikow 1993; see also supporting informa- Penguins (Aves: Sphenisciformes), highly specialized tion in Mu¨ ller et al. 2010), form the caudalmost portion seabirds, have 47Á48 vertebrae (Stephan 1979). Accord- of the presacral spinal column within which some median- ing to Stephan (1979) and Guinard et al. (2010; but see plane mobility is still possible (Stephan 1979). The neck Watson 1882), 13 of these bones are the neck (cervical) folding into an S-shaped curve (and straightening if vertebrae and eight moveable-rib-bearing elements form necessary) is important for penguins on both land and Polar Research 2014. # 2014 P. Jadwiszczak. This is an Open Access article distributed under the terms of the Creative Commons Attribution-Noncommercial 3.0 1 Unported License (http://creativecommons.org/licenses/by-nc/3.0/), permitting all non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. Citation: Polar Research 2014, 33, 23861, http://dx.doi.org/10.3402/polar.v33.23861 (page number not for citation purpose) Penguin cervicodorsal spines from the Eocene of Antarctica P. Jadwiszczak sea for a number of reasons, for example, to improve aforementioned fragments of the spinal column, heretofore hydrodynamics and body balance, and as an extendable documented only as an unannotated black-and-white ‘‘arm’’ for feeding and nest-building to name just two photograph and a two-sentence description. Further- activities, and more (e.g., Guinard & Marchand 2010). more, I report on the new rare finding represented by The spinal column of early penguins (i.e., those from three articulated vertebrae from the transitional cervico- the Palaeocene and Eocene epochs, 66.0Á33.9 Mya) is dorsal series belonging to a ‘‘non-giant’’ Eocene penguin poorly known. This is especially true for the Antarctic from the La Meseta Formation (Seymour Island, Antarc- penguins, for their remains (found solely on Seymour tica). Both specimens provide a unique opportunity to Island, Antarctic Peninsula; Fig. 1) are, with few excep- expand our knowledge of the axial skeleton in basal tions (Ksepka & Bertelli 2006; Acosta Hospitaleche & Sphenisciformes. Di Carlo 2010; Acosta Hospitaleche & Reguero 2010; Jadwiszczak 2010, 2012; Jadwiszczak et al. 2013), isolated Geological setting bones (Wiman 1905; Myrcha et al. 2002; Jadwiszczak 2006; Jadwiszczak & Chapman 2011; Jadwiszczak & Seymour (Marambio) Island is located in the Weddell Acosta Hospitaleche 2013, references therein). The oldest Sea, within the James Ross Basin, a back-arc feature Antarctic penguin vertebra ever collected is a single poorly that developed east of a magmatic arc of the Antarctic preserved thoracic element described by Jadwiszczak et al. Peninsula (Fig. 1; e.g., Pore˛bski 2000). The siliciclastic (2013), a part of the incomplete skeleton of late Palaeocene sedimentary rocks of the EoceneÁ?early Oligocene La Crossvallia unienwillia Tambussi, Reguero, Marenssi & Meseta Formation crop out in the north-eastern part of Santillana, 2005 from the Cross Valley Formation (Tambussi the island (Fig. 1). These uplifted nearshore, estuarine et al. 2005). So far, there have been only two reported and deltaic, mostly poorly consolidated, sediments are series of articulated vertebrae of Palaeogene (66.0Á23.0 very fossiliferous, and have yielded a diverse marine and Mya) Antarctic Sphenisciformes. These are a segment terrestrial fauna (including numerous penguins) and flora comprising three anterior thoracic skeletal elements (Pore˛bski 2000; Myrcha et al. 2002; Marenssi 2006; Tatur assignable to Palaeeudyptes gunnari (Wiman, 1905), a ‘‘giant’’ et al. 2011; Reguero et al. 2013, references therein). A penguin from the EoceneÁ?early Oligocene La Meseta number of studies revealed that the sediments had come Formation (Fig. 1; Acosta Hospitaleche & Reguero 2010) from the westÁnorth-west, hence the source rocks must and a very poorly preserved fragment of a supposedly have been located within the nearby northern Antarctic caudal thoracic series of a very large sphenisciform from Peninsula (Reguero et al. 2013, references therein). The the same formation (Acosta Hospitaleche & Di Carlo recent micromorphological investigation of quartz sand 2010). grains (Kirshner & Anderson 2011) supports the domi- The purpose of this paper is to thoroughly describe, nant view that the La Meseta Formation is devoid of any illustrate and eventually reinterpret the former of two evidence of glaciation. Hence, it represents the youngest preglacial succession on the Seymour Island (e.g., Marenssi et al. 2002). Cape Wiman The base of the uppermost (or the youngest) unit of N the La Meseta Formation, Telm7/Submeseta Allomember, Cape was interpreted by Pore˛bski (2000) as a basinward facies Bodman Cross shift (supposedly bay-head delta deposits over those 64˚15'00''S Valley interpreted as the estuary mouth) followed by ‘‘transgres- sive reactivation’’ (alternately: flooding surfaces of trans- gressive marine origin and shorefaces). According to Marenssi Penguin Bay (2006), the lower part of Telm7 represents the inner estuary channels/shoals, its upper levels the shallow Fossil Seymour Island La Meseta Formation marine environment. For detailed stratigraphy of this Bay Telm6-7 Telm1-5 unit, consult Myrcha et al. (2002, figure 2) and Reguero Collecting localities et al. (2013, figure 3.1). ‘shelves’ Unit Telm7 of the La Meseta Formation has been tradi- 56˚40'00''W DPV10/84 tionally regarded as late Eocene in age (Myrcha et al. 1 km airstrip 2002; Jadwiszczak 2006; Marenssi 2006; Acosta Hospita- Fig. 1 Map showing the location of Seymour (Marambio) Island, the La leche & Reguero 2010). Some recent works, however, Meseta Formation and sampling sites. have adopted the view that its lower levels can be dated 2 Citation: Polar Research 2014, 33, 23861, http://dx.doi.org/10.3402/polar.v33.23861 (page number not for citation purpose) P. Jadwiszczak Penguin cervicodorsal spines from the Eocene of Antarctica to the middle Eocene (Reguero et al. 2013, references The calculation of the Bayes factor (see Rouder et al. therein). The youngest documented ages (34.78Á33.57 2009 for a readable introduction), used to validate the Mya) from the formation are based on strontium isotope taxonomic position of the latter specimen, was based on stratigraphy and suggest
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