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Determining the Ontogenetic Variation of Lower Cheek Teeth Occlusal Surface Patterns in Lagomorphs Using Micro-Ct Technology

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Determining the Ontogenetic Variation of Lower Cheek Teeth Occlusal Surface Patterns in Lagomorphs Using Micro-Ct Technology Palaeontologia Electronica palaeo-electronica.org Determining the ontogenetic variation of lower cheek teeth occlusal surface patterns in lagomorphs using micro-ct technology Chiara Angelone, Julia A. Schultz, and Margarita A. Erbajeva ABSTRACT Micro CT-scanning has been performed on the lower jaws of some selected lago- morph taxa in order to reconstruct unequivocally their controversial ontogenetic devel- opment. The analyses were concentrated on the development of p3, and on the sequences of lobe connections and on the sequence of appearance/disappearance of flexids/fossettes of p4-m2. This is the first time that this approach has been applied to lagomorphs and opens promising perspectives especially for the taxonomy and phylogeny of this complex order. Chiara Angelone. Institut Català de Paleontologia, Universitat Autònoma de Barcelona, 08193 Cerdanyola del Vallès, Barcelona, Spain, [email protected] Julia A. Schultz. Steinmann-Institut für Geologie, Mineralogie und Paläontologie, Universität Bonn, Nussallee 3, 53115 Bonn, Germany, [email protected] Margarita A. Erbajeva. Geological Institute, Siberian Branch, Russian Academy of Sciences, Ulan-Ude, Russia 670047, [email protected] Keywords: micro-ct; Lagomorpha; ontogeny; lower teeth INTRODUCTION has led to taxonomic misinterpretation and thus incorrect phylogenetic reconstructions. The occlusal surface of lagomorph cheek Micro CT-scanning is a non-destructive teeth, especially stem lagomorphs, drastically var- method of obtaining x-ray images of objects and ies with ontogeny. Until now ontogenetic variation the reconstructed image stack can be used for vir- has usually been inferred by extrapolation from tual cutting of the object at different levels. In the specimens in different wear stages (e.g., Tobien, case of high-crowned teeth this method gives 1974, pp. 110-122; López Martínez, 1989, p. 189), insight in the variation of tooth crown pattern by vir- with the implicit a priori assumption that these tually imitating different wear stages/levels. stages pertained to the same taxon. This approach PE Article Number: 17.1.5A Copyright: Society for Vertebrate Paleontology January 2014 Submission: 8 July 2013. Acceptance: 16 January 2014 Angelone, Chiara, Schultz, Julia A., and Erbajeva, Margarita A. 2014. Determining the ontogenetic variation of lower cheek teeth occlusal surface patterns in lagomorphs using micro-ct technology. Palaeontologia Electronica Vol. 17, Issue 1;5A; 13p; palaeo-electronica.org/content/2014/659-tooth-ontogeny-from-ct ANGELONE, SCHULTZ, & ERBAJEVA: TOOTH ONTOGENY FROM CT Using Micro CT-technology we attempted to isotropic voxels, meaning the same thickness in reconstruct the ontogenetic development of the each direction. As a result the resolution of the lower cheek teeth of some lagomorph genera of scan is equal with the slice thickness. The CT-data different families, ages and continents. The aim of were processed with the reconstruction software this study is to describe the ontogenic development datos x res of the device and VGStudioMax and of the lower tooth row of different lagomorph gen- Avizo 6.1 in order to visualize virtual slices and 3D era, which previously were only inferred from the surfaces. In addition single tooth positions were occusal surface of different wear stages by differ- reconstructed with datos x res software in “resolu- ent authors. This is the first time that CT-scanning tion 2” mode in order to virtually halve the resolu- is used for this purpose in lagomorphs. tion and as a result achieve higher resolved scans. Once integrated in the analysis of other lago- Therefore the software automatically interpolates morph genera, our results will provide (1) a new between the grey values of two adjacent voxels approach for reconstructing the relationships and calculates the virtual grey value for an addi- among lagomorph genera; and (2) tools for investi- tional voxel between them using the integrated gating the differences in the patterns of evolution algorithm. between different lineages as well as between dif- In this preliminary study we decided to anal- ferent continents. yse the lower cheek teeth, because enamel pat- terns and structures are simpler than in upper MATERIAL AND METHODS cheek teeth. In addition, lower cheek teeth in con- trast to upper ones do not undergo a patent non- In order to investigate the un-erupted parts of parallel abrasion during ontogeny. Rather, the wear the hypsodont teeth, jaws were scanned using the plane of upper teeth follows a very complex pat- Micro-CT scanner v/tome/xs (GE sensing Inspec- tern, especially in archaic lagomorphs (Bair, 2007). tion technologies GmbH phoenix/X-ray) housed in The specimens have been virtually sectioned at the Steinmann-Institut für Geologie, Mineralogie several heights parallel to a plane approximating und Paläontologie, Rheinische Friedrich-Wilhelms the occlusal surface (Figure 1). Universität Bonn, Germany. The scans were car- We chose taxa of different families and from ried out with the 240 kV tube of the Phoenix|x-ray different continents (“Amphilagus”, stem lago- v|tome|x s 240 (GE Electronics) with varying set- morph, ~25-22 Ma, Europe; Desmatolagus, stem tings (see below). The Micro-CT device produces FIGURE 1. Example of how the selected Micro-CT scanned specimens were virtually sectioned at several heights parallel to a plane approximating the occlusal surface. 2 PALAEO-ELECTRONICA.ORG TABLE 1. Sequences of appearance/development of p3-m2 of lobe connections, and of appearance/disappearance of flexids/fossettes in the studied specimens. Desmatolagus “Amphilagus” antiquus Piezodus branssatensis Palaeolagus sp. cf. youngi p3 development deep hypoflexid --> hypoflexid + mesoflexid --> shallowing of hypoflexid + mesoflexid --> posterior flexids/ hypoflexid+mesofossettid --> hypoflexid + mesofossettid--> hypoflexid hypoflexid + mesofossettid --> fossettes shallowing of hypoflexid + shallowing of hypoflexid + hypoflexid alone disappearance of mesoflexid disappearance of mesoflexid p4-m2 trigonid-talonid m2-m1-p4 m2-m1-p4 m2-m1-p4 m1-m2-p4 connection hypoconulid --> m1-m2-p4 m1-p4-m2 - - fossette fossette m1-m2-p4 m1-m2-p4 - - disappearance lagomorph, ~45-25 Ma in Asia; Palaeolagus, lep- Abbreviations: NHMW- Naturhistorisches orid, ~38-26 Ma, North America; Piezodus, ochot- Museum Wien; NMB- Naturhistorisches Museum onid, ~24-20 Ma, Europe; age data from López Basel; STIPB- Steinmann-Institut für Geologie, Martínez, 2008) in order to have a wide range of Mineralogie und Paläontologie Bonn. different ontogenetic developmental stages of the selected characters of the dental occlusal surface, RESULTS such as trigonid-talonid connections in p3-m2 and In the virtual Micro CT-scan “sections” of the flexids/fossettes in p4-m2. Nomenclature of fea- studied genera we observed different timings in the tures of teeth occlusal surfaces follows Palacios appearance/development in p3-m2 of lobes con- and López Martínez (1980). nections, and in the appearance/disappearance of The analyzed specimens are (see also Koe- flexids/fossettes (sequences are summarized in nigswald et al., 2010): Table 1). “Amphilagus” antiquus, Coderet-Branssat, France, uppermost Oligocene MP 30 (NMB - BST Development of p3 45-3c [CT scan resolution = 0.0154 mm, Being the most important tooth for diagnosis virtual resolution = 0.0077 mm, settings = at the species level, the reconstruction of ontoge- 140 kV and 110 mA]; netic development of the p3 is of primary impor- Desmatolagus cf. youngi (quite senile individual), tance for the phylogeny of lagomorphs, especially Tatal Gol D/0, Mongolia, Late Oligocene rooted ones, in order to avoid taxonomic mistakes. (NHMW_2011-0209-0003) [CT scan reso- Controversies and errors in the taxonomy of lution = 0.025851 mm, settings = 150 kV European stem lagomorphs have already been and 130 mA]; highlighted in literature, as for example the long debated synonymy between Titanomys visenovi- Palaeolagus sp., Toadstool Park Area, Nebraska, ensis and “Amphilagus” antiquus (Álvarez-Sierra et Late Eocene/Early Oligocene (col. K. al., 1990 and references therein) or the different Hirsch; STIPB - KOE 668) [CT resolution = genera lumped in “Amphilagus” (cf. Angelone, 0.02102 mm, settings = 150 kV and 130 2009). mA]; The ontogenetic development of the p3 of Piezodus branssatensis, Coderet-Branssat, “Amphilagus” antiquus is shown in Animation 1. France, uppermost Oligocene MP 30 The occlusal surface of the p3 of the analyzed (NMB - BST 3899 [CT scan resolution = specimen is C-shaped, with a single labial flexid 0.02264 mm, p4 and m1 virtual resolution (hypoflexid) and a lingually directed hypoconulid = 0.01134 mm, settings 130 kV and 110 (Figure 2.1). With wear the hypoconulid shallows mA]; up until disappearing, whereas the hypoflexid is Piezodus branssatensis (juvenile individual), interrupted by a central connection that develops Coderet-Branssat, France, uppermost Oli- between trigonid and talonid; such a connection gocene MP 30 (NMB - Cod 273) [CT scan determines the formation of a mesofossettid (Fig- resolution = 0.0147 mm, settings = 150 kV ure 2.2-3). These steps correspond to the recon- and 140 mA]. struction of the ontogenetic development of the p3 in “Amphilagus” antiquus based on the relatively 3 ANGELONE, SCHULTZ, & ERBAJEVA: TOOTH ONTOGENY FROM CT rich collections from Coderet (MP 30) (López Martínez, 1989, p. 189, figure 50 B 3-4-5). Our results show that further wear of the tooth leads to the simultaneous shallowing up of the hypoflexid
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