Swiss J Geosci DOI 10.

1007/s00015-011-0076-y

Survival of the thinnest: rediscovery of Bauers (1898) ichthyosaur tooth sections from Upper Jurassic lithographic limestone quarries, south Germany
Torsten M. Scheyer Markus Moser

Received: 12 July 2011 / Accepted: 12 September 2011 Swiss Geological Society 2011

Abstract The re-discovery of nine petrographic slides from the late 19th century at the palaeontological collections of the University of Zurich, showing thin-sectioned ichthyosaur teeth, revealed these slides be the only preserved remains of the historical collection of Upper Jurassic ichthyosaurs from the Bavarian State Collection for Palaeontology and Geology; fossil material which, up to now, was thought to have been completely destroyed during World War II. Here the history of these slides, from their origin in Munich as part of the doctoral thesis of Franz Bauer (1898) to their rediscovery in Zurich in 2010 is presented. Furthermore, a complete overview of all slides is given to elucidate their scientic value with the background of up-to-date knowledge of ichthyosaur dentition and tooth histology, including aspects of tissue and growth mark identication. As such, the sectioned teeth show an exposed layer of acellular cementum at the tooth neck, and sets of short and long period growth lines in the orthodentine. The slides of one tooth are part of the original syntype material of Aegirosaurus leptospondylus (WAGNER). They reveal an oval rather than a rectangular shape of the root, as well as the presence of peculiar vascular canals,

interpreted as secondary osteodentine deposition, in the peri-pulpal orthodentine. Keywords History of science Ichthyopterygia Ichthyosaurus trigonus var. posthumus Aegirosaurus leptospondylus Nannopterygius Orthodentine Growth increments Institutional abbreviations BSPG Bavarian State Collection for Palaeontology and Geology, Munich = Bayerische Staatssammlung fur Palaontologie und Geologie, Munchen, Formerly Bayerische Palaontologische Staatssammlung PIMUZ Palaontologisches Institut und Museum, University of Zurich, Switzerland

Introduction General introduction In 2010, a tray with nine petrographic thin-sections of two ichthyosaur teeth was discovered during renovation works in the Palaeontological Institute and Museum of the University of Zurich. From the labels still attached to the slides it was apparent that these slides were part of the doctoral thesis materials of Dr. phil. Franz Bauer in Munich, published in Palaeontographica in 1898. Besides the nine slides still preserved (Fig. 1), all other remains of the ichthyosaur specimens gured on plates 2527 by Bauer (1898) were completely destroyed during World War II (WWII) on the 2425. April 1944 during a bombing raid of the old Academy building in Munich, which housed the Bayerische Palaontologische Staatssammlung (now

Editorial handling: Michael J. Benton & Daniel Marty.

Electronic supplementary material The online version of this article (doi:10.1007/s00015-011-0076-y) contains supplementary material, which is available to authorized users.
T. M. Scheyer (&) Palaontologisches Institut und Museum der Universitat Zurich, Karl Schmid-Strasse 4, 8006 Zurich, Switzerland e-mail: tscheyer@pim.uzh.ch M. Moser Bayerische Staatssammlung fur Palaontologie und Geologie, Richard-Wagner-Strae 10, 80333 Munich, Germany

T. M. Scheyer, M. Moser

Fig. 1 Photographs of all recovered slides of Bauer (1898). a1a8 BSPG AS XIX 504ah (slides 17 and 8), tooth from the Oberndorfer specimen, syntype material of Aegirosaurus leptospondylus (WAGNER, 1853) in Wagner (1853a), sectioned transversely and longitudinally. b BSPG AS I 1656, longitudinal section of a tooth from the Solnhofen specimen, referable to Ichthyosauria indet., aff. Nannopterygius

BSPG) at that time. The specimens were stored and displayed in public exhibition then (Dehm 1978). Similarly, all paper documents, including inventory, loan documents, eld books, working notes, manuscripts, and theses, were lost to re. Bauers monograph (1898) work was often cited in the palaeontological literature (e.g. Bauer 1900b; Broili 1907; von Huene 1922; Kuhn 1934; Camp 1942). Especially references to the osteological description of the caudal region and tail n referred to Ichthyosaurus trigonus var. posthumus by Bauer (1898) already appeared shortly after the original print in a review by Koken (1901, p. 476), the

monographic work on Triassic ichthyosaurs by Merriam (1908, p. 40), and in palaeobiological textbooks (e.g. Abel 1919, pp. 461, 463). The histological sections of the teeth shown in this work, however, went without further notice for some decades. Peyer (1945) was the rst to mention and reproduce part of the histological details, i.e. gure 28 of plate 26, of Bauer (1898), when discussing the presence of minute bore canaliculi of Mycelites ossifragus Roux, 1887 (lamentous fungi = kalkfressende Algen of Bauer) in ichthyosaur teeth and other vertebrate hard tissues. In the same context, the gure was also reproduced in Peyers (1968) book on comparative odontology. Bauer (1898, p. 290) already interpreted the bore canaliculi as secondary disease symptoms instead of primary or post-mortem structures, a view which was validated by microstructural and ultrastructural studies of fungoid lesions in extant sh teeth (Schmidt 1954; Kerebel et al. 1979). In 1947, Albert Besmer, published a doctoral thesis on the dentition of ichthyosaurs under the supervision of Prof. Bernhard Peyer. For this work, Besmer borrowed the original slides of Bauer (1898) for comparison to his material, which was composed of ichthyosaurs from the Middle Triassic of Monte San Giorgio, Ticino, Switzerland and from the Lower Jurassic (Lias e) of Holzmaden in Baden Wurttemberg, southwestern Germany. Although the slides had to be sent to Besmer prior to April 1944, it is unclear in which year exactly the slides were sent to Zurich, because there are no written loan forms or receipts from that time. Besmer, however, in his thesis acknowledged the late Prof. Broili, who apparently was responsible for the loan. Broili retired in 1939 but he was working as a volunteer for the State Collection until 1942, so probably the slides were sent to Zurich before that time. Besmer noted the prolonged time it took to complete his thesis due to obstacles caused by WWII (Besmer 1947, preface on p. 2, p. 7). After 1947, the slides remained rst in the Zoologischvergleichend anatomische Institut and were stored and nally forgotten for seven decades in an unnamed drawer of the PIMUZ, which was formally established under that name in 1956. Besides the two passages mentioned above, Besmer (1947) included two more references to the slides, namely on page 3 (Bauer 1898, plate 26, gs. 2830) when discussing how Bauer (1898) already identied Kiprijanoffs (1881) misidentication of bore canaliculi of Mycelites ossifragus as larger dentine tubules, as well as on page 11 (Bauer 1898, plate 26, gs. 2830) when noting the overall similarity of his tooth sections to previously published accounts by Bauer (1898), Fraas (1891), Kiprijanoff (1881) and Owen (18401845). The osteological descriptions of Bauer (1898) are furthermore referenced in Bardet and Fernandez (2000) and McGowan and Motani (2003), when discussing the assignment of the species studied by Bauer

Ichthyosaur tooth thin-sections

(see Materials and methods below), but the thin-sections were not, for the reasons described above. Further mention of Aegirosaurus leptospondylus (WAGNER) material either in faunal or systematic overviews or in comparison to other material is found in Kuhn (1957, 1961, 1968, 1971), McGowan (1976), Buchy and Lopez Oliva (2009), Maisch (2010), Angst et al. (2010), Fischer (2011) and Fischer et al. (2011). The ichthyosaur material described and gured by Bauer (1898) as Ichthyosaurus trigonus Owen var. posthumus Wagner belongs, according to Bardet and Fernandez (2000), to three different specimens, which represent three separate taxa: (1) the ophthalmosaurid Aegirosaurus leptospondylus (the Oberndorfer specimen), (2) Ichthyosauria indet. close to Caypullisaurus or Oph thalmosaurus (the Haberlein specimen), (3) Ichthyosauria indet. aff. Nannopterygius (the Solnhofen specimen). Concerning the Oberndorfer specimen, Bardet and Fernandez (2000, p. 510) noted that the specimen roughly shares the same character combination that the two new skeletons [the specimen in private col. Schwegler, designated as neotype and referred to as SM unnumbered; and BSPG 1954 I 608, designated as referred specimen by Bardet and Fernandez] here described. The specic name leptospondylus is thus kept as valid and newly combined to Aegirosaurus []. Concerning the Solnhofen specimen, they further write that because of the scarcity of the illustrations given by Bauer (1898) it is not possible to assign the Solnhofen specimen with certainty to this genus [to Nannopterygius] and it is thus only considered as an indeterminate ichthyosaur, possibly close to Nannoptery gius. (Bardet and Fernandez 2000, p. 511). According to McGowan and Motani (2003, p. 118), Aegirosaurus leptospondylus (WAGNER, 1853), originally described in Wagner (1853a) and as described and newly combined by Bardet and Fernandez (2000, p. 504), is still valid, and both Aegirosaurus BARDET AND FERNANDEZ, 2000 from the Solnhofen Formation, Upper Jurassic (Early Tithonian), Bavaria, Germany, and Nannopterygius von Huene 1922 from the Kimmeridge Clay; Upper Jurassic (Kimmeridgian), Dorset, UK, belong to Ophthalmosauridae BAUR, 1887. After 70 years of storage in Zurich (PIMUZ), all slides are again stored in the Bavarian State Collection for Palaeontology and Geology, Munich (BSPG). Material of the Haberlein specimen, the oldest nding of an Upper Jurassic ichthyosaur, which was already described by Quenstedt (18511852), was not sectioned histologically by Bauer (1898), and is not further considered here. The importance of the rediscovery and notication of some of the oldest, still preserved sections of ichthyosaur teeth from the 19th century, being part of the doctoral thesis documentation on one side and being the sole survivors of the original specimens of Upper Jurassic

ichthyosaurs from the Bavarian lithographic limestone quarries stored in Munich on the other, is obvious. Given the fact that eight of these slides belong to the purportedly lost original type material of Ichthyosaurus leptospondylus WAGNER, 1853 (Wagner 1853a, b), a note on the taxonomic status of the material is warranted and is in preparation elsewhere. Besides reconstructing the history of the slides, the purpose of this contribution is to give a complete overview of all slides preserved, and to elucidate the value of the slides with the background of up-to-date knowledge of ichthyosaur dentition and tooth histology, including aspects of tissue and growth mark identication. Biographical data on Bauer and Besmer Franz Bauer Franz Bauer (see Electronic Supplementary Material for a more exhaustive biography and bibliography of Franz Bauer) was born in Dollnstein near Solnhofen, studied theology in Eichstatt and worked there as a priest in pastoral care before studying natural sciences in Munich in 1895. He completed a doctoral thesis under Prof. Karl Alfred von Zittel in 1897. From 1901, Bauer had an appointment as associate professor of Geology and Palaeontology at the Chemistry Department, Technical University of Munich. He published several scientic articles and abstracts, including anthropological themes (Bauer 1900a) and three on palaeontological issues (Bauer 1898, 1900b, 1901 [the latter being his postdoctoral/habilitation thesis]), before having a tragic accident in 1903 on the Risserkogel Mountain near Tegernsee, south of Munich. Albert Besmer Albert Besmer (19001981) was born in Zurich, Switzerland, studied at the Universities of Zurich and Fribourg, and since 1928 worked as a dentist in Zurich. He nished his doctoral thesis under the supervision of Prof. Dr. Bernhard Peyer (Zoological Institute) at the Medical Faculty of the University of Zurich in 1947. To our knowledge, Besmer published only his thesis, for which the original slides from Bauer (1898) were sent to him by Prof. Dr. Ferdinand Broili (18741946).

Materials and methods Slides with petrographic thin-sections Nine slides with petrographic thin-sections of ichthyosaur teeth were identied as being part of the doctoral thesis materials of Bauer (1898). Previous to this date, the

T. M. Scheyer, M. Moser

collection of Upper Jurassic ichthyosaurs from Bavarian lithographic limestone quarries in the Bayerische Palaontologische Staatssammlung (now BSPG) was thought to have been completely destroyed during WWII (Dehm 1978). Eight of the nine slides (Figs. 1a1a8, 2a) belong to a single ichthyosaur tooth (BSPG AS XIX 504ah), which was sectioned both transversally through the crown (serial sections: slides 17, Fig. 1a1a7) and longitudinally through the root base (slide originally without number, here referred to as slide 8, Figs. 1b, 2b, 4). In the original caption to plate 26, Bauer (1898) noted that the transverse sections seen in gures 2830 belong to a tooth from the Oberndorfer specimen, which is part of the syntype material of Ichthyosaurus leptospondylus WAGNER, 1853. Following the discussion of Bardet and Fernandez 2000, the longitudinal sections would now be referable to Aegirosaurus leptospondylus. The remaining slide (Figs. 1b, 2b) shows an ichthyosaur tooth (BSPG AS I 1656) longitudinally sectioned from crown to root. The plane of section lays slightly parasagittally, thus the cusp of the crown is missing. Bauer (1898, p. 285) mentioned that gure 27 of plate 26 shows a longitudinal section of a tooth belonging to the Solnhofen specimen, which was excavated at the Maxbruch, Solnhofen, in 1894. According to Bardet and Fernandez (2000), this tooth
Fig. 2 Overview of microscopic images of all sections. a1a8 BSPG AS XIX 504ah (slides 17 and 8), tooth from the Oberndorfer specimen, referable to Aegirosaurus leptospondylus, sectioned transversely and longitudinally. Note resorption pit and remnant of enamel probably belonging to a replacement tooth at lower right margin of root in a8. b BSPG AS I 1656, longitudinal section of a tooth from the Solnhofen specimen, referable to Ichthyosauria indet. aff. Nannopterygius

would be referable to an indeterminate ichthyosaur close to the ophthalmosaurid Nannopterygius. To sum up, BSPG AS XIX 504ah belongs to the Oberndorfer specimen = Aegirosaurus leptospondylus; whereas BSPG AS I 1656 belongs to the Solnhofen specimen = Nannopterygius. Analysis of the slides The slides were rst photographed with a Nikon D2x with Nikkor 105 mm micro lens (Fig. 1). The tooth microstructure was then analysed in normal transmitted and polarized light using a Leica DM2500 M composite microscope mounted with a Leica DFC 420 C digital camera.

Results and discussion Matching slides and original drawings Due to the level of detail (especially of the pattern of micro-cracks in the histological tissues) in the drawings of Bauer (1898), it was possible to relocate and match the exact areas (Fig. 3) in the original images of gure 2830 of plate 26: gure 28 shows part of slide 6; gure 29 part of slide 5 and gure 30 part of slide 4. The planes of

Ichthyosaur tooth thin-sections Fig. 3 Matching slides 46 (BSPG AS XIX 504df) with gs. 2830 of plate 26 of Bauer (1898); (g. 28 = slide 6, no. 504fg. 29 = slide 5, no. 504eg. 30 = slide 4, no. 504d). Note that the new microscopic images (slides 46) have been rotated 90 clockwise for technical reasons

sectioning of the gured slides are thus situated at midcrown height (gures 29, 30) and at the neck of the tooth (gure 28). As such, we can be positively sure that these slides were indeed used to create the original gures. In all three slides, the shown orthodentine and outer enamel layer is in various stages of deterioration due to borings of Mycelites ossifragus. Furthermore, it is plausible to assume that the slides 2 and 3 are those Bauer (1898, p. 287) referred to when mentioning the presence of Furchen (=grooves) in two cross sections, which were apparently cut 2.0 and 3.0 mm below the tip of the crown of a small ichthyosaur tooth.

The slide with the longitudinal section (Fig. 4) is more difcult to match with the original gure 27 of plate 26 of Bauer (1898), for the following reasons: 1. The image of the sectioned tooth in the gure does not express the same level of detail as seen in gures 2830 due to the lower level of magnication. The image presents a tooth sectioned exactly through the tip of the crown complete with undamaged borders of the tissues. The image lacks any indication of micro-cracks, changes in coloration of the orthodentine, the enamel

2.

3.

T. M. Scheyer, M. Moser

Fig. 4 Matching the longitudinal section (BSPG AS I 1656) (on the left) with g. 27 of plate 26 of Bauer (1898) (on the right)

or the cementum layers, as well as differences in radicular and pulpal mineral inllings. The slide on the other hand, shows a similar but not congruent shape and distribution of tissues seen in the image. Otherwise, the crown, as well as complete tissue borders is not visible. The slide furthermore shows histological details such as a pattern of micro-cracks or changes in coloration, which apparently have not been used in the production of the original gure. There are two possibilities: rst, the slide is indeed the original on which gure 27 is based, in which case Bauer would have presented a highly stylized, reconstructed depiction of this ichthyosaur tooth, not uncommon at the time of lithographic printing; second, the slide does indeed belong to the sectioned tooth shown by Bauer, but is not the section on which gure 27 is based. In this case, we would have to assume the presence of another section on which the crown might have been completely present, but which, unfortunately, was not preserved. Because the present slide represents the lower section of the tooth only, the second possibility, also less appealing, cannot be ruled out completely. The slides 17 and 8 (BSPG AS XIX 504ah) The tip of the crown (slide 1; Fig. 2a1) shows a concentric circular core of orthodentine ringed with a thin layer of

enamel. Enamel striation is absent. The dentine tubules are arranged perpendicular to the outer surface of the tooth; the central tubules extend towards the tip of the crown, whereas the tubules surrounding the central area show the radial arrangement, as seen in the other transverse sections. The junction between enamel and orthodentine is smooth without any dentine infolding. Slides 26 (Fig. 2a2a6) show an oval shape of the enamel and orthodentine. Coarse striation of the enamel is present in slides 2 and 3. At the enamel-dentine junction a hypomineralised interglobular zone of the orthodentine is visible (Fig. 5a, b). Slides 3 and 5 also reveal the presence of peculiar scattered vascular canals (vaguely reminiscent of primary osteons in bone) in the peri-pulpal orthodentine (Fig. 5cf). The vascular canals, which appear to be directly connected with the pulp cavity, are here interpreted to have resulted from secondary deposition of pulpal osteodentine, leading to a decrease in size of the pulp cavity in these slides. Based on the scarcity of material, the origin of this osteodentine tissue remains unknown. The other transverse slides did not show similar vascular canals. Where preserved, slides 46 again show a smooth outer enamel layer, as well as a smooth enamel-dentine junction. In slide 7 (Fig. 2a7), a small central core of pulpal cellular osteocementum is surrounded by the orthodentine, which itself is surrounded by cellular osteocementum vascularised by mostly radially oriented canals. At the dentinecementum junction (Fig. 5g, h) the hypomineralised layer, i.e. the granular layer of Tomes (e.g. Tomes 1876; see also Peyer 1968; Dean 2000), is present. Furthermore, a thin layer of acellular cementum, recently identied for the rst time in ichthyosaur teeth, i.e., in Platypterygius australis from the Cretaceous of Australia (Maxwell et al. 2011), is also identiable between the orthodentine and the outer cellular osteocementum. Slide 8 (Fig. 2a8) shows the remainder of the root of the tooth, i.e. dentine surrounded by pulpal and external osteocementum and the thin layer of acellular cementum, in longitudinal section. A resorption pit and a small part of dentine, possibly of a replacement tooth are visible at the lower right margin of the section. The longitudinal section (BSPG AS I 1656) On this slide (Figs. 1b, 2b) the root and the neck of the tooth in longitudinal section is preserved. The orthodentine is bordered by pulpal osteocementum and external osteocementum (Fig. 6a, b). As in the transverse sections, a thin layer of acellular cementum lies between the orthodentine and the external cellular cementum (Fig. 6b). The layer extends from the proximal tips of orthodentine up to the neck of the tooth, at which it is exposed as in P. australis (Maxwell et al. 2011). The enamel and associated part of the orthodentine of the crown are not preserved on the slide.

Ichthyosaur tooth thin-sections

Fig. 5 Histological details of the transverse sections on slides 2, 3, 5, 7 (BSPG AS XIX 504b, c, e, g). a, c, f and g seen in normal transmitted light, d, in polarised light and b, e and h in polarized light with lambda compensator. a, b Close-up of striated enamel and orthodentine (slide 2). Note interglobular zone of dentine at enamel dentine junction. c, d Central pulpal area of tooth showing vascular canals in the orthodentine (slide 5). e Central area of tooth with vascular canals surrounded by orthodentine (slide 3). f Close-up of

vascular canal seen in c (slide 5). Note how dentine tubules deviate around the canal. g, h Close-up of the cementumdentine junction at the level of the neck of the tooth (slide 7). Note zones of acellular cementum and the granular layer of Tomes. AC acellular cementum, CC cellular cementum, DT dentine tubules, E enamel, GT granular layer of Tomes, IGZ interglobular zone of orthodentine, OD orthodentine, VC vascular canals

The pulpal osteocementum reaches up to ca. 1/3 of the pulp cavity. The vascularisation is higher than in the external osteocementum layer. The brous arrangement of

the pulpal osteocementum (Fig. 6a) is more ordered (yellow and blue colours in polarised light with Lambda compensator) towards the pulp cavity and grades into an

T. M. Scheyer, M. Moser

Fig. 6 Histological details of the longitudinal section BSPG AS I 1656. Images in a and b are both seen in polarized light with lambda compensator. a Complete view of the sectioned tooth. At the root the orthodentine (blue colours) is embedded in vascularised pulpal and external cellular cementum of the osteocementum type. Note the differences in structural orientation between the ordered upper pulpal cementum (yellow and blue colours) and the irregular external and basal root cementum (pink colours). At the cementumdentine

junction, a thin layer of acellular cementum is visible (yellow colour). b Close-up of the orthodentine pillar. Coarse parallel Sharpeys bres are visible in the external cellular osteocementum. The orthodentine shows a weak banding often obscured by dark dentine tubules. The pulp cavity is lled with sparitic calcite minerals. AC acellular cementum, CC cellular cementum, DT dentine tubules, GT granular layer of Tomes, OD orthodentine, P pulp cavity, SC sparitic calcite, ShF Sharpeys bres

unordered diffuse tissue towards the root base (pink colour). Similarly the external osteocementum has an unordered structure (pink colour). The external border of both the pulpal and external osteocementum is irregular due to cavities and canals. Furthermore, coarse parallel Sharpeys bres are present in the external osteocementum at the tooth neck (Fig. 6b), whereas they appear less ordered and conspicuous in the root base. The Sharpeys bres are the remnants of the non-mineralised periodontal ligament anchoring the cementum to the periradicular bone in the living animal. Incremental growth in dentine In both the transverse and longitudinal sections of both teeth sectioned by Bauer (1898), incremental growth marks of the orthodentine can be identied in polarized light (Fig. 7), although they are often obscured by the dentine tubules, micro-cracks or diagenetic colouration. In the transverse sections, the markings appear as sub-circular light and dark banded rings in polarised light, whereas they appear as light and dark band lines in longitudinal sections, paralleling the outer dentine border. The lines can be separated into two sets (Fig. 7), a very ne short period line set where the spaces between lines varies around 23 lm
Fig. 7 Detail of the orthodentine showing incremental lines of growth in polarised light in the longitudinal section (BSPG AS I 1656, Ichthyosauria indet. aff. Nannopterygius). Two sets of incremental lines are visible: the rst set is composed of very ne dark and light lines (indicated by arrow heads), interpreted as daily lines of von Ebner. The second set consists of broader dark and light bands, indicating longer time periods, i.e. Andresen lines. DB dark band, DT dentine tubules, LB light band

(=0.0020.003 mm) and a coarser, long period line set where spaces vary around 1520 lm (=0.0150.02 mm). This is in contrast to the ndings of Maxwell et al. (2011),

Ichthyosaur tooth thin-sections

who did not nd evidence for incremental lines in the orthodentine of P. australis. As discussed by Dean (1998, 2000) it is much debated how to identify the incremental growth marks in dentine, so that similarly spaced lines might be interpreted either as long or short period lines by different authors. Dean (1998, g. 7) showed that in human dentine seven to eight short period lines (i.e. daily lines von Ebner spaced\3 lm apart; after von Ebner 1902, 1906) are situated between long period lines of Andresen (after Andresen 1898), which are spaced approximately 20 lm apart. In reptiles, Erickson (1996a), when studying the dentine deposition in juvenile Crocodylus mississippiensis using double uorochrome staining, found daily growth lines of von Ebner in the crocodylian dentine similar to those found in most mammals. Erickson (1996b: p. 14625) also described the presence of lines of von Ebner in dinosaur teeth, and noted that: dinosaur incremental line widths revealed a range of 628 lm and mean values ranging from 10.1 to 19.8 lm with a general increase in incremental line widths through ontogeny in the taxa for which multiple age specimens were available, leading to calculated tooth replacement rates in dinosaurs between 46 and 777 days. Following Ericksons (1996b) approach, Sereno et al. (2007) and DEmic et al. (2009) calculated even higher tooth replacement rates for sauropod dinosaurs, based on counts of lines of von Ebner. Both works, however, did not explicitly discuss Deans (1998, 2000) articles and the possibility that the growth lines found in the sauropod teeth might also present long period lines instead of daily increments, in which case the replacement rate in sauropod dinosaurs would increase signicantly. Based on the two sets of lines found in Bauers (1898) slides, we follow Dean (1998, 2000) in identifying the ner set of growth lines as lines of von Ebner, whereas the coarser set of growth lines (or rather bands) are tentatively referred to as long period Andresen lines. The shape of ichthyosaur teeth in cross-section Bardet (1990), who analysed characteristic dental crosssections, indicated that a circular shape of the crown and a quadrangular outline in the lower part of the tooth root apparently unites all Cretaceous ichthyosaurs. Later, Maxwell and Caldwell (2006) noted that not only the Cretaceous Platypterygius and Maiaspondylus possess this character, but also the Upper Jurassic Brachypterygius, whereas the character state remained unknown in the Upper Jurassic Aegirosaurus. Given the fact that the series of transverse sections can be referred to Aegirosaurus leptospondylus (sensu Bardet and Fernandez 2000), this character can also be addressed in the serial transverse sections on slides 17 and the longitudinal section of slide

8: the sections indicate that, while the crown has a circular outline, the root of the sectioned tooth did not have a quadrangular but an oval outline. Regarding the second tooth (possibly aff. Nannoptery gius, according to Bardet and Fernandez 2000), Bauer (1898) discussed the sub-quadratic shape of the root (the shape of the external cementum) and refers to his plate 26, gure 27. Without knowing the original outer shape of the second tooth, however, the preserved longitudinal section does not yield the information necessary to address the shape of the root for this taxon.

Conclusions Despite their advanced age, the slides prepared for Bauers (1898) dissertation, are in exceptionally good condition. All histological features of the ichthyosaur teeth could be studied in the slides and all slides were matched with the original gures and descriptions of Bauer. Following Bardet and Fernandez (2000), the transversely sectioned tooth (slides 18; BSPG AS XIX 504ah) belongs to the ophthalmosaurid Aegirosaurus leptospondylus, whereas the longitudinally sectioned tooth is derived from an indeterminate ichthyosaur close to the ophthalmosaurid Nannopterygius. Studying the slides with up-to-date knowledge of ichthyosaur dentition and tooth histology further led to the recognition of an exposed layer of acellular cementum (sensu Maxwell et al. 2011) at the tooth neck, and daily and long period growth lines in the orthodentine in both specimens. Some slides of the tooth of A. leptospondylus furthermore revealed a more oval than rectangular shape of the root and the presence of peculiar vascular canals, interpreted as secondary osteodentine deposition, in the peri-pulpal orthodentine. It is therefore documented that dental characters of advanced ichthyosaurs are taxonomically useful for distinguishing at least the genera.
Acknowledgments Winand Brinkmann, Heinz Furrer, Christian Klug, Ingmar Werneburg and Jerome Gapany (all PIMUZ) are thanked for their varied help in acquiring information and discussions. Irene Schwendimann (Zivilstandsamt Zug, Switzerland), Franz Moser (Kirchheim, Germany) and the Diozesanarchiv Eichstatt (Germany) are thanked for help in further acquiring biographical data. We further would like to express our gratitude to the two reviewers Nathalie Bardet and Vivian de Buffrenil, as well as to the editors Michael J. Benton and Daniel Marty. This work was partly funded by the Swiss National Science Foundation (No. 31003A_127053 to TMS), MM is supported by the Volkswagenstiftung.

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Ichthyosaur tooth thin-sections Schmidt, W. J. (1954). Uber Bau und Entwicklung der Zahne des Knochensches Anarrhichas lupus L. und ihren Befall mit Mycelites ossifragus. Zeitschrift fu Zellforschung, 40, 2548. r Sereno, P. C., Wilson, J. A., Witmer, L. M., Whitlock, J. A., Maga, A., Ide, O., & Rowe, T. A. (2007). Structural extremes in a Cretaceous dinosaur. PLoS ONE, 2(11), e1230, 19. doi: 10.1371/journal.pone.0001230. Tomes, C. S. (1876). A Manual of Dental Anatomy: Human and Comparative (406 pp.). London: J. & A. Churchill. von Ebner, V. (1902). Histologie der Zahne mit Einschluss der Histogenese. In J. Scheff (Ed.), Handbuch der Zahnheilkunde (pp. 243299). Vienna: A. Holder. von Ebner, V. (1906). Uber die Entwicklung der leimgebenden Fibrillen, insbesondere im Zahnbein. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Klasse der Kaiserlichen Akademie der Wissenschaften in Wien, 115(Abteilung 111), 281347. von Huene, F. (1922). Die Ichthyosaurier des Lias und ihre Zusammenha nge (VIII ? 114 pp.) Berlin: Gebruder Borntraeger. Wagner, A. (1853a). Die Charakteristik einer neuen Art von Ichthyosaurus aus den lithographischen Schiefern und eines Zahnes von Polyptychodon aus dem Grunsandsteine von Kelheim. Gelehrte Anzeigen der ko niglich bayerischen Akademie der Wissenschaften, 36(3), 2532. Wagner, A. (1853b). Beschreibung einer fossilen Schildkrote und etlicher anderer Reptilien-Uberreste aus den lithographischen Schiefern und dem grunen Sandstein von Kehlheim. Abhandlungen der mathematisch-physikalischen Classe der ko niglich bayerischen Akademie der Wissenschaften, 7(1. Abtheilung), 239264.

Electronic Supplementary Material

Biography of Franz Bauer Franz Xaver Bauer was born on April, 28th 1870 in Dollnstein, a village between Solnhofen and Eichsttt, Middle Franconia, as son of the smallholder Jakob Bauer and Kreszenz Heim (Dizesanarchiv Eichsttt 1866-1884). After visiting Elementary School and Middle School he successfully completed the Catholic High School in Eichsttt in 1889. Until 1894 he studied Theology there and graduated as a priest. The following year he worked as a local pastor but then decided to begin studies in natural sciences at the Royal Ludwig-Maximilians University of Munich with a focus on geology and palaeontology (Anonymous 1903c). His supervisor became Prof. Karl Alfred von Zittel, the first conservator (director) of the Bavarian State Collection for Palaeontology and Geology (BSPG). At that time, the palaeontological collection and museum was housed in the "Wilhelminum", the old building of the Bavarian Academy of Sciences, Neuhauser Strae 51, in the city centre of Munich (Dehm 1978). The student Franz Bauer had a chamber in the neighbourhood in just five minutes walking distance (Amtliches Verzeichnis des Personals der Lehrer, Beamten und Studierenden an der kniglich bayerischen Ludwig-Maximilians-Universitt zu Mnchen, Sommersemester 1895: 44). In 1897 Franz Bauer finished his Ph. D. with magna cum laude based on a dissertation on the then known ichthyosaur material from the Upper Jurassic Solnhofen Limestone (Bauer 1898; this material is the subject of the present contribution). Afterwards in danger of unemployment, Franz Bauer had to take the job of a private teacher in Madrid for the family of the former Spanish ambassador in Washington (Anonymous 1903c, Weber 1904). However, in 1899 Ferdinand Broili, then assistant for mineralogy and geology at the Chemical Department of the Royal Technical High School (later Technical University of Munich TUM), left his position to become assistant at the BSPG (Peyer 1946; Dehm and

Schrder 1948) and the open position at TUM was offered to and filled with Franz Bauer. Immediately he became productive with scientific contributions on anthropological (Bauer 1900a,d) and palaeontological topics (Bauer 1900c). Already in 1900 he finished his habilitation thesis on a new ichthyosaur species from the Lower Jurassic of Bamberg (Bauer 1900b) and thereupon was appointed as associate professor at the Chemical Department. In this position he offered courses, which were well visited, and especially numerous field courses, in palaeontology and geology. He was also in charge of the geological collection, which he constantly improved with instructive rock and fossil samples for educational purposes (Bericht ber die Knigliche Technische Hochschule zu Mnchen fr das Studienjahr 1899-1900 ... 1902-1903; the collection unfortunately did not survive World War II, Prof. H. Scholz oral comm. Feb.2011). Somewhat surprising, during this period of his life as a natural scientist, Franz Bauer did not abandon his interest and connections to theology and so some of his writings appeared in theological journals. In an essay [transl.] "On the evolution of organic life with special consideration of vertebrates", Bauer (1900f) probably to the utmost embarrassment of some of his fellow theologists clearly took position in favour of the Darwinian theory (which he obviously had accepted as undisputable truth not even necessary to discuss). In another essay, Bauer (1900e) summarized the evidence for an older age of the Earth of probably several to 100 million years. In September 1900 he served as secretary of the section for mathematics and natural sciences at the Fifth International Congress of Catholic Savants, which was held in Munich under the presidency of the famous French geologist Albert de Lapparent. In the years from 1900 to 1903, Bauer was mostly occupied with his duties as a lecturer, and so publications during this time were limited to a number of abstracts and reviews (Bauer 1901a-e, 1902a-c), although he had several projects in progress (Weber 1904). He also became a member of the Deutsche Geologische Gesellschaft (Zeitschrift der Deutschen

Geologischen Gesellschaft. 54, Mitgliederverzeichnis 1.1.1903). In 1903, at the age of 33, Franz Bauer, now Second Order Assistant and Associate Professor for Mineralogy and Geology of the Mineralogical Laboratory of the Chemical Department at the Royal Technical High School of Munich, came to a sudden death by an unfortunate accident (Anonymous 1903c,d): On 20th of June, he joined a group of nine of his students to celebrate midsummer night on the top of the 1826 m high Risserkogel Mountain south of Tegernsee (Bavaria). When the group arrived at the summit late in the night, Franz Bauer happened to change his shoes to wear more comfortable slippers. Together with the group of young people Franz Bauer sat around a bonfire before at 1 a.m. he stood up, made a few steps aside and, probably due to the moisture of the grass, suddenly slipped over the steep northeastern mountain side and fell down a 150 meters. The students immediately undertook every effort under risk of their own lives, but could only rescue the shattered corpse on the next day. Franz Bauer's death was the subject of several short notes in newspapers (Anonymous 1903ad) and scientific journals (Anonymous 1903e,f). An official obituary (Weber 1904), although with only few biographical details, shortly characterized Franz Bauer as a highly respected colleague and proficient vigorous teacher with a bright potential. Biographical sketches were published by Wolff (1905) and Quenstedt & Quenstedt (1938). The most proliferous biographical source, however, proved to be two consecutive articles of the Mnchner Zeitung (Anonymous 1903c,d), which were obviously not known to other biographers.

Obituaries of Franz Bauer Anonymous. (1903a). Todesfall. [Dr. Franz Bauer]. Allgemeine Zeitung, 1903 (Beilage 139): 536; (Verlag der Allgemeinen Zeitung) Mnchen. (23.6.1903). Anonymous. (1903b). Unglck in den Bergen. [Dr. Franz Bauer]. Mnchener Post, 17 (138): 6; Mnchen. (23.6.1903).

Anonymous. (1903c). Das Unglck auf dem Risserkogel. [Dr. Franz Bauer]. Mnchener Zeitung Unabhngige Tageszeitung, 1903 (140): 4; Mnchen. (23.6.1903). Anonymous. (1903d). Zu dem Absturz des Herrn Dr. Bauer. Mnchener Zeitung Unabhngige Tageszeitung, 1903(141): 3. Mnchen. (24.6.1903). Anonymous. (1903e). Der Privatdozent der Geologie an der Techn. Hochschule in Mnchen Dr. Franz Bauer verunglckte am 21. Juni d. J. auf einer Exkursion bei Tegernsee. Centralblatt fr Mineralogie, Geologie und Palaeontologie, 1903(12), 394. Stuttgart: E. Schweizerbart. [7.1903]. Anonymous. (1903f). Dr. Franz Bauer [obituary]. Leopoldina, 39(8), 100. (Kaiserlich Leopoldino-Carolinische Deutsche Akademie der Naturforscher) Halle a. S. (8.1903). Quenstedt, W., & Quenstedt, A. (1938). Bauer, Franz. In K. Lambrecht, W. Quenstedt, & A. Quenstedt (Eds.), Palaeontologi Catalogus bio-bibliographicus; Fossilium Catalogus, 1: Animalia, Pars 72 (p. 25). s'Gravenhage: W. Junk. [495 pp.] Weber, M. (1904). Franz Bauer. Bericht ber die Knigliche Technische Hochschule zu Mnchen fr das Studienjahr 1902-1903, Beilage Nekrologe, 6. Mnchen: F. Straub Kgl. Techn. Hochschule. Wolff, G. (1905). Bauer, Franz, Dr. phil., Privatdozent fr Geologie und Palontologie in der Chem. Abteilung d.Techn. Hochschule in Mnchen [obituary]. Biographisches Jahrbuch und Deutscher Nekrolog, 8 (1. Januar bis 31. Dezember 1903), col. 10*; (G. Reimer) Berlin.

Other biographical sources Dehm, R. & Schrder, J. (1948). Ferdinand Broili 1874-1946. Neues Jahrbuch fr Mineralogie, Geologie und Palontologie, Monatshefte, 1945-1948, Abteilung B, Geologie und Palontologie, 1948(9-12), 257271.

Dizesanarchiv Eichsttt (1866-1884). Taufmatrikel Dollnstein, Bd. 11 (Taufen 1866-1884), 168 (No. 29). Peyer, B. (1946). Ferdinand Broili, 1874-1946. Verhandlungen der Schweizerischen Naturforschenden Gesellschaft, 1946, 358360.

Extended bibliography of Franz Bauer Bauer, F. (1898). Die Ichthyosaurier des oberen weissen Jura. [Ph. D. thesis 11.3.1897]. Palaeontographica, 44(5-6), 283328. [pls. 25-27; Stuttgart: E. Schweizerbart; 5.1898] Bauer, F. (1900a). ber den Schwund der Diploe an einem Philippinenschdel. Anatomischer Anzeiger [Centralblatt fr die gesamte wissenschaftliche Anatomie], 17(2-3), 5862. [1 fig.; Jena: G. Fischer; 15.1.1900] Bauer, F. (1900b). Ichthyosaurus bambergensis nov. sp. Beschreibung einer neuen Ichthyosaurus-Art aus dem oberen Lias vom Geisfeld nebst einigen vergleichendanatomischen Bemerkungen ber den Schultergrtel. [Habilitiationsschrift]. Bericht der naturforschenden Gesellschaft in Bamberg, 18, 1-56. [6 figs., 2 pls.; Bamberg] Bauer, F. (1900c). Osteologische Notizen ber Ichthyosaurier. Anatomischer Anzeiger [Centralblatt fr die gesamte wissenschaftliche Anatomie], 18(24), 574588. [18 figs.; Jena: G. Fischer; 31.12.1900] Bauer, F. (1900d). Ueber Schdel von den Philippinen. Archiv fr Anthropologie [Zeitschrift fr Naturgeschichte und Urgeschichte des Menschen], 27(1), 107116. [3 figs.; Braunschweig: Fr. Vieweg; Deutsche Gesellschaft fr Anthropologie, Ethnologie und Urgeschichte; 9.1900] Bauer, F. (1900e). Geologische Zeitrume. Monatsbltter fr den kathol. Religionsunterricht an hheren Lehranstalten, 1(8), 229236. [Kln: J. P. Bachem; 8.1900] Bauer, F. (1900f). Die Entwickelung des organischen Lebens mit specieller Bercksichtigung

der Wirbeltiere. Monatsbltter fr den kathol. Religionsunterricht an hheren Lehranstalten, 1(9), 257262. [Kln: J. P. Bachem; 9.1900] Bauer, F. (1901a). [Review of] Spandel, E., Eine fossile Holothurie (Synaptareste aus den oberoligocnen Cerithienschichten des Mainzer Beckens). Abh. der naturhist. Ges. Nrnberg, Bd. XIII, 1900. Geologisches Zentralblatt, 1(21), 667. [Leipzig: Borntraeger; 1.11.1901] Bauer, F. (1901b). [Abstract of] Bauer, Fr., Ichthyosaurus bambergensis nov. sp. Beschreibung einer neuen Ichthyosaurus-Art aus dem oberen Lias vom Geisfeld nebst einigen vergleichend-anatomischen Bemerkungen ber den Schultergrtel. XVIII. Ber. d. naturforsch. Ges., Bamberg, 1900, 56 Seiten, 2 Tfln. u. 6 Fig. i. Text. (Habilitationsschrift.). Geologisches Zentralblatt, 1(21), 670. [Leipzig: Borntraeger; 1.11.1901] Bauer, F. (1901c). [Abstract of] Bauer, Fr., Osteologische Notizen ber Ichthyosaurus. Mit 18 Abbildungen. Anatom. Anzeiger, XVIII. Bd., 1900. Geologisches Zentralblatt, 1(21), 670. [Leipzig: Borntraeger; 1.11.1901] Bauer, F. (1901d). [Abstract of] Ichthyosaurus bambergensis nov. sp. [Berichte der naturforschenden Gesellschaft zu Bamberg, 18]. Akten des fnften Internationalen Kongresses katholischer Gelehrten[r] zu Mnchen vom 24. bis 28. September 1900. Compte rendu du Ve Congrs scientifique international des Catholiques, 1901, 438. [Mnchen: Herder] Bauer, F. (1901e). [Abstract of] Osteologische Notizen ber Ichthyosaurier. [Anatomischer Anzeiger, 18]. Akten des fnften Internationalen Kongresses katholischer Gelehrten[r] zu Mnchen vom 24. bis 28. September 1900. Compte rendu du Ve Congrs scientifique international des Catholiques, 1901, 439-440. [Mnchen: Herder] Bauer, F. (1902a). [Review of] Roger, Otto, Ueber Rhinoceros Goldfussi Kaup und die

anderen gleichzeitigen Rhinocerosarten. Mit 2 Tafeln. 34. Bericht d. Naturwiss. Vereins f. Schwaben u. Neuburg, Augsburg, 1900, 70 S. Geologisches Zentralblatt, 2(2), 5859. [Leipzig: Borntraeger; 15.1.1902] Bauer, F. (1902b). [Review of] Spandel, Erich, Die Foraminiferen des Permo-Karbon von Hooser, Kansas, Nordamerika. Mit 10 Abbildungen. Festschr. z. Skularfeier d. nat. Ges. in Nrnberg, 1901. Geologisches Zentralblatt, 2(15), 476. [Leipzig: Borntraeger; 1.8.1902] Bauer, F. (1902c). [Review of] Spandel, Erich, Untersuchungen an dem Foraminiferengeschlecht Spiroplecta im Allgemeinen, und an Spiroplecta carinata D'Orb. im Besonderen. Mit 6 Abbildungen. Festschr. z. Skularfeier d. nat. Ges. in Nrnberg 1901. Geologisches Zentralblatt, 2(15), 476. [Leipzig: Borntraeger; 1.8.1902]

Reviews of Bauers works Huene, F. von (1903). [Review of] F. Bauer: Osteologische Notizen ber Ichthyosaurier. (Anatom. Anz. 18. 1900. 574588. 18 Fig.). Neues Jahrbuch fr Mineralogie, Geologie und Palaeontologie, 1903, II, Ref.(3), 441442. [Stuttgart: E. Schweizerbart] Keilhack, K. (1901). [Review of] Bauer, Fr., Osteologische Notizen ber Ichthyosaurier. Anatomischer Anzeiger, XVIII, 1900, 574588. Geologisches Zentralblatt, 1(23), 734. [Leipzig: Borntraeger; 1.12.1901] Koken, E. (1901). [Review of] Fr. Bauer: Ichthyosaurus bambergensis sp. n. Beschreibung einer neuen Ichthyosaurus-Art aus dem oberen Lias von Gaisfeld nebst einigen vergleichend anatomischen Bemerkungen ber den Schultergrtel. (XVIII. Ber. d. naturforsch. Ges. in Bamberg. 1900. 56 p. 2 Taf.). Neues Jahrbuch fr Mineralogie, Geologie und Palaeontologie, 1901, II, Ref.(2), 314. [Stuttgart: E. Schweizerbart] Koken, E. (1901). [Review of] Fr. Bauer: Die Ichthyosaurier des oberen weissen Jura.

(Palaeontogr. 44. 1898. 283328. 3 Taf.). Neues Jahrbuch fr Mineralogie, Geologie und Palaeontologie, 1901, II, Ref.(3), 476. [Stuttgart: E. Schweizerbart].