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Palaeontologia Electronica palaeo-electronica.org

Late jaw bones of Halecomorph (: ) studied with X-ray microcomputed tomography

Błażej Błażejowski, Paul Lambers, Piotr Gieszcz, Daniel Tyborowski, and Marcin Binkowski

ABSTRACT

New finds of Halecomorphi fish remains from limestones of Owadów-Brzezinki (Poland) are investigated using X-ray microcomputed tomography (XMT) revealing details of jaw bones for actinopterygian fish histology. Three-dimen- sional (3-D) reconstruction allows for correct taxonomical verification. The possibility of 3-D printing gives the opportunity for work with a model in any desired scale without risk of damaging the specimen.

Błażej Błażejowski. Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warszawa, Poland. [email protected] Paul Lambers. Universiteitsmuseum Utrecht, Lange Nieuwstraat 106, 3512 PN Utrecht, The Netherlands, [email protected] Piotr Gieszcz. Independent Center for Advanced Research machinic.it, Stawki 4c/10, 00-193 Warszawa, Poland, [email protected] Daniel Tyborowski. Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland, [email protected] Marcin Binkowski. Department of Computer Science and Materials Science, University of Silesia, Katowice, Poland, [email protected]

Keywords: Halecomorph fish; Late Jurassic; X-ray microcomputed tomography (XMT)

Submission: 14 October 2014. Acceptance: 21 October 2015

INTRODUCTION sils of latest Jurassic (Late ) terrestrial and marine organisms (including also remains of This paper presents the recently discovered , ammonites, horseshoe crabs, decapod jaw bones of halecomorph fish studied with X-ray crustaceans and insects) were discovered (Błaże- microcomputed tomography. The bones came from jowski, 2015; Kin and Błażejowski, 2012, 2014; new paleontological site of the Sławno limestones Błażejowski et al., 2015). at the Owadów-Brzezinki quarry (Figure 1) about The most interesting feature of the Owadów- 19 km south-east of Tomaszów Mazowiecki (Cen- Brzezinki quarry (Figure 2) is some similarity to the tral Poland), where unusually well- preserved fos- classic Solnhofen lithographic limestone (e.g.,

Błażejowski, Błażej, Lambers, Paul, Gieszcz, Piotr, Tyborowski, Daniel, and Binkowski, Marcin. 2015. Late Jurassic jaw bones of Halecomorph fish (Actinopterygii: Halecomorphi) studied with X-ray microcomputed tomography. Palaeontologia Electronica 18.3.53A: 1-10 palaeo-electronica.org/content/2015/1354-jurassic-fish-x-ray

Copyright: Palaeontological Association December 2015 BŁAŻEJOWSKI, ET AL.: JURASSIC FISH - X-RAY

Kimmeridgian lithographic limestone in Nusplin- gen (Baden-Württemberg, ) (Heineke, 1906; Heimberg, 1949; Dietl and Schweigert, 2004), Cerin (south-eastern ) (Wenz et al., 1993) and the nearby locality of Orbagnoux (Sau- vage, 1893). The fauna from the lower Tithonian lithographic limestone of Canjuers, France (Fabre et al., 1982; Peyer et al., 2014) appear to be very simliar. More or less complete and articulated fish specimens are also known from Upper Jurassic strata of England (e.g., Woodward, 1893, 1895, 1897, 1915-1917; Jain and Robinson, 1963). An overview is given in Dineley and Metcalf (1999). FIGURE 1. Map of Poland (1) with the location of the From other regions in Europe mainly fragmen- Owadów-Brzezinki Quarry (2B) near Tomaszów tary remains, such as vertebra and skull fragments, Mazowiecki. but more often scales and teeth have been described. From Switzerland the small Tithonian Munnecke et al., 2008). The identified at fish fauna of Solothurn (teeth, scales) is known, both localities indicate a similar geological and which is comparable to the fauna’s from Southern environment. The similarities refer to both marine Germany (Müller, 2011). Pictet and Jaccard (1860) and terrestrial organisms and allow comparative described fish remains (teeth, scales) from the paleontological studies at a previously unattainable Upper Jurassic of Neuchatel in Switzerland. Fricke level of high taxonomic resolution (Bechly and Kin, (1876) described a fish fauna (pycnodont teeth and 2013; Kin et al., 2012; 2013). a single articulated specimen; see also Kriwet, The best preserved, often complete and artic- 2008; Licht, 2011) from the Upper Jurassic near ulated, specimens of Upper Jurassic actinoptery- Hannover, Northwestern Germany. Kriwet (1998, gian fish fossils are known from the late 2000, 2002, 2005) and Klug and Kriwet (2012) to early Tithonian lithographic lime- reported fish remains from the Upper Jurassic of stones in the Altmühltal and nearby region (Soln- Portugal and . Fragmentary fossils are known hofen-Eichstätt area and Wattendorf, Bavaria, from several localities in France (e.g., Sauvage, Germany) (Chellouche et al., 2012; Ebert and 1902; Priem, 1912, 1917; Pharisat, 1975; Cuny et Kölbl-Ebert, 2012; Lambers, 1999), and the late al., 1991; Wenz et al., 1987; Vullo et al., 2014,

FIGURE 2. Panoramic view of the highest level of exploitation in Owadów-Brzezinki quarry (i.e., unit III and most fos- siliferous ‘Corbulomima horizon’ occurring in the middle of the quarry wall).

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FIGURE 3. Jaw bone of osteichthyan fish sp. (ZPAL P.16/O-B/1): 1-2. 3-D model of ‘virtual fossils’—a different view of the same specimen after digital processing and analysis of tomographic data. 4. vertical sections of Furo sp. teeth. 5. 3-D printed dentary – ‘virtual fossils’. 6. specimen in piece of limestone (scale bars equal 10 mm).

Cavin, 2010) and also from the Tithonian of allows us to look into the internal structure of inves- (Gemmellaro, 1871; Bassani, 1885). Most of these tigated specimens, and after computer processing, descriptions deal with caturid teeth, lepidotid to render a 3-D model (Figures 3, 4) with further scales and teeth and pycnodont teeth. Mudroch possibility of generating virtual cross-sections in and Thies (1996) and Thies and Mudroch (1996) any plan through teeth bones (Figures 3.4, 4.3). were the first to present detailed descriptions The quality of XMT imaging revealed details based on fish teeth from the Kimmeridgian of Oker important for histology studies, which may be a in Northwestern Germany. ground for later comparative studies. The Owadów- Brzezinski quarry near Tomaszow Mazowiecki (Central Poland) is an out- GEOLOGICAL AND PALEONTOLOGICAL crop of Late Tithonian strata. It may be considered SETTINGS as a close stratigraphic equivalent to the Early The study area is located approximately 18 Tithonian sediments of the Solnhofen area in Bay- km southeast of Tomaszów Mazowiecki (Central ern, Germany. Fragmentary fish fossils from Poland). At the moment, the Owadów-Brzezinki Owadow-Brzezinski have previously been figured quarry is the only place in extra-Carpathians (Kin and Błażejowski, 2012, 2013). In the present Poland where the Upper Tithonian strata are avail- paper two jaw bones will be described in more able to study (the classic locality Brzostówka is detail. now within the Tomaszów Mazowiecki town limits Since the preparation of specimens from the and inaccessible; quarries in Pomerania are rock in any conventional means was almost impos- flooded) (Kin et al., 2013). The exposed carbonate sible due to risk of unrecoverable destruction, we sequence belongs to the Kcynia Formation and used X-ray microcomputed tomography (XMT), a can be divided into four successive units. Unit I is non-invasive tool (Błażejowski et al., 2011) that

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FIGURE 4. Maxillary bone of osteichthyan fish sp. (ZPAL P.16/O-B/2): 1-2. reconstruction of 3-D ‘virtual fos- sils’— the same specimen after digital processing and analysis of tomographic data (scale bars equal 10 mm). 3. ver- tical sections of Caturus sp. teeth. 4. just after discovery.

composed of indistinctly laminated massive fine- nistic bivalve Corbulomima. It follows that the grained limestone (~6.6 m total thickness) with Del- ‘Corbulomima horizon’ in the higher portion of the toideum delta, which forms a few beds of 40–80 section (i.e., unit III), from which jaw bones of acti- cm thickness. The overlying c. 2 meters thick unit II nopterygian originate, was laid down in a is represented by thinly-bedded, fine-grained lime- very shallow marine basin, which had rather limited stones with occasional distinctive parallel lamina- links with the open sea (Kin et al., 2013). The con- tion and mass occurrence of calcareous stant proximity of the open sea is indicated by polychaete tubes in one horizon. Within this part of occasional finds of ammonites of the Zara- the profile quite common specimens of decapod iskites (Semenov). Other fossils are rare and rep- crustaceans (lobsters) were found. Unit III, c. 15.6 resented by exceptionally well-preserved m thick, is highly fossiliferous and has yielded the horseshoe crabs, disarticulated remnants of vari- specimens that are the subject of this study. Unit ous marine and land arthropods (including deca- IV, c. 2.3 m in thickness, the top being not pods, beetles, and grasshoppers) and exposed, is developed as organodetrital limestone moulds of ammonite shells. Moreover, teeth, bones rich in Nanogyra oysters, bryozoans and serpulids. and rarely partially articulated skeletons of various They often form small bioherms. In general, units I, occur. II and III probably represent a transition from an offshore to nearshore, perhaps lagoonal, setting, METHODOLOGY whereas unit IV bears evidence of a return to more X-ray microcomputed tomography (XMT), is open marine conditions. Below the Kcynia Fm yel- an imaging technique, which detects differences in lowish marls and marly clays of the Pałuki Forma- the attenuation of an X-ray beam propagating tion occur. through a solid object, and as a result detailed The uppermost part of the unit (III) is highly information about internal structure is obtained. fossiliferous, with a horizon of finely bedded fine- This data after computer processing allows us to grained limestones at its base (also called the ‘Cor- reconstruct a virtual 3-D model of studied bones bulomima horizon’), dominated by small opportu-

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(Figures 3, 4), and after removing the background The triangular crown is markedly narrower than the from the images by thresholding, a set of virtual shaft. The bases of the teeth are broad and flat- cross-sections through teeth can be generated tened laterally. The teeth are about 2 mm long. (Figures 3.4, 4.3) (Błażejowski et al., 2013; Sutton Seven teeth are complete and eight are broken off et al., 2014). We used the most effective methods or incomplete, and only their bases are preserved. for digital processing and analysis of tomographic These are the teeth in the positions 1-3, 5, 7, 9, 11 data, enabling the construction of isosurface-based and 14. Lane and Ebert (2012) describe a similar and volume-based 3-D ‘virtual fossils’, which can situation in a specimen of Furo muensteri, in which, be manipulated and dissected interactively. The besides complete teeth, incomplete teeth or tooth resulting images are similar to traditional ones bases are present and also isolated acrodin caps. obtained by destructive slicing, and the resolution The dentigerous border bends down slightly at the here is limited by voxel size of computed model, level of the eighth tooth and forms a round anterior reaching 22.4 × 22.4 × 22.4 μm. The collected edge. In the lateral side of the bone, below the six material is housed at the Institute of Paleobiology, anterior teeth six pores of the sensory canal are Polish Academy of Science in Warsaw (ZPAL P.16/ present, the canal running parallel to the dentiger- O-B). ous border. From the level of the seventh tooth the sensory canal runs more ventrally in the dentary. DESCRIPTION Posteriorly from about the level of the tenth tooth the canal is housed in a groove. Medially there is a Specimen A meckelian groove that runs form posterior to ante- Specimen (A) is the left dentary of the lower rior. jaw of an actinopterygian fish. It has previously Identification. The original identification in Kin and been figured in Kin and Błażejowski (2012, figure Błażejowski (2012) and Kin et al. (2013) of the jaw 10) and Kin et al. (2013, figure 5A) as a ?Pleuro- bone as a ?Pleurosoaurus sp. is erroneous. The saurus sp. The bone is slender and elongate, 3.4 general shape of the dentary, elongated and with a cm long and slightly curved inwards (Figure 3). The relatively long dentigerous border and pronounced ventral border slightly undulates. The outer surface coronoid process, is as known from halecomorph is striated and wrinkled near the dentigerous bor- fishes (e.g., see figures in Grande and Bemis, der. The pores of the sensory canal are obvious. 1998; Lambers, 1994, 1998). To the Halecomorphi The bone forms a blunt, rounded anterior sym- belong the Parasemionotiformes (e.g., Watsonulus physial region. The dentary consists of an anterior from Madagascar, from East Greenland and dentigerous portion and posteriorly a pronounced from , see Quanguo, 2009), Ionos- coronoid process. The dentigerous portion is 2.1 copiformes (e.g., Ionoscopus, Furo), and Amii- cm long and about 6 mm high halfway its length. formes, the latter including the The coronoid process is 4 mm high. The dorsal- (Amblysemius, Caturus) and , (recent most point of the coronoid process is at the level of calva and its closest relatives, such as dorsal tips of the teeth. It seems that the coronoid Pachyamia, Solnhofenamia and Amiopsis) (Chalifa process is damaged, and a part is broken off pos- and Tchernov, 1982; Grande and Bemis, 1998; terodorsally. The posterior border that sutures with Martín-Abad and Poyato-Ariza, 2013). Throughout the angular is deeply V-shaped invaginated and the Mesozoic many halecomorph genera have has a posterior ventral process that projects further been described worldwide, but from the posteriorly than the posterior border of the coro- onwards they are restricted to the Amiinae noid process. (Cyclurus and Amia) only (Martín-Abad and Poy- There are seven spaced, pointed and conical ato-Ariza, 2013; Poyato-Ariza and Martín-Abad, teeth that are strongly recurved inwards almost 2013). bending over, and eight broken or incomplete The height of the coronoid process might be a teeth. distinctive feature. In the present specimen the cor- Dimensions, outlook, visible details, internal onoid process and the teeth are of approximately structure, composition. All teeth have an acrodin the same height. This is comparable to the lower crown, without lateral keels. The acrodin is a typi- jaw figured in a specimen of ‘Furo’ microlepidotes cal enameloid tissue in actinopterygians. It is usu- (Lambers, 1998, figure 5). In Solnhofenamia, Ami- ally found at the tip of the tooth and is called “cap opsis and Furo muensteri the coronoid process enameloid” or “acrodin cap” (Ørvig, 1978; seems higher than the tips of the teeth (Grande Sasagawa et al., 2009; Shellis and Miles, 1974). and Bemis, 1998; Lambers, 1994, pers.obs.; Lane

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and Ebert, 2012). In Amblysemius the tips of the lepidotes is known from the limestones of Soln- teeth and the coronoid process are approximately hofen (Lambers, 1998). at the same level. But in all these figured speci- Specimen B mens the outline of the posterior part of the dentary and coronoid is partly obscured by other bones, This fossil is figured in Kin and Błażejowski such as the maxilla, which makes an exact obser- (2012, figure 9) and Kin et al. (2013, figure 5B) as a vation difficult. dentary bone of Caturus sp. The specimen is inter- In figures 43-45 of Grande and Bemis (1998) preted as a fragment of a left maxilla, anteriorly the height of the coronoid process of the dentary of and posteriorly incomplete (Figure 4). The frag- Amia, with respect to the much lower teeth, is ment is around 5,9 cm long. This indicates a large clearly visible. fish of at least 1 meter long. Nine teeth are more or The deep V-shaped invagination of the poste- less completely preserved, and in three teeth only rior border and the long ventral posterior portion the sockets are visible. The teeth are roughly are conspicuous. In a specimen of ‘Furo’ microlepi- between 1 cm long and 0,5 cm broad. All teeth are dotes a deeply invaginated posterior border of the longer than the depth of the supporting bone. They dentary, with a ventral process that extends slightly are conical, with a broad and laterally compressed, further posteriorly than the posterior border of the slightly indented base. All teeth have an acrodin coronoid process has been figured (Lambers, cap. The most anterior tooth is best preserved and 1998, figure 5). shows a small, arrow cap with anterior and poste- In Solnhofenamia the posterior border is only rior keels. The dorsal border of the bone slightly a little invaginated, or in some specimens not undulates. invaginated at all and rather convex (Grande and Identification. The arrow shaped crowns of the Bemis, 1998). In Amiopsis lepidota the posterior teeth with anterior and posterior flanges resembles border is deeper invaginated than in Soln- those of the halecomorph Caturus (Mudroch and hofenamia, especially in the specimen in figure 361 Thies, 1996). in Grande and Bemis (1998). But the ventral por- The specimen resembles the maxilla of the tion of the dentary does not proceed beyond the large Portlandian C. cliftoni figured by Woodward posterior border of the coronoid process. In Ambly- (1895, pl. VII, 5, pers. obs.) in that the teeth are semius the dentary is slightly invaginated (Lam- longer than the depth of the supporting bone and bers, 1994, figure 3; Grande and Bemis, 1998, somewhat slender, but it differs in that they are figure 405). In Furo muensteri the dentary sutures more widely spaced. From the contemporaneous posterodorsally along an interdigitating zig-zag Upper Jurassic German localities C. giganteus is structure with a long posteriorly projected ventral known, of which specimens of up to 1 meter are portion (Lane and Ebert, 2012, figure 4). collected. The maxillary teeth of C. giganteus are The shape of the teeth, with a crown clearly however not much longer than the depth of the narrower than the shaft, is similar to that figured of supporting bone (Lambers, 1994, figure 3). The teeth of Ionoscopus (Mudroch and Thies, 1996). specimen cannot be determined more precisely But the crown lacks keels, whereas Ionoscopus than as Caturus sp. teeth have two small mesial and distal keels (see also Kriwet, 1998; Prasad et al., 2004). Caturus DISCUSSION teeth have a distinctive arrowhead shaped crown Besides irreversible damage of investigated with large lateral keels (Mudroch and Thies, 1996). specimens, one of the most important disadvan- Among amiids only the Vidalamiinae have keeled tages of traditional sectioning is very limited infor- crowns. Keeled crowns have not been described in mation about volumetric dependencies within the species of Furo. internal structure (histology). X-ray tomography The number of teeth ‘Furo’ microlepidotes, with proper images post processing allows genera- Furo muensteri Solnhofenamia elongata, Amiopsis tion of a 3-D model in a form of “virtual fossil” can lepidota, Amblysemius pachyurus, is comparable, be manipulated, measured and dissected interac- all these have about 13-15 teeth in the lower jaw. tively (Błażejowski et al., 2011). It is also possible Considering the shape and relative dimen- to print a reconstructed shape on a 3-D printer in sions of the dentary and the characteristics of the actual size or any desired magnification (Figure dentition as outlined. The specimen is tentatively 3.5), with all internal structure retained. Sample identified as cf. ‘Furo’ microlepidotes. ‘Furo’ micro- visualization (3-D model) of the discussed dental bones are shown in Supplemental Materials.

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A detailed projection of investigated fossil, δ18O) and X-ray microtomography. Polish Polar including parts that are hidden in rock, (Figure 3.6) Research, 34(1):23-38. allows us to identify the to the genus Błażejowski, B., Gieszcz, P., Brett, C.E., and Binkowski, level. It turns out that first, preliminary identification M. 2015. A moment from before 365 Ma frozen in after examining visible fragments of specimen as a time and space. Scientific Reports, 5, 14191; doi: mandible of aquatic sphenodontian Pleurosaurus 10.1038/srep14191 (2015). Cavin, L. 2010. Diversity of Mesozoic semionotiform ex gr. goldfussi (Kin et al., 2013) was wrong. Cur- fishes and the origin of (Lepisosteidae). Natur- rent study indicates that the specimen is a dentary wissenschaften, 97:1035-1040. bone of halecomorph fish Furo sp. (Figure 3). Chalifa, Y. and Tcherenov, E. 1982. Pachyamia latimaxil- Both jaw fragments belong to halecomorph laris, new genus and species (Actinopterygii: Amii- fishes known from the Upper Jurassic deposits dae), from the Cenomanian of Jerusalem. Journal of from Europe and from the Tithonian limestone of Paleontology, 2:269-285. Solnhofen in particular. The limestone of Owadów- Chellouche, P., Fürsich, F.T., and Mäuser, M. 2012. Brzezinski is stratigraphically close to Solnhofen, Taphonomy of neopterygian fishes from the Upper being about 5 Ma older. Preliminary analysis of its Kimmeridgian Wattendorf Plattenkalk of Southern fauna has revealed that the land and marine mac- Germany. Palaeobiodiversity and Palaeoenviron- rofauna of both localities is very similar. The pres- ments, 92(1):99-117. Cuny, G., Buffetaut, E., Cappetta, H., Martin, M., Mazin, ently described jaw bones that most probably J.M., and Rose, J.M. 1991. Nouveaux restes de belong to species known previously only from Sol- Vertébrés du Jurassique terminal du Boulonnais nhofen strongly support this view. (Nord de la France). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 180(3):323-347. ACKNOWLEDGMENTS Dietl, G. and Schweigert, G. 2004 . The Nusplingen litho- graphic limestone - a "fossil lagerstaette" of late Kim- This work was supported by the Polish meridgian age from the Swabian Alb (Germany). National Science Centre (grant number 2012/07/B/ Rivista Italiana di Paleontologia e Stratigrafia, ST10/04175). The funders had no role in study 110(1):303-309. design, data collection and analysis, decision to Dineley, D. and Metcalf, S. 1999. Fossil Fishes of Great publish, or preparation of the manuscript. No addi- Britain, Geological Conservation Review , No. tional external funding was received for this study. 16, Joint Nature Conservation Committee, Peterbor- We sincerely thank the three anonymous reviewers ough, 675 pp. for their insight full comments on our manuscript. Ebert, M. and Kölbl-Ebert, M. 2012. Grabunsbericht Ettling 2012: Fund-Highlights und neue For- REFERENCES schungsergebnisse. , 30:23-37. Fabre, J.A., de Broin, F., Ginsburg, L., and Wenz, S., Bassani, F. 1885. Avanzi di pesci oolitici nel Veronese. 1982. Les vertebres du Berriasien de Canjuers (Var, Atti della Società Italiana di Scienze Naturali, France) et leur environnement. Geobios, 15(6):891- 23(1):142-163. 923. Bechly, G. and Kin, A. 2013. First record of the fossil Fricke, K. 1876. Die fossilen Fische aus den oberen family Eumorbaeschnidae from the Upper Juraschichten von Hannover. Palaeontographica, Jurassic of Poland. Acta Palaeontologica Polonica, 22:45-87. 58(1):121-124. Gemmellaro, G.G. 1871. Studi paleontologici sulla fauna Błażejowski, B. 2015. The oldest species of the genus del Calcare a Terebratula janitor del nord di Sicilia. Limulus from the Late Jurassic of Poland. 3-14. In Giornale di Scienze Naturali ed Economiche 6:153- Carmichael, R.H., Botton, M.L., Shin, P.K.S., and 187. Cheung, S.G. (ed.), Changing global perspectives on Grande, L. and Bemis, W.E. 1998. A comprehensive biology, conservation, and management of horse- phylogenetic study of amiid fishes (Amiidae) based shoe crabs. Springer, New York 2015. on comparative skeletal anatomy. An empirical Błażejowski, B., Binkowski, M., Bitner, M.A., and search for interconnected patterns of natural history. Gieszcz, P. 2011. X-ray microtomography (XMT) of Society of Vertebrate Paleontology Memoir 4: i-x, 1- fossil brachiopod shell interior for taxonomy. Acta 690; supplement to Journal of Vertebrate Paleontol- Palaeontologica Polonica, 56(2):427-428. ogy, 18(1). Błażejowski, B., Duffin, C., Gieszcz, P., Małkowski, K., Heimberg, G. 1949. Neue Fischfunde aus dem Weissen Binkowski, M., Walczak, M., McDonald, S.A., and Jura zeta von Württemberg. Palaeontographica, A, Withers, P. 2013. Lower Saurichthys (Pisces, 97:75-98. Actinopterygii) teeth from Spitsbergen, with com- Heineke, E. 1906. Die Ganoiden und Teleostier des ments on their stable isotope composition (δ13C and lithografischen Schiefers von Nusplingen. Geolo-

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Geobios, 27, Supple- Jurassic and of India, p. 625-638. In ment 1:91-99. Arratia, G. and Tintori, A. (ed.), Mesozoic Fishes 3 – Lambers, P.H. 1998. The genus Furo (Pisces, Haleco- Systematics, Palaeonvironments and Biodiversity. morphi) from the Upper Jurassic Plattenkalke of Ger- Priem, F. 1912. Sur des poissons des terrains Secon- many. Oryctos, 1:23-35. daires du sud de la France. Bulletin de la Societé Lambers, P.H. 1999. The actinopterygian fish fauna of géologique de France, sér. 4, 12:250-271. the Late Kimmeridgian and Early Tithonian ‘Platten- Priem, F. 1917. Sur des poissons des terrains Secon- kalke’ near Solnhofen (Bavaria, Germany: state of daires du sud de la France. (Note complémentaire) the art.). Geologie en Mijnbouw, 78:215-229. Bulletin de la Societé géologique de France, sér. 4, Lane, J.A. and Ebert, M. 2012. Revision of Furo muen- 16:286-297. steri (Halecomorphi, Ophiopsidae) from the Upper Quanguo, LI, 2009. A new parasemionotid-like fish from Jurassic of Western Europe, with comments on the the Lower Triassic of Jurong, Jiangsu province, genus. Journal of Vertebrate Paleontology 32(4):799- South China. Palaeontology, 52(2):369-384. 819. Sasagawa, I., Ishiyama, M., Yokosuka, H., Mikami, M., Licht, M. 2011. A short contribution about the pycnodont and Uchida, T. 2009. Tooth enamel and enameloid in fishes (Actinopterygii, Neopterygii) from Lower Sax- actinopterygian fish. Frontiers of Materials Science in China, 3:174-182.

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Sauvage, H.E. 1893. Note sur quelques Poissons du dus Palevol. http://dx.doi.org/10.1016/ Calcaire bitumineux d'Orbagnoux (Ain). Bulletin de j.crpv.2014.03.003 Societé d’Histoire Naturelle d’Autun 7:427-443. Wenz, S., Bernier, P., Barale, G., Bourseau, J.-P. Buffe- Sauvage, H.E. 1902. Recherches sur les vertébrés du taut, E., Gaillard, C., and Gall, G.-C. 1993. L’ ichthyo- Kimméridgien supérieur de Fumel (Lot-et-Garonne). faune des calcaires lithographiques du Kimméridgien Mémoires de la Société géologique de France. Palé- supérieur de Cerin (Ain, France). Geobios, Mém. ontologie, 25, 32 pp. Spec. 16:61-70. Shellis, R.P. and Miles, A.E.W. 1974. Autoradiographic Wenz, S., Breton, G., and Lepage, G. 1987. Presence du study of the formation of enameloid and dentine genre Caturus (Pisces, Actinopterygii, Caturidae) matrices in fishes using tritiated amino acid. dans le Kimmeridgien superieur des environs du Proceedings of the Royal Society of London B. Havre (Normandie, France). Bulletin trimestriel de la 185:51-72. Société géologique de Normandie et des Amis du Sutton, M.D., Rahman, I., Garwood, R.J. 2014. Tech- Muséum du Havre, 74(1):21-27. niques for Virtual Palaeontology. Wiley-Blackwell, Woodward, A.S. 1893. On some British Upper-Jurassic 208 pp. fish-remains, of the genera Caturus, Gyrodus and Thies, D. and Mudroch, A. 1996. Actinopterygian teeth Notidanus, Annals and Magazine of Natural History, from the Late Jurassic (Kimmeridgian) of N-Ger- series 6, 12:398-402. many, p.105-114. In Arratia, G. and Viohl, G. (ed.), Woodward, A.S. 1895. Catalog of fossil fishes in the Brit- Mesozoic Fishes - Systematics and Paleoecology. ish Museum (Natural History), 3. xlii + 554 pp. Lon- Proceedings of the international meeting, Eichstätt don: British Museum (Natural History). 1993. Woodward, A.S. 1897. A contribution to the osteology of Vullo, R., Abit, D., Ballèvre, M., Billon-Bruyat, J.-.P., the Mesozoic amioid fishes Caturus and Osteora- Bourgeais, R., Buffetaut, E., Daviero-Gomez, V., chis. II. Osteorachis leedsi, sp. n., from the Oxford Garcia, G., Gomez, B., Mazin, J.-M., Morel, S., Clay of Peterborough. Annals and Magazine of Natu- Nèraudeau, D., Pouech, J., Rage, J.-C., Schnyder, ral History ser. 6, 19:379-387. J., and Tong, H. 2014. Palaeontology of the Purbeck- Woodward, A.S. 1915-1917. The fossil fishes of the type (Tithonian, Late Jurassic) bonebeds of Chas- English Wealden and Purbeck Formations. London. siron (Oléron Island, western France). Comptes Ren- 148 pp.

9 BŁAŻEJOWSKI, ET AL.: JURASSIC FISH - X-RAY

SUPPLEMENTAL MATERIAL

Caturus sp. (top) and Furo sp. (bottom). See palaeo-electronica.org/content/2015/1354-jurassic- fish-xray for animation.

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