Echinoid bite traces on Late Cretaceous (lower Maastrichtian) sea
Transcription
Echinoid bite traces on Late Cretaceous (lower Maastrichtian) sea
Papers in Press “Papers in Press” includes peer-reviewed, accepted manuscripts of research articles, reviews, and short notes to be published in Paleontological Research. They have not yet been copy edited and/or formatted in the publication style of Paleontological Research. As soon as they are printed, they will be removed from this website. Please note they can be cited using the year of online publication and the DOI, as follows: Humblet, M. and Iryu, Y. 2014: Pleistocene coral assemblages on Irabu-jima, South Ryukyu Islands, Japan. Paleontological Research, doi: 10.2517/2014PR020. doi:10.2517/2016PR015 Echinoid bite traces on Late Cretaceous (lower Maastrichtian) sea lilies from southern Poland Tomasz Brachaniec1, Rafał Lach2, Mariusz A. Salamon3* and Krzysztof R. Brom3 Ac 1 pt ce Department of Geochemistry, Mineralogy and Petrography, Faculty of Earth Sciences, University of Silesia, Będzińska Street 60, 41-200 Sosnowiec, Poland; e-mail: ed tomasz.brachaniec@o2.pl. Centre for Polar Studies KNOW (Leading National Research Centre) WNoZ UŚ m Department of Palaeontology and Stratigraphy, Faculty of Earth Sciences, University of c us an 2 Silesia, Będzińska Street 60, 41-200 Sosnowiec, Poland; e-mail: rafal_lach86@o2.pl 3 Department of Palaeontology and Stratigraphy, Faculty of Earth Sciences, University of rip Silesia, Będzińska Street 60, 41-200 Sosnowiec, Poland; paleo.crinoids@poczta.fm, t kbrom@us.edu.pl. Centre for Polar Studies KNOW (Leading National Research Centre) WNoZ UŚ * corresponding author Abstract: Echinoid bite traces on Late Cretaceous (lower Maastrichtian) bourgueticrinids and isocrinids of southern Poland (Miechów Trough) were documented. The bitten sea lilies co- occurred with Goniopygus, a regular echinoid possessing an Aristotle's lantern. This is the first record of Goniopygus in the lower Maastrichtian deposits of Poland. Considering former studies, as well as direct in situ observations of extant sea lilies and sea urchin behavior, the traces at hand could be most likely linked with predatory actions of Goniopygus echinoid. Such studies on predatory phenomena are crucial and could provide baseline data concerning the evolutionary trends among organisms engaged in the “arms race”. Introduction ed pt ce Ac Key words: Echinoidea, Crinoidea, Maastrichtian, Cretaceous, predation, Poland Along one of the longest Cretaceous epicontinental sedimentary sections in Central Europe m (almost 30 km), exposed during the construction of European route E77 between Cracow and c us an Warsaw (Poland), a few hundred of fossils represented by numerous crinoids, irregular echinoids, ammonites, belemnites, inoceramids, sponges and coral specimens were recently documented (details in tables 1-3 in Salamon et al., 2016). A bourgueticrinid mass rip accumulation, described as a concentration Lagerstätte, was also discovered (Salamon et al., t 2016). Its origin is probably related to a storm event (details in Salamon et al., 2016 and literature cited therein). During the last field works, carried out in a previously (first half of 2015) unexposed part of the road in construction (Jędrzejów city), lower Maastrichtian sediments had been found which yielded numerous crinoid specimens. Among them, the occurrence of bourgueticrinids and isocrinids was confirmed. Oval pits on the lateral surface of pluricolumnals and columnals were observed and subsequently interpreted as predation traces probably left by regular echinoids belonging to genus Goniopygus. The remains of this latter were found in the same layer as those of crinoids. It is worth mentioning that records of Cretaceous echinoids, possessing an Aristotle's lantern enabling them to prey on crinoids, are relatively rare in Poland. Mączyńska (1984 and literature therein) listed almost 60 echinoid taxa observed in Cretaceous epicontinental sediments, but only five represent regular taxa, although no Goniopygus specimen has ever been observed. Somewhat earlier, Halicki (1939) documented this taxa in the Cenomanian of Poland. Thereby, the specimen at hand is the first documented Goniopygus species from the lower Maastrichtian sediments of Poland. Ac Material and methods pt ce Field studies have been carried out in October 2015 during the construction of European route E77 in Jędrzejów city, in the Miechów Trough area in southern Poland (Fig. 1). The Miechów ed Trough is located between the Mesozoic margin of the Holy Cross Mountains in the east and the Kraków-Częstochowa Homocline in the west. It is filled by Upper Cretaceous sediments m (Albian-lower Maastrichtian), with a total thickness of which ranged from 800 to 1000 m (for c us an details see Bieńkowska-Wasiluk et al., 2015; Lach, in press and literature therein). The sedimentary series of Miechów Trough starts with Albian deposits generally covering t et al., 2016). rip Kimmeridgian rocks or oldest part of Jurassic and Maastrichtian deposits at the top (Salamon Figure 1 around here During the road construction, 3 m of lower Maastrichtian lithological section was exposed, composed of gaizes, opokas and locally intercalated by marls (Fig. 1). The Cretaceous sediments of this previously unknown section were documented and sampled. In one layer (see Fig. 1), 8 cups, over 100 pluricolumnals, columnals, radicular cirri of bourgueticrinid crinoids, 43 pluricolumnals and columnals of isocrinids and 34 echinoid fragments of Goniopygus (test plates from 1 isolated to up to 10 connected elements) were collected (Fig. 2J-M). They co-occurred with irregular echinoids (Micraster, Echinocorys) preserved as more or less complete tests, often compacted. Additionally, ammonite specimen of Acanthoscaphites tridens (Kner) indicative an early Maastrichtian age (Kin, 2009; Machalski, 2010), were also observed in the field. For further studies, two bulk samples (ca. 8 and 10 kg) were taken to the Laboratory of Ac Palaeontology and Stratigraphy Department, University of Silesia. The first one was soaked in pt ce Glauber's salt (sodium sulfate), frozen up to 8 times at a temperature -16 oC, and then boiled in a Glauber's salt diluted solution. The sample was washed and sieved through a screen ed column (mesh size: 1.0, 0.5 and 0.315 mm). The invertebrate faunal elements were picked m under a stereoscopic microscope (SM800T). On the whole, over 40 faunal remains were c us an retrieved. Among them crinoids (columnals, pluricolumnals, radicular cirri) and fragments of irregular echinoids were evidenced. Some of these elements, probably due to maceration procedure, were strongly corroded. rip The second sample was macerated following Boczarowski’s (2001) method. All rocks t were etched in buffered acetic acid (a mixture of ½ supersaturated acetate, ¼ 30% acetic acid, and ¼ water). After 5 days, the resulting residues was washed out and sieved through the standard screen column (mesh size: 1.0, 0.5 and 0.315 mm). The material was then dried at 150 oC. Some material was further subjected to an additional dissolution-washing procedure described above for up to about 15 days. Specimens treated this way did not display any trace of corrosion. 33 crinoid specimens (bourgueticrinid and isocrinid columnals and pluricolumnals) and echinoid test plates (from 1 to 3 plates) were also evidenced. The described material is housed in the collections of the Laboratory of Palaeontology and Stratigraphy Department, Faculty of Earth Sciences at the University of Silesia, Sosnowiec, Poland (acronymed: GIUS). Results On one cup specimen three perpendicular scratches were observed. Similar traces have been illustrated by Jagt and Salamon (2007, pl. 3, fig. 4), Salamon and Gorzelak (2010, pl. 6D) and Ac Salamon et al. (2016, fig. 2a) and ascribed to fish predation traces. On the other hand, such pt ce traces could have been produced by echinoids (detailed discussion in Salamon et al., 2016; Fig. 2G, H). The other specimens did not bear similar traces. However, bourgueticrinid and ed isocrinid pluricolumnals and columnals are displaying numerous clear pits (frequency estimate: 9%). These traces are comparable in shape and size to those produced by recent m echinoids (Baumiller et al., 2008, 2010); Fig. 2A-F. Similar bite traces on fossil crinoids have c us an been illustrated and ascribed to echinoid predation (e.g., Gorzelak and Salamon, 2009). They are oval or drop-like in shape , ranging from 0.2 to 1.1 mm in diameter. The pit depth ranges rip up to 1.1 mm. All observed marks are sharp and occur on the external part of the lateral t surfaces of columnals that are distinct from abrasion. This suggests that the crinoid material was ‘fresh’ and alive when the marks were made. No overgrowth, regeneration or repair signs have been documented which imply that the predation was lethal (comp. Fig. 2A-I). In addition, on the ossicle surface, both the lateral and articular surfaces, abrasion traces were observed (ca. 4% of all investigated ossicles). It is most probably the mere sign of transporting or deposition of these remains. Majority of ossicles are, however, perfectly preserved suggesting that the transport was not very long (details in Gorzelak and Salamon, 2013). Mineralization (iron oxides) is observed on only 2% of the documented specimens. Slightly more abundant is the chemical dissolution (5% of investigated ossicles), which might be evidence of post-mortem decomposition, diagenesis or digestion by predators. Bioerosion was observed on 8% specimens. Mostly irregular “holes”, ranging from 0.3 to 3.4 mm, probably produced by sponges, were documented. This indicates that majority of crinoid remains were resting on the sea bottom for a short period of time before being burried in the sediments. The absence of elements with incrustation traces also supports this hypothesis. Discussion Ac pt ce Praedichnia are one of the ichnofossil types (trace fossil) that directly evidence the interactions between a predator and its prey (Ekdale, 1985). However, distinguishing ed predation traces from those induced by other factors, i.e. biotic or abiotic (e.g., abrasion, bioerosion, corrosion, post-diagenetic fracturing or pressure dissolution) is a permanent m challenge for the palaeontologists’ community. Furthermore, identification of what kind of c us an potential predator could leave such given predation trace is often not obvious (Zatoń et al., 2007; Gorzelak and Salamon, 2009; Gorzelak et al., 2010; Lach et al., 2015 and literature rip therein). However, such differentiations are crucial to obtain sound and reliable conclusions concerning palaeoecology. Additionally, studies on the predatory phenomenon might provide t secondary baseline data concerning the evolutionary trends of organisms running an “arms race” (Baumiller and Gahn, 2004; Baumiller et al., 2008; Gorzelak and Salamon, 2009; Baumiller et al., 2010; Gorzelak et al., 2012). Distinguishing predation traces from those resulting of any other process or action relies on criteria listed by Zatoń et al. (2007), Gorzelak and Salamon (2009), Baumiller et al. (2010), Gorzelak et al. (2012) and Lach et al. (2015). As mentioned above, the collected specimens exhibit a low level of abrasion and are relatively well preserved. Thus, they were probably not transported for a long time and over a long distance; otherwise, the traces of predation would become blurred and unidentifiable. Most likely, most of the material was buried soon after death of the individuals (Gorzelak and Salamon, 2013; Salamon et al., 2016). Presence of a clear well-preserved stereom without a stair-step like surfaces disqualifies abrasion as the cause of the pits (Villier, 2008). The traces at hand were also not procuded by bioerosion, pressure dissolution or post-diagenetic fracturing as there is no evidence for the presence of deeply and irregularly penetrating (not Ac tapering) holes towards the interior (bioerosion), stylolites or “fitted” fabrics (pressure pt ce dissolution) and rhombohedral depressions along the calcite-cleveage (post-diagenetic fracturing); details in Salamon and Gorzelak (2010). ed The pits conform in size and shape to biometric parameters of traces left by regular m echinoids (Results section). Indeed, recent, regular echinoids exhibit wide spectrum of food c us an type. The Aristotle's lantern is adaptet for biting, tearing and rasping and can function as a "grab". Food resorces available for them include inter alia soft- and hard bodies animals, mainly corals and shelly fauna (de Ridder and Lawrence, 1982). Additionally it has been rip showed that cidaroid sea urchins are feeding actively on stalked crinoids (direct in situ t observations by Baumiller et al. 2008). Crinoid ossicles with Aristotle's lantern traces were extracted from cidaroid guts (Baumiller et al. 2008, fig. 7c, f). They are in the form of shallow pits up to about 1mm in diameter. These observations support the interpretation of the fossil crinoid material with bite marks, co-occurring with echinoids provided with an Aristotle's lantern and that were not probably left by vertebrates. Similar conclusions were reached by Gorzelak and Salamon (2009) with Late Jurassic stalked crinoids. The latter authors illustrated similar shallow bite marks with circular, oval or droplike outline. Their size ranges from 0,15 to 0,8 mm. Such traces are also known from the Middle Triassic or Late Jurassic crinoids of Poland (Gorzelak et al., 2012). However, they are slightly more elongated (e.g., Lach et al., 2015), although not as long as those observed in the vertebrate predations (Nebelsick, 1999; Neumann, 2000 and literature therein). In the present study, the latter type of praedichnia was documented on a single cup specimen. Three straight, sharply-edged depressions, parallel to each other, are probably the traces of a toothed predator (e.g., Salamon et al., 2016). It is not possible to determine which phylum this predator belongs to, besides no vertebrate fossil has been found so far. However, they might belong to chimaeras, rays or juvenile? reptiles. Sharks and other durophagous fishes are discarded as potential predators, ed Acknowledgements pt ce pers.comm.). Ac because of their pre-Maastrichtian extinction, at least locally (Dr. R. Niedźwiedzki, m The comments of two reviewers, Dr. Andreas Kroh (Natural History Musuem, Vienna, c us an Austria) and an anonymous reviewer that helped to improve the manuscript. Dr. Andreas Kroh (Natural History Musuem, Vienna) is thanked for help with the echinoid identification rip (Goniopygus). This project was partly supported by NCN grant no. DEC- 2012/07/N/ST10/04103. The project has also been granted by the Leading National Research t Centre (KNOW) to the Centre for Polar Studies for the period 2014-2018. References Baumiller, T.K. and Gahn, F.J., 2004: Testing predation-driven evolution using mid-Paleozoic crinoid arm regeneration. Science, vol. 305, p. 1453-1455. Baumiller, T.K., Mooi, R. and Messing, C.G., 2008: Urchins in the meadow: paleobiological and evolutionary implications of cidaroid predation on crinoids. Paleobiology, vol. 34, p. 2234. Baumiller, T.K., Salamon, M.A., Gorzelak, P., Mooi, R., Messing, C.G. and Gahn, F.J., 2010: Post-Paleozoic crinoid radiation in response to benthic predation preceded the Mesozoic marine revolution. Proceedings of the National Academy of Sciences of the United States of America, vol. 107, p. 5893-5896. Bieńkowska-Wasiluk, M., Uchman, A., Jurkowska, A., Świerczewska-Gładysz, E., 2015: The Ac trace fossil Lepidenteron lewesiensis: a taphonomic window on diversity of Late Cretaceous pt ce fishes. Paläontologische Zeitschrift, vol. 89, p. 795-806. Boczarowski, A., 2001: Isolated sclerites of Devonian non−pelmatozoan echinoderms. ed Palaeontologia Polonica, vol. 59, p. 3-220. m Ekdale, A.A., 1985: Palaeoecology of the marine endobenthos. Palaeogeography, c us an Palaeoecology, Palaeoclimatology, vol. 50, p. 63-81. Gorzelak, P. and Salamon, M.A., 2009: Signs of benthic predation on Late Jurassic stalked t rip crinoids, preliminary data. Palaios, vol. 24, p. 70-73. Gorzelak, P. and Salamon, M.A., 2013: Experimental tumbling of echinoderms-taphonomic patterns and implications. Palaeogeography Palaeoclimatology Palaeoecology, vol. 386, p. 569-574. Gorzelak, P., Salamon, M.A. and Baumiller, T., 2012: Predator induced macroevolutionary trends in Mesozoic crinoids. Proceedings of the National Academy of Sciences of the United States of America, vol. 109, p. 7004-7007. Gorzelak, P., Rakowicz, Ł., Salamon, M.A. and Szrek, P., 2010: Inferred placoderm bite marks on Devonian crinoids from Poland. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, vol. 259, p. 105-112. Halicki, B., 1939: Materiały do znajomości budowy podłoża Polski północno-wschodniej. Rocznik Polskiego Towarzystwa Geologicznego, vol. 15, p. 86-128. Jagt, J.W.M. and Salamon, M.A., 2007: Late Cretaceous bourgueticrinid crinoids from southern Poland – preliminary observations. Scripta Geologica, vol. 134, p. 61-76. Ac Kin, A., 2009: Early Maastrichtian ammonites and nautiloids frorm Hrebenne, southeast pt ce Poland, and phenotypic plasticity of Acanthoscaphites tridens (Kner, 1848). Cretaceous Research, vol. 31, p. 27-60. ed Lach, R., (in press): Late Cretaceous sea lilies (Crinoids; Crinoidea) from the Miechów m Trough, Southern Poland. Palaeontographica Abt. A. c us an Lach, R., Brom, K. and Leśko, K., 2015: Bite marks and overgrowths on crinoids from the Štramberk-type limestones in Poland. Neues Jahrbuch für Geologie und Paläontologie, t rip Abhandlungen, vol. 276, p. 151-154. Machalski, M., 2010: Early Maastrichtian ammonites and nautiloids from Hrebenne, southeast Poland, and phenotypic plasticity of Acanthoscaphites tridens (Kner, 1848): A commentary. Cretaceous Research, vol. 31, p. 593-595. Mączyńska, S., 1984: Gromada Echinoidea, Leske 1778. In, Malinowska, L. ed., Budowa geologiczna Polski. Atlas skamieniałości. Tom 2c, p. 435-461. Wydawnictwa Geologiczne. Warszawa. Nebelsick, J.H., 1999: Taphonomic comparison between Recent and fossil sand dollars. Palaeogeography, Palaeoclimatology, Palaeoecology, vol. 149, p. 349-358. Neumann, C., 2000: Evidence of predation on Cretaceous sea stars from northwest Germany. Lethaia, vol. 33, p. 65-70. de Ridder and Lawrence, 1982: Food and feeding mechanisms: Echinoidea. In, Jangoux M. and Lawrence J.M. eds., Echinoderm nutrition, p. 57-116. A.A. Balkema Publishers. Rotterdam. Ac Salamon, M.A. and Zatoń, M., 2007: A diverse crinoid fauna from the Middle Jurassic (Upper pt ce Bajocian-Callovian) of the Polish Jura Chain and Holy Cross Mountains (south-central Poland). Swiss Journal of Geosciences, vol. 100, p. 153-164. ed Salamon, M.A. and Gorzelak, P., 2010: Late Cretaceous crinoids (Crinoidea) from Eastern c us an m Poland. Palaeontographica Abteilung A, vol. 291, p. 1-43. Salamon, M.A., Lach, R., Wieczorek, A., Ferré, B., Brachaniec, T., Trzęsiok, D. and Brom, K.R., (2016): A bourgueticrinid crinoid concentration Lagerstätte in the Lower Maastrichtian (Late Cretaceous) of the Miechów Trough (southern Poland). Neues Jahrbuch für t rip Paläontologie und Geologie Abhandlungen (in press). Villier, L., 2008: Sea star ossicles from the Callovian black clays of the Łuków area, eastern Poland. Neues Jahrbuch für Paläontologie und Geologie Abhandlungen, vol. 247, p. 147-160. Zatoń, M., Villier, L. and Salamon, M.A., 2007: Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy. Lethaia, vol. 40, p. 139-151. Captions Figure 1. A, Map of Poland (slightly modified from Salamon and Zatoń, 2007) and enlargement of investigated area (the Miechów Trough). The map of Miechów Trough (middle map) is slightly modified from Bieńkowska-Wasiluk et al. (2015). B, Investigated section and its enlargement. Line on the lower figure shows the upper surface of the investigated layer. Ac Figure 2. Crinoids showing predation-prey interactions collected in southern Poland. Scale pt ce bars equal 1 mm. A-B, lateral view of Isocrinus columnals. Miechów Trough. Maastrichtian. GIUS 9-3651/Je/1-2. C-F, lateral view of Bourgueticrinus columnals. Miechów Trough. ed Maastrichtian. GIUS 9-3651/Je/3-6. G, lateral view of isocrinid pluricolumnal. Kruhel Wielki, Tithonian. GIUS 8-3651/KrWlk/2. H, lateral view of isocrinid pluricolumnal. Wojkowice, m Middle Triassic (Muschelkalk). GIUS 7-3651/Wjk/12. I, lateral view of isocrinid c us an pluricolumnal. Andrychów, Tithonian. GIUS 8-3651/And/22. J, apical disk of Goniopygus. Miechów Trough. Maastrichtian. GIUS 9-3651/Je/12. K-M, Goniopygus test plates. t predation. rip Miechów Trough. Maastrichtian. GIUS 9-3651/Je/8-11. Arrows indicate signs of echinoid ed pt ce Ac t ip cr us an m ed pt ce Ac cr us an m t ip