Echinoid bite traces on Late Cretaceous (lower Maastrichtian) sea

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Humblet, M. and Iryu, Y. 2014: Pleistocene coral assemblages on Irabu-jima,
South Ryukyu Islands, Japan. Paleontological Research,
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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
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Department of Geochemistry, Mineralogy and Petrography, Faculty of Earth Sciences,
University of Silesia, Będzińska Street 60, 41-200 Sosnowiec, Poland; e-mail:
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[email protected]. Centre for Polar Studies KNOW (Leading National Research
Centre) WNoZ UŚ
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Department of Palaeontology and Stratigraphy, Faculty of Earth Sciences, University of
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2
Silesia, Będzińska Street 60, 41-200 Sosnowiec, Poland; e-mail: [email protected]
3
Department of Palaeontology and Stratigraphy, Faculty of Earth Sciences, University of
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Silesia, Będzińska Street 60, 41-200 Sosnowiec, Poland; [email protected],
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[email protected]. 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
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Key words: Echinoidea, Crinoidea, Maastrichtian, Cretaceous, predation, Poland
Along one of the longest Cretaceous epicontinental sedimentary sections in Central Europe
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(almost 30 km), exposed during the construction of European route E77 between Cracow and
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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
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accumulation, described as a concentration Lagerstätte, was also discovered (Salamon et al.,
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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.
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Material and methods
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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
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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
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(Albian-lower Maastrichtian), with a total thickness of which ranged from 800 to 1000 m (for
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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
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et al., 2016).
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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
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Palaeontology and Stratigraphy Department, University of Silesia. The first one was soaked in
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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
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column (mesh size: 1.0, 0.5 and 0.315 mm). The invertebrate faunal elements were picked
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under a stereoscopic microscope (SM800T). On the whole, over 40 faunal remains were
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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.
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The second sample was macerated following Boczarowski’s (2001) method. All rocks
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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
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Salamon et al. (2016, fig. 2a) and ascribed to fish predation traces. On the other hand, such
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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
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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
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echinoids (Baumiller et al., 2008, 2010); Fig. 2A-F. Similar bite traces on fossil crinoids have
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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
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up to 1.1 mm. All observed marks are sharp and occur on the external part of the lateral
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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
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Praedichnia are one of the ichnofossil types (trace fossil) that directly evidence the
interactions between a predator and its prey (Ekdale, 1985). However, distinguishing
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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
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challenge for the palaeontologists’ community. Furthermore, identification of what kind of
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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
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therein). However, such differentiations are crucial to obtain sound and reliable conclusions
concerning palaeoecology. Additionally, studies on the predatory phenomenon might provide
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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
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tapering) holes towards the interior (bioerosion), stylolites or “fitted” fabrics (pressure
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dissolution) and rhombohedral depressions along the calcite-cleveage (post-diagenetic
fracturing); details in Salamon and Gorzelak (2010).
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The pits conform in size and shape to biometric parameters of traces left by regular
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echinoids (Results section). Indeed, recent, regular echinoids exhibit wide spectrum of food
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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
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showed that cidaroid sea urchins are feeding actively on stalked crinoids (direct in situ
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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,
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Acknowledgements
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pers.comm.).
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because of their pre-Maastrichtian extinction, at least locally (Dr. R. Niedźwiedzki,
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The comments of two reviewers, Dr. Andreas Kroh (Natural History Musuem, Vienna,
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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
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(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
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Centre (KNOW) to the Centre for Polar Studies for the period 2014-2018.
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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.
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Figure 2. Crinoids showing predation-prey interactions collected in southern Poland. Scale
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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.
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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,
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Middle Triassic (Muschelkalk). GIUS 7-3651/Wjk/12. I, lateral view of isocrinid
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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.
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predation.
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Miechów Trough. Maastrichtian. GIUS 9-3651/Je/8-11. Arrows indicate signs of echinoid
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