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A Bollettino della Società Paleontologica Italiana, 56 (2), 2017, 171-179. Modena

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A (, Polyplacophora) at the end of its trail: a unique find from the

Radek Mikuláš, Andrew K. Rindsberg, Ana Santos & Martin Pavela

R. Mikuláš, Institute of Geology, Czech Academy of Sciences, Rozvojová 269, CZ-16502 Praha 6, Czech Republic; [email protected] A.K. Rindsberg, Biological & Environmental Sciences, Station 7, University of West Alabama, Livingston, AL-35470, USA; [email protected] A. Santos, Departamento Estratigrafía y Paleontología, Facultad de Ciencias, Campus de Fuentenueva s/n, Universidad de Granada, ES-18002 Granada, Spain; [email protected] M. Pavela, Czech Paleontological Society, Na pastvisku 10, CZ-74705 , Czech Republic; [email protected]

KEY WORDS - Viséan, Recent, locomotion traces, tracemakers, Proleptochiton.

ABSTRACT - An exceptional find of a chiton preserved at the end of its locomotion trace comes from the Culm facies, that is, from a succession of turbidity-controlled dark shales, greywackes and sandstones. The chiton trace is a smooth, rather indistinct, 7-8 mm wide bilobate ridge (convex epirelief), forming an incompletely preserved loop of estimated extent 50 x 80 mm. At its end is preserved a completely articulated chiton Proleptochiton sp., 4.0 mm wide and 11.5 mm long, oriented congruently with the trace. The neighbouring strata provide relatively common ichnofossils, including Chondrites isp., Planolites isp., Dictyodora liebeana (Geinitz) and Diplocraterion isp. The chiton’s trace corresponds to ichnotaxa that have historically been compared to modern gastropod trails. The gastropods are well-known for two different monotaxic locomotion techniques, one for hard substrates such as glass, and another for soft substrates, where the move through muscular waves of much higher amplitude than observed on glass. Thereby, adhesion useable for movement on the hard surface is functionally replaced with friction. Loosening of the sediment by rapid movements of foot muscles is the cause of the structure’s convexity, that is, increasing volume. Similar behaviour is documented for the first time in . The studied specimen is the first locomotion attributed to polyplacophorans. The find documents the burrowing technique of chitons in deep-marine, turbidity-influenced soft substrate during the Viséan (330 Ma). It demonstrates the similarity of chiton and some gastropod traces in soft substrates, and adds to the lengthening list of animals that were fossilized within their traces.

RIASSUNTO - [Un chitone (Mollusca, Polyplacophora) alla fine della sua pista: una scoperta unica nel Carbonifero della Repubblica Ceca] - Questo studio descrive l’eccezionale scoperta di un chitone alla fine della sua traccia di locomozione. L’esemplare è conservato in facies Culm, rappresentata da una successione di scisti scuri, grovacche e arenarie torbiditiche. La traccia di chitone consiste in una cresta bilobata liscia, preservata come epirilievo convesso; è piuttosto indistinta, ampia 7-8 mm e forma un circuito, conservato parzialmente, esteso per circa 50 x 80 mm. Alla fine di questa traccia è conservato il chitoneProleptochiton sp. L’esemplare di chitone è completamente articolato, orientato in maniera congruente con la traccia; misura 4,0 mm di larghezza e 11,5 mm di lunghezza. Gli strati adiacenti forniscono icnofossili relativamente comuni, tra cui Chondrites isp., Planolites isp., Dictyodora liebeana (Geinitz) e Diplocraterion isp. La traccia fossile di chitone corrisponde ad icnotaxa che sono tipicamente comparati alle piste di gasteropodi moderni. I gasteropodi sono noti per due diverse tecniche di locomozione monotassica, una per substrati duri come il vetro e l’altra per substrati soffici, dove gli animali si muovono attraverso onde muscolari di ampiezza molto più elevata di quanto osservato sul vetro. In questo modo l’attrito sostituisce funzionalmente l’adesione utilizzabile per il movimento sulle superfici dure. L’allentamento del sedimento da parte di rapidi movimenti dei muscoli del piede è la causa della convessità della struttura, comportando un aumento di volume. Il presente studio è il primo a documentare questo comportamento nei chitoni. L’esemplare studiato costituisce la prima traccia fossile di locomozione attribuita a poliplacofori. La scoperta qui descritta documenta la tecnica di scavo utilizzata dai chitoni durante il Viseano (330 Ma) in ambienti marini profondi, torbiditici ed a substrato soffice. Questo studio dimostra la somiglianza tra tracce di chitoni e di gasteropodi prodotte in substrati soffici, fornendo un nuovo contributo alla crescente lista di animali fossilizzati assieme alle loro tracce.

INTRODUCTION for diminutive infauna living in deep-sea sediments, even such experiments are problematic. As a result, tracemakers Ichnology is the study of trace and biogenic of even such common traces as Paleodictyon and Zoophycos structures in solid substrates, and is often confronted with its (and partly also Chondrites) remain unrecognized. Catching inability to answer the question: “What made it?” The list of the tracemaker “in the act” has not succeeded yet despite trace fossils that are typically preserved altogether with their attempts to remove voluminous samples of unlithified tracemakers is limited (including, for example, dwelling substrate from deep-sea bottoms (Gaillard, 1991). chambers of bivalves drilling in limestone rockgrounds; The aim of the present contribution is to describe an Wilson, 2007). Actually, the problem is much worse; even exceptional find of a fossil chiton preserved at the end of its direct observations of organisms in their natural environment locomotion trace. The value of this find is increased by the fact cannot always enable researchers to assign a trace to its that it comes from a broadly extended facies with well-known maker. The reason is chiefly the fact that most of the traces trace fossil assemblages (Stepanek & Geyer, 1989), that is, that are preserved on bedding planes originate within the the Culm facies of the Lower Carboniferous (Mississippian) substrate (most often the seafloor), whereby the process in central Europe. Moreover, the finding is remarkable since forming the traces remains out of view. This handicap can be it broadens the ichnological record of chitons that nowadays in many cases overcome by aquarium experiments; however, are just known from structures.

ISSN 0375-7633 doi:10.4435/BSPI.2017.15 172 Bollettino della Società Paleontologica Italiana, 56 (2), 2017

Fig. 1 - Paleogeography of the world during the sedimentation of the Moravo-Silesian Culm Basin. Adapted from Isaacson et al. (2008, fi g. 12).

GEOGRAPHY, PALEOGEOGRAPHY AND Massif (Chlupáč & Vrána, 1994) belong to the Moravo- GEOLOGIC SETTINGS Silesian Zone which was connected to the Brunovistulian Terrane, located - contrary to other Bohemian terranes As stated above, locality which yielded the described - probably at the passive continental margin of the sample is in the Culm (Kulm in German and Czech) facies, paleocontinent Laurussia (Fig. 2). The Moravo-Silesian composed of dark shales, greywackes and sandstones Culm is divided into numerous blocks, sub-basins, related historically and regionally to the Variscan plate formations and members. convergence (Bábek et al., 2015; Fig. 1). Most of the The fi nding place of the chiton and its trace is the Culm deposits of the chiefl y Gondwana-related Bohemian Lhotka slate mine at - Region, Czech

Fig. 2 - Paleogeography of the central Europe during the Viséan. Adapted from Hartley & Otava (2001, fi g. 1). The Moravia-Silesia Region is marked by the rectangle. R. Mikuláš et alii - Carboniferous chiton trail, Czech Republic 173

have been reworked to produce minor artefacts from the slate. At the studied locality, the following trace fossils were identifi ed: Chondrites cf. intricatus (Brongniart, 1822), Planolites isp., Dictyodora liebeana (Geinitz, 1867) and Diplocraterion parallelum Torell, 1870 (see Pavela, 2013). Localities in the close neighbourhood of the mine also provided Gordia isp., Phycosiphon isp. and Nereites missouriensis (Weller, 1899) (Lehotský, 2016). The fossil animals found and determined represent (other than the discussed chiton) trilobites Liobole sp. and Phillibole sp.; crinoids Lophocrinus sp., Poteriocrinus sp. and Cyclocaudiculus edwardii Prokop & Pek, 1998; Arnsbergites sp., Girtyoceras sp., Goniatites sp., Hibernicoceras sp., Metadimorphoceras sp., Nomismoceras sp., Paraglyphioceras spp., Sulcogirtyoceras sp., Brachycycloceras sp., Dolorthoceras sp., Liroceras sp., Rayonoceras sp., Rineceras sp., Stroboceras sp. and Culmoteuthis sp.; Orbiculoidea sp. and Propriopugnus sp.; bivalves Citothyris spp., Janacekia sp., Polidevcia sp., Posidonia (ca. ten ), Selenimyalina sp. and Streblochondria sp.; and Listracanthus sp., Cladodus sp., and Ctenacanthiformes gen. et sp. ind. Fossil plants are represented by Lepidodendron sp., Sublepidophloios sp., Ulodendron sp., Archaeocalamites sp., Asterophyllites sp., Aneimites sp., Dyotheca sp., Zeilleria sp., Adiantites sp., Anisopteris sp., Aphlebia spp., Diplotmema spp., Hexagonocarpus sp., Holcospermum sp., Lyginodendron sp., Lyginopteris sp., Rhodeopteridium spp., Spathulopteris sp., Sphenopteridium spp., Sphenopteris spp., Trigonocarpus sp. and Imparipteris spp. (Pavela, 2013). The mixture of terrestrial plants, marine nekton and diverse demonstrates that sedimentation was rapid near the continent margin. Besides this, the rapid changes of environment are also indicated by trace fossils found on the basin. Three ichnocoenoses have been recognized in the Moravice Formation (Bábek et al., 2001), each refl ecting a distinct environment: 1) diverse Dictyodora-Planolites; 2) simple Dictyodora-Planolites; and 3) Diplocraterion- Nereites. Among them, ichnocoenosis 1 best fi ts the studied locality. The diverse Dictyodora-Planolites ichnocoenosis consists mostly of fodinichnia accompanied by agrichnia, pascichnia and traces showing complex feeding strategies. The most common ichnogenera are Chondrites, Dictyodora, Phycosiphon, Zoophycos and Planolites. In the classical concept of Seilacher’s (1967), this ichnocoenosis can be Fig. 3 - Stratigraphic position of the Moravice Formation. From considered as a transitional Zoophycos-Nereites ichnofacies Bábek et al. (2004, fi g. 3). indicating typically bathyal, aphotic, low-energy, oxygen- depleted environments, which are relatively unfavourable for benthic organisms to live in and evolve (Frey & Republic (GPS: 49.809194N, 17.736184E - main surface Pemberton, 1984). Such conditions appeared on the top building). The stratum is within the Moravice Formation of megacycles corresponding to tectonic quiescence in the (upper Viséan), which typically consists of fi ne-grained hinterland (Fig. 4). sandstones, siltstones and mudstones with only a minor proportion of thicker sandstone and conglomerate bodies (Bábek et al., 2015; Fig. 3). The whole formation is ca. DESCRIPTIONS AND INTERPRETATIONS 1,800-2,500 m thick. The Lhotka locality, a slate mine OF THE FIND with a large dump of waste material, provides mostly mudstones and siltstones, in places also with greywacke The body fossil that may have carbonate cement. Despite the long history of the mine, most fi nds of body fossils and ichnofossils PolYPlaCoPHoRa de Blainville, 1816 have been made during the last ten years, as waste dumps Family Dall, 1889 174 Bollettino della Società Paleontologica Italiana, 56 (2), 2017

Fig. 4 - Tectonic and sedimentary model of the study area. Adapted from Bábek et al. (2004, fig. 9B).

Genus Proleptochiton Sirenko & Starobogatov, 1977 Material - “Aulichnites” is an infrequent trace fossil Proleptochiton sp. in the Lhotka slate mine and neighbouring localities; (Fig. 5) however, the low relief and weak colour contrast of the trace make it easy to overlook or dismiss. The rock sample Material - The sole specimen found at the end of the stored in the collection of the Czech Paleontological trace fossil, described below. Stored in the collection of Society, Inv. No Bo172 provides a couple of cross-cutting the Czech Paleontological Society, Opava; Inventory trails connected to the above-described Proleptochiton Number Bo172; maximum length of the rock sample is sp. near the sample margin; on the opposite part of the 0.32 m. sample, further three very poorly preserved sections of “Aulichnites” isp. can be recognized. Description - The specimen preserves a mould of the dorsal side of the valves. No shell material is preserved. Description - Low, smooth, rather indistinct bilobate Head and tail valves are articulated together with six ridge (convex epirelief) on a bedding plane composed intermediate valves. The specimen is 11.5 mm long and of slate. The sample preserves two separated parts. 4.0 mm wide. The valves are carinate and the median ridge Through cross-cutting relations, it is evident that the part is pronounced; the side slopes of the intermediate valves deposited on Fig. 5a centre left originated later than the are slightly concave. The body is elongate (more than part oriented vertically in the middle of the photograph. five times long as wide) and the head plate is distinctively The right end of the younger part of the trace continues large. The margin of the head is nearly circular. The by the above-described body fossil of Proleptochiton sp. intermediate valves are narrow and ca. 1 mm in length and These are oriented congruently; the axes of the trace and 4 mm in width. The tail valve shows a parabolic margin body fossils are nearly parallel, shifted about 1 mm one and is concave; it is 2.5 mm long and 4.1 mm wide. There from another. is no evidence of dorsal sculpture. Both parts of the trace are moderately curved; if they formed a circle, its diameter would be 8-10 cm. Remarks - The specimen is placed in the The configuration of both parts indicates that the trace Proleptochiton according to the diagnosis by Sirenko & originally formed a loop; thus, both parts represent the Starobogatov (1977). same specimen. Analogous configurations are known, such as for Gordia (e.g., McCann & Pickerill, 1988) and The trace fossil Plagiogmus (Seilacher, 2007); overall, the loop was most probably about 4-5 cm wide and 8 cm long. The trace Aulichnites Fenton & Fenton, 1937 intersects itself at an angle close to 90°. Width of the ridges “Aulichnites” isp. is nearly constant at 7-8 mm, which is about 150% of the (Fig. 5a, c) width of the preserved chiton . R. Mikuláš et alii - Carboniferous chiton trail, Czech Republic 175

Fig. 5 - “Aulichnites” isp., a low convex epirelief terminated by an articulated chiton specimen Proleptochiton sp. Lhotka slate mine, Czech Republic; GPS: 49.809194N, 17.736184E. a, c) Planar views of the slab at different lighting. b) Proleptochiton sp. (the same specimen as 5a and 5c). Scale bar corresponds to 1 cm.

Lithology - The sample of the slate is 2×7×32 cm above-mentioned weakly preserved trails that resemble in size. Petrographically, the sample is a dark grey the trace fossil Aulichnites are preserved on the opposite micaceous silty shale. The Ichnofabric Index (Droser & side of the sample, 7-18 cm from the chiton. Bottjer, 1986) is 2. Besides the described specimen(s) of “Aulichnites”, a complete, regularly radial horizontal Remarks - The specimen is placed in open nomenclature, cross-section of Chondrites intricatus is also present; the as there is at present no well-established ichnogeneric diameter of whose system is 30 mm. Moreover, three name for a smooth, bilobated, tape-like, convex epirelief 176 Bollettino della Società Paleontologica Italiana, 56 (2), 2017 trace. The ichnogenus Aulichnites in the sense of the activity. Some individuals seek for protection inside large diagnosis and illustrations by Fillion & Pickerill (1990) shells of bivalves such as Glycymeris. was, especially in the 1980s and 1990s, used commonly for bilobate convex epirelief trace fossils (see the quotations below). However, it has been shown that INTERPRETATION the holotype of Aulichnites is in fact a poorly preserved specimen of Psammichnites (D’Alessandro & Bromley, Modern chitons (Polyplacophora) are a minor, but 1987; Mangano et al., 2002; Seilacher et al., 2005). relatively well-studied group of molluscs (e.g., Slieker, 2000; Sigwart & Lindberg, 2015). This satisfactory degree of knowledge is partly enabled by the frequent occurrence NEOICHNOLOGY of chitons on tidal and shallow subtidal zones on rocky coasts, their most easily observable niches (cf. Slieker, Modern chitons have been repeatedly observed, 2000). Based on limestone rocky coast, feeding bioerosion alongside with their locomotion traces, on the Rompido traces attributable to chitons (classified asRadulichnus by beach near Huelva (southern Spain). The specimens have Voigt, 1977) have been reported; recent observations are been identified as belonging to angulata ubiquitous but finds of corresponding fossils are infrequent (Spengler, 1797). They are common free-living in this (Bromley, 2004 and references therein). area in shelly coarse sand and have been observed during Chitons also excavate ovate resting traces over a period low tide. Their locomotion trace consists of a gently of years, which may be utilized by successive generations curved, linear burrow forming a couple of rather thin of chitons (MacGinitie & MacGinitie, 1949, p. 386, fig. ridges with a central flat zone between Fig.( 6b). Spacing 241). Mikuláš (1992) described and interpreted smooth, between the lateral ridges is about 3 mm. Traces may be concave areas on limestone rockgrounds at terminated with a broad and high horseshoe ridge, i.e. a Štramberk (east Czech Republic) as feeding/resting traces resting trace (Fig. 6a). Traces are preserved as convex of echinoids and/or chitons, naming them Planavolites to concavoconvex epireliefs. Usual length of the trace homolensis Mikuláš, 1992. ranges from 10 to 20 cm. Material in burrows is similar Consequently, the mechanics of locomotion in chitons to surrounding sediment. On the flanks of the traces, there has been studied only in species living on hard substrates are conspicuous ridges that derive from the locomotion (e.g., Vlès, 1907; Parker, 1914; Trueman, 1983). Parker (1914) reported that, in Bermuda, Linnaeus, 1758 moves by generating monotaxic pedal waves from front to back. An whose foot is 70 mm long can move up to 10 mm forward per wave and a new wave may begin at the anterior end before the last wave has completed at the posterior. The animals can also reverse direction. Lateral shifts can be accomplished by pivoting the two ends of the animal around a fixed medial part of the foot. Chitons that were stimulated by a noxious substance on one side moved away from it by an abrupt lateral wave. Sampson (1895) reported that the animals moved slowly and could turn only through a gentle arc. Vlès (1907), Parker (1914), Trueman (1983) and Lai et al. (2010; herein on Fig. 7) noted similarly simple behaviour in some gastropods. The chiton foot is anatomically quite similar to the gastropod foot. Thereby, it can be presumed that chiton traces in soft substrates are very similar to gastropod traces.

Fig. 6 - Chaetopleura angulata (Spengler, 1797) at the end of the locomotion traces at Rompido beach near Huelva (S Spain). a) Individual within its resting trace. b) Continuously burrowing specimen. Photos by Eduardo Mayoral. Scale bar corresponds to Fig. 7 - Principle of movement of a gastropod on a hard substrate. 5 cm. Adapted from Lai et al. (2010, fig. 2). V = velocity. R. Mikuláš et alii - Carboniferous chiton trail, Czech Republic 177

extremely rare or unknown for certain groups including polyplacophorans. Nevertheless, numerous trace fossils including Plagiogmus, “Aulichnites”, and Didymaulichnus have been referred to “possible gastropod traces” (Häntzschel, 1975). Our find, though showing a different mollusc class than gastropods as a tracemaker, generally supports the idea of “gastropod” traces of the above- mentioned form. Recent chitons are known mostly from shallow marine environments, but they occur also in deep sea. Approximately 1,000 extant species have been described so far (Brusca & Brusca, 1990). Polyplacophorans have been found in rocks as old as late (Smith, 1960) although their fossil record is generally poor, which may be due to the low preservation potential of chiton valves Fig. 8 - A trail made by the gastropod (Say, 1822) (Puchalski & Johnson, 2009) and because chiton fossils on a sandy beach at Sapelo Island, Georgia, USA. Photograph by Anthony Martin. are rare and often fragmentary in sedimentary deposits; thereby, they are frequently overlooked (Puchalski et al., 2008). Therefore burrows produced by chitons may not Gastropod locomotion traces are well-known in recent be as rare as discernible chiton body fossils. The problem substrate. They have numerous morphologic analogues in for their future use as a paleoenvironmental tool is based the fossil record (Psammichnites, Aulichnites; Häntzschel, on the close similarity with gastropod crawling traces. 1975; Seilacher, 2007). Bilobate convex epireliefs are one Nevertheless, the described find proves straightforward of forms of gastropod locomotion traces, either smooth evidence that chitons belong to those groups (Fenton & Fenton, 1937) or irregularly transversely creating the ichnogenus “Aulichnites” (i.e., Aulichnites segmented (Fig. 8). in the sense of the following authors). This ichnogenus The above-described locomotion techniques on has been reported, for example, from the marine Upper hard substrates cannot be used on or within loose Pennsylvanian strata in Kansas (Hakes, 1977), from the substrates. Seilacher (2007; herein Fig. 9) showed through presumably shallow marine fine-grained sandstones of experiments with the modern gastropod Polinices that on the Upper Cretaceous of the Western Interior Seaway soft substrates the animals move through muscular waves (Frey & Howard, 1982), from the marine to nonmarine of much higher amplitude than those observed on a glass Fountain Formation, Colorado (Maples & Suttner, 1990), plate. Thereby, adhesion useable for movement on the hard from the upper Cambrian(?) to Lower of surface is functionally replaced by friction. Loosening of eastern Newfoundland (Fillion & Pickerill, 1990) and the sediment by rapid motion of foot muscles is the cause from clastic shelf settings of the Upper Cretaceous of of the structure’s convexity, thus increasing volume. The Utah (Frey & Howard, 1990). The future re-examination trace fossil is broader than the shell of the tracemaker, of traces attributable to Aulichnites should take chitons which corresponds to observations of extant organisms. into consideration as probable tracemakers. Such a configuration is consistent with the enlargement of the foot during the locomotion, along with the presumed effect of this burrowing technique on the substrate (that CONCLUSIONS is, loosening and volume increase). Although fugichnia terminated with a bivalve or 1. For the first time, a chiton was found together gastropod are not rare and bioerosion structures with with its locomotion trace, this documents the burrowing bivalves are common (Mikuláš et al., 2003), locomotion technique of chitons in deep-marine, turbidity-influenced traces found along with the molluscan tracemakers are soft substrates during the Viséan (330 Ma). Moreover, although it is well known that chitons can be responsible for bioerosion, this discovery suggests that these animals also produce bioturbation structures in unconsolidated substrates. 2. The find demonstrates the similarity of chiton and some gastropod traces in soft substrates; in the same way, it proves that Paleozoic molluscs with a broad “gastropod” foot produced bilobate ridges (Fig. 8).

ACKNOWLEDGEMENTS

This work is a contribution to the call for papers arising from the Fourth International Congress on Ichnology “Ichnia 2016 - Ichnology for the 21st century: (palaeo)biological traces towards sustainable development”, held in Idanha-a-Nova (Portugal), 6-9 May 2016. Fig. 9 - Movement of a modern gastropod Polinices sp. on soft The work of RM was supported by the RVO67985831 Project of substrates (adapted from Seilacher, 2007, pl. 27). the Institute of Geology of the Czech Academy of Sciences. AS was 178 Bollettino della Società Paleontologica Italiana, 56 (2), 2017 supported by the Research Group RNM-276 (Junta de Andalucía). Frey R.W. & Pemberton S.G. (1984). Trace fossil facies models. In The authors gratefully acknowledge Eduardo Mayoral (University Walker R.G. (ed.), Facies Models, 2nd ed. Geoscience Canada of Huelva) for providing valuable photographs of recent chiton Reprint Series: 189-207. locomotion traces and Anthony J. Martin (Emory University) for Gaillard C. (1991). 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