The ichnogenus Rhizocorallium: Classification, trace makers, palaeoenvironments and evolution

    The ichnogenus Rhizocorallium: Classification, trace makers, palaeoenvironments and evolution Dirk Knaust PII: DOI: Reference:

S0012-8252(13)00081-0 doi: 10.1016/j.earscirev.2013.04.007 EARTH 1851

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Earth Science Reviews

Received date: Accepted date:

22 January 2013 15 April 2013

Please cite this article as: Knaust, Dirk, The ichnogenus Rhizocorallium: Classification, trace makers, palaeoenvironments and evolution, Earth Science Reviews (2013), doi: 10.1016/j.earscirev.2013.04.007

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ACCEPTED MANUSCRIPT The ichnogenus Rhizocorallium: classification, trace makers, palaeoenvironments and evolution

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Dirk Knaust, Statoil ASA, 4035 Stavanger (e-mail: [email protected])

Abstract

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Rhizocorallium is one of the oldest known trace fossils, with wide distribution through the Phanerozoic and all over the world. Originally introduced from the epicontinental Triassic of central

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Germany, its high morphological lability gaves reason for the subsequent erection of about twenty ichnospecies. The study of newly collected material from the type area and many specimens from

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various collections permits the conclusion that R. jenense and R. commune are the only valid ichnospecies of Rhizocorallium. The type ichnospecies, R. jenense, is a comparatively small, inclined and heavily scratched firmground burrow with passive fill, while R. commune consists of extensive,

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more or less horizontal burrows with occasionally scratched marginal tubes and an actively filled spreite between the tubes. The faecal pellets Coprulus oblongus are typically associated with R.

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commune. Morphological variations of R. commune are captured in ichnosubspecies and varieties of this ichnospecies and can aid a refined reconstruction of palaeoenvironments. A review of more than 180 records from the literature reveals the common confusion of both ichnospecies, which has

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consequences for the application of Rhizocorallium in facies interpretations. The end members of both ichnospecies may be linked by transitional forms, which suggests the same kind of trace maker. Polychaetes are the most likely producers of marine Rhizocorallium, based on their long-ranging occurrence, morphological features, appearance of faecal pellets, associated soft-body remains, and modern analogues. R. commune occurs from Early Cambrian to Holocene, while R. jenense just appears after the end-Permian mass extinction, probably as a consequence of an adapted firmground burrowing lifestyle of its producer. Fluvial R. jenense are probably produced by mayfly larvae in homology to marine polychaete burrows. A consequent application of the newly established classification scheme allows for a more rigorous application of Rhizocorallium in the reconstruction of palaeoenvironments. Thus, Palaeozoic and Mesozoic R. commune are restricted to the Cruziana ichnofacies of shallow-marine environments, while in Cenozoic time similar forms are also found in

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ACCEPTED MANUSCRIPT deep-marine deposits. R. jenense, on the other hand, is a constituent of the widespread Glossifungites ichnofacies and, aside from continental environments, occurs in peritidal to deep-marine deposits. Several studies have demonstrated the value of Rhizocorallium for interpreting sequence-

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stratigraphical surfaces, current directions, and fluctuations in salinity and oxygen.

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Key words: Trace fossils, ichnology, Rhizocorallium, ichnotaxonomy, polychaetes,

Introduction

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palaeoenvironment, Germanic Triassic.

The ichnogenus Rhizocorallium is one of the earliest described trace fossils, with cosmopolitan

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distribution from the Early Cambrian to the Holocene. It is among the most cited trace fossils, with an increasing record of citation (according to GeoRef). Rhizocorallium has been often used in the

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characterization of shallow-marine depositional systems. Since the introduction of the type ichnospecies R. jenense by Zenker (1836), additional ichnospecies were attributed later, of which only three were regarded as valid in a review performed by Fürsich (1974a), R. jenense, R. irregulare

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Mayer, 1954 and R. uliarense Firtion, 1958. Fürsich’s proposal has been followed by the majority of ichnologists and sedimentologists, although some workers have continued to introduce new ichnospecies of Rhizocorallium, mainly based on material from India and China. A restudy of material from the type area of Rhizocorallium (the Germanic Basin) let Knaust (2007a) and Knaust et al. (2012) conclude that Fürsich’s classification essentially may be retained but concurrently offers room for improvement because of its oversimplification in part. Especially confusion between R. jenense and similar small spreite burrows referred herein to R. commune var. auriforme (Hall, 1843) has weakened the potential of Rhizocorallium for palaeoenvironmental interpretations. Recently, Schlirf (2011) proposed a new classification concept for U-shaped spreite trace fossils and recommended the abandonment of Rhizocorallium commune Schmid, 1876 in favour of Ilmenichnus devonicus (Hecker, 1930). This procedure is not followed herein for reasons outlined below.

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ACCEPTED MANUSCRIPT Since the work done by Seilacher (1963, 1967), much has been known about the palaeoecological value of trace fossils, although the ichnofacies model is still in debate (e.g. Seilacher, 2007; Buatois and Mángano, 2011; MacEachern et al., 2012a). A proper determination of ichnospecies

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of Rhizocorallium is crucial to the interpretation of both the Cruziana and Glossifungites ichnofacies and augments their value for palaeoenvironmental interpretation and sequence stratigraphy. Trace

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fossils are ‘only’ the fossilized traces left by organisms living in the sediment but nonetheless are covered by the rules of the ICZN (Rindsberg, 2012). One reason for this procedure is their utilization

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for reconstructions of facies and palaeoenvironments in connection with sedimentological studies. On the other hand, a strict classification scheme is needed to exploit the full value of trace fossils. Weak

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diagnoses or ‘lumping’ of different ichnospecies with a reduced number of diagnostic features decreases the chance of interpreting the right producer and consequently leads to a dilution of the ichnofacies model (Knaust, 2012b).

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For this review, thousands of specimens of Rhizocorallium were investigated from many natural and industrial outcrops, primarily in central Germany (Thuringia), such as valleys, hill slopes,

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abundant and active quarries, and also from collections over the last two decades. The substantial material from the type area of Rhizocorallium demonstrates that the ichnospecies, occurring in conjunction with other palaeontological and sedimentological criteria, are suitable for the

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differentiation of subenvironments in shallow-marine carbonate systems with an otherwise homogeneous appearance (Knaust et al., 2012; Knaust and Costamagna, 2012). Following the description and interpretation of diagnostic features of Rhizocorallium, a differential diagnosis from ichnogenera with similar elements as those in Rhizocorallium is briefly outlined. Selected examples from the type area of its type ichnospecies are described and interpreted to underpin the newly proposed classification scheme. Moreover, a careful analysis of significant features sheds new light on the interpretation of potential trace makers. In a comparison with published data, the established facies model from the Germanic Triassic is tested and general facies relationships and environmental constraints of Rhizocorallium are worked out. Finally, the evolution of Rhizocorallium is briefly discussed.

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Early reports of Rhizocorallium

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Rhizocorallium is a conspicuous biogenic sedimentary structure which has been recognized and documented for more than 250 years. One of the oldest quotations is from Schütte (1761), who

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mentioned “Beinstein, Rhizolithi, versteinte Wurzeln oder Wurzelsteine” from Jena and obviously related these ‘bones and fossil root casts’ partly to Protovirgularia, Planolites and Rhizocorallium.

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Batsch (1802) meticulously described “Wurm- und Zungen-kalkstein” [“worm- and tonguelimestone”] from the Muschelkalk, the latter of which can clearly be related to Rhizocorallium

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commune. Schlotheim (1822, pl. 5, fig. 2) illustrated Rhizocorallium jenense from its type area (Fig. 1) well before its formal introduction by Zenker (1836) and interpreted it as fossil algae or coral (Conferve, Coralline). Hall (1843, 1852) described and figured crowded Fucoides auriformis from the

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Early Silurian of North America (Fig. 2), while Jugler (1853) discussed Thier-Fährten (animal traces) from the Lower Cretaceous Bentheim Sandstone in NW-Germany (Fig. 3). Both records can now be

Morphological characteristics (ichnotaxobases)

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attributed to R. commune.

Several criteria will serve for an ichnotaxonomic treatment of the ichnogenus Rhizocorallium and its ichnospecies, although their application to the level of ichnogeneric or ichnospecific discrimination is a matter of consideration and can be debated (Fig. 4). The ichnofamily Rhizocoralliidae Richter, 1926 consists of U-shaped spreiten burrows with a specific set of characteristics (ichnotaxobases; Bromley, 1990; Bertling et al., 2006; Buatois and Mángano, 2011), of which orientation has been used to distinguish between Diplocraterion (vertical) and Rhizocorallium (horizontal to oblique). Furthermore, the main types of substrate (e.g. hardground versus firmground/softground) result in contrasting behaviour of the trace maker (e.g. boring versus burrowing) and, consequently, serve to discriminate morphologically similar ichnogenera such as Caulostrepsis and Rhizocorallium. In

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ACCEPTED MANUSCRIPT contrast, softground and firmground belong to the same catergory of substrate and share a continuous transition between both end members with the same principal manner of penetration (burrowing). Together with other features such as fill, faecal pellets and scratches, these substrate differences can be

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regarded as ranking on the ichnospecies level. Lower in the hierarchy, errant morphologies serve distinguishing ichnosubspecies, while burrow size may be used delineating varieties. Although names

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published after 1960 with the term “variety” are not regulated by the International Code of Zoological Nomenclature, varieties were frequently introduced and applied to delineate different infrasubspecific

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characteristics (e.g. Hester and Pryor, 1972 for Ophiomorpha; Ekdale and Lewis, 1991b for Diplocraterion; Uchman, 1995 for Scolicia; Miller, 2011 for Phymatoderma). The following features

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help to classify ichnospecies of Rhizocorallium (listed in accordance to their importance in a descending order).

Burrow orientation

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Rhizocorallium burrows are orientated parallel or oblique to the bedding plane and thus differ from vertical spreite burrows as included in the ichnogenus Diplocraterion (Fig. 5). Similarly, the average

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and variation of burrow orientation (measured in degrees of angle to the bedding plane) help to subdivide Rhizocorallium on an ichnospecies level. Notably, the classification scheme proposed by

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Schlirf (2011) also includes steeply inclined spreite burrows into Diplocraterion, a procedure that illustrates the close affinity of both ichnogenera. 3.2

Burrow morphology and fill

Rhizocorallium typically consists of a U-shaped marginal tube with more or less parallel limbs (or arms), enclosing an area occupied by a spreite. The spreite refers to the numerous tightly arranged tunnels spreading between the limbs to be arranged subparallel to the distal U-shape of the marginal tube. The overall shape of the spreite tunnels is either semicircular with a symmetrical appearance, or asymmetrically produced with a J-shaped form. In the latter case, the sediment probing originates from one (merging) limb but the spreite is truncated on the opposite (truncating) limb (cf. Basan and Scott, 1979). The process of sediment probing either results in completely reworked sediment with a diffuse

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ACCEPTED MANUSCRIPT spreite pattern (soft sediment; tecticol after Reis, 1910, p. 245) or, with increasing substrate cohesion, with spreiten that are well defined by tunnels or corresponding grooves and ridges (firm sediment; nudicol after Reis, 1910, p. 245) (Fig. 6). Reworking of soft substrate with ploughed sediment

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preserved within the spreite (active spreite) differs from spreite burrows in firm substrates, where the excavated sediment is displaced and the spreite area together with the marginal tube is passively filled

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with sediment (passive spreite) (Fig. 7). In the latter case, only the casts or fills of casts (steinkerns) of the U-shaped spreite tunnels are preserved. Most Rhizocorallium spreite burrows are protrusive but in

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some cases retrosive spreite elements can be observed.

Faecal pellets

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It has been well recognized that faecal pellets are a common constituent of Rhizocorallium but they have received little attention in their usage as an ichnotaxobase. This analysis shows the importance of

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faecal pellets for the classification of ichnospecies of Rhizocorallium. Two ichnospecies of the

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ichnofamily Coprulidae Knaust, 2008 are associated with Rhizocorallium, both having an ellipsoidal shape and a homogeneous composition: Coprulus oblongus Mayer, 1952 (common), ca. 1.0-1.5 mm

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long and 0.5-0.8 mm wide; and C. bacilliformis Mayer, 1955 (rare), ca. 3.0 mm long and 0.5 mm wide (Fig. 8). Faecal pellets are common in R. commune but lacking in R. jenense.

Scratches

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Scratch traces (bioglyphs sensu Ekdale and Gibert, 2010) are important fingerprints of Rhizocorallium and originate from the burrowing activity in firm substrate. Their presence alone is not a reliable criterion for ichnotaxonomic subdivision; however, subtle differences in the morphology of scratches constitute suitable ichnotaxobases and may aid in the discrimination of ichnospecies (Ekdale and Gibert, 2010). For instance, R. jenense is commonly scratched with net-like, criss-crossing and closely spaced bioglyphs, whereas R. commune contains more or less parallel, rarely crossing and sparse scratches (Fig. 9).

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ACCEPTED MANUSCRIPT 3.5

Size

As demonstrated by morphometric analysis (see below), burrow dimensions are suitable for clustering specimens of Rhizocorallium in correspondence to varieties, and thus dimensions such as length,

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width and tube diameter together with other criteria are valuable in their separation. This discrimination is important for the use in palaeoenvironmental reconstructions but yields only limited

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information for an ichnotaxonomical treatment.

Branching

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Branching is commonly regarded as main ichnotaxobase to define ichnogenera (e.g. Bromley, 1996; Bertling et al., 2006; Buatois and Mángano, 2011) but remains controversial as many ichnogenera

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comprise both branched and unbranched forms (e.g. Palaeophycus, Planolites, Rosselia, Schaubcylindrichnus, Teichichnus, Trichichnus and others; see Knaust, 2012b). In the ichnogenus

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Rhizocorallium, R. jenense is unbranched (or shows false branching in crowded occurrences), while

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primary successive branching of the entire spreite burrow may occur in R. commune var. irregulare

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(terminology after D’Alessandro and Bromley, 1987).The latter is facultative and probably a response to food supply or phobotaxic behaviour of the trace maker, while other features remain unchanged. It is therefore unlikely that branching will serve as a robust ichnotaxobase for Rhizocorallium, although

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‘Ilmenichnus multilobatus’ (=R. commune) Schlirf, 2011 is based on it.

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Proposed classification Ichnotaxonomy

Ichnofamily Rhizocoralliidae Richter, 1926

This ichnofamily contains spreite burrows of the ichnogenera Rhizocorallium and Diplocraterion. It differs from the ichnofamily Alectoruridae Schimper and Schenk, 1879 (which mainly includes

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ACCEPTED MANUSCRIPT Zoophycos and Spirophyton; see Fuchs, 1895) by having a proportionately much wider marginal tube and a more constrained burrow outline instead of a widely rounded curve (Seilacher, 2007).

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Type ichnospecies: Rhizocorallium jenense Zenker, 1836

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Ichnogenus Rhizocorallium Zenker, 1836

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Emended diagnosis: Horizontal to oblique, U-shaped spreite burrow.

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Ichnospecies Rhizocorallium jenense Zenker, 1836

Rhizocorallium commune uliarense (Firtion, 1958)

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Rhizocorallium commune problematicum (Gümbel, 1861)

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Rhizocorallium commune var. auriforme (Hall, 1843)

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Rhizocorallium commune var. irregulare (Mayer, 1954b)

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Ichnospecies Rhizocorallium commune Schmid, 1876

Discussion

Rhizocorallium jenense Zenker, 1836 from the marginal-marine Upper Buntsandstein of Jena, Germany, differs in size, orientation, length/width-ratio and scratches from the spreite burrows of the shallow-marine Muschelkalk Group in the same area, which was the reason for the erection of R. commune Schmid, 1876. Although neither ichnospecies was originally figured, their descriptions and listed occurrences are sufficient to regard them as available according to Article 12 of the International Code of Zoological Nomenclature. R. devonicum Hecker, 1930, as originally described from the St. Petersburg region in Russia (Fig. 10), and R. irregulare Mayer, 1954b from the Upper Muschelkalk of southern Germany, are identical with R. commune and are thus junior synonyms.

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ACCEPTED MANUSCRIPT In the Germanic Triassic, early workers (e.g. Schmidt, 1936, p. 20) recognized a morphological continuation between R. commune and R. jenense in dependence of the substrate condition from soft to firm: “A complete transition from one form to the other would not be surprising

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given the current view of the supposed origin of the net-like scratches. The differences between both forms must be developed and preserved in accordance with the physical properties of the substrate in

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which the producers of these tunnels burrowed. The current way in the discrimination of the species obviously has little or nothing to do with the differences between their producers.” (Translated from

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German).

Hecker (1980, p. 20) regarded the differences of the characteristic features (such as

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orientation, length, width and scratches) between R. jenense and R. devonicum as great enough to justify a new taxon at the ichnogenus level and so introduced Ilmenichnus with type ichnospecies I. devonicus Hecker, 1930. Although this procedure recently was followed by Schlirf (2011), in this

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proposal it is suggested to retain the differentiation on the ichnospecies level of Rhizocorallium. R. commune Schmid, 1876, as known from the Muschelkalk Group, includes large and

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winding burrows as well as relatively small and tongue-shaped burrows with otherwise similar features. In order to differentiate these two forms, R. commune var. auriforme and R. commune var. irregulare are reassigned as varieties. R. uliarense Firtion, 1958 was established for trochospiral U-

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shaped spreiten burrows from the Upper Jurassic of SW France but is herein regarded as ichnosubspecies R. commune uliarense. Moreover, ichnosubspecies R. commune problematica (Gümbel, 1861) represents R. commune with a vertically retrusive spreite element. This newly proposed classification of Rhizocorallium is illustrated in Figure 11, while Table 1 lists the main characteristics of both ichnospecies. 4.3

Interrelationship with other trace fossils

Several trace fossils are closely related to Rhizocorallium, either as gradational transition forms or as part of compound and composite trace fossils (for definitions, see Pickerill, 1994). The most common are briefly described in order of importance, with the aim to clarify ichnotaxonomical problems and from the point of view of ethological interpretations.

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4.3.1

Morphological transitions and similar trace fossils

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Diplocraterion Torell, 1870

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Belonging to the same ichnofamily, Diplocraterion has close affinity to Rhizocorallium and some (if

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not all) ichnospecies could even be regarded as synonymous to the latter if orientation were disregarded as ichnotaxobase (Fig. 12). Indeed, orientation of R. jenense varies from subhorizontal to steeply inclined (e.g. Fürsich et al., 1981; Uchman et al., 2000) and thus creates a continuum with

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vertically orientated Diplocraterion (Figs. 4, 5). This fact has led some authors (e.g. Kvale et al., 2001; Rodríguez-Tovar et al., 2007; Rodríguez-Tovar and Pérez-Valera, 2008) to assignsuch specimens to

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both ichnogenera. In other cases, vertically orientated U-shaped spreiten burrows contain faecal pellets similar to R. commune and were actually attributed to Rhizocorallium instead to Diplocraterion (e.g.

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Howard and Singh, 1985). Schlirf (2011) proposed an emended diagnosis for Diplocraterion to

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accommodate vertical and oblique U-shaped spreiteburrows, a procedure that certainly will lead to

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confusion between Diplocraterion and Rhizocorallium and therefore is not followed herein.

Zoophycos Massalongo, 1855

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Zoophycos is another famous spreite burrow, which today includes a variety of morphological forms that would better be separated at the ichnogenus level (Bromley and Hanken, 2003). In general, a much lower tube diameter to burrow width ratio, and lobes with a semicircular outline, are used as criteria to distinguish Zoophycos and Rhizocorallium (Häntzschel, 1960; Fürsich, 1974a; Seilacher, 2007). However, gradation between the two ichnogenera is not uncommon and hampers an unambiguous assignment. This can be well illustrated based on examples from Oligocene deep-marine turbidite deposits of SE-France, where transitions between simple R. commune and lobate Z. insignis occur (Fig. 13). Remarkably, both forms contain abundant faecal pellets (Coprulus oblongus), suggesting the same kind of producer. Lateral shift and progressive deepening of individual Rhizocorallium-like burrows may gradually lead to the development of a Zoophycos-like spreite system as described by Häntzschel (1960), Bradley (1973), Miller and D’Alberto (2001), Bromley and

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ACCEPTED MANUSCRIPT Hanken (2003) and Neto de Carvalho and Rodrigues (2003). In the Muschelkalk, Zoophycos is known as a rare component of a compound burrow system most likely produced by the same animal as R.

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commune (Knaust, 2004, and references therein).

Echinospira Girotti, 1970

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Echinospira is regarded as a member of the Zoophycos group and is characterized by a semicircular outline with numerous long, narrow U-shaped spreite burrows similar to R. commune, which are

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related to a central cylindrical structure. Echinospira is common in Upper Cretaceous to Miocene deep-sea deposits (e.g. Ekdale and Lewis, 1991a; Uchman and Demírcan, 1999; Fig. 14), and similar

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forms were also described from the Carboniferous of the United Kingdom (McIlroy and Falcon-Lang, 2006).

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Lophoctenium Richter, 1850

Superficially, the spreite burrow Lophoctenium resembles R. commune but differs from it by a highly

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variable lobate pattern surrounded by an external tube (Fu, 1991; Seilacher, 2007). The similarity between the two ichnogenera is reflected in the ichnospecies name of Lophoctenium rhizocoralloides

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Hundt, 1931 (pp. 59, 68).

Beaconites Vialov, 1962 Stanistreet et al. (1980) described an about 20 cm wide trace with spreite from an Early Permian bedding plane of South Africa and assigned it to a supposed new ichnospecies of Rhizocorallium. However, Gevers et al. (1971) pointed out, the lack of a marginal tube does not allow an assignment to Rhizocorallium but rather to a large segmented burrow, Beaconites antarcticus Vialov, 1962, which is common in Palaeozoic deltaic rocks of Gondwana (e.g. Buatois and Mángano, 2011).

Phycodes Richter, 1850 Tightly packed, very shallow U-shaped burrows with spreiten and a rope-like appearance are attributed to Phycodes parallelum Seilacher, 2000. Superficially, P. parallelum mimics R. jenense, but

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ACCEPTED MANUSCRIPT differs from it by a preferred subvertical orientation, a much smaller length/width ratio and greater sinuosity. Scratches were not documented from P. parallelum, although this could be an effect of syndepositional erosion. In addition to Cambrian (e.g. Baldwin et al., 2004, fig. 8), Ordovician

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(Seilacher, 2000, 2007) and Carboniferous specimens (Eagar et al., 1985, pl. 14D; Pollard, 1988, fig. 2F), similar forms are also known from fluvial to lagoonal, marginal-marine sandstone beds of the

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Lower Keuper (Lettenkeuper, Ladinian) above the Muschelkalk succession in the Germanic Triassic

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(Fig. 15).

Fuersichnus Bromley and Asgaard, 1979

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R. jenense may grade into loosely clustered, J- and U-shaped burrows commonly referred to Fuersichnus communis Bromley and Asgaard, 1979. This can lead to confusion as both ichnotaxa may occur in the same facies (e.g. red beds). In contrast, F. striatus Buatois, 1995 from the marine

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Cretaceous strongly resembles R. jenense in terms of shape, incipient spreite and striation pattern, and thus is better accommodated within this ichnotaxon as its junior synonym. The crescentic F.

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commerfordi Garvey and Hasiotis, 2008 from the continental Lower Carboniferous represents another ichnospecies. Some specimens of the above-mentioned Phycodes parallelum from the fluvial to

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lagoonal Lettenkeuper actually belong in the Fuersichnus-Rhizocorallium transition field (Fig. 15).

Hoplichnus Hitchcock, 1848 and Furculosus Roniewicz and Pieńkowski, 1977 The ichnogenera Hoplichnus and Furculosus are quite similar in morphology and constitute hoofshaped, semiovate reliefs and cylindrical burrows forming tight, fork-like loops with parallel or divergent terminations. Both ichnogenera superficially resemble Rhizocorallium, although flute casts were also called Hoplichnus (Lockley et al., 1994). However, they lack the diagnostic spreite of Rhizocorallium. Examples include Rhizocorallium praecursor Hundt, 1940 from the Lower Triassic Buntsandstein of Germany and Rhizocorallium sp. as described by Spörli and Grant-Mackie (1976) from the Late Jurassic of New Zealand (see also discussion by Ballance, 1976).

Tisoa de Serres, 1840

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ACCEPTED MANUSCRIPT Superficially, the morphology of Tisoa may resemble that of R. jenense, but differs from it by several criteria. It has a narrow U-shape with tightly arranged, elongate limbs without significant curvature (Frey and Cowles, 1972; Fig. 16). Thus, the length/width ratio of the burrows is extremely high

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(typically in the range of 20-50 or more). Cross-sections appear to have a dumbbell or figure- eight shape. Tisoa has no spreite and its fill is passive, commonly including sulphides. A striking feature is

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its frequent occurrence in the central part of carbonate concretions related to hydrocarbon seep deposits (Breton, 2006). Given the extreme length of the burrows, their U-turn is rarely preserved,

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which may give the impression that vertically orientated specimens are not trace fossils but conduits

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for escaping seep gases (van de Schootbrugge et al., 2010).

Caulostrepsis Clarke, 1908

Caulostrepsis consists of pouch-shaped borings (Bromley and D’Alessandro, 1983), and its similarity

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to some R. jenense (despite their generally smaller size) has long been recognized (Seilacher, 1969).

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Some such borings have been attributed to ‘Glossifungites saxicava’ (= R. jenense, e.g. Zatoń et al.,

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2006, fig. 6). In modern hard substrates (e.g. oyster shells), Caulostrepsis originates from the bioeroding activity of spionid polychaetes (e.g. Polydora sp.; Douvillé, 1908). Spionidae include both boring and non-boring species (Sato-Okoshi 1999, 2000), commonly distributed in mud-dominated

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firmgrounds of marginal-marine environments (Gingras et al., 2001). Various incipient spionid borings (such as grooves) are commonly preserved on omission surfaces in the Muschelkalk succession (Knaust, 2008) and it is tempting to argue that Caulostrepsis and R. jenense result from the same producer, a spionid. Following the recommendation of Bertling et al. (2006), the occurrence of similar trace fossils in contrasting substrate (such as hard- and firmground) implies different behaviour (boring versus burrowing) and requires ichnotaxonomical differentiation.

Asthenopodichnium Thenius, 1979 Similar to Caulostrepsis, U-shaped pouches within ligneous or bone substrates are called Asthenopodichnium and resemble R. jenense (see MacEachern et al., 2012a, fig. 6C). They occur in

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ACCEPTED MANUSCRIPT Neogene freshwater strata and are interpreted as produced by insect larvae such as mayflies (Uchman et al., 2007), although wood-rotting fungi were recently considered by Genise et al. (2012).

Compound and composite trace-fossil elements

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Balanoglossites Mägdefrau, 1932

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In the Germanic Basin, both ichnospecies of Rhizocorallium are intimately associated with traces that may be attributable to Balanoglossites. The holotype of B. triadicus from the Lower Muschelkalk of Jena is a ramified tunnel system intergrading with pouch-shaped elements attributable to R. jenense

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(Knaust, 2008; Fig. 17A). This type of compound trace fossil is common in firm- to hardgrounds of the Lower Muschelkalk, which are amply perforated with shallow to deep, U-shaped pouches and

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irregularly winding grooves and tunnels (Fig. 17B). They occur in different size classes and show morphological transitions to R. jenense. Similar forms were also described from the Jurassic of

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Germany (Hölder and Hollmann, 1969) and from oyster shells (e.g. Reis, 1922), and result from

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boring and scratching spionids (see Sato-Okoshi and Okoshi, 2000). The type material of ‘R. irregulare’ (= R. commune) from the Upper Muschelkalk of southern Germany is also closely

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associated with Balanoglossites (Mayer, 1954b). Moreover, grooves on the surface or winding burrows (e.g. Basan and Scott, 1979) typically intergrade with B. ramosus in the subsurface (e.g.

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Mayer, 1954b; Knaust, 2008; Fig. 17C, D). The occurrence of the faecal pellets Coprulus in both Rhizocorallium and Balanoglossites, along with their intimate linkage, suggests that they result from different activities (e.g. feeding and dwelling) of the same producer, acting under changing substrate conditions. However, commensalism of various organisms is another possibility to explain compound Balanoglossites/Rhizocorallium trace fossils.

Radichnus Allington-Jones, Braddy and Trueman, 2010 Some firmground surfaces of the Muschelkalk Group not only contain simple Rhizocorallium-like burrows but also exhibit associated delicate scratch patterns (Fig. 18A). The same trace maker apparently behaved in different ways as reflected in contrasting trace fossils, e.g. burrowing into the subsurface and scraping on the surface (or a subsequently eroded bedding plane). Allington-Jones et

14

ACCEPTED MANUSCRIPT al. (2010) introduced Radichnus allingtona for similar trace fossils from the Rhaetian, where it occurs with ‘R. irregulare’ (= R. commune var. irregulare) in a restricted or lagoonal palaeoenvironment. Given a firm substrate, lobate burrows with a R. commune outline within the sediment (as described in

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connection with Balanoglossites triadicus; see Knaust, 2008) may remain open and show scratches all

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over their surface (Fig. 18B).

Thalassinoides Ehrenberg, 1944 and Spongeliomorpha de Saporta, 1887

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Ichnospecies of Rhizocorallium, Thalassinoides and Spongeliomorpha share various marine subenvironments and, consequently, occur together in particular stratigraphical successions (e.g.

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Farrow, 1966; Ager and Wallace, 1970; Dawson and Reaser, 1996). Superimposition of Rhizocorallium and Thalassinoides or Spongeliomorpha as composite trace fossils may result in their co-occurrence on the same bedding plane and, where closely associated, leave the impression of a

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compound trace fossil (Fig. 19; Schlirf, 2011, fig. 5). Furthermore, the effect of commensalism and subsequent inhabitation of empty burrow tubes may be responsible for aberrant characteristics such as

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pustules and protuberances (e.g. Schlirf, 2011; Rodríguez-Tovar et al., 2012), but their significance is

5

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yet poorly understood.

Rhizocorallium from the Germanic Basin

In this section, examples of the two valid ichnospecies of Rhizocorallium are described from their type area with emphasis to delineate their characteristic features. The coordinates of all mentioned localities are listed in Table 2, whereas Figure 20 displays a stratigraphical overview. The synonymy lists include junior synonyms together with key records from the Germanic Triassic. 5.1

*

Rhizocorallium jenense Zenker, 1836 1822

Conferve, Coralline – Schlotheim, p. 49; pl. V, fig. 2

1836

Rhizocorallium jenense – Zenker, pp. 219-220

15

ACCEPTED MANUSCRIPT Rhizocorallium jenense Zenk. – Schmid and Schleiden, pp. 45-46; pl. IV, fig. 9

1846

Spongia Rhizocorallium Gein. – Geinitz, p. 695; pl. XXV, fig. 21

1852

Rhizocorallium Jenense Zenk. – Bronn, p. 44; pl. XII, fig. 1

1853

Rhizocorallium Jenense Zenk. – Schmid, p. 27

1864

Rhizocorallium Jenense Zenker – Alberti, p. 51

1881

Taonurus saportai – Dewalque, pp. 43-44; pl. 1, fig. 1, 2

1881

Taonurus ultimus Sap. et Mar. – Saporta and Marion, pp. 89-90; fig. 28

1881

Taonurus Panescorsii Sap. et Mar. – Saporta and Marion, pp. 89-90; fig. 27

1883

Taonurus ultimus Sap. et Mar. – Saporta and Marion, pp. 104-106; fig. 28

1883

Taonurus Panescorsii Sap. et Mar. – Saporta and Marion, pp. 105-106; fig. 27

1885

Glossifungites saxicava Lomn. (Rhizocorallium) – Fuchs, pp. 419-423; pl. VII, fig. 1-2

1885

Rhizocorallium jenense Zenker – Fuchs, pp. 416-417; pl. VII, fig. 3

1886

Glossifungites saxicava – Łomnicki, p. 99

1887

Cancellophycus Marioni – Saporta, pp. 289-290; pl. IV, fig. 2-3

1887

Taonurus ultimus – Saporta, pp. 290-295; pl. V, fig. 1-2; pl. VI, fig. 1

1887

Taonurus ruellensis – Saporta, pp. 295-298; pl. VII, fig. 1-2

1893

Rhizocorallium Hohendahli – Hosius, pp. 42-49; pl. II, III

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D

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Rhizokorallium hildesiense n. sp. – Menzel, pp. 4-6; fig. 1; pl. 1, fig. 1

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partim 1902

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1846

1905

Spongillopsis triadica n. sp. – Fliche, pp. 33-34; pl. II, fig. 2

1905

Spongillopsis recurva n. sp. – Fliche, pp. 34-35; pl. II, fig. 3-4

1906

Rhizocorallium jenense – Walther, pl. 1, fig. 1

1928

Rhizocorallium jenense Zenk. – Schmidt, p. 120; fig. 230

1929

Rhizocorallium jenense – Weigelt, pp. 28-29; pl. II, fig. 3, 6-8

1935

Rhizocorallium jenense Zenker – Abel, pp. 448-452; fig. 374

1935

Rhizocorallium weigelti n. sp. – Abel, pp. 451-452

1954a Rhizocorallium jurense n. sp. – Mayer, p. 25, fig. 1 1959

Rhizocorallium jenense – Faber, p. 31; fig. 10

1965

Corophioides – Hecker, p. 42; pl. XI, fig. 1-2

16

ACCEPTED MANUSCRIPT ?

?

1965

Rhizolith – Hecker, p. 42; pl. XI, fig. 3

1965

Rhizocorallium jenense Zenker – Hoppe, pp. 282-283; pl. I, fig. 2

1969

U-förmige Bohrungen von Polydorites – Hölder and Hollmann, pp. 82-85; fig. 2,

T

4, pl. 5

Rhizocorallium hunanensis – Zhang and Qing, p. 256; pl. 1, fig. 1

1995

Fuersichnus striatus isp. nov. – Buatois, pp. 260-262; fig. 2-5

1999

Rhizocorallium jenense – Werneburg, p. 253; fig. 6

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1988

Shallow to deep U-shaped pouches – Knaust, p. 364; fig. 2A, D, 5D, 7F

2011

Diplocraterion parallelum Torell, 1870 – Schlirf, pp. 38-39; fig. 3

2011

Diplocraterion ultimum (Saporta & Marion, 1883), n. comb. – Schlirf, pp. 39-41

2011

Rhizocorallium jenense Zenker, 1836 – Schlirf, pp. 48-51; fig. 13, 14

2012

Rhizocorallium jenense – Knaust et al., p. 725; fig. 8C, D

D

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2008

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v

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2007a Rhizocorallium jenense – Knaust, p. 230; fig. 8A

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Original diagnosis: “Wurzelförmige schlingenartige (hakenförmig) gebogene am Rande abgerundete zusammengedrückt-walzenförmige Gebilde mit verlängerten maschenförmigen schmalen Längsstreifen (ungefähr wie in der Lindenrinde sich verbindende Baströhren).”

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[“Rooty, loop-like (hook-shaped) arcuate, on the margin rounded, compressed-cylindrical structures with elongate stitch-shaped narrow longitudinal striae (like the connecting bast canals within basswood bark)”].

Emended diagnosis: Unbranched burrows with varying oblique orientation. The short to elongate, straight, bow- or U-shaped excavations have a system of subparallel or net-like scratches on the wall, and are open or passively filled.

Locus typicus: Hausberg near Jena (Zenker, 1836, p. 202).

Stratum typicum: Rhizocorallium-Dolomit (Upper Buntsandstein, Lower Pelitröt).

17

ACCEPTED MANUSCRIPT

Description 1: Upper Buntsandstein, Lower Pelitröt The type bed of R. jenense in the vicinity of Jena, as introduced by Zenker (1836), is a 10-40 cm thick

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sandy dolomite bed within a 3-5 m thick horizon of greenish marlstone with gypsum (Wackenroder, 1836; Passarge, 1892; Fig. 21). The dominance of marlstone with intercalated gypsum, dolomite and

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sandstone is typical of the Pelitröt succession and results from the deposition on a coastal plain with fluvial influence (Backhaus, 1981). The Rhizocorallium-Dolomit contains marine bivalves and

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vertebrate remains (Mägdefrau, 1957), indicating the transgression of the Muschelkalk sea. R. jenense occurs in positive hyporelief with a high abundance and burrow density between 30 and 70 specimens

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per 100 cm2, resulting in up to 100% coverage of the bedding plane (Fig. 21). Burrow orientation with respect to the bedding ranges from 10° to 80°, with a tendency towards steep inclination (>50°). The burrow length/width ratio rarely exceeds 1, with a mean of 0.7. This is not a result of erosion, because

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the proximal parts of the burrows are preserved. The mean burrow length and width are 14 mm and 20 mm, respectively. The average tube diameter is 5 mm and the tube diameter/width ratio is 0.25. All

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examined burrows are straight in plan view and without faecal pellets, but are densely scratched. Sets with parallel scratches are intersecting each other at various angles, thus generating a net-like pattern along the tube margins and in the enclosed spreite area. The original firm sediment between the

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marginal tubes was successively excavated and the resulting empty space was subsequently filled with sediment from the overlying sandy carbonate bed.

Description 2: Basal Muschelkalk, Gelbe Plattenkalke In Rittersdorf, the basal portion of the Lower Muschelkalk consists of dolomitic platy limestone (Gelbe Plattenkalke) with polygonal desiccation cracks originating on an upper intertidal flat. Some of the biolaminated beds contain R. jenense preserved as endichnia with dolomitic limestone fill (Fig. 22A). Most burrows occur in areas between the cracks, while some penetrate the filled cracks. They are steeply inclined and have a straight or curved dumbbell cross-section. The average density is 8 specimens per 100 cm2. Burrow width ranges between 15 and 30 mm and the average tube diameter is 3 mm.

18

ACCEPTED MANUSCRIPT

Description 3: Lower Muschelkalk, Obere Oolithbank The basal part of the Obere Oolithbank contains a micritic firmground to hardground with burrows of

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the Glossifungites ichnofacies and borings of the Trypanites ichnofacies (mainly Balanoglossites triadicus and Trypanites weisei; see Mägdefrau, 1932), indicative of an omission surface with gradual

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substrate hardening. Fracturing and initial brecciation of this bed is attributed to calichification on a peritidal mud flat, whereas the overlying oolitic limestone resulted from shoal deposition (Knaust et

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al., 2012). R. jenense occurs as an accompanying trace fossil, and may be intergrade with the burrow systems of B. triadicus (Knaust, 2008; Fig. 17A). In Blankenhain-Lohma, this surface contains open

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or loosely filled burrows with an average density of 26 specimens per 100 cm2 and with a burrow orientation between 10° and 80° (mean 56°, Fig. 22B). The mean burrow length/width ratio is 1.4. Striae are well preserved along the burrow margins. Most commonly exposed are cross-sections with

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the characteristic dumbbell-like outlines and a width between 1 and 3 cm (Fig. 22C), whereas the

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length of the oblique to subhorizontal structures may reach several centimetres.

Description 4: Lower Muschelkalk, Untere Terebratelbank The interval around the Terebratelbank is assumed to contain the maximum flooding surface within

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the third-order Lower Muschelkalk sequence (Aigner and Bachmann, 1992; Knaust, 1998). The skeletal Untere Terebratelbank rests above a micritic limestone with an omission surface riddled by numerous specimens of R. jenense. In Großmonra-Burgwenden, the dolomite-filled burrows are 10-15 mm wide and few centimetres long, and penetrate the surface with an angle of 10-30° (Fig. 22D). Only the lower burrow portions are preserved due to subsequent erosion.

Description 5: Lower Muschelkalk, upper part R. jenense also occurs on some micritic firmground surfaces, especially within the Upper Wellenkalk and between the Untere and Obere Schaumkalkbank. Many excavations have a large spread in size, ranging from few centimetres in width down to only some millimetres (Fig. 23A, B). They are open (Fig. 23C) or filled with dolomitic soil (Fig. 23D) and are often associated with B. triadicus and

19

ACCEPTED MANUSCRIPT elongated to winding grooves on the bedding surface from which they originate (Fig. 23E). Closely associated fractures, probably a result of calichification in a supratidal environment, indicates incipient

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hardground conditions.

Description 6: Lower Muschelkalk, lower and upper part (quarry Winterswijk)

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In contrast to the basinal Muschelkalk facies as developed in central Germany (descriptions 1-5), the quarry Winterswijk in The Netherlands exposes the marginal Muschelkalk facies. It is dominated by

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thin-bedded platy limestone with abundant microbial structures and desiccation cracks owing deposition on extensive tidal flats (Diedrich, 2001). Horizons with R. jenense occur in the proximal

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lower and upper part of the succession (Faber, 1959). The burrows are preserved in hyporelief and occur together with numerous casts of the bivalve Neoschizodus orbicularis. R. jenense is abundant on several bedding planes within the biolaminites (10 specimens per 100 cm2), where the burrows have a

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highly variable orientation (ranging from 0° to 80°, mean 44°, Fig. 23F). Their size is comparatively small with burrow length of 3-52 mm and burrow width of 10-38 mm. The length/width ratio of the

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burrows ranges from 0.1 and 1.0 (mean 0.55). In some instances, the burrows originate from the margins of narrow cracks which serve as weak zones for penetration. Interpenetration of burrows is

Discussion:

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common. The burrows exhibit clear networks of scratches and are passively filled.

R. jenense has been identified from marginal-marine deposits of the Germanic Basin by numerous workers (e.g. Schmid and Schleiden, 1846; Walther, 1906; Hoppe, 1965; Knaust et al., 1999; Werneburg, 1999). However, because of its nature and appearance, its recognition has not always been straightforward and Fuchs (1895, p. 417) noted: “The densely packed, gregarious or nest-like accumulation, which the Rhizocorallium Jenense for the most part shows at the lower surface of the Rhizocorallium Dolomit, but particularly the mutual intermingling of single individuals, generally creates such a tangle of fibrous bulges and skins that it becomes difficult to identify the actual basic form, and this is certainly the reason that this simple basic form is known to the fewest specialists, and discrete but well developed individuals of Rhizocorallids, if occurring in isolation, usually are not

20

ACCEPTED MANUSCRIPT recognized as such.” [Translated from German]. As a result of the difficulty of recognition, various ichnospecies of Rhizocorallium were described subsequently despite being within the morphological and size range of R. jenense or R. commune. In the review of Fürsich (1974a), only three ichnospecies

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are regarded as valid, an approach that was respected by most subsequent workers. Even so, the emended diagnosis and figures of R. jenense given by Fürsich (1974a) do not

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reflect this ichnotaxon as it was originally introduced and as it is known from its type area, and no paratype was evaluated, probably owing to the political situation of the time (the type locality was part

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of East Germany). Thus, the diagnosis given by Fürsich (1974a) is more applicable for R. commune var. auriforme instead of R. jenense. The approach followed by Fürsich (1974a) can be traced back to

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the influential work done by Richter (1926) and Seilacher (1955), although early field workers such as Wahnschaffe (1899, p. 13) underscore the discrimination of R. commune and R. jenense as proposed herein. It has had wide-ranging consequences as these two ichnotaxa differ considerably in their

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morphology, inclination and fill and therefore also by their ethological interpretation. Subsequent workers mainly referred to R. jenense sensu Fürsich (1974a) but not to the original description from

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Zenker (1836)!

The outline and shape of some R. jenense may resemble those known from R. commune, which gives reason for confusion. However, R. jenense consists of open or passively filled burrows

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(bored “Kalk-Rhizocoralliden” sensu Richter, 1924, p. 134), whereas R. commune has an actively filled spreite and a passively filled marginal tube (“Sand-Rhizocoralliden” in loose sediment; Richter, 1924). This is due to different substrate consistency (firm versus soft) as a result of different subenvironments, sequence-stratigraphic positions and animal behaviours. As previously realized by Weigelt (1929, p. 27), the principal types of substrate consistency are useful ichnotaxobases (Bertling et al., 2006) and thus discrimination of R. jenense and R. commune with similar morphology but different fill is regarded as essential. Bromley and Asgaard (1979) introduced the ichnogenus Fuersichnus, which somewhat resembles R. jenense. F. communis shows a gradational series of transitions to R. jenense (e.g. Eagar et al., 1985) and might be produced by the same kind of organism. F. striatus Buatois, 1995 has

21

ACCEPTED MANUSCRIPT characteristics that are more consistent with R. jenense (particularly in comparison with its type material) than with Fuersichnus and therefore F. striatus is regarded as junior synonym of R. jenense. R. jenense was from the first recognized as a component of the Glossifungites ichnofacies (e.g.

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Seilacher, 1967), of which its junior synonym Glossifungites saxicava is the namesake (Fig. 24). R. jenense commonly demarcates major marine flooding surfaces and thus is a valuable trace fossil for

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sequence-stratigraphical analysis (MacEachern et al., 2012b). Only the separation of morphologically similar but structurally different R. jenense and R. commune allows maintaining the sedimentological

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importance of R. jenense as an indicator of flooding surfaces within the Glossifungites ichnofacies. The descriptions given above demonstrate the stratigraphical and palaeoenvironmental

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occurrence of R. jenense in the Germanic Basin, where it is related to marginal-marine dolomitic deposits preferably in a supratidal to upper intertidal environment. This fact is supported by additional occurrences given in the literature: Hohenstein (1913, pp. 17, 52) mentioned Rhizocorallium from the

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dolomitic facies below the Hornstein-Schichten in the upper part of the Middle Muschelkalk Group of the eastern Black Forest, and R. jenense occurs within the same stratigraphic horizon in the abandoned

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quarry Troistedt (personal observation). Heller (1930, pp. 69-70) reports R. jenense from the dolomitic Steinmergelbänke of the Grundgips of the Middle Keuper of southern Germany.

Rhizocorallium commune Schmid, 1876

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5.2

?partim 1761

Beinstein, Rhizolithi, versteinte Wurzeln oder Wurzelsteine – Schütte and Merckel, p. 105

partim 1802

Wurm- und Zungen-Kalkstein – Batsch, pp. 55-58

?

1820

Bitubulites irregularis – Schlotheim, p. 376

1843

Thier-Fährthen? – Becks, pp. 188-190

1843

Fucoides auriformis – Hall, p. 47; pl. I, fig. 2

1846

Korallenähnliche Bildungen, wurmförmige (schlangen- und hufeisenförmige) Wülste – Schmid and Schleiden, pp. 46-47

1852

Fucoides auriformis – Hall, p. 7; pl. III, fig. 4

1853

Thier-Fährthen – Jugler, pp. 150-152; pl. II-IV

22

ACCEPTED MANUSCRIPT

1876

Rhizocorallium commune n. sp. – Schmid, pp. 5, 14, 17

1904

Melophycus curvatum – Ulrich, pp. 145-146, pl. XIII, fig. 2

1905

Cancellophycus – Fliche, pp. 20-21; pl. I, fig. 1

1909

Rhizocorallium jenense – Reis, p. 630; pl. XVII, fig. 8

1911

Rhizocorallium jenense Zenk. – Wurm, p. 129; pl. VII, fig. 12

1922

Rhizocorallium – Reis, pp. 229-230; fig. 1

1926

Rhizocorallium jenense Zenker – Richter, p. 217; pl. 3, fig. 1, 2

1928

Rhizocorallium commune Schmid – Schmidt, pp. 120-121; fig. 231

1929

Cavernaecola bärtlingi – Bentz, fig. 1, 2; pl. 74

1930

Rhizocorallium devonicum nov. sp. – Hecker, pp. 152-153, 156; pl. XVI, fig. 1, 2

1935

Rhizocorallium jenense Zenker – Abel, pp. 448-452; fig. 373

1936

Rhizocorallium jenense Zenk. – Schmidt, pp. 19-20; pl. III, fig. 5

1941

Upsiloides permiana, gen. et sp. nov. – Byrne and Branson, p. 261; fig. 1-6

D

MA

NU

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Lithochela problematica Guemb. – Gümbel, p. 411

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*

1861

v

1956

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1954b Rhizocorallium irregulare n. sp. – Mayer, pp. 82-83; pl. 2-3, fig. 1-4 Spiral eingedrehte Bauten von Rhizocorallium – Müller, pp. 405-407; pl. 1, 2 1958

Rhizocorallium uliarensis – Firtion, p. 111; fig. 1, 3; pl. 1, 2

?

1958

Rhizocorallium ? striatum nov. spec. – Kühn, pp. 447-449; fig. 3

1959

Rhizocorallium jenense – Faber, p. 31; fig. 11

1965

Rhizocorallium devonicum Gekker – Hecker, p. 42; pl. X, fig. 1

1971

Rhizocorallium variété Glossifungites Lomnicki, 1886 – Gall, p. 69; pl. XXV,

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+

fig. 3, 4 1974a Rhizocorallium uliarense Firtion, 1958 – Fürsich, p. 24 ?

1975

Rhizocorallium mongraensis sp. nov. – Chiplonkar and Ghare, pp. 78-79; fig. 2I, 4

1979

Rhizocorallium karaiensis sp. nov. – Chiplonkar and Ghare, p. 104; pl. 1, fig. 7

1980

Ilmenichnus devonicus (Hecker, 1930) – Hecker, p. 20; pl. 3, fig. 2

23

ACCEPTED MANUSCRIPT partim 1982

Rhizocorallium kueichowensis n. ichnospec. Yang et Sun – Yang and Sun, p. 373; pl. 1, fig. 1-6; pl. 3, fig. 1

1986

Rhizocorallium karaiensis Chiplonkar and Ghare – Ghare and Kulkarni, p. 51; pl. 3,

Rhizocorallium kutchensis ichnosp. nov. – Ghare and Kulkarni, pp. 50-51; pl. 6, fig. 2

1988

Gumatagichnus unguliformis Gab., Kurb. et Senn., ichnosp. nov. – Gabunia et

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1986

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?

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fig. 2

al., p. 196; fig. 2, 3

Rhizocorallium lhasaensis (ichnosp. nov.) – Yao et al., pp. 221-222; pl. 1, fig. 1, 7a;

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1992

pl. 2, fig. 1, 2a, b, 5, 7

Rhizocorallium shexingensis (ichnosp. nov.) – Yao et al., p. 222; pl. 1, fig. 5, 6

1993

Rhizocorallium wanghucunensis Xia, Zhong, Fan et al. (sp. nov) – Xia et al., pp. 61-

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1992

62; pl. 1, fig. 1-6

Rhizocorallium commune Schmid, 1896 – Helms, pp. 301-303; fig. 1; pl. I

1996

Rhizocorallium lixianensis ichnosp. nov. – Zhang and Wang, pp. 485-486; pl. 3, fig.

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1995

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15 1997

Rhizocorallium yelamensis ichnosp. nov. – Sanganwar and Kunda, p. 51; fig. 3/7

2001

Rhizocorallium karaiensis – Borkar and Kulkarni, pp. 16-17; fig. 2-5

2007a Rhizocorallium commune – Knaust, p. 230; fig. 8B

v

2007a R. uliarensis – Knaust, p. 230; fig. 8D

?

2007

Rhizocorallium uliarense? Firtion – Lanés et al., p. 66

2008

Rhizocorallium jenense – Rodríguez-Tovar and Pérez-Valera, pp. 79-81; fig. 5-7

2008

Rhizocorallium irregulare – Rodríguez-Tovar and Pérez-Valera, p. 81; fig. 8

2011

Ilmenichnus atherus n. isp. – Schlirf, pp. 41-42; fig. 6-10

2011

Ilmenichnus multilobatus n. isp. – Schlirf, p. 45; fig. 5, 11-12

2012

Rhizocorallium jenense spinosus – Rodríguez-Tovar et al., pp. 115-117; fig. 4, 5

2012

Rhizocorallium commune – Knaust et al., pp. 725-726; fig. 8E, F

v

AC

v

24

ACCEPTED MANUSCRIPT Original diagnosis: “…wurmförmige[n] Concretionen auf den Schichtflächen der Kalkschiefer des Muschelkalks …zeigen eine netzartige Zeichnung der Oberfläche, wenn auch minder deutlich und in sehr vergrössertem Maasstabe, wie sie Rhizocorallium jenense auszeichnet” (Schmid, 1876, p. 17).

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“…rätselhafte[n], wurmförmig-verschlungene[n], auch wohl sich verzweigende[n] Concretionen” (Schmid, 1876, p. 5).

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[“…worm-like concretions on the limestone bedding planes of the Muschelkalk …displaying a net-like pattern on the surface, although less pronounced and on a larger scale than developed in

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branching concretions” (Schmid, 1876, p. 5)].

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Rhizocorallium jenense” (Schmid, 1876, p. 17). “…enigmatic, worm-like intermingled, obviously also

Emended diagnosis: Unbranched or branched burrows with a preferred subhorizontal orientation. The burrows are elongate, band-like, straight or winding, and may have subparallel longitudinal scratches

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on the wall. Faecal pellets (Coprulus isp.) are common within the actively filled spreite and the

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marginal tube.

Locus typicus: Eastern Thuringia.

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Stratum typicum: Basal part of the Lower Muschelkalk (Lower Wellenkalk) near Jena.

Description 1: The type horizon of Rhizocorallium commune The type horizon of R. commune from eastern Thuringia, as introduced by Schmid (1876), lies within the basal part of the Lower Muschelkalk (Lower Wellenkalk), where this ichnotaxon occurs on several bedding planes within wavy and platy limestone facies. In Dorndorf-Steudnitz, a bedding plane with large R. commune was exposed a few metres above the Muschelkalk base but still below the Konglomeratschicht d2. The burrows occur as endichnia and occupy about 30% of the bedding plane. The long, loosely winding spreiten burrows are relatively narrow and robust (Fig. 25A). A better exposure was available in Bad Berka-Tannroda, where the succession between the uppermost Buntsandstein and lowermost Muschelkalk Group revealed numerous bedding surfaces

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ACCEPTED MANUSCRIPT containing R. commune with a length of 5-15 cm and a widt of 4-8 cm (Knaust, 1996). A high burrow concentration occurs on discrete bedding surfaces within the dolomitic and platy limestone of the Gelbe Plattenkalke (Fig. 25B). Some burrows have short and abandoned side-branches (primary

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successive) resulting from an earlier generation of burrows, whereas others exhibit retrusive burrow elements. In one specimen, this horizontal retrusive burrow part is obviously related to interference

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from another burrow, whereas in others the producer simply cross-cut pre-existing burrows. All burrows are densely scratched and filled with Coprulus oblongus.

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In addition, R. commune is found in the Lower Pelitröt near the Rhizocorallium-Dolomit (with R. jenense), where it occurs on the upper bedding plane of a sandstone bed (Fig. 25C; see also Gall,

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1971). It also occurs in the Upper Buntsandstein (Röt) of other regions (e.g. Rücklin, 1934), and is common throughout the Lower Muschelkalk succession (Fig. 25D-F).

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Description 2: The type bed of ‘Rhizocorallium irregulare’ (= R. commune)

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The type bed of ‘R. irregulare’ occurs about 2 m above the Spiriferinabank, a regional marker bed

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within the Germanic Basin (Schäfer, 1973). In addition to the original account by Mayer (1954b), Mundlos and Urlichs (1987) provide a detailed description of the bed with Rhizocorallium. The same bed was exposed in Troistedt, from which substantial material could be studied in situ and collected

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for detailed evaluation (Fig. 26). This bed is accompanied by other R. commune-bearing beds below and above it, all within the Anisian evolutus Zone (Fig. 20). The R. commune bed in Troistedt is a ca. 5-10 cm thick rudite bed with bioclasts (mainly bivalves) within a marlstone-limestone alternation (Fig. 26A). Its base shows orientated flute casts and its micritic top surface preserves large R. commune. The bed is yellowish to brownish due to its dolomitic content. The bedding surface is extensively bioturbated and R. commune typically covers half or more of the available area. In rippled areas, the burrows preferentially follow the troughs between the ripples. The burrows are mostly preserved in negative epirelief but occasionally a combination of negative and positive epirelief occurs where parts of the burrow are filled with sediment. Burrow orientation is horizontal. Some bedding planes are microbially modified, while

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ACCEPTED MANUSCRIPT others were diagenetically overprinting with various caliche features such as crystallaria, fractures and chalky limestone reprecipitation. Most measured burrows are 10-20 cm (maximum up to 59 cm) long and 5- 7 cm (maximum

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more than 17 cm) wide. The average tube diameter is ca 1.5 cm with a range between 0.9 and 2.5 cm, whereas the average tube diameter/width ratio is 0.22 (ranging from 0.09 to 0.3). R. commune shows a

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tendency toward a sinuosity index with an average of 1.15 and a maximum of 2.7. All burrows are scattered with the faecal pellets Coprulus oblongus. Elongated and subparallel scratches are poorly

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preserved along some marginal tubes (Fig. 26B). In contrast to the actively fillede spreite area, the marginal tubes are passively filled with micritic sediment or with loosely arranged bioclasts. One tube

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contained an accumulation of the oyster Placunopsis ostracina. Some specimens have one or more side branches (Fig. 26C), evidently as a reaction of the producer to avoid obstacles such as pre-existing spreiten. The marginal tubes are repeatedly reworked

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and consist of several generations of burrows with the same diameter as the grooves or tunnels of the spreite (Fig. 26E, F). Some of these burrows contain a distinct fill of C. oblongus as is typical for R.

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commune, and thus indicate the same producer for burrows and pellets. Asymmetrically produced sinistral and dextral spreiten are common, with an origin of the tunnel from the merging tube and truncation on the other.

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Similar to the Spurenbank described by Mayer (1954b), R. commune is closely associated with other trace fossils such as loosely winding burrows, shallow grooves and ichnospecies of Balanoglossites. Winding burrows are a common constituent of this ichnocoenosis and in several instances are linked with the spreiten in a similar manner as described by Basan and Scott (1979). Some of them branch off and display an incipient spreite. Grooves occur abundantly on some wellpreserved bedding surfaces and are also linked to the spreiten (Fig. 26B, D). They reach several centimetres in length and their width is in the same size range as the tube diameter of R. commune. Repeated excavation and lining with C. oblongus is common for both, the spreite burrows and the grooves. Furthermore, a genetic relationship between the grooves and R. commune is obvious in a spreite which consists of a groove-like extended marginal tube and is associated with other isolated grooves on the same surface. Finally, winding burrows and shallow grooves typically intergrade with

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ACCEPTED MANUSCRIPT complex burrow systems down to several centimetres, which are referred to Balanoglossites ramosus (Knaust, 2008). Other limestone beds above and below the described bed also preserve R. commune with

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similar features as described above. As well as negative epirelief, preservation in positive hyporelief is common, where the spreiten burrows overprint a softground suite consisting of Planolites,

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Protovirgularia and Lockeia, all attributed to the burrowing activity of benthic bivalves (Knaust,

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2007a).

Discussion: R. commune has been a well established and commonly used ichnotaxon for the last 137

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years in the realm of the Germanic Triassic (Schmid, 1876; Wagner, 1897; Wurm, 1911; Schmidt, 1936; Hörauf, 1958; Faber, 1959; Freyberg, 1963; Wagenbreth, 1968; Schulz, 1972; Hoppe and Seidel, 1974; Aigner, 1979, 1984; Wetzel and Aigner, 1986; Lukas, 1989; Gensel et al., 1990; Dünkel

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and Vath, 1990; Klotz, 1991; Helms, 1995; Rosenfeld and Thiele-Papke, 1995; Hagdorn, 2001; Diedrich, 2002; Gall and Grauvogel-Stamm, 2005; Knaust, 2007a, 2012b; Salamon et al., 2012). In

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contrast, it is poorly recognized elsewhere (e.g. Hagdorn, 1996; Mørk and Bromley, 2008; Desjardins et al., 2010; Boyd and Lillegraven, 2011; Buatois and Mángano, 2011; Knaust and Costamagna,

(Table 3).

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2012; Knaust et al., 2012), where it has been gradually replaced by its junior synonym R. irregulare

Fucoides auriformis (Hall, 1843) (Fig. 2) appears to be the senior synonym of Rhizocorallium commune Schmid, 1876. Hofmann (1979) re-examined the material of Hall (1843, 1852) and “...consider[ed] it to represent burrows with protrusive spreiten”, consequently attributing it to the ichnogenus Rhizocorallium. In order to maintain ichnotaxonomical stability and in accordance with the conditions of Article 23.9.1 of the International Code of Zoological Nomenclature, Rhizocorallium commune Schmid, 1876 (nomen protectum) is valid and has precedence over Fucoides (= Rhizocorallium) auriformis (Hall, 1843) (nomen oblitum). This reversal of precedence can be done because the conditions of Article 23.9.1.2. are met (references to the usage of the junior synonym R. commune as presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years are listed in the

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ACCEPTED MANUSCRIPT previous paragraph). To the author’s knowledge, the condition in Article 23.9.1.1. applies, and the senior synonym F. auriformis has not been used as a valid name after 1899. Exceptions include very few mere listing of that name in a catalogue, index and list of fossils traceable till 1915 (Clarke and

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Ruedemann, 1903; Grabau, 1010; Bassler, 1915), which in accordance to Article 23.9.6. must not be taken into account in determining usage under Articles 23.9.1.1. and 23.9.1.2.

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Schweigert (1998) and Schlirf (2011) have questioned the validity of R. commune but their reasoning is not followed here: Given the time of publication, Schmid (1876) properly described this

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ichnotaxon, as also demonstrated by its subsequent unambiguous application. Although originally unfigured, the holotype was deposited in the Museum of Jena, where parts of the Schmid collection

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are still available in today’s Mineralogical Museum of the University of Jena. Unfortunately, the type material of R. commune could be found neither in the Museum nor in the University’s Geosciences collection and likely has been lost. In contrast, Mayer (1954b) established R. irregulare for the same

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form with proper figures but no designation of a holotype; therefore a lectotype was defined subsequently by Schweigert (1998).

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Thirty years before its formal introduction, Schmid and Schleiden (1846) characterized R. commune from the Upper Muschelkalk (Trochitenkalk): “Vermiform bulges – It is very likely that the meandering and horseshoe-shaped bent bulges, which occur throughout the whole Muschelkalk

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succession, have a common and in fact organic cause. In most instances they lie firm on the bedding surfaces and consist, even if they can be separated, of the same substance as the rock; their length and thickness is very variable. I cannot notice an outer and inner structure as reported from Quenstedt (Das Flötzgebirge Würtembergs S. 70.) and Geinitz (G.v.S.S. 109).” [Translated from German]. Many workers have realized that size differences (i.e. small and large forms) of R. commune exist (e.g. Ager and Wallace, 1970; Worsley and Mørk, 2001; Bann and Fielding, 2004) and this fact has palaeoenvironmental implications. Because small and large forms have gradual transitions, a subdivision on the ichnosubspecies level cannot be justified but distinction of varieties seems to be meaningful with respect to their value as facies indicators. Therefore, a relatively small form, R. commune var. auriforme, is discriminated from a relatively large form, R. commune var. irregulare, in

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ACCEPTED MANUSCRIPT addition to the rare trochospiral ichnosubspecies R. commune uliarense and the vertically retrusive spreite burrow R. commune problematica. Rhizocorallium uliarense Firtion (1958), from the Upper Jurassic of SW-France, consists of

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trochospiral U-shaped spreite burrows (Fürsich, 1974a, p. 24). This form is rare in the Muschelkalk and only few specimens were found by the author (Knaust, 2007a) in addition to those described by

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Reis (1910) and Müller (1959). All material is from the Lower Muschelkalk and two new specimens were collected just 2-3 m below the Untere Terebratelbank in Bad Berka-Tannroda. The diameter of

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the tight spirals is about 8 cm with a compacted depth of ca. 2 cm. A marginal tube is only weakly developed, but scratches are well pronounced and mimic the spreiten. Specimens with weakly

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developed spreiten show affinity to the ichnogenus Gyrolithes. Separation of R. uliarense from the ichnogenus Rhizocorallium, as proposed by Schlirf (2011), is not recommended here because the type material from SW-France contains a long spreite structure with the scratches and faecal pellets

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Coprulus oblongus typical of R. commune, but only differs from it by trochospiral morphology. Gümbel (1861, p. 411) described Lithochela problematica from the Upper Triassic (Kössener

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Beds, Rhaetian) of the Bavarian Alps : “Bodies of horseshoe-like bent form, which are similar to the Rhizocorallium of the Muschelkalk [Röth], commonly occur in fairly consistent size; the bulges are rounded, more flattened towards the inner side and in that direction accompanied by figures on the

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rock, similar to a body which has moved in the mud and left the bulges back as a trace.” [Translated from German]. Because no type material could be found in the Gümbel collection of the Paläontologische Staatssammlung of Munich, it must either have never deposited or was subsequently destroyed during World War II (Winfried Werner, personal communication 2010). However, Fuchs (1895, p. 427; pl. VII, fig. 4-7) described and figured Rhizocorallium with a vertical retrusive spreite element from the same beds, indicating that this form seems to be common in the Kössener Beds and therefore the original ichnospecies name is applied in R. commune problematica (Fig. 25F). Later workers have described this form (partly as Teichichnus) from different geological periods and regions of the world (e.g. Chisholm, 1970; Sellwood, 1970; Fürsich, 1974, 1981; Chamberlain, 1977; Buckman, 1994; Bhattacharya and Bhattacharya, 2007; Kowal-Linka and Bodzioch, 2011; Schlirf, 2011).

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ACCEPTED MANUSCRIPT 5.3

Morphometric analysis

A morphometric analysis of hundreds of specimens from the German Triassic was performed in order to test the proposed classification. The prominent morphology and size of ichnospecies and varieties

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of Rhizocorallium offer ideal morphometric data for comparison, such as burrow length/width ratio, width/tube diameter ratio and inclination to the bedding. The graphs of these parameters suggest a

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close relationship between the different forms with broad transition fields (Fig. 27). The length/width ratio (Fig. 27A) clearly separates R. commune var. irregulare from the much

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smaller R. commune var. auriforme and R. jenense. The wide transition field suggests the activity of the same or similar producer in different ontogenetic stages for all three forms, although

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palaeoenvironmental conditions such as depositional energy, substrate consistency etc. may also play a role. A comparison of R. jenense and R. commune var. auriforme shows their similarity in size, a fact

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that explains the widespread confusion of both forms and the general application of the ichnotaxon

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name R. jenense for both forms (Fig. 27B). However, the length/width ratio of R. jenense is less than half of the one observed in R. commune var. auriforme, a fact that may be related to different feeding

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styles (e.g. suspension- versus deposit-feeding). Further, a more resistant substrate (e.g. firmground) needs to be penetrated by the R. jenense producer, which results in the development of more shallow and broadened bow-shaped burrows, while R. commune tends to be more elongate.

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The ratio of burrow width and tube diameter roughly varies between 4 and 5, and a roughly linear trend between R. commune var. auriforme and R. commune var. irregulare is obvious (Fig. 27C). R. jenense differs from R. commune var. auriforme by a slightly higher mean value (4.6 versus 3.8). Effective discrimination of R. jenense and R. commune is accomplished by the burrow inclination to the bedding (Fig. 27D). R. jenense is typically inclined with highly variable angle ranging from 0° to 90°, whereas R. commune is only slightly inclined, with angles about 10° (R. commune var. auriforme), or are completely horizontal (R. commune var. irregulare). Only rarely does the deviation of R. commune reach 70°, which is the case when incipient or minute burrows of juvenile producers are taken into consideration.

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6

Rhizocorallium outside the Germanic Basin

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In addition to the above mentioned and described records of Rhizocorallium from the Germanic Basin, this trace fossil has been reported from Phanerozoic deposits all over the world. Following the newly

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established classification, Table 3 contains Rhizocorallium records from outside the Germanic Basin as compiled from the literature. Although this overview is by no means complete, it includes about

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180 published records of described and/or figured occurrences which could be designated accordingly

The Rhizocorallium producers

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7

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and analysed with respect to their age, environment, inferred producer, etc.

The interpretation of the Rhizocorallium trace maker has long been debated, a fact that is related to the

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heterogeneous nature of the ichnogenus that implies several producers. End members of both ichnospecies differ in morphology, orientation, size ratio, and presence or absence of faecal pellets, and each has its specific scratch pattern calling for distinct trace maker behaviour (Table 1). After a

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brief historical review, the most likely trace makers of Rhizocorallium are discussed based on the evidence of significant features and by ruling out unlikely producers.

7.1

History of interpretation

R. jenense was originally regarded as a peculiar alga or coral (Schlotheim, 1822), and the name Rhizocorallium as introduced by Zenker (1836) reflects an interpretation as coral. Following workers presumed R. jenense to be a sponge (Geinitz, 1846; Schmid and Schleiden, 1846; Bronn, 1852; Alberti, 1864) or alga (Saporta and Marion, 1881; Saporta, 1887). Łomnicki (1886) came very close to a trace fossil interpretation of ‘Glossifungites saxicava’ (= R. jenense) and broadly interpreted it as boring sponges whose excavations were subsequently filled with sand. Eventually, Fuchs (1895)

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ACCEPTED MANUSCRIPT suggested worm-like animals as the producers of those burrows, a view that was shared subsequently (Sarle, 1906: sedentary polychaetes; Douvillé, 1908: Polydora; Fuchs, 1909: annelids; Richter, 1924: Polydora; Prell, 1925: Polydora). Based on comparison with modern crab burrows, Weigelt (1929)

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concluded that the abundant scratches in R. jenense must result from the activity of crustaceans and since then, this opinion has a major influence on the interpretation of Rhizocorallium (e.g. Seilacher,

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1955, 1967, 2007; Fürsich, 1974a; Rodríguez-Tovar and Pérez-Valera, 2008; Neto de Carvalho et al., 2010). However, scratches can be produced by many groups of animals other than crustaceans and

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therefore do not necessarily prove crustaceans as trace makers!

In its original description of R. commune, Schmid (1876) followed the interpretation given by

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Zenker (1836) for R. jenense and also introduced R. commune as a coral, although already Gümbel (1861) was aware of the trace fossil nature of ‘Lithochela problematica’ (= R. commune problematica). Apart from one description as a body fossil (i.e. alga; Fliche, 1905), R. commune was

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commonly interpreted as a trace fossil produced by (annelid) worms (Reis, 1910; Richter, 1927; Schmidt, 1928; Heller, 1930; Hecker, 1965). Seilacher (1955) described and figured R. commune from

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the Lower Cambrian as R. jenense, and generally considered worms, crustaceans and insect larvae as potential trace makers. He hesitated to apply Weigelt’s (1929) brachyuran crustacean interpretation for such old burrows and favoured long-tailed crustaceans or other arthropods instead. Subsequent

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workers have either accepted the crustacean interpretation for R. commune because of the associated scratches (e.g. Rodríguez-Tovar and Pérez-Valera, 2008), favoured the worm interpretation (e.g. Hamm, 1957; Knaust, 2007a; Knaust and Costamagna, 2012), or considered both as possibilities (Mundlos and Urlichs, 1987). Three different groups of organisms are now commonly regarded as potential producers of Rhizocorallium: decapod crustaceans, annelids and mayflies (e.g. Prell, 1925; Seilacher, 1986). 7.2

Decapod crustaceans

Many of the delicately scratched surfaces of R. jenense offer detailed insights into the behaviour of their occupants, and the nicely developed bioglyphs are often cited as evidence of crustacean activity (e.g. Seilacher, 2007; Rodríguez-Tovar and Pérez-Valera, 2008; Neto de Carvalho et al., 2010). Aside

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ACCEPTED MANUSCRIPT from single scratches, paired scratches and sets with six or more parallel scratches typically criss-cross the burrow margin and produce the typical net-like pattern. Similar rhomboidal bioglyphs are also known from Spongeliomorpha, which is attributed to the activity of crustaceans (cf. Seilacher, 2007;

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Gibert and Ekdale, 2010) and can occur together with R. jenense (Reis, 1922; Weigelt, 1929; Fürsich et al., 1981). Some of the R. jenense material from Thuringia and the Ukraine (‘Glossifungites

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saxicava’) reveals an intimate co-occurrence of normally sized burrows and very tiny burrows only 1 mm or less in diameter (Figs. 23B). This feature is best explained by different ontogenetic stages, such

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as adults and juveniles, burrowing concurrently. Different development strategies with brooding and care are known from crustaceans and would support such an interpretation (Linke, 1939, p. 304;

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Brusca and Brusca, 2003).

Neoichnological evidence for crustaceans as producers of U-shaped burrows comes from amphipods of the genus Corophium, which comprises several species. The well-studied C. volutator

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creates U-shaped and simple vertical to subvertical burrows in sand, silt and mud of soft and firm consistency (Häntzschel, 1939; Schäfer, 1962, p. 346). These burrows can be several centimetres deep

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and have a smooth, mucus-stabilized burrow wall. Lateral branches or a repetition of the U-tube may mimic a spreite, however, result from vertical adjustment instead of continuous sediment mining and processing as in a true spreite. Overall, modern Corophium burrows may serve as an analogue for

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Arenicolites and Diplocraterion (e.g. Dashtgard and Gingras, 2012, p. 281, Fig. 5C, D) but only to a lesser degree for Rhizocorallium. The appearance of burrowing decapod crustaceans in the Triassic (probably already by the end of the Palaeozoic; Carmona et al., 2004) coincides with the occurrence of R. jenense and thus would support its interpretation as crustacean burrow, but probably discounts Palaeozoic occurrences of R. commune. 7.3

Annelids

Although decapod crustaceans have been quite fashionable in the interpretation of the Rhizocorallium producers, worm-like organisms (particularly annelids) are a second large but heterogeneous target group. Many lines of evidence make annelids the most likely group of producers, while their responsible systematic position may still be debatable.

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ACCEPTED MANUSCRIPT The extensive material from the Muschelkalk and individual published reports indicate a continuum between Rhizocorallium developed in softground (R. commune) and those in firmground (R. jenense), with similar overall morphology but contrasting fill (active versus passive). This fact

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suggests that the same kind of organism may have acted in various substrates. Moreover, shallow to deep U-shaped pouches in firm- and hardground (Figs. 17B, C, 23A-E) meet the diagnosis of R.

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jenense and intergrade with Balanoglossites triadicus, which suggests that both ichnotaxa were produced by the same trace maker. There is good evidence that B. triadicus results from the burrowing

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and boring activity of polychaete worms (Knaust, 2008) and the same may be true for R. jenense. Further support for a polychaete interpretation comes from the frequent occurrence of scolecodonts,

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the jaws of polychaete annelids, from the Muschelkalk, particularly the intervals with abundant Rhizocorallium (e.g. spinosus Zone; Kozur, 1967, 1974). In fact, the eunicid polychaete Marphysa sanguinea produces similar excavations (Warme,

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1970; Warme and McHuron, 1978, fig. 7C), and paired scratch traces could indicate chewing polychaetes (Warme and McHuron, 1978, fig. 6D) or, more likely, be the result of various stiff and

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scratching chaetae. Hooked chaetae are found in at least some members of all major clades of polychaetes and function as anchors to support the animal within its burrow (Merz and Woodin, 2006). Other potential candidates include species of the Spionidae (e.g. Polydora sp.) and Capitellidae

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(e.g. Dasybranchus glabrus).

Some species of spionid worms such as Polydora ciliata are able to burrow into soft to firm substrate and boring rocks or shells (e.g. Linke, 1939; Schäfer, 1962). The resulting U-shaped burrows/borings are orientated in various directions (from horizontal to vertical) and morphologically resemble the burrows of Corophium (Gingras et al., 2001). However, P. ciliata produces open pouches which subsequently may be filled actively with a spreite instead of vertical adjustment (Schäfer, 1962, fig. 177). Burrow excavation of Spionidae in firm and hard substrates takes place by means of scraping with the fifth chaetae. Lateral movement of the worm along the substrate surface creates an initially reniform burrow, before an arcuate burrow leads into the substrate (Prell, 1925; Hempel, 1957; Sato-Okosho-Okoshi, 2000). All these burrow stages are common on omission surfaces in the Muschelkalk (Knaust, 2008). This mechanical excavation is also responsible for the development of

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ACCEPTED MANUSCRIPT scratches (Sato-Okosho-Okoshi, 2000), which are best preserved along the inner part of the U-tube (Hempel, 1957, p. 119). In calcareous substrates, a combination of mechanical and chemical excavation is employed for boring (Sato-Okoshi, 1999, 2000). After a short planktonic phase,

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Polydora larvae settle in the vicinity of adults and immediately start to burrow with their fifth chaetae. This behaviour is consistent with the co-ocurrence of Rhizocorallium burrows of different size classes

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(Figs. 17B, C, 23A-E). Polydora species are mainly suspension-feeders. The faecal pellets of P. ciliata are similar to Coprulus oblongus as known from R. commune, and are deposited within their dwellings

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(Schäfer, 1962, fig. 187).

Terebellidae (e.g. Terebellides stroemi) are nicely documented as the producers of

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Rhizocorallium in Holocene sediments of the Western Baltic Sea based on X-ray images from box cores and vibra cores (Winn, 2006: Diplocraterion on plate 2; Fig. 28). These burrows penetrate bioturbated sandy clay in a shallow oblique direction, and their emptiness or passive sand fill indicates

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a firm substrate. They show a figure- eight cross-section, are ca. 2 cm wide and are traceable over several centimetres. In some instances, even elongated scratches are revealed by the radiographs. This

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finding clearly supports the polychaete origin of Rhizocorallium. The evidence that modern T. stroemi constructs inclined spreite burrows with scratched marginal tubes and Coprulus-like faecal pellets (Winn, 2006, p. 165) strongly suggests polychaetes as the producer of marine Rhizocorallium.

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As demonstrated on the comprehensive material studied from the Muschelkalk, R. commune has various features that allow their preferred assignment to the work of worm-like animals: 

In dependence of the kind of substrate, the spreite area reveals interesting features. For instance, originally stiff to firm substrates preserve traces of single movements of the trace maker in form of discrete grooves or tubes, which are progressively stacked into each other (U-in-U) to construct the spreite (Fig. 6). These small grooves are also evident in some better preserved marginal tubes (cf. Häntzschel and Reineck, 1968, pl. 16, fig. 4) and result from the repeated movement of the trace maker back and forth (Fig. 26E, F). The rounded cross section and the narrow but constant diameter of these grooves or tubes are most consistent with the work of a vermiform animal. In even stiffer substrates, only grooves are developed, which are clearly separated by narrow

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ACCEPTED MANUSCRIPT ridges. In soft sediment, burrowing results in a diffuse and completely reworked spreite structure. 

In many instances, the spreite burrows are closely associated with tunnels or grooves,

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which wind loosely along the bedding plane (Figs. 10, 17C, 26B, D). Their same appearance with respect to diameter and fill, as well as the fact that the tunnels may

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intergrade with the spreiten burrows, suggests that the same organism has produced these distinct traces. Most likely, a worm-like animal with different behaviour

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produced these compound trace fossils, in which the elongated and rounded tubes reflect movement through the sediment (repichnia), whereas the spreite burrows

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indicate deposit-feeding (fodinichnia). This relationship has frequently been pointed out in R. commune (e.g. Reis, 1910; Weigelt, 1929; Hecker, 1930; Farrow, 1966;

Faecal pellets (Coprulus) typically occur within the spreite and marginal tube of R.

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

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Fürsich, 1974a).

commune (Figs. 8, 13, 26D-F). These ellipsoidal coprolites with a rounded shape and a

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homogeneous composition have a length/width-ratio between 1.5 and 2.0 (Knaust, 2008). Instead of being produced by crustaceans (as suggested by, e.g., Seilacher, 2007; Patel and Desai, 2009), these faecal pellets differ from the typically cylindrical

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and internally structured crustacean faecal pellets. In contrast, they closely resemble faecal pellets produced by annelids (e.g. Martens, 1978) and fall into the same size range of modern Heteromastus filiformis pellets (Linke, 1939, fig. 67; Knaust, 2008). Furthermore, the diameter of the pellets and the tunnels within the spreiten are similar, indicating a sediment-processing worm-like producer.

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Coprulus ichnospecies are common in ichnospecies of Rhizocorallium, Diplocraterion, Zoophycos and Balanoglossites. R. commune and Balanoglossites ramosus are both closely associated and known elements of a compound trace fossil (Mayer, 1954b; Knaust, 2008; Figs. 17, 22C). A range of trace fossils from the Muschelkalk has exceptionally preserved producers (Knaust, 2010a), including a B. ramosus system with (Eunice-like) annelid fragments (Knaust, 2008, fig. 9).

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One of the bedding planes of the Upper Muschelkalk contains numerous limonitized remains of various trace makers. Incipient spreite burrows, assignable to R. commune, notably contain elongate limonite aggregates resembling a diagenetically modified

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Mayflies

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7.4

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cast of its vermiform producer (Fig. 29).

Fürsich and Mayr (1981) described non-marine (fluvial) Rhizocorallium from the Upper Miocene of southern Germany and classified them as R. jenense based on their morphology. These forms, most

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likely excavated by the larvae of commonly burrowing mayflies (Ephemeridae), suddenly weakened the ichnogenus Rhizocorallium as a facies indicator of marine environments. U-burrows with closely

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spaced tubes (“Rhizocorallium”) were also described by Goldring et al. (2005) from non-marine Early Cretaceous strata but failed to exhibit the diagnostic spreite. Burrows of mayflies have long been

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known in modern rivers, lakes and ponds (Abel, 1935). Intriguingly, the gregarious occurrence of R.

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jenense at the base of its type stratum (uppermost Lower Triassic, coastal plain environment) coincides with the record of a rich mayfly fauna in the same area (Bashkuev et al., 2012, and

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references therein). Sinitshenkova et al. (2005) described several hundred specimens of Ephemeridae from the early Middle Triassic Grès à Voltzia in NE-France and attributed U-shaped burrows to the

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activity of Voltziaephemera fossoria, the most abundant mayfly in the Triassic of the Vosges Mountains. Little is known about the morphological variability of fossil mayfly burrows. However, modern mayfly burrows may resemble R. jenense superficially, although a closer comparison reveals numerous subtle differences which allow for their proper assignment, with consequences for a correct palaeoenvironmental interpretation (compiled after Abel, 1935; Fürsich and Mayr, 1981; Charbonneau and Hare, 1998; and De, 2002): 

Gregarious occurrence of predominantly horizontal burrows, commonly penetrating the sediment from a vertical surface (e.g. cliff) parallel to each other in the same direction.



Commonly U- or J-shaped burrows, in some cases with discontinuous spreiten present.

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ACCEPTED MANUSCRIPT 

The length/width ratio of the burrows is much larger than in R. jenense from its type area, resulting in narrower marginal tubes. Consequently, the width/tube diameter is much smaller than in R. jenense.

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Bioglyphs, if present, consist of sets with scratches as produced by their gills and

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

differ from the net-like pattern characteristic for R. jenense.

Finally, their association with fluvial or lacustrine deposits aids in separation, although

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

this criterion is independent from ichnotaxonomy.

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Given these features, Abel (1935) erected the ichnogenus Ephemerites for such burrows, a procedure that was not followed by Fürsich and Mayr (1981). Ephemerites is a nomen nudum because no type

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ichnospecies was fixed. The above mentioned characteristics may warrant a different ichnogenus, and indeed, the ichnogenus Fuersichnus Bromley and Asgaard, 1979 shows gradational transition with R.

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jenense. The type ichnospecies F. communis is known from Upper Triassic red beds, and Garvey and

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Hasiotis (2008) introduced F. commerfordi from the Lower Carboniferous for poorly organized members of this ichnogenus. F. commerfordi is the oldest known Fuersichnus, and its first occurrence

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coincides with the appearance of mayflies during the Carboniferous. Similarly, the pouch-shaped and passively filled boring Asthenopodichnium Thenius, 1988 is attributed to the burrowing and boring activity of mayfly larvae or other arthropods in woody material and bones (Uchman et al., 2007). The

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variable behaviour of burrowing ephemeroptera was described by Edmunds and McCafferty (1996). Accordingly, more advanced burrowing with U-shaped burrows independent from adjacent rock interfaces has only developed in a late evolutionary lineage of Polymitarcyidae and Ephemeridae and thus could count for the Miocene freshwater R. jenense described by Fürsich and Mayr (1981).

8

Behaviour of the trace maker

Close agreement exists in the literature regarding the interpretation of the trace maker’s behaviour in both ichnospecies of Rhizocorallium. Producers belonging to different higher taxa with a contrasting behaviour are responsible for the origin of both ichnospecies. Fürsich (1974a) regarded R. jenense as

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ACCEPTED MANUSCRIPT the dwelling trace (domichnion) of a suspension feeder, whereas R. commune (‘R. irregulare’ therein) was interpreted as the feeding trace (fodinichnion) of a deposit feeder. In the light of the reinterpretation of the Rhizocorallium producer as a polychaete, and the existence of a continuum

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between R. jenense and R. commune, a combined suspension- and deposit-feeding is likely. R. jenense with its passive fill and inclination is best explained as the burrow of a suspension-feeder, while the

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extensive course, actively created spreite and faecal pellets make a deposit-feeding interpretation tempting for R. commune. Nevertheless, introduction of sediment material from the surface for storage

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(cache behaviour, Löwemark et al., 2004) or gardening (Fu and Werner, 1995) in the form of pellets similar to those observed in Zoophycos remains to be tested by compositional analysis of the pellets of

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R. commune and the surrounding sediment.

The mode of burrow construction remains uncertain. The most obvious method of spreite construction is progressive sediment-reworking on the surface or shallow subsurface by a worm

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positioned head-down (posteriorward) in its burrow, a concept that may apply for Eunice-like producers. Terebellidae, on the other hand, are known to lie head-up (anteriorward) in their dwellings

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and feed with their long tentacles on the surface. In this scenario, the spreite would result from a continuous backward movement of a growing worm, which uses the spreite for defaecation. This scenario is documented in numerous cases where the spreite displays an increase in size distally and

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thus can be regarded as the dwelling of a gradually growing vermiform organism (e.g. Bromley and Hanken, 1991). Jumars et al. (2007) pointed out that many burrowing polychaetes have been erroneously classified as surface deposit-feeders, feeding on sediment/water interface, but actually burrow in the shallow subsurface (e.g. the burrowing terebellid polychaete Eupolymnia heterobranchia).

9

Rhizocorallium as facies indicator

Trace fossils can serve as good facies indicators if a robust determination of ichnospecies in concert with sedimentological characteristics is applied (Knaust and Bromley, 2012). Rhizocorallium has a

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ACCEPTED MANUSCRIPT long tradition as facies indicator of shallow-marine deposits (Reis, 1910; Seilacher, 1955; Farrow, 1966; Ager and Wallace, 1970; Jansa, 1972; Heinberg and Birkelund, 1984; Worsley and Mørk, 2001; Rodríguez-Tovar et al., 2007; Rodríguez-Tovar and Pérez-Valera, 2008). However, the existing

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classification of Rhizocorallium, together with reports of deep-marine and fluvial specimens, has limited its full potential for interpreting sedimentary environments, a misperception which is also

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reflected in some textbooks. For instance, Bjørlykke (2010) adapted a scheme from Collinson and Thompson (1982), wherein Rhizocorallium characterizes the neritic zone (outer shelf) “... below the

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Cruziana facies”. This scheme only concerns one expression of Rhizocorallium typical for a shelf setting, but disregards abundant occurrences in other shallow-marine and marginal-marine

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environments as well as its contribution to the widespread Glossifungites ichnofacies.

Ichnofacies and bathymetry

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R. jenense is a firmground burrow characterizing the substrate-controlled Glossifungites ichnofacies.

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R. jenense is restricted to Mesozoic and Cenozoic deposits, where it occurs in all environments from peritidal (supralittoral) to the shelf and also in deep-marine (from Early Cretaceous) and fluvial

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(Miocene) deposits (Fig. 33). The type stratum of R. jenense was deposited by the gradual incursion (transgression) of the Muschelkalk Sea, where the burrows were produced in clay-rich red beds of

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continental origin and filled with marginal-marine fossiliferous limestone. This situation suggests a peritidal environment during colonization. In the Muschelkalk succession, R. jenense mainly occurs along omission surfaces in the shallowest part of shallowing-upward sequences (surfaces of maximum regression/onset of transgression, see MacEachern et al., 2012b; Fig. 30), chiefly corresponding to an upper intertidal to supratidal environment. Prell (1925, p. 376) already stated: “From the presence of Hühnerfährten [= U-shaped spreite burrows] it can be concluded with confidence that a marine transgression took place.” R. commune is a constituent of the Cruziana ichnofacies, which is characterized by predominantly horizontal feeding traces in softgrounds on the shelf, although a gradual transition exists to the colonization of firmground. It must be stressed that the occurrence of no particular tracefossil type is important in the interpretation of sedimentary environments, but the dominance of

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ACCEPTED MANUSCRIPT ichnospecies and the entire trace-fossil association needs to be considered. Moreover, the occurrence of Rhizocorallium is dependent on specific conditions (such as substrate type, sedimentary energy,

Morphotypes and palaeoenvironments

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bottom oxygen, nutrient and salinity) and thus its morphology and size may vary from basin to basin.

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The two R. commune varieties from the Germanic Triassic clearly show a facies relationship, and material from different basins and ages suggests a relationship between the shape and the size of burrows and palaeoenvironment. Large, winding, horizontal and partly branched R. commune var.

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irregulare typically occur in the intertidal or littoral zone, while in shallow subtidal (sublittoral) deposits the burrows appear to be straighter, shorter and inclined. Deeper subtidal environments

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usually contain short and tongue-shaped burrows assignable to R. commune var. auriforme (Fig. 30). Similar bathymetric trends were also recognized by Farrow (1966) in the Jurassic of England, while

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Ager and Wallace (1970) documented horizontal Rhizocorallium in shallow and deeper subtidal

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environments and inclined specimens in intermediate subtidal environments in Jurassic rocks of

Germanic Basin.

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France. The analyzed global data set (Table 3) also confirms the overall trend as observed in the

In late Cretaceous and Cenozoic time, R. commune var. irregulare has also invaded deep-

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marine environments (Papp, 1962; Książkiewicz, 1977; Uchman, 1991, 1992; Kappel, 2003; Demírcan and Toker, 2003; Giannetti and McCann, 2010; Kotlarczyk and Uchman, 2012), which complicates the differentiation of those forms from their shallow-marine counterparts. However, in addition to their characteristic trace-fossil association with deep-marine trace fossils, those deepmarine forms differ from shallow-marine R. commune var. irregulare by the tendency to build lobate burrow systems (similar to Zoophycos) or systematically overlapping burrow systems (similar to Echinospira), their usual occurrence as endorelief, and a reduced size of the marginal tube (expressed by the tube/burrow width ratio) (Fig. 13).

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ACCEPTED MANUSCRIPT 9.3

Faecal pellets as facies indicators?

The occurrence of different ichnospecies of the coprolite ichnogenus Coprulus seems to offer a refinement of the established facies scheme. Of the three originally erected ichnospecies, C. oblongus

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and C. bacilliformis are regarded as valid based on their length/width ratio (Knaust, 2008). C. oblongus is a common constituent of R. commune, where it is accumulated in the spreite and the

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marginal tube. R. commune bearing C. oblongus occurs in intertidal and deeper environments. In contrast, R. commune with C. bacilliformis is rare (see Freyberg, 1963, fig. 2; Seilacher,

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2007, p. 60) and has so far only been reported from upper intertidal to shallow supratidal deposits. This observation is exemplified by the Middle Triassic spinosus Zone (Upper Muschelkalk) of

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Troistedt, where metre-scale shallowing-upward cycles are stacked on top of each other (Knaust and Langbein, 1995). The uppermost part of these cycles contain secondary calcite precipitations with a

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chalky or powdery appearance, including nodular, honeycomb, crystallaria, mottled, hardpan and

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laminar fabrics (Knaust, 2007b). These features are consistent with an interpretation as caliche (calcrete) and originated as alpha fabrics abiogenically within calcareous soils at surfaces exposed to

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the atmosphere, and influenced by microbial activity, lichens or cyanobacteria (Wright, 2007). Ammonoids, bivalves, gastropods and brachiopods are common and indicate shallow-marine conditions before these beds were subaerially exposed. Most R. commune specimens are associated

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with C. oblongus and were subsequently affected by calichification, as indicated by overprinting caliche nodules. However, rare examples of R. commune in the uppermost part of the cycles contain C. bacilliformis, in which caliche is cross-cut by R. commune (Fig. 31). This relationship can be taken to indicate that C. bacilliformis-bearing R. commune originated in the upper intertidal to supratidal zone. The evidence of C. bacilliformis for inter- to supratidal environments is further supported by its occurrence in peritidal facies along the margin of the Germanic Basin, e.g. on microbially modified bedding planes in Winterswijk (Fig. 8B) and in the mixed siliciclastic/carbonate system of the Muschelsandstein of Luxembourg (Hary, 1974), Nevertheless, similar pellets were also reported from distal-bay deposits (Desjardins et al., 2010).

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ACCEPTED MANUSCRIPT 9.4

Depositional energy and sequence stratigraphy

The epicontinental Germanic Basin during Triassic time was dominated by low-energy deposition (marlstone and muddy limestone) interrupted by high-energy event deposits (bio- and intraclastic

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wackestone and rudstone) (Aigner, 1979, 1984; Knaust, 2000). The distribution of Rhizocorallium needs to be differentiated on the established ichnospecies level and is a response to existing

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environmental conditions.

The R. jenense producer preferential colonized firm and muddy substrates along discontinuity

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surfaces, either in response of starved sedimentation or due to exhumation of substrates by erosion. R. jenense typically occurred at the margins of shoals and is associated with high-energy event

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deposition, which accentuated the burrow preservation by filling it with contrasting sediment. Submarine shoals and nearshore high-energy areas have also been documented from other areas (e.g.

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Fürsich, 1998), while in the deep-marine setting R. jenense occurs along canyon margins (e.g.

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Hayward, 1976). R. jenense also occurs on peritidal mudflats, as exemplified by the Rhizocorallium type stratum (Fig. 20). In a sequence-stratigraphical context, those occurrences demarcate ravinement

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surfaces during marine transgression (e.g. Rodríguez-Tovar et al., 2007) and thus R. jenense is most common in transgressive systems tracts and subordinately occurs in highstand systems tracts. In contrast, R. commune is a common constituent of low-energy background sediments, where

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it preferred fine-grained sediment. The simple and short R. commune var. auriforme can be found in muddy deposits of outer and mid-ramp position as well as protected lagoons, whereas various and partly complex burrow morphologies of R. commune var. irregulare occur in intertidal areas. R. commune also occurs frequently as post-event trace fossil in association with storm deposits (Jensen, 1997; Głuszek, 1998; Worsley and Mørk, 2001; Bann et al., 2004; Fig. 32A). Response to storm deposition is well expressed by vertically retrusive spreiten in R. commune problematica (e.g. KowalLinka and Bodzioch, 2011). Although it may be related to deposits of transgressive systems tracts, R. commune is most characteristic of highstand systems tracts.

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ACCEPTED MANUSCRIPT 9.5

Currents

Alignment of the long axes of burrows in response to prevailing currents is a common feature of various trace fossils and may assist in the reconstruction of palaeocurrent regimes and shoreline

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directions. Bilateral, quasi-symmetric burrows such as Rhizocorallium are well suited for palaeocurrent analysis, and Farrow (1966) documented the alignment of long axes of R. commune

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within intertidal deposits and interpreted this as a response to tidal currents and consequently alignment perpendicular to the palaeoshoreline. In the Lower Triassic of Spitsbergen, R. commune

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partly shows “… unimodal distribution, with the curved distal ends usually directed onshore”, probably related to longshore currents (Worsley and Mørk, 2001). An orientation of the tunnel

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openings with respect to (tidal?) currents was also observed by Mørk and Bromley (2008). Similar observations were made in the Germanic Muschelkalk, where long axes of minute R. commune run

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parallel to a gutter cast (Fig. 32B), a fact which is consistent with more quantitative data collected in

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the Spanish Muschelkalk (Rodríguez-Tovar and Pérez-Valera, 2008). A set of over 1000 measurements of R. commune from different beds and localities in the Late Jurassic of northern France

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only shows slight preference in orientation (Schlirf, 2000). The largest data set with over 2000 measurements from numerous sites was collected by Cotillon (2010) in the French Aptian. The results show that the line of the two burrow openings is perpendicular to the prevailing current direction,

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while the long axes of the burrows trend oblique to it. This data also constrained the interpretation of various current regimes operating in the basin. However, care must be taken in interpreting aligned spreite burrows such as R. commune, as this alignment may result from the exploitation of nutrientrich ripple troughs by the trace maker instead of current orientation (Fürsich, 1975). 9.6

Salinity

Deviations from mean seawater salinity are limiting factors for many marine organisms, and only few animals can cope with such conditions (see Buatois and Mángano, 2011). As documented in Table 3 and Figure 34, Rhizocorallium is reported from a wide range of salinity from hypersaline seawater on the one side to mesohaline brackish water on the other. Fürsich and Mayr (1981) reported R. jenense

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ACCEPTED MANUSCRIPT from fluvial (freshwater) environments, but these nonmarine forms are supposedly produced by mayflies instead of polychaetes in the marine realm. The type stratum of R. jenense cast burrows in a sabkha-like environment with increased

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salinity. In the Muschelkalk succession, faunal composition, geochemical evidence (e.g. celestine occurrence, affinity with evaporite and dolomite) and a restricted carbonate system suggest intervals

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with hypersaline conditions, in which Rhizocorallium occurred. Similar conditions also obtained in the Middle Triassic of the Tethyan domain (e.g. Jaglarz and Uchman, 2010; Knaust and Costamagna,

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2012). Lagoonal environments without freshwater influx are prone to increased evaporation, particularly under arid and semi-arid conditions. Examples of lagoonal R. commune occurrences are

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given, among others, by Carey (1978), Fürsich et al. (1980), Archer (1984), Gaillard et al. (1994) and Schweigert (1998).

On the other end of the scale, the marine Rhizocorallium producer was quite tolerant with

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respect to brackish-water conditions. In many cases, R. commune is reported from glaciomarine (e.g. Eyles et al., 1992; Bhattacharya and Bhattacharya, 2007; Sarkar et al., 2009) and other brackish

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deposits (e.g. Winn, 2006; Desjardins et al., 2010), fluviodeltaic environments (including delta front, interdistributary bay, distributary channel and point bar; e.g. Archer and Maples, 1984; Gouramanis et al., 2003; Mørk and Bromley, 2008), and estuarine deposits (Basan and Scott, 1979; Greb and

9.7

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Chesnut, 1996; MacEachern et al., 2007, 2012b).

Oxygenation

Trace fossils and bioturbation have long been recognized as reliable indicators of the degree of oxygenation of pore water within the sediment (e.g. Savrda and Bottjer, 1986; Wignall, 1991; Martin, 2004; Savrda, 2007). R. commune was described from various oxygen-depleted deposits where it occurs under dysoxic conditions with increasing oxygenation of the sediment (e.g. Jordan, 1985; Wignall, 1991; Martin, 2004; Kotlarczyk and Uchman, 2012). Fluctuating dysoxic conditions can be assumed for many portions of the Muschelkalk succession. The Lower Muschelkalk in the western part of the Germanic Basin shows signs of restricted circulation within a starved basin, for instance in the reduced content of shelly fossils. In the

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ACCEPTED MANUSCRIPT Lower Muschelkalk of the Germanic Basin, R. commune var. auriforme occurs together with Planolites montanus and Protovirgularia isp. in marlstone with nodular bedding of an outer ramp origin (Knaust et al., 2012; Fig. 30), while the same lithofacies type in the Upper Muschelkalk of the

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more restricted (low oxygen, high salinity) Catalan Basin in Spain (e.g. in the section Colldejou) is dominated by P. montanus and seems to lack R. commune (Calvet and Tucker, 1988; personal

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observations). Despite frequent event deposition in the Upper Muschelkalk, some intervals of the background deposits consist of laminated mudstone, and microbial structures contributed to the

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preferential preservation of soft-bodied organisms under oxygen-reduced conditions (Knaust, 2010a). Beside nematodes (preserved as sulphide mineral), nemerteans and platyhelminths, such biolaminites

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also contain R. commune. The occurrence of Rhizocorallium in dysoxic deposits is consistent with a polychaete interpretation of their producer, because many polychaetes (including spionids) are morphologically adapted to a life in sediments with permanently oxygen-depleted conditions (Levin,

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2003).

10 Rhizocorallium evolution

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The ichnospecies Rhizocorallium commune is one of the longest-ranging fossils and is known from the Early Cambrian to the Holocene (Fig. 33). During that long Phanerozoic eon (542 Ma), R. commune has stayed remarkably constant with respect to its morphology, burrow orientation, spreite development and faecal pellets, suggesting that it was produced by the same kind of organism (probably a polychaete worm). However, these spreite burrows were subject to significant size variation during that period, which may be related to changing palaeoenvironmental conditions and probably also evolutionary aspects of their producers. The differently sized groups are accounted for by two variations: R. commune var. auriforme (small) and R. commune var. irregulare (large). Although reasonably sized R. commune already occur in the lowermost Cambrian, the early Palaeozoic is generally dominated by rather smaller forms, while the burrow size increases in the late Palaeozoic. Following the end-Permian mass extinction, early Triassic R. commune was considerably reduced in

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ACCEPTED MANUSCRIPT size but recovered to its pre-extinction burrow size at the latest by the middle Triassic. It must be noted that middle and late Permian as well as late Triassic records of R. commune are very rare or even absent, disregarding those records with an unspecified stratigraphical range. An optimum for the R.

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commune producer, coinciding with a size maximum, however, can be recognized in Jurassic and Cretaceous strata, in which reports of R. commune var. irregulare clearly dominate over R. commune

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var. auriforme. In the Cenozoic, this trend seems to be reversible and is perhaps related to a change in habitat of the producer.

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Palaeozoic and Mesozoic R. commune is an effective indicator of shallow-marine environments, although individual outliers were reported from deep-sea deposits, such as a poorly

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preserved R. commune var. irregulare from the Ordovician (Pickerill et al., 1982) and a poorly developed R. commune var. auriforme from the Triassic (Carvalho et al., 2005). The so-called ‘Rhizocorallium’from the Early Carboniferous flysch of the Czech Republic (Mikuláš et al., 2002)

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does not qualify as such. There are many reports of deep-marine R. commune from Palaeogene deposits through Miocene strata, a trend that started in the Cretaceous (e.g. Papp, 1962; Kappel, 2003).

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This is somewhat comparable to the development observed in the heterogeneous ichnogenus Zoophycos, to which Rhizocorallium is partly similar (cf. Savary et al., 2004, fig. 6-7). Neto de Carvalho and Rodrigues (2003) have shown that Palaeozoic Zoophycos occurred sporadically as a

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pioneer in some deep-marine niches, while deep-marine colonization was extensive from the Jurassic onwards, with the ichnogenus being restricted to the deep sea in Cenozoic time. R. jenense (according to the nomenclature used herein) appeared after the end-Permian mass extinction (Fig. 33) and is continuously reported from shallow-marine deposits throughout the Mesozoic until the early Miocene, although it commonly penetrated older rocks along “Glossifungites” surfaces. This distribution pattern suggests the interpretation that firmground burrowing by the Rhizocorallium producer developed after the end-Permian mass-extinction when firm substrates were widespread and competing benthic organisms were strongly diminished. Finally, R. jenense from late Miocene and younger freshwater deposits were probably produced by mayfly larvae with a convergent evolution of burrowing behaviour with that attributed to polychaetes.

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ACCEPTED MANUSCRIPT Mass extinctions, such as the one at the Permian/Triassic boundary, have been well documented by means of their ichnological signature. It has been argued that certain ichnotaxa reappeared in the Early Triassic in a stepwise manner with Rhizocorallium not before Spathian

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(Olenekian) time (Twitchett and Wignall, 1996; Twitchett, 1999; Twitchett and Barras, 2004; Chen et al., 2011). However, as more data became available, other studies have demonstrated that

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Rhizocorallium reappears right above the Permian/Triassic Boundary already in Griesbachian (Induan) time (e.g. Fraiser and Bottjer, 2009; Zonneveld et al., 2010; Hofmann et al., 2011). A comprehensive

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dataset from well cores across that boundary in the Persian Gulf consistently shows a much faster recovery than earlier anticipated (ca. 1.2 Ma instead of >5 Ma; Knaust, 2010b).

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Gong and Si (2002) attempted to present a morphological classification system of trace fossils based on both topological and Euclidean geometric characteristics. Accordingly, Rhizocorallium belongs to the group (“ichnoorder”) with the highest behavioural complexity and evolutionary level.

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Similarly, Ekdale and Lamond (2003) developed a cladogram of spreite burrows, in which an evolutionary progression beginning with a simple vertical shaft (Skolithos) led to a vertical U-shaped

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burrow (Arenicolites) with spreite (Diplocraterion), and then to inclination (Rhizocorallium) and finally sequential radiating (Zoophycos). A modification of this cladogram includes Rhizocorallium uliarense and Lapispira at the end (Lanés et al., 2007). Such cladograms remain theoretical and highly

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speculative not simply because the sequence of different stages of development is hard to prove (although it is testable), but, more cogently, because the schemes ignore the fact that the evolution of behaviour occurs in living species with a phylogenetically contrasting position (Seilacher, 1986).

11 Summary and conclusions

The ichnogenus Rhizocorallium is one of the first trace fossils to have been recognized and has a global distribution throughout the Phanerozoic. Many workers have treated it with respect to its morphological features, substrate relationship, responsible producer and inferred ethology. The introduction of numerous ichnospecies has led to a complicated situation, although most workers

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ACCEPTED MANUSCRIPT today follow the classification provided by Fürsich (1974) and subdivide Rhizocorallium into two common (R. jenense and R. irregulare) and one rare (R. uliarense) ichnospecies. In a recent proposal, Schlirf (2011) challenged the existing classification of Rhizocorallium and other U-shaped spreite

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burrows. However, a careful review of the original descriptions together with the analysis of comprehensive new material from the type area of the type ichnospecies reveals problems with both

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classifications. These problems have consequences for a strict application of ichnospecies of Rhizocorallium for palaeoenvironmental interpretations and weaken their value for facies and

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sequence-stratigraphical analaysis.

The results of this study show that Rhizocorallium from the Germanic Triassic justifies a

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subdivision into two ichnospecies. Following recent trends in ichnotaxonomy (e.g. Bertling et al., 2006), substrate consistency promises to be a major ichnotaxobase for the discrimination at the ichnogenus and ichnospecies levels. Consequently, R. jenense is restricted to passively filled burrows

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in firmground, while R. commune exhibits a spreite actively produced chiefly in softground. Both ichnospecies can have gradational transitions as exemplified by R. commune with scratched marginal

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tube or spreite area. Burrow morphology can be quite variable and is obviously a function of the trace maker’s behaviour (e.g. feeding and dwelling) and external circumstances (e.g. sedimentation rate, current activity), but only serves as ichnotaxobase at the ichnosubspecies level (R. commune

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problematica with a vertically retrusive spreite element, R. commune uliarense with a spiral morphology). In contrast, burrow size is not justified as an ichnotaxobase because a numerical analysis results in a wide range of size parameters without showing distinct clusters. However, judging from a stratigraphical and sedimentological analysis, absolute burrow sizes reflect different positions with regard to facies and palaeoenvironment. Therefore, a subdivision of R. commune into the varieties R. commune var. auriforme and R. commune var. irregulare is proposed to support sedimentological analysis. Three groups of organisms qualify as potential producers of Rhizocorallium: decapod crustaceans, annelids and mayflies. Although Rhizocorallium in some earlier studies was interpreted as the product of annelids, since most workers have favoured crustaceans as trace makers because of the presence of scratches. However, numerous groups of animals are known to produce scratches. Several

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ACCEPTED MANUSCRIPT lines of evidence suggest annelids (e.g. spionids, eunicids) as the producer of Rhizocorallium, an interpretation which is supported by constructional morphology, accompanying features (e.g. faecal pellets), neoichnological comparison and remains of the fossilized producers preserved in situ. An

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interpretation of marine Rhizocorallium as a polychate burrow is straightforward, while the larvae of emerging mayflies probably account for R. jenense produced in fluvial settings.

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Combined suspension- and deposit-feeding behaviour is favoured by most workers, although tests for gardening and cache behaviour are pending a detailed analysis of the included faecal pellets.

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The two ichnospecies of Rhizocorallium and their varieties may serve as good facies indicators with the potential of supporting sedimentological and palaeontological interpretations. R. commune is

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a component of the Cruziana ichnofacies and typically occurs in shelf and nearshore environments, while R. jenense is part of the widespread Glossifungites ichnofacies. Data from the Germanic Basin confirm the preferential occurrence of R. commune var. irregulare in intertidal and shallow subtidal

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environments, whereas R. commune var. auriforme occurs in deeper parts of the basin and in lagoonal environments. Furthermore, R. commune-associated faecal pellets, C. bacilliformis, occur in inter- to

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supratidal environments and C. oblongus is evident in intertidal and deeper environments. R. jenense typically occurs along omission surfaces in high-energy areas and is often documented together with ravinement surfaces excavated during marine transgressions. It is a common constituent of

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transgressive systems tracts.

Several studies have utilized Rhizocorallium as current indicator, with its long axis running parallel or oblique to the inferred palaeocurrent (or the line of the two burrow openings perpendicular to it), and the distal (curved) end directed onshore. This pattern can be obscured by a more random alignment in response to the exploitation of nutrient-rich ripple troughs by the trace maker. Many marine Rhizocorallium are documented as tolerating a wide range of salinity from hypersaline to mesohaline water. R. commune occurs in various oxygen-depleted deposits with dysoxic conditions and increasing oxygenation of the sediment. A review of over 180 Rhizocorallium records from the literature, in accordance with the newly proposed classification, reveals a long range of R. commune from the Early Cambrian to the Holocene,

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ACCEPTED MANUSCRIPT while R. jenense is restricted to Mesozoic and Cenozoic strata. Evidently, only after the end-Permian

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mass extinction did polychaetes became adapted as firmground burrowers.

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Acknowledgements

The initiation of this review reaches back more than 20 years and has benefited from discussing

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ichnotaxonomical aspects with Andrew K. Rindsberg (Livingston), Alfred Uchman (Kraków), Jordi de Gibert (Barcelona, deceased), Francisco J. Rodríguez-Tovar (Granada), Luis A. Buatois (Saskatoon)

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and Richard G. Bromley (Copenhagen). For assistance during field work and offering access to collected material I am indebted to Henk Oosterink (Winterswijk). Help with, and information about material in historical collections was received from Hagen Hopf, Thomas Kammerer and Hermann

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Huckriede (all Weimar), Birgit Kreher-Hartmann, Thomas Voigt and Rolf Beutel (all Jena), Winfried Werner (Munich), Richard G. Bromley and Finn Surlyk (both Copenhagen), Alfred Uchman

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(Kraków), and Andrei V. Dronov (Moscow). Nina Sinitshenkova (Moscow) and Jörg Ansorge (Stralsund) provided valuable information about fossil mayflies. Sincerely thanks go to Andrew K.

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area. Acta Palaeont. Sinica 35, 475-489, pl. 1-3. (In Chinese with English abstract.) Zonneveld, J.-P., Gingras, M.K., Beatty, T.W., 2010. Diverse ichnofossil assemblages following the P-T mass extinction, Lower Triassic, Alberta and British Columbia, Canada: Evidence for shallow marine refugia on the northwestern coast of Pangaea. Palaios 25, 368-392. Zonneveld, J.-P., Gingras, M.K., Beatty, T.W., Bottjer, D.J., Chaplin, J.R., Greene, S.E., Martindale, R.C., Mata, S.A., McHugh, L.P., Pemberton, S.G., Schoengut, J.A., 2012. Mixed siliciclastic/carbonate systems. In: Knaust, D., Bromley, R.G. (Eds.), Trace Fossils as Indicators of Sedimentary Environments. Dev. Sediment. 64. Elsevier, Amsterdam, pp. 807-833. Zonneveld, J.-P., Gingras, M.K., Pemberton, S.G., 2001. Trace fossil assemblages in a Middle Triassic mixed siliciclastic-carbonate marginal marine depositional system, British Columbia. Palaeogeogr., Palaeoclimatol., Palaeoecol. 166, 249-276.

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ACCEPTED MANUSCRIPT Figure captions

Fig. 1: Historical figures of Rhizocorallium jenense Zenker, 1836 as introduced from various areas in

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the 19th century.

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Fig. 2: Fucoides auriformis (= Rhizocorallium commune) from the Lower Silurian of North America

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(from Hall, 1843).

Fig. 3: Historical figures of Thier-Fährten (animal traces, = Rhizocorallium commune) from the

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Lower Cretaceous Bentheim Sandstone of NW-Germany (from Jugler, 1853).

Fig. 4: Concept of trace-fossil taxonomy and classification based on a defined hierarchy of

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ichnotaxobases and exemplified by the ichnogenus Rhizocorallium.

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Fig. 5: Orientation of U-shaped spreite burrows as a feature for discriminating the ichnogenera Rhizocorallium and Diplocraterion. A: Lower bedding plane with casts of oblique to sub-horizontal R. jenense. Early Triassic (Upper Buntsandstein, Rhizocorallium Dolomit), Jena, coll. Mägdefrau (TLGU

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5035-701-202). B: Section with two vertically orientated D. parallelum. Early Triassic (Middle Buntsandstein), Jena, coll. Mägdefrau (TLGU 5135-701-140). Arrow pointing up.

Fig. 6: Different expressions of Rhizocorallium commune in response to substrate consistency. Middle Triassic (Upper Muschelkalk, evolutus-spinosus Zone), Troistedt. A: Soft sediment, hyporelief. Note the preservation of minute burrows backfilled with Coprulus oblongus faecal pellets within the marginal tube due to repeated sediment reworking by a vermiform producer. B: Firm sediment, epirelief.

Fig. 7: Contrasting burrow fill in ichnospecies of Rhizocorallium in response to substrate consistency. A: Vertical section of a firm- and hardground below an omission surface and overlain by a

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Obere Oolithbank), Bad Berka-Gutendorf. B: Vertical thin section of a horizontal R. commune displaying an actively created spreite (active spreite) between the passively filled marginal tube. Early

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Jurassic, Grimmen (see Knaust, 2012a).

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Fig. 8: Faecal pellets in association with Rhizocorallium commune. A: Coprulus oblongus Mayer, 1952. Middle Triassic (Upper Muschelkalk, evolutus-spinosus Zone), Troistedt. B: C. bacilliformis

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Mayer, 1955. Note the diagnostic difference in the length/width ratio of the pellets. Middle Triassic (Lower Muschelkalk), Winterswijk (The Netherlands) (Henk Oosterink collection, Winterswijk).

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Fig. 9: Scratches (bioglyphs) along the marginal tube of Rhizocorallium. A: R. jenense with net-like, crossing and closely spaced scratches. Early Triassic (Upper Buntsandstein, Rhizocorallium Dolomit),

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Jena-Ziegenhain, coll. Mägdefrau (TLGU 5035-701-202) B: R. commune with subparallel, rarely crossing and sparse scratches. Middle Triassic (Lower Muschelkalk, Gelbe Plattenkalke), Bad Berka-

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Tannroda.

Fig. 10: Type specimens of Rhizocorallium devonicum Hecker, 1930 from the Late Devonian of the Syas River section near St. Petersburg, NW Russia. A: Holotype from the St. Petersburg State Mining Institute (photo courtesy of Andrey Dronov). B: Paratype from the Paleontological Museum in Moscow.

Fig. 11: Newly proposed classification of Rhizocorallium. For discussion, see text.

Fig. 12: Rhizocorallium commune showing subvertical (aff. Diplocraterion) and subhorizontal (aff. Rhizocorallium) burrow components. Early Jurassic, Punta Rodiles, Asturias (northern Spain). See Rodríguez-Tovar et al. (2012).

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Fig. 13: Examples of Rhizocorallium–Zoophycos transitional spreite burrows from Oligocene deepsea turbidite sandstone of Braux/Annot (southern France). All specimens contain abundant Coprulus

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oblongus (faecal pellets). A: Incipient R. commune, slightly oblique to bedding surface. B: Elongate, tongue-shaped R. commune with relatively thick marginal tube on bedding plane. C: Winding and

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branched, lobate Z. insignis on bedding plane.

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Fig. 14: Echinospira pauciradiata from Miocene deep-sea deposits of the Mohakatino Formation along the Mohakatino estuary north of New Plymouth (New Zealand). A: Circular burrow system

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consisting of numerous progressively overlapping Rhizocorallium-like spreite burrows. B: Spiral-like trace of spreite burrows on a bedding plane.

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Fig. 15: Loosely clustered, shallow U- and J-shaped burrows with incipient spreiten and a rope-like appearance, attributed to aff. Phycodes parallelum and aff. Fuersichnus communis. Fluvial to

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lagoonal, marginal-marine sandstone bed of the Upper Triassic (Lower Keuper, Lettenkeuper), collected from the now refilled clay pit ‘Schleyersche Ziegelei bei Bhf. Weimar’, coll. Mägdefrau

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(TLGU 5033-701-467).

Fig. 16: Tisoa siphonalis with long and narrow U-shaped burrows, occurring in connection with seep deposits. Late Miocene continental slope deposits, Urenui Formation, Waiau coastal section north of New Plymouth (New Zealand). A: Elongate T. siphonalis (arrows) exposed slightly oblique to the bedding. B: Cross-sections of passively filled T. siphonalis with a dumbbell to figure- eight shape.

Fig. 17: Compound Balanoglossites/Rhizocorallium trace fossils. A: Holotype of B. triadicus in vertical section, displaying integral U-shaped pouches (R. jenense, R) (Mägdefrau, 1932; Knaust, 2008). B: Upper bedding plane with ‘B. eurystomus’ (= B. triadicus, elliptical openings) from its type locality (Jena-Göschwitz), closely associated with an incipient spreite burrow with collapsed and brecciated roof due to a hardened surface. C: Upper bedding plane with R. commune and numerous

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ACCEPTED MANUSCRIPT grooves and U-shaped tunnels leading into a B. ramosus burrow system. Note the similarity in shape and size of the grooves and tunnels to the grooves within the Rhizocorallium spreite, from which one tunnel arises (arrow). Middle Triassic, Upper Muschelkalk (spinosus Zone), Troistedt. D: Lower

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bedding plane of the same bed as in C displaying a B. ramosus burrow systems and its intimate

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association with Rhizocorallium-like elements (R).

Fig. 18: Scratched firmground surfaces as an expression of different behaviour of the Rhizocorallium

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commune producer. A: Bedding surface with a compound trace fossil consisting of incipient Rhizocorallium-like pouches, from which radiating scratches (resembling Radichnus allingtona) run

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all over the surface. Middle Triassic (Lower Muschelkalk), Schkölen-Tünschütz. B: Lobate U-shaped burrow resembling R. commune transitional to R. jenense, with an extensively scratched burrow wall.

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Middle Triassic (Lower Muschelkalk, Konglomeratschicht d2), Dorndorf-Steudnitz.

Fig. 19: Composite trace fossil consisting of Rhizocorallium commune reburrowed with Ophiomorpha

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rudis. Deep-marine turbidite deposits, Oligocene, Grand Coyer near Annot (SE-France). Photo courtesy of Michal Warchoł.

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Fig. 20: Stratigraphy of the Germanic Triassic with palaeoenvironmental interpretation and representative sections with occurrences of ichnospecies of Rhizocorallium as described and figured. The numbers in brackets refer to figured specimens.

Fig. 21: Rhizocorallium jenense from its type stratum and type area in the vicinity of Jena. A: JenaGöschwitz, coll. Phyletisches Museum Jena (P1821). B: Engerda, University of Jena, Geosciences collection (T4338).

Fig. 22: Rhizocorallium jenense from the Middle Triassic Lower Muschelkalk. A: Dolomitic marlstone bedding plane displaying desiccation cracks (C) and cross sections of small and inclined R. jenense (R). Basal Muschelkalk, Rittersdorf. B: Bedding plane of a firmground with numerous

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ACCEPTED MANUSCRIPT inclined and scratched open burrows. Obere Oolithbank, Blankenhain-Lohma. C: Cross-section of a mudstone (micrite) with a firm- to hardground suite, exhibiting dolomite-filled R. jenense (R), a fragment of a Balanoglossites triadicus system (B), and superimposed needle-like Trypanites weisei

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borings (T) from the top margin. Obere Oolithbank (type area and type stratum for both, T. weisei and B. triadicus), Dorndorf-Steudnitz. D: Omission surface on top of a mudstone (cross-section) with

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dolomite-filled R. jenense (R), overlain by a packstone. Lower Terebratelbank, Großmonra-

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Burgwenden.

Fig. 23: Rhizocorallium jenense of various shapes and sizes from the Middle Triassic Lower

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Muschelkalk (Upper Wellenkalk). A: Rittersdorf. B: Bad Berka-Gutendorf. C, D: BlankenhainNeckeroda. E: Plaue. F: Bedding plane with desiccation cracks, casts of Neoschizodus orbicularis and

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R. jenense in hyporelief. Winterswijk.

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Fig. 24: ‘Glossifungites saxicava’ (= R. jenense) from the Miocene of Lviv, Ukraine (type area).

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Institute of Geological Sciences of the Jagiellonian University in Kraków, Poland (see Uchman et al., 2000). Note the pronounced scratch traces over entire surface, intersectioning with a smaller specimen

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and the diagnostic passive fill (passive spreite). Photo courtesy of Alfred Uchman.

Fig. 25: Rhizocorallium commune from the Early Triassic (Upper Buntsandstein) and Middle Triassic (Lower Muschelkalk). A: Robust R. commune var. irregulare in marly limestone. Between Muschelkalk base and Konglomeratschicht d2 (type horizon), Dorndorf-Steudnitz. B: Partly retrusive spreite elements of R. commune var. irregulare on a bedding plane of a dolomitic and platy limestone bed. Lower Muschelkalk (Gelbe Plattenkalke), Bad Berka-Tannroda. C: Large winding R. commune var. irregulare together with incipient individuals (stacked arcs, middle) on top of a sandstone bed. Upper Buntsandstein (Salinarröt so1, near the Rhizocorallium Dolomit), clay pit Jena-Göschwitz. Coll. Phyletisches Museum Jena (P556). D: Robust R. commune var. irregulare in marly limestone. Lower Muschelkalk (Upper Wellenkalk), Plaue. E: R. commune var. auriforme originating from desiccation cracks and migrating along the bedding plane. Lower Muschelkalk, Winterswijk. F: R. commune

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Fig. 26: Rhizocorallium commune var. irregulare in epirelief preservation on limestone beds from the

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type bed of ‘R. irregulare’ (B-F) in the Middle Triassic (Upper Muschelkalk, spinosus Zone) of

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Troistedt. A: Marlstone/limestone alternation containing the type bed of ‘R. irregulare’ (arrow heads). Hammer (encircled) for scale. B: Retrusive spreite (top), reburrowed spreite with groove-like

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excavated marginal tube (lower right), isolated deep grooves (middle) and scattered circular burrow openings. C: Multiply branched burrow. D: Retrusive spreite with partly preserved marginal tube,

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which intensely scratched surface indicates burrowing of the trace maker along the marl (now weathered) / limestone interface. Note the densely packed Coprulus oblongus faecal pellets within the spreite. The bow-shaped grooves (top right, bottom right) define initial spreiten of the same producer

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and are best explained by sediment removal by a vermiform animal. E, F: Spreite fragments showing grooves and ridges with scattered C. oblongus. The marginal tube contains smaller burrows of the

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same diameter as the grooves, which are densely packed with faecal pellets, and originated from the repeated movement of their producers.

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Fig. 27: Morphometric analysis of ichnospecies and varieties of Rhizocorallium from the German Triassic. See text for explanations.

Fig. 28: X-ray radiographs (negative) of Holocene cores from the Western Baltic Sea. (Courtesy of the radiography data base, University of Kiel, http://www.ifg.unikiel.de/Radiographien/radiographien.phtml). A: Strongly bioturbated sandy clay with numerous empty burrows, partly showing a figure- eight cross-section similar to that of Rhizocorallium jenense. These firmground burrows were probably made by the terebellid polychaete Terebellides stroemi. Vibracore sample GIK 15391-1, Kiel and Mecklenburg Bight, 21.0 m water depth. B: Mud/clay with sandy patches and single and composite polychaete tubes. lw = lined worm tube, hb = hollow burrow, fb = filled burrow, pb = echinoid burrow (Scolicia) overprinting a mollusc burrow, mb = composite

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ACCEPTED MANUSCRIPT spreiten burrow similar to R. commune, made by various organisms, including Pectinaria sp., T. stroemi (both polychaetes) and Echiurus echiurus (echiuroid). Boxcore sample GIK 12549-1, Great

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Belt, 27.2 m water depth. From Winn (2006, pl. 2).

Fig. 29: Incipient Rhizocorallium commune on bedding surfaces from the Middle Triassic) Upper

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Muschelkalk, evolutus Zone) of Troistedt, together with their producers preserved in situ. The vermiform animals occur as limonitic aggregates (brown colour, black arrow heads) together with

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meiobenthic trace fossils and their trace makers preserved at trace termination (white arrow heads). See Knaust (2007b, 2010) for more information. A: Horizontal spreite and foraminifer traces (to the

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right). B: Oblique spreite. C: Oblique spreite together with tiny Cochlichnus isp. trails produced by nematodes (white arrow heads) and nemertean trails partly preserving their producers (lower right). D:

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Oblique spreite with a robust limonite aggregate representing remains of its producer.

Fig. 30: The distribution of Rhizocorallium jenense and R. commune (and varieties) and their

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palaeoenvironmental relationships in the Middle Triassic (Lower Muschelkalk) of Thuringia (type area). A: Idealized shallowing-upward sequence with the occurrence of Rhizocorallium. B: Conceptual model of the Lower Muschelkalk carbonate ramp with characteristic depositional

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environments and preferred occurrences of Rhizocorallium. Modified after Knaust et al. (2012).

Fig. 31: Rhizocorallium commune var. irregulare on bedding surfaces from the Middle Triassic (Upper Muschelkalk, spinosus Zone) of Troistedt, and their relationship with caliche structures. A: That the spreite burrow predates calichification is indicated by its overprinting by caliche nodules. Coprulus oblongus is typically associated with these burrows. An intertidal origin of the burrow with subsequent subaerial exposure is likely. B: The spreite burrow partly postdates calichification and crosscut a chalky crust with crystallaria. Note the continuation of some scratches from the burrow tube onto the crust and the abundant occurrence of C. bacilliformis. An upper intertidal to shallow supratidal environment can be inferred for the burrow origin.

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Coprulus oblongus, which is partly eroded by the contemporaneous storm event (waning stage). Middle Triassic (Upper Muschelkalk, spinosus Zone), Weimar-Holzdorf. B: Cluster of minute R.

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commune var. auriforme aligned with their long axes more or less parallel to a gutter cast (lower part) on the base of a limestone bed. Middle Triassic (Lower Muschelkalk, Upper Wellenkalk),

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Blankenhain-Lohma. C: R. commune in biolaminite. Middle Triassic (Upper Muschelkalk, evolutus

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Zone), Troistedt.

Fig. 33: Stratigraphical distribution of R. jenense (A) and R. commune (B) with respect to their overall environment. Numbers refer to entries given in Table 3. Ichnosubspecies and varieties of R. commune,

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reported more than once from the same area and stratigraphical unit, are not repeatedly plotted. The

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numerous occurrences from the Germanic Triassic are schematically indicated. See text for discussion.

Table 1: Summary and comparison of significant characteristics of the two valid ichnospecies from

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their type area, the Triassic Germanic Basin.

Table 2: Coordinates of all studied localities.

Table 3: Rhizocorallium records from outside the Germanic Basin as compiled from the literature.

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Substrate Preservation

Firmground Dominantly hyporelief (positive), subordinate epirelief (negative) and endorelief Intersecting, crisscrossing common

Inter-burrow relationship Orientation

Size

Small to medium

Outline

Bow- to U-shaped, short

Cross-section Sinuosity Burrow length / width ratio Faecal pellets

Ellipsoidal-elongate, dumbbell-like No to low Small

Scratches

Abundant, net-like

Fill

Passive

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Branching

Varying, subhorizontal to subvertical (0-90°) No

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Balanoglossites triadicus, Fuersichnus communis Polychaetes (marine realm), crustaceans?, mayflies (fluvial settings) Suspension feeder (domichnion) Ethology Glossifungites Ichnofacies Palaeoenvironment Upper intertidal to supratidal, fluvial Table 1

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Transitional ichnotaxa Inferred producer

R. commune Typically isolated or loosely associated (scattered, grouped) Softground, partly firmground Dominantly epirelief (positive and negative), subordinate hyporelief (positive) and endorelief Isolated, segregated, coexistent (side by side), cross-cutting rare Subhorizontal (0-10°)

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R. jenense Typically clustered (gregarious)

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Ichnospecies Occurrence

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Primary successive branching may occur (rare) Medium to large, subordinate small (juvenile forms) U- to tongue-shaped, elongate band-like Dumbbell-like Low to high, irregularly winding Large Abundant (Coprulus oblongus, C. bacilliformis) Occasional, sub-parallel (mainly covering the marginal tube) Active (spreite)/passive (marginal tube) Balanoglossites ramosus, Radichnus allingtona Polychaetes Deposit feeder (fodinichnion) Cruziana Intertidal to subtidal (sublittoral)

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Table 2

50°47'35.00'' N, 11°52'58.40'' E 50°47'55.30'' N, 11°18'32.50'' E 50°49'51.70'' N, 11°13'17.10'' E 50°50'46.30'' N, 11°15'29.80'' E 50°52'41.00'' N, 11°35'15.50'' E 50°52'41.60'' N, 11°22'49.20'' E 50°52'50.20'' N, 11°34'38.40'' E 50°54'49.60'' N, 11°13'00.50'' E 50°56'33.10'' N, 11°15'04.70'' E 50°56'45.60'' N, 11°16'29.80'' E 50°57'29.00'' N, 11°38'50.20'' E 50°59'41.90'' N, 11°49'44.30'' E 51°00'51.80'' N, 11°41'55.20'' E 51°13'44.50'' N, 11°18'03.40'' E 51°57'59.60'' N, 06°46'49.00'' E 54°07'54.33" N, 13°03'31.42'' E

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Plaue (quarry) Blankenhain-Neckeroda (quarry, abandoned) Rittersdorf (quarry) Bad Berka-Tannroda (quarry) Jena-Göschwitz (clay pit, abandoned) Blankenhain-Lohma (quarry) Jena-Göschwitz (quarry) Bad Berka-Gutendorf (quarry) Troistedt (quarry, abandoned) Weimar-Holzdorf (excavation, refilled) Jena (Gleißberg, slope) Schkölen-Tünschütz (quarry, abandoned) Dorndorf-Steudnitz (quarry) Großmonra-Burgwenden (quarry) Winterswijk (quarry) Grimmen (clay pit, abandoned)

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