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Early Jurassic “worm holothurians” (Echinodermata) as faecal traces of a worm-like holothurian-eater reflecting the consumed species

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Abstract

“Worm holothurians” from a sequence of Early Liassic (Hettangian) shales at Göttingen (Germany) are string- and strip-shaped relics of faeces of a worm-like holothurian-eater. They consist of three typical associations of morphotypes of microscopic calcitic ossicles of the body wall skeletons of holothurians: (1) mostly buttons and a few larger hooks; (2) polyperforate spectacles and many tiny hooks; or, rarely, (3) sieve platelets only. The faeces were discarded by the predator partly as it rapidly advanced, leaving an elongated, irregularly curved string at the sediment surface, but mostly during its slow cree** and crawling at and below the sediment/water interface, when it left a narrowly meandering string in its moulded or tunnel-like trail. The elongated strings are generally well preserved, whereas the meandering ones are generally totally disintegrated, possibly due to ventilating activities of the predator. Upon the collapse of the trail wall, the biostratinomic state of disintegration was fixed as a strip of ossicles. Such a strip, therefore, represents the faecally documented part of a cree** or crawling trace in or just below the sediment surface. All holothurian calcareous ring segments found in some of the two kinds of strip-shaped ossicle associations with hooks belong to the same new type of ring form. According to a feasible scenario for the predator–prey relationship, this form should be defined as a new genus of chiridotid Apodida that contains the ossicle associations with hooks as two biological species. The strings of concentrated sieve platelets should also be interpreted as a biological species from a new genus within the Dendrochirotida.

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Acknowledgements

The first aggregates and worm-like ossicle associations were shown to me for determination by Peter Wüstemann (Göttingen), who also provided permission to visit this protected site at Göttingen. Hartmut Scholz (Göttingen) gave insight into his microcollections from Jurassic shales in southern Germany. Burkhard Schmidt (Göttingen) and Tanja R. Stegemann (Göttingen, Munich) enabled the application of digital microphotography. Charlotte Kniest (Göttingen) helped with the acquisition of rare literature. Jes Rust (Bonn) commented on some remnants of arthropods. Walter Riegel (Göttingen) improved earlier versions of the English text. Alexei Smirnov (St. Petersburg) commented positively on a previous version of the manuscript. As reviewers, Tanja R. Stegemann (see above), Alexander M. Kerr (Guam), and Mike Reich (the editor-in-chief, Munich) eliminated a multitude of formal deficiencies, and made constructive suggestions. I am extremely grateful for all this support.

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    Reimund Haude

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Appendix

Appendix

Type I/II-CA specimens (often only fragments; I/II indicated; contents of hooks very variable, generally small, in a few specimens no or only 1–2 hooks externally visible, indicated by [0] or [1]; possibly mixed with BA in italics):

GZG.INV.112001_3 (I), 112005_1 (I), 112005_2 (I), 112006 (I), 112010 (II?, 2), 112018_1 (II), 112020_1 (I), 112021_1 (I), 112021_2 (II [1]), 112022 (I? [1]), 112025 (I), 112034 (II), 112047 (I), 112051 (I), 112057 (I, II?), 112063_2 (I?), 112066 (I), 112071 (I?), 112072_1 (I), 112073 (I ?), 112074 (I + II), 112081 (I), 112083 (I), 112084 (II), 112090 (I), 112094_1 (I), 112094_2 (I), 112094_3 (I), 112094_4 (II), 112096_1 (I), 112106 (I), 112117_1 (I), 112121 (I or II?), 112126_1 (I), 112143 (II), 112146_1 (II), 112146_2 (II), 112160_1 (II [0]), 112164 (II?), 112166 (II?), 112169 (I?), 112174 (II), 112175 (I or II?), 112188 (I), 112190 (II), 112191 (II), 112192 (II), 112194 (II), 112197 (I), 112198_1 (I), 112200_1, 112200_2 (I or II?), 112202 (I?), 112205 (I), 112206_1 (I), 112206_2 (I), 112208 (I), 112211_1 (II), 112213 (I), 112225 (I), 112216 (I), 112226_1 (I?), 112251 (I?), 112259_1 (I), 112277 (II), 112278_1 (I), 112279 (I?), 112282 (I?), 112284_1 (I), 112318_1 (I), 112319 (I?), 112328 (I?), 112337_2 (II), 112397 (I?, 0), 112408 (II).

Type I/II-BA specimens (mostly only fragments; I/II indicated; some containing relative large polyperforate spectacles in addition to normal ones, indicated by [L]):

GZG.INV.112004_2 (II), 112007_1 (I), 112009 (I?), (?)112014_1 (II), 112041 (II?), 112050 (I?), 112063_1 (I), 112095 (I?), 112105 (I), 112120 (II), 112163_1 (I?), 112182_1 (I?), 112186 (II), 112196_1 (I [L]), 112203_1 (?), 112203_2 (I), 112211_2 (II), 112211_3 (II), 112223_1 (I? [L]), 112223_2 (I? [L]), 112262 (I), 112284_2 (II), 112306 (?), 112331 (II).

Ring segments

In CA associations: GZG.INV.112001_3 (in MS); 112020_1 (partly in MS: GZG.INV.113020_2, _3, _4, _5; partly in SEM: 113020/1,/2,/3,/4,/5,/6), GZG.INV.112047_1 (partly in MS: GZG.INV.113047_2, _3, _4; partly in SEM: 113047/7,/8), in 112071 (not extracted), 113073 (MS), 113106_1 (MS), 113117_1 (MS), 113126_1 (MS), 113205_1, 113205_2 (MS), 112208_1 (cluster, not extracted);

In BA associations: GZG.INV.112062_1/1 (spot-like association; partly not extracted, partly in SEM 113062_1/9, _1/10), 112063_1, 112063_2 (partly in SEM: GZG.INV.112063_1/1, partly in MS: 112063_1/2, _1/3 [aggregate]), 112100_1 (MS), 112105_1, 112203_1/1 (not extracted), 112262_1 (cluster, not extracted).

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Haude, R. Early Jurassic “worm holothurians” (Echinodermata) as faecal traces of a worm-like holothurian-eater reflecting the consumed species. PalZ 93, 265–283 (2019). https://doi.org/10.1007/s12542-018-0428-2

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